Alien terrestrial arthropods of Europe

Fotini Koutroumpa

This book provides the fi rst comprehensive review of the fauna of alien terrestrial arthropods that have colonized the European continent and its associated islands. Directly ensuing from the DAISIE project, this is the result of the joint work of 89 authors from 27 different European countries. The book summarizes present knowledge of the arthropod invasion process, from temporal trends and biogeographic patterns, to pathways and vectors, invaded habitats, and ecological and economical impacts. A total of 1590 species alien to Europe, including crustaceans, myriapods, mites, spiders, and insects, are listed in two volumes and 21 separate chapters that detail the different taxonomic groups. For each species, all key information - feeding regime, date and country of fi rst record in Europe, invaded countries, invaded habitats, plant or animal host - is supplied. More detailed factsheets are provided for the 80 species considered to be most representative of the different pathways of...

Unlike other groups of animals and plants, no checklist of alien terrestrial invertebrates was available in any of the European countries until recently. Since 2002, such checklists were successively provided by Austria (Essl and Rabitsch 2002), Germany (Geiter et al. 2002), the Czech Republic (Šefrová and Laštů vka 2005), Scandinavia (NOBANIS 2007), the United Kingdom (Hill et al. 2005), Switzerland (Wittenberg 2006) and Israel (Roll et al. 2007). However, most European regions remained uncovered and, furthermore, comparisons between the existing lists were inherently difficult because they used different definitions of alien. Thus, estimating the importance of terrestrial alien invertebrates at the European level remained impossible, mostly because of poor taxonomic knowledge existed for several groups. By gathering taxonomists and ecologists specialised on most invertebrate taxa together with collaborators working at the national level in 35 European countries, the DAISIE project intended to fill this gap. However, a lack of European expertise in some taxonomic groups did not allow coverage of all the terrestrial invertebrates with the same level of precision. Data on insects were more reliable than those of other taxa, and consequently the analyses presented below will mostly refer to this group.

Alien terrestrial arthropods of Europe Edited by Alain ROQUES, Marc KENIS, David LEES, Carlos LOPEZ-VAAMONDE, Wolfgang RABITSCH, Jean-Yves RASPLUS and David B. ROY Sofia–Moscow 2010 BioRisk 4(1) (Special Issue) Alien terrestrial arthropods of Europe Edited by Alain Roques, Marc Kenis, David Lees, Carlos Lopez-Vaamonde, Wolfgang Rabitsch, Jean-Yves Rasplus And David B. Roy This work was supported by a grant from the Sixth Framework Programme of the European Commission under the project DAISIE (Delivering Alien Species Inventories in Europe), contract SSPI-CT-2003-511202. We thank very much Jean-Marc Guehl (INRA department of «Ecologie des Forêts, Prairies et Milieux Aquatiques») and Olivier Le Gall (INRA department of «Santé des Plantes et Environnement ») for their financial help which allowed to publish this book. We are also very grateful to all colleagues who gently supplied us photos to illustrate the alien species: Henri-Pierre Aberlenc, C. van Achterberg, Daniel Adam, G. Allegro, J.J. Argoud, Margarita Auer, Juan Antonio Ávalos, Ab Baas, Antony Barber, Claude Bénassy, Christoph Benisch, C. van den Berg, Mark Bond, Nicasio Brotons, Gert Brovad, Peter J. Bryant, David Capaert, Jérôme Carletto, Rémi Coutin, David Crossley, Györgi Csóka, Massimiliano Di Giovanni, Joyce Gross, L. Goudzwaard, Jan Havelka, Jean Haxaire, Franck Hérard, R. Hoare, R. Kleukers, Zoltán Korsós, Gernot Kunz, Jørgen Lissner, Jean-Pierre Lyon, Mike Majerus†, Kiril Makarov, Chris Malumphy, Erwin Mani, Paolo Mazzei, Tom Murray, Louis-Michel Nageilesen, Laurence Ollivier, Jean-Pierre Onillon, Claude Pilon, Francesco Porcelli, Jean-Paul Raimbault, Urs Rindlisbacher, Gaëlle Rouault, Gilles San Martin, R.H. Scheffrahn, Vaclav Skuhravý, John I. Spicer, Massimo Vollaro, Jordan Wagenknecht, Beate Wermelinger, Alex Wild, Vassily Zakhartchenko, and the Montpellier Station of the Laboratoire National de Protection des Végétaux, France. Olivier Denux did a great job in realizing all the distribution maps. First published 2010 ISBN 978-954-642-554-6 (paperback) Pensoft Publishers Geo Milev Str. 13a, Sofia 1111, Bulgaria Fax: +359-2-870-42-82 info@pensoft.net www.pensoft.net Printed in Bulgaria, July 2010 Contents 1 DAISIE and arthropod invasions in Europe Philip E. Hulme & David B. Roy 5 Chapter 1. Introduction Wolfgang Nentwig & Melanie Josefsson 11 Chapter 2. Taxonomy, time and geographic patterns Alain Roques 27 Chapter 3. Pathways and vectors of alien arthropods in Europe Wolfgang Rabitsch 45 Chapter 4. Invaded habitats Carlos Lopez-Vaamonde, Milka Glavendekić & Maria Rosa Paiva 51 Chapter 5. Impact of alien terrestrial arthropods in Europe Marc Kenis & Manuela Branco 73 Chapter 6. Future trends Jean-Yves Rasplus 81 Chapter 7.1. Alien terrestrial crustaceans (Isopods and Amphipods) Pierre-Olivier Cochard, Ferenc Vilisics & Emmanuel Sechet 97 Chapter 7.2. Myriapods (Myriapoda) Pavel Stoev, Marzio Zapparoli, Sergei Golovatch, Henrik Enghoff, Nesrine Akkari & Anthony Barber 131 Chapter 7.3. Spiders (Araneae) Wolfgang Nentwig & Manuel Kobelt 149 Chapter 7.4. Mites and ticks (Acari) Maria Navajas, Alain Migeon, Agustin Estrada-Peña, Anne-Catherine Mailleux, Pablo Servigne & Radmila Petanović 193 Chapter 8.1. Longhorn beetles (Coleoptera, Cerambycidae) Christian Cocquempot & Åke Lindelöw 219 Chapter 8.2. Weevils and Bark Beetles (Coleoptera, Curculionoidea) Daniel Sauvard, Manuela Branco, Ferenc Lakatos, Massimo Faccoli & Lawrence R. Kirkendall 267 Chapter 8.3. Leaf and Seed Beetles (Coleoptera, Chrysomelidae) Ron Beenen & Alain Roques 293 Chapter 8.4. Ladybeetles (Coccinellidae) Helen Roy & Alain Migeon 315 Chapter 8.5. Coleoptera families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae Olivier Denux & Pierre Zagatti 407 Chapter 9.1. True bugs (Hemiptera, Heteroptera) Wolfgang Rabitsch 435 Chapter 9.2. Aphids (Hemiptera, Aphididae) Armelle Cœur d’acier, Nicolas Pérez Hidalgo & Olivera Petrović-Obradović 475 Chapter 9.3. Scales (Hemiptera, Superfamily Coccoidea) Giuseppina Pellizzari & Jean-François Germain 511 Chapter 9.4. Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea) and Hemiptera Auchenorrhyncha David Mifsud, Christian Cocquempot, Roland Mühlethaler, Mike Wilson & Jean-Claude Streito Abbreviations and glossary of technical terms used in the book Alain Roques & David Lees List of Authors David Agassiz Department of Entomology, The Natural History Museum, Cromwell Road, London, SW7 5BD, UK (dcl@nhm.ac.uk); The Garden House, Stafford Place, Weston-super-Mare, BS23 2QZ, UK; (agassiz@btinternet.com) Nesrine Akkari Research Unit of Biodiversity and Biology of Populations, Institut Supérieur des Sciences Biologiques Appliquées de Tunis, 9 Avenue Dr. Zouheir Essafi, La Rabta, 1007 Tunis, Tunisia; (nesrineakkari@gmail.com) Sylvie Augustin INRA UR633 Zoologie Forestiere, 2163 Av. Pomme de Pin, 45075 Orléans, France; (sylvie. augustin@orleans.inra.fr) Yuri Baranchikov Department of Forest Zoology, V.N.Sukachev Institute of Forest, Siberian Branch, Russian Academy of Science, 50 Akademgorodok, Krasnoyarsk 660036, Russia; (baranchikov_ yuri@yahoo.com) Anthony Barber Rathgar, Exeter Road, Ivybridge, Devon, PL21 0BD, UK; (tony@barber-jones.com) Ron Beenen Universiteit van Amsterdam, Zoölogisch Museum Amsterdam, Plantage Middenlaan 64, 1018 DH Amsterdam, The Netherlands; (r.beenen@wxs.nl) Manuela Branco Centro de Estudos Florestais, Instituto Superior de Agronomia, Technical University of Lisbon, Tapada da Ajuda, 1349-017 Lisboa, Portugal; (mrbranco@isa.utl.pt) Pierre-Olivier Cochard 113 Grande rue Saint-Michel, 31400 Toulouse, France; (pierre-olivier.cochard@wanadoo.fr) Christian Cocquempot Institut National de la Recherche Agronomique, UMR CBGP (INRA/IRD/Cirad/Montpellier SupAgro), Campus International de Baillarguet, CS 30016, F-34988, Montferrier-sur-Lez Cedex, France; (cocquemp@supagro.inra.fr) Armelle Cœur d’acier Institut National de la Recherche Agronomique, UMR CBGP (INRA/IRD/Cirad/Montpellier SupAgro), Campus International de Baillarguet, CS 30016, F-34988, Montferrier-sur-Lez Cedex, France; (coeur@supagro.inra.fr) Ejup Çota Plant Protection Department, Faculty of Agriculture and Environment, Agriculture University of Tirana, Albania; (ejupcota@gmail.com) Gérard Delvare Institut National de la Recherche Agronomique, UMR CBGP (INRA/IRD/Cirad/Montpellier SupAgro), Campus International de Baillarguet, CS 30016, F-34988, Montferrier-sur-Lez Cedex, France; (delvare@supagro.inra.fr) Olivier Denux INRA - Centre de recherche d’Orléans, Unité de Zoologie Forestière, 2163 Avenue de la Pomme de Pin - CS 40001 ARDON, 45075 Orléans Cedex 2, France; (olivier.denux@ orleans.inra.fr) Jurate De Prins Entomology Section, Royal Museum for Central Africa, Leuvensesteenweg 13, B-3080 Tervuren, Belgium; (jurate.de.prins@africamuseum.be) Willy De Prins Entomology Section, Royal Museum for Central Africa, Leuvensesteenweg 13, B-3080 Tervuren, Belgium; (willy.deprins@gmail.com) Henrik Enghoff Natural History Museum of Denmark (Zoological Museum), University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen, Denmark; (henghoff@snm.ku.dk) Agustin Estrada-Peña Faculty of Veterinary Medicine, Department of Parasitology, Miguel Servet 177, 50013-Zaragoza, Spain; (aestrada@unizar.es) Massimo Faccoli Universita di Padova, Department of Environmental Agronomy and Crop Sciences, Agripolis, Viale dell’Università 16, 35020 Legnaro, Italy; (massimo.faccoli@unipd.it) Jean-François Germain Laboratoire National de la Protection des Végétaux, Station de Montpellier, CBGP Campus international de Baillarguet CS 30016, 34988 Montferrier-sur-Lez Cedex, France; (germain@supagro.inra.fr) Milka M. Glavendekić University of Belgrade, Faculty of Forestry, Kneza Viseslava 1, 11030 Belgrade, Serbia; (milka. glavendekic@nadlanu.com) Sergei Golovatch Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky prospekt 33, Moscow 119071 Russia; (sgol@orc.ru) Stanislav Gomboc Siskovo naselje 19, SI-4000 Kranj, Slovenia; (stane.gomboc@gov.si) Philip E. Hulme The Bio-Protection Research Centre, Lincoln University, Christchurch, New Zealand; (hulmep@lincoln.ac.nz) Povilas Ivinskis Nature Investigation Centre, Institute of Ecology, Akademijos str. 2, Lt 08412 Vilnius, Lithuania; (ivinskis@ekoi.lt) Melanie Josefsson Swedish Environmental Protection Agency, c/o Department of Environmental Monitoring, P.O. Box 7050, SE 750 07 Uppsala, Sweden; (melanie.josefsson@snv.slu.se) Ole Karsholt The Natural History Museum of Denmark, Zoologisk Museum, Universitetsparken 15, DK2100 København Ø, Denmark; (okarsholt@snm.ku.dk) Marc Kenis CABI Europe-Switzerland, 1, Rue des Grillons, CH- 2800, Delémont, Switzerland; (m.kenis@ cabi.org) Lawrence Kirkendall University of Bergen, Biology Institute, Postbox 7803, N-5020, Bergen, Norway; (lawrence. kirkendall@bio.uib.no) Manuel Kobelt Community Ecology, Institute of Ecology and Evolution, University of Bern, Baltzerstrasse 6, CH-3012 Bern, Switzerland. Zoltán Korsós Zoological Department, Hungarian Natural History Museum H-1088 Budapest, Hungary (korsos@zoo.zoo.nhmus.hu) Fotini Koutroumpa 98 Zahou st 38333, Volos, Greece; (fotini.koutroumpa@gmail.com) Athanasios Koutroumpas 98 Zahou st 38333, Volos, Greece; (fotini.koutroumpa@gmail.com) Ferenc Lakatos University of West-Hungary, Institute of Sylviculture and Forest Protection, Bajcsy-Zs. u. 4., H-9400 Sopron, Hungary; (flakatos@emk.nyme.hu) Zdeněk Laštůvka Department of Zoology, Fisheries, Hydrobiology and Apidology, Faculty of Agronomy, Mendel University in Brno, Zemědělská 1, CZ-613 00 Brno, Czech Republic; (last@mendelu.cz) Yves Le Conte Institut National de la Recherche Agronomique (INRA), UMR0406 AE Abeilles et Environnement, Domaine Saint-Paul - Site Agroparc 84914 Avignon, France; (yves.leconte@avignon.inra.fr) David Lees INRA UR633 Zoologie Forestiere, 2163 Av. Pomme de Pin, 45075 Orléans, France; (david. lees@orleans.inra.fr) Åke Lindelöw Swedish university of agricultural sciences, Department of ecology. P.O. Box 7044, S-750 07 Uppsala, Sweden; (Ake.Lindelow@ekol.slu.se) Carlos Lopez-Vaamonde INRA UR633 Zoologie Forestiere, 2163 Av. Pomme de Pin, 45075 Orléans, France; (carlos. lopez-vaamonde@orleans.inra.fr) Anne-Catherine Mailleux Université catholique de Louvain, Unité d’écologie et de biogéographie, local B165.10, Croix du Sud, 4-5 (Bâtiment Carnoy), B-1348 Louvain-La-Neuve, Belgium; (Anne-Catherine. Mailleux@uclouvain.be) Eduardo Marabuto CBA - Centro de Biologia Ambiental, Faculdade Ciências Universidade de Lisboa, Campo Grande, edificio C2 - Lisboa, Portugal; (edu_marabuto@netcabo.pt) Michel Martinez INRA Centre de Biologie pour la Gestion des Populations (CBGP), Campus International de Baillarguet, 34988 Montferrier-sur-Lez, France; (martinez@supagro.inra.fr) David Mifsud Junior College, Department of Biology, University of Malta, Msida MSD 1252, Malta; (david.a.mifsud@um.edu.mt) Alain Migeon Institut National de la Recherche Agronomique, UMR CBGP (INRA/IRD/Cirad/Montpellier SupAgro), Campus International de Baillarguet, CS 30016, F-34988, Montferrier-sur-Lez Cedex, France; (migeon@supagro.inra.fr) Ljubodrag Mihajlović University of Belgrade, Faculty of Forestry, Kneza Viseslava 1, 11030 Belgrade, Serbia; (mljuba@EUnet.rs) Leen Moraal Alterra, Wageningen UR, Centre Ecosystems, PO Box 47, NL-6700 AA Wageningen, The Netherlands; (Leen.Moraal@wur.nl) Roland Mühlethaler Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity, Humboldt University Berlin, Invalidenstrasse 43, 10115 Berlin, Germany; (roland.muehlethaler@mfn-berlin.de), Franck Muller Museum National d‘Histoire Naturelle Entomologie CP50, 45 rue Buffon, 75005 Paris, France Maria Navajas Institut National de la Recherche Agronomique, UMR CBGP (INRA/IRD/Cirad/Montpellier SupAgro), Campus International de Baillarguet, CS 30016, F-34988, Montferrier-sur-Lez Cedex, France; (navajas@supagro.inra.fr) Wolfgang Nentwig Community Ecology, Zoological Institute University of Bern, Baltzerstrasse 6, CH-3012 Bern, Switzerland; (wolfgang.nentwig@iee.unibe.ch) Elisenda Olivella Museu de Ciències Naturals de Barcelona (Zoologia), Passeig Picasso s/n, E-08003 Barcelona, Spain; (eolivell@xtec.cat) Maria Rosa Paiva DCEA, Faculdade de Ciências e Tecnologia, Universidade Nova de Lisboa, 2829-516 Campus de Caparica, Lisbon, Portugal; (mrp@fct.unl.pt) Giuseppina Pellizzari Università di Padova - Dipartimento Agronomia Ambientale e Produzioni Vegetali, Agripolis - Viale dell’Università 16, 35020 Legnaro Padova, Italia; (giuseppina.pellizzari@unipd.it) Nicolas Pérez Hidalgo Universidad de León, Facultad de Ciencias Biológicas y Ambientales, Universidad de León, 24071, León, Spain (nperh@unileon.es) Radmila Petanović Department of Entomology and Agricultural Zoology, Faculty of Agriculture University of Belgrade, Nemanjina 6, Belgrade-Zemun,11080 Serbia; (rpetanov@agrif.bg.ac.rs) Olivera Petrović-Obradović University of Belgrade, Faculty of Agriculture, Nemanjina 6, SER-11000, Belgrade, Serbia; (petrovic@agrif.bg.ac.rs) Lukasz Przybylowicz Polish Academy of Sciences, Institute of Systematics and Evolution of Animals, Slawkowska 17; 31-016 Krakow; Poland; (lukasz@isez.pan.krakow.pl) Wolfgang Rabitsch Environment Agency Austria, Dept. Biodiversity & Nature Conservation, Spittelauer Lände 5, 1090 Vienna, Austria; (wolfgang.rabitsch@umweltbundesamt.at) Jean-Yves Rasplus Institut National de la Recherche Argonomique, UMR Centre de Biologie et de Gestion des Populations, CBGP, (INRA/IRD/CIRAD/Montpellier SupAgro), Campus international de Baillarguet, CS 30016, 34988 Montferrier-sur Lez, France; (rasplus@supagro.inra.fr) Hans Peter Ravn Forest & Landscape Denmark, University of Copenhagen, Hoersholm Kongevej 11, DK-2970 Hoersholm, Denmark; (hpr@life.ku.dk) Philippe Reynaud Laboratoire national de la protection des végétaux, Station d’Angers, 7 rue Jean Dixméras, 49044 Angers Cedex 01, France; (philippe.reynaud@agriculture.gouv.fr) Quentin Rome Museum National d’Histoire Naturelle Entomologie CP50, 45 rue Buffon, 75005 Paris, France (vespa.velutina@gmail.com) Alain Roques Institut National de la Recherche Agronomique (INRA), UR 0633, Station de Zoologie Forestiere, 2163 Av. Pomme de Pin, 45075 Orléans, France; (alain.roques@orleans.inra.fr) David B. Roy Centre for Ecology & Hydrology, Crowmarsh Gifford, Oxfordshire, OX10 8BB, United Kindgom; (dbr@ceh.ac.uk) Helen Roy NERC Centre for Ecology & Hydrology, Biological Records Centre, Crowmarsh Gifford, Oxfordshire, OX10 8BB, United Kindgom; (hele@ceh.ac.uk) Nils Ryrholm Department of Natural Sciences, University of Gävle, S-801 76 Gävle, Sweden; (Nils.Ryrholm@ hig.se) Daniel Sauvard INRA UR633 Zoologie Forestiere, 2163 Av. Pomme de Pin, 45075 Orléans, France; (daniel. sauvard@orleans.inra.fr) Martin H Schmidt-Entling University of Bern, Institute of Ecology and Evolution, Community Ecology, CH-3012 Switzerland; (martin.schmidt@zos.unibe.ch) Nico Schneider 79, rue Tony Dutreux, L-1429 Luxembourg-Bonnevoie, Luxemburg; (nico.schneider@education.lu) Emmanuel Sechet 20 rue de la Résistance, 49125 Cheffes, France; (e-sechet@wanadoo.fr) Hana Šefrová Department of Crop Science, Breeding and Plant Medicine, Faculty of Agronomy, Mendel University in Brno, Zemědělská 1, CZ-613 00 Brno, Czech Republic; (sefrova@mendelu.cz) Pablo Servigne Service d’Ecologie Sociale, Université libre de Bruxelles, CP231, Avenue F. D. Roosevelt, 50, B-1050 Brussels, Belgium; (pablo.servigne@ulb.ac.be) Peter Šima Koppert Biological Systems, Komárňanská cesta 13, 940 01 Nové Zámky, Slovakia; (psima@ koppert.sk) Ian Sims Syngenta International Research Centre, Jealott’s Hill, Bracknell, Berkshire RG42 6EY; (ian. sims@syngenta.com) Sergey Sinev Zoological Institute RAS, Universitetskaya nab. 1, 199034 St.Petersburg, Russia; (sinev@zin.ru) Marcela Skuhravá Bítovská 1227/9, 140 00 Praha 4, Czech Republic; (skuhrava@quick.cz) Bjarne Skulev Brøndsted 411, DK-3670 Veksø, Denmark; (uglebo@mail.dk) Pavel Stoev National Museum of Natural History, Tsar Osvoboditel Blvd. 1, 1000 Sofia, Bulgaria; (pavel.e.stoev@gmail.com) Jean-Claude Streito Laboratoire national de la protection des végétaux, CBGP Campus international de Baillarguet, CS 30016, 34988 Montferrier-sur-Lez cedex, France; (streito@supagro.inra.fr) Rumen Tomov University of Forestry, 10 Kliment Ohridski blvd., 1756 Sofia, Bulgaria; (rtomov@yahoo.com) Georgyi Trenchev University of Forestry, 10 Kliment Ohridski blvd., 1756 Sofia, Bulgaria; (k_trencheva@yahoo. com) Katia Trencheva University of Forestry, 10 Kliment Ohridski blvd., 1756 Sofia, Bulgaria; (k_trencheva@yahoo. com) Katalin Tuba University of West-Hungary, Institute of Silviculture and Forest Protection, Bajcsy-Zs. u. 4. 9400 Sopron, Hungary; (tubak@emk.nyme.hu) Ferenc Vilisics Szent István University, Faculty of Veterinary Sciences, Institute for Biology, H-1077, Budapest, Rottenbiller str. 50., Hungary; (vilisics.ferenc@gmail.com) Claire Villemant Museum National d’Histoire Naturelle, UMR Origine, Structure et Evolution de la Biodiversite, OSEB, (MNHN/CNRS) CP50, 45 rue Buff on, 75005 Paris, France; (villeman@mnhn.fr) Mike Wilson Department of Biodiversity & Systematic Biology, National Museum Wales, Cathays Park, Cardiff CF10 3NP, United Kingdom; (mike.wilson@museumwales.ac.uk) Pierre Zagatti INRA – Centre de recherche de Versailles, Unité PISC, Route de Saint-Cyr, 78026 Versailles Cedex, France; (pierre.zagatti@versailles.inra.fr) Marzio Zapparoli Universita degli Studi della Tuscia, Dipartimento di Protezione delle Piante, via S. Camillo de Lellis s.n.c., I-01100 Viterbo, Italy; (zapparol@unitus.it) Jürg Zettel University of Bern, Institute of Ecology and Evolution Baltzerstrasse 6, CH-3012 Bern, Switzerland; (juersi.zettel@bluewin.ch) Alberto Zilli Museo Civico di Zoologia, Via U. Aldrovandi 18, I-00197 Rome, Italy; (alberto.zilli@comune. roma.it) A peer reviewed open access journal BioRisk 4(1): 1–3 (2010) doi: 10.3897/biorisk.4.41 EDITORIAL BioRisk www.pensoftonline.net/biorisk DAISIE and arthropod invasions in Europe Philip E. Hulme1, David B. Roy2 1 The Bio-Protection Research Centre, Lincoln University, Christchurch, New Zealand 2 Centre for Ecology & Hydrology, Crowmarsh Gifford, Wallingford, United Kingdom Corresponding authors: Philip E. Hulme (philip.hulme@lincoln.ac.nz), David B. Roy (dbr@ceh.ac.uk) Academic editor: Alain Roques | Received 21 January 2010 | Accepted 18 May 2010 | Published 6 July 2010 Citation: Hulme PE, Roy DB (2010) DAISIE and arthropod invasions in Europe. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 1–3. doi: 10.3897/biorisk.4.41 A milestone in the knowledge of alien species in Europe has been achieved by the DAISIE (Delivering Alien Invasive Species Inventories for Europe) project. Through the Sixth Framework Programme of the European Union, DAISIE has delivered a major portal for information on biological invasions that is publicly available at http://www. europe-aliens.org. The rationale was to develop a pan-European inventory of invasive alien species by integrating existing databases, to describe patterns and evaluate trends in biological invasions in Europe, identify priority species and assess their ecological, economic and health risks and impacts. Although an on-going process, the foundation, scope, and technological architecture of DAISIE was established through a consortium of leading researchers of biological invasions in Europe from 19 institutions across 15 countries and delivered through the cooperation of experts in ecology and taxonomy from throughout Europe that in total amounted to 182 contributors. The inventory, accounts, and distribution maps today provide the first qualified reference system on invasive alien species for the European region. The information presents an outstanding resource to synthesise current knowledge and trends in biological invasions in Europe. The data will help identify the scale and spatial pattern of invasive alien species in Europe, understand the environmental, social, economic and other factors involved in invasions, and can be used as a framework for considering indicators for early warning. A key component of DAISIE is The European Alien Species Database, an inventory of all alien species in Europe, and resulted from compiling and peer-reviewing national and regional lists of alien fungi, bryophytes, vascular plants, invertebrates, fish, amphibians, reptiles, birds and mammals. Data were collated for all 27 European Union member states (and separately for their significant island regions), other European states (Andorra, Iceland, Liechtenstein, Moldova, Monaco, Norway, the European Copyright P.E. Hulme, D.B. Roy. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 2 Philip E. Hulme & David B. Roy / BioRisk 4(1): 1–3 (2010) part of Russia, Switzerland, Ukraine, former Yugoslavia states) and Israel. Marine lists are referenced to the appropriate political region with administrative responsibility. To have full coverage of the European marine area, the data for countries bordering the Mediterranean Sea in North African and Near East countries are included. By November 2008, records of 10,771 alien species, were included in the database, belonging to 4492 genera and 1267 families. Both species of exotic origin and species of European origin introduced in European regions outside their native range were considered. Plants are most represented accounting for 55% of all taxa (5789 species), terrestrial invertebrates 23% (2477 species), followed by vertebrates (6%), fungi (5%), molluscs (4%), Annelida (1%) and Rhodophyta (1%). In total, the database includes records of 45,211 introduction events to particular regions (plants: 28,093; terrestrial invertebrates: 11,776; aquatic marine species: 2777, terrestrial vertebrates: 1478; aquatic inland species: 1087). Due to unprecedentedly thorough assessment, DAISIE substantially improved the accuracy of estimates of alien species numbers derived from previous datasets. The information accumulated by DAISIE has been summarized in the Handbook of Alien Species in Europe (DAISIE 2009), which contains analytical chapters on each taxonomic group, and fact sheets of the 100 most invasive alien species in Europe with distribution maps and images. The book also lists all alien species recorded, ranked taxonomically; this list can be used as a reference for future assessment of trends in biological invasions in Europe. The current volume “Alien terrestrial arthropods of Europe” largely follows the lead set by the Handbook of Alien Species in Europe but provides much needed detail on one of the largest and most complex taxonomic groups, the arthropods. Unlike other groups of animals and plants, no checklist of alien terrestrial invertebrates was available in any of the European countries until the beginning of this century. Thus more than any other taxonomic group, creating an inventory of invasive alien arthropods in Europe proved to be a major challenge. Consequently, an estimate of the importance of terrestrial alien invertebrates at the European level remained impossible, largely due to the limited taxonomic knowledge regarding several major arthropod groups. As a result, the initial analyses in DAISIE were drawn from the most reliably studies group, the insects. Even with such a partial picture, the new evidence emphasised the need for more detailed assessment of alien arthropods. For example, the initial work in DAISIE has shown that approximately 90% of terrestrial insects having arrived into Europe unintentionally (75% associated with a commodity, 15% as stowaways). The highest numbers of insects occur in human-made habitats (ruderal, cultivated land, parks and gardens) and invasions are concentrated to these few highly invaded habitats. Not surprisingly insects are one of the taxonomic groups with the most species causing impacts in Europe, and most of these impacts are on the economic rather than environmental sectors. In this regard, Alien terrestrial arthropods of Europe extends the initial work in DAISIE and develops a clearer picture of arthropod invasions across a much larger taxonomic range than insects. This substantial work will set the benchmark for authoritative assessments of invasive terrestrial invertebrates. DAISIE and arthropod invasions in Europe 3 Through DAISIE, Europe is today the continent with the most complete information on its alien biota. The continent has been working towards implementing an effective strategy on invasive alien species and DAISIE is considered as one of the major instruments towards achieving this goal. An internet-accessible knowledge base, such as DAISIE, can provide crucial information for the early detection, eradication, and containment of invasive aliens —which is most achievable for species that have just arrived. As a result of DAISIE, managers and policy-makers addressing the invasive alien species challenge can easily obtain data on which species are invasive or potentially invasive in particular habitats, and use this information in their planning efforts. Agencies responsible for pest control can quickly determine if a species of interest has been invasive elsewhere in Europe. Importers of new alien species can access data to make responsible business choices. Land managers can learn about control methods that have been useful in other areas, reducing the need to commit resources for experimentation and increasing the speed at which control efforts can begin. DAISIE is potentially a model for other continents which currently have much less detailed information on their alien biota. References DAISIE (2009) Handbook of Alien Species in Europe. Dordrecht: Springer. A peer reviewed open access journal BioRisk 4(1): 5–9 (2010) doi: 10.3897/biorisk.4.43 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Introduction Chapter 1 Wolfgang Nentwig1, Melanie Josefsson2 1 Community Ecology, University of Bern, Baltzerstrasse 6, CH-3012 Bern, Switzerland 2 Swedish Environmental Protection Agency, c/o Department of Environmental Monitoring, P.O. Box 7050, SE 750 07 Uppsala, Sweden. Corresponding authors: Wolfgang Nentwig (wolfgang.nentwig@iee.unibe.ch), Melanie Josefsson (melanie. josefsson@snv.slu.se) Academic editor: Alain Roques | Received 7 January 2010 | Accepted 18 May 2010 | Published 6 July 2010 Citation: Nentwig W, Josefsson M (2009) Introduction. Chapter 1. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 5–9. doi: 10.3897/biorisk.4.43 Dispersal of organisms is among the most important conditions that has enabled the development of life on earth and the high diversity of species we encounter today. This natural process is guided by biogeographical barriers which subdivide the accessible space of the Earth into compartments: species are limited to islands, summits, lakes, or oceans and shorelines, mountain ridges or climate zones. Such natural boundaries reduce competition, create conditions for speciation, and form the basis for the evolutionary centre where a given species has originated. This species is then native (indigenous) to this area. These natural biogeographical barriers have increasingly been overcome by human dispersal and humans now inhabit all parts of the world. This process of human dispersal started in Africa more than 100,000 years ago, and is an intrinsic part of human history. At first, this slow but continuous conquest was performed by walking, at the natural speed of humans, and was limited by the physical condition of individuals. The speed of movements increased in the last centuries and today, we can reach virtually any spot on earth by airplane within 24 h. The turning point was certainly, when sailing ships circumnavigated the world and connected continents. With such big carriers, mass transportation of materials, animals and plants over large distances was also possible. Christopher Columbus was the second European in the New World (the first discovery of North America by the Vikings some 500 years earlier had no long-lasting consequence, other than the introduction of the North American bivalve Mya arenaria Copyright W. Nentwig, M. Josefsson This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 6 Wolfgang Nentwig & Melanie Josefsson / BioRisk 4(1): 5–9 (2010) to Scandinavia in the 1200s (Petersen 1992), and with him the global race duel to connect all parts of the world faster and tighter began. Thus, the year 1492, when Columbus set foot on the first Caribbean island was the starting point of this self-accelerating process later called globalisation. This process had serious consequences because man did not travel alone. His entourage comprised crop plants and domesticated animals and pets, including all the pests, pathogens and parasites which usually adhere to them. In other words: in the last 500 years hundreds and thousands of species have been spread worldwide both intentionally and unintentionally. Through this human aided spread the biogeographical barriers have become more and more permeable and more and more species are no longer restricted to their native areas. Species living outside of their natural range and outside of their natural dispersal potential are alien species. Their presence in the new habitat is due to intentional or unintentional human activities and without this human support they would never have reached their new area. Thus, there is an important difference between natural dispersal of species that, e.g., allows Mediterranean species to spread north of the Alps because the summers are becoming warmer and man-mediated transport of American, African, Asian or Australian species which then suddenly show up in European harbours or airports and disperse into the hinterland. These last species are called alien to Europe. Obviously, species of European origin may also be translocated by man outside of their natural range, e.g. Mediterranean species to Northern Europe or species of continental Europe to Atlantic and Mediterranean islands. In this case, they are called alien in Europe. However, in many cases it appears highly difficult to disentangle the effect of human-mediated transport from that of natural dispersion when a native European species is suddenly found outside its range. But why is it disadvantageous to increase the number of naturally occurring species (the native fauna and flora) by some alien species? In most (if not all) natural ecosystems the given set of species is the result of a long adaptation and co-evolution to the physical and biotic environment. The higher the natural biological diversity is, the greater the biotic resistance is against additional, foreign species. If ecosystems are disturbed (e.g. by fire, flooding or erosion) or are artificial ecosystems (such as agricultural habitats or urban areas), alien species have a much higher chance to establish. An alien species will interact with resident species or the abiotic environment in a different manner than a native species and therefore such an additional species is usually neither an enrichment of the ecosystem nor any amelioration of a process. Alien species are usually somehow different from the resident species since they have evolved in a different environment. They may represent a new type of predator, they may have novel weapons, or they may have other new properties which may enable them to alter habitats or even ecosystem functioning. They can fill hitherto empty niches, they may change matter flux or impact energy flow. Such changes affect the resident species most often in a negative way and native species may become less common or even disappear. At this stage, the alien species impacts the invaded ecosystem and becomes an invasive Introduction. Chapter 1 7 species. Usually the term “alien” is used in the sense of “not wanted here” but calling it invasive is a clearly negative attribute. The consequences of an alien species can be manifold: Most obvious is direct competition with native species, an increasing abundance in the new environment until a complete replacement of native residents occurs. Alien species may be associated with pathogens and parasites or they are pathogens and parasites, which may transfer onto and affect a new host. If the new host is susceptible to the new pathogen or parasite, a strong reduction in the population of this native species will result or even local extinction is possible: The alien species has thus caused a loss of biodiversity. Further consequences of an alien and invasive species may concern water flux, e.g. by increasing consumption or contamination. Matter flux (primarily carbon or nitrogen) may be influenced by an altered decomposition of plant litter and wood or via nitrogen-fixating symbionts. Besides such environmental impacts many alien species cause enormous economic impacts or directly influence human or animal health. Many alien invertebrates, especially insects, cause great damage to agriculture and forestry. Many protozoans and “worms” are human parasites and many insects are vectors of bacteria and viruses which cause numerous serious diseases. Today, such super-pests are cosmopolitan but this term camouflages that in most parts of the world, where they occur today, they are alien and invasive species. In the case of humans and on a global scale, they cause millions of fatalities each year. Not all alien species are invasive and it is in fact strange to observe some aliens for years and decades at a given location that show no signs of obvious spread. The process from the first introduction of an alien species into a new environment until aggressive invasiveness is characterised by several steps and an alien species may fail at each of these steps. After a first introduction, it is decisive if the new environment fits the need of this species. Usually, if the number of individuals is low, the species has a rather small chance of establishing reproducing populations. But the higher this number is or the longer the introduction process lasts, the better the chances are of the new species establishing. Establishment means survival and reproducing viable populations on the spot, which is called the lag phase. The next step is when the alien species produces a surplus reproduction which allows modest migration. In this period an alien species may adapt in some way to its new environment and this phase is often called bottleneck with a transition from the lag phase to the log phase. In the log phase, the alien species reaches more suitable habitats which allow a higher reproduction. By continuous population growth, the population pressure on adjacent areas is increased and impacts on the ecosystem also become evident and increase: now the alien has become invasive. Observing an alien in a non-invasive status does not mean that it will not become invasive (and thus can be tolerated as harmless), it rather means that it is not (yet) invasive but it could be just a matter of time until it becomes invasive. Changes in land use or climate can also enable previously harmless alien species to begin to spread uncontrollably and become invasive. 8 Wolfgang Nentwig & Melanie Josefsson / BioRisk 4(1): 5–9 (2010) Roughly 50 years ago, the British ecologist Charles Elton published his Ecology of invasions by animals and plants, already then warning of the danger arising from alien and invasive species: “The whole matter goes far wider than any technological discussion of pest control, though many of the examples are taken from applied ecology. The real thing is that we are living in a period of the world’s history when the mingling of thousands of kinds of organisms from different parts of the world is setting up terrific dislocations in nature. We are seeing huge changes in the natural population balance of the world” (Elton 1958). Elton was among the first to describe the typical pattern of an alien species establishment. That what he called “biological explosion” is today known as biological invasion (Nentwig 2008). He was also among the first to investigate why and how species were dispersed by human activities and he analysed even then the negative impacts of species in a new environment. He was among the first to ask how this could be prevented. Astonishingly, the hazards provoked by alien species did not cause that much concern among scientists, nor did it attract public awareness as much as would have been expected (Hulme et al. 2009). However, the ultimate reason for the loss of more than 5% of the world GNP, one main reason for the loss of biodiversity, for millions of human deaths, and for the loss of more than 20% of the world’s food production cannot be ignored. Prevention has multiple faces leading from raising awareness in the public to better scientific knowledge and documentation. More regulations and guidelines must to be put into place and existing regulations must be applied more consequently and carefully. Further import of aliens should be avoided; current aliens should be confined, controlled and even eradicated. We must face this challenge through changes in world trade, adoption of regional strategies and regulations, improved national legislation and better administration, but also through improvements in general education and awareness and the improved spread of information through the media. Science is also absolutely required in order to manage the problems that alien species may cause. How can they be detected and identified? What is their population development and habitat requirement? What is their impact in the invaded area? How can they be controlled, reduced, or eradicated? How can we predict which species that may become invasive and how can we manage the risks? For most alien species there are yet no answers to most of these questions. Even the seemingly simple question on the number of alien species in Europe could not been answered a few years ago. Therefore, the European Commission, in its Sixth Framework Programme, launched a call for an inventory of alien invasive species. The successful application was awarded to a consortium of leading researchers of biological invasions in Europe, drawn from 19 institutions across 15 countries. The resulting project, DAISIE (Delivering Alien Invasive Species Inventories for Europe), was launched in February 2005 and ran for three years, until the end of January 2008. The main objectives of DAISIE were (1) the creation of an inventory of all known alien species in the European terrestrial, freshwater and marine environments; (2) to describe the worst alien and invasive species in Europe and to describe their envi- Introduction. Chapter 1 9 ronmental, economic and health risks impacts; and (3) to compile a directory of experts on alien species. Since February 2008, the DAISIE information system is freely available at http://www.europe-aliens.org. In 2009 a condensed version of the DAISIE information system was published in a Handbook of Alien Species in Europe (DAISIE 2009). Invertebrates, and among them arthropods, comprise the largest proportion of alien animals and are of pronounced importance, e.g. in agriculture, horticulture and forestry, the cultural environment and for human and animal health. Despite the far reaching and serious effects that alien invertebrate species have on biological diversity, health and society, knowledge of their effects and potential risks is still insufficient. This knowledge is crucial for managing the risks involved with the transfer of species both intentionally and unintentionally. Based on the expert knowledge of 78 scientists from 25 European countries, this book will present for the first time in a comprehensive way the alien arthropods having established in Europe, including detailed information on taxonomy, pathways, invaded habitats, impacts and trends. The book will focus on the 1590 terrestrial arthropod species presently identified as aliens to Europe. They will be presented by taxonomic rank. For each group, additional information will be provided about the species alien in Europe whenever the actual status of such species can be considered as ascertained with regard to the difficulties mentioned above. Moreover, the 80 most important alien invasive species are presented in factsheets in more detail in order to raise awareness and provide information upon which to base measures to prevent and control these species. References DAISIE (2009) Handbook of alien species in Europe. Dordrecht: Springer. 399 pp. Elton CS (1958) The ecology of invasions by plants and animals. London: Methuen & Co. Ltd. 181 pp. Hulme PE, Pyšek P, Nentwig W, Vilà M (2009) Will threat of biological invasions unite the European Union? Science 324: 40–41. Nentwig W (Ed) (2008) Biological Invasions. Ecological Studies 193. Heidelberg: Springer. 441 pp. Petersen KS, Rasmussen KL, Heinemeier J, Rud N (1992) Clams before Columbus? Nature 359: 679. A peer reviewed open access journal BioRisk 4(1): 11–26 (2010) doi: 10.3897/biorisk.4.70 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Taxonomy, time and geographic patterns Chapter 2 Alain Roques INRA UR633 Zoologie Forestière, 2163 Av. Pomme de pin, 45075 Orléans, France Corresponding author: Alain Roques (alain.roques@orleans.inra.fr) Academic editor: David Lees | Received 15 April 2010 | Accepted 20 May 2010 | Published 6 July 2010 Citation: Roques A (2010) Taxonomy, time and geographic patterns. Chapter 2. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 11–26. doi: 10.3897/biorisk.4.70 Abstract A total of 1590 species of arthropods alien to Europe have already established on the continent, including 226 more or less cosmopolitan species of uncertain origin (cryptogenic). These alien species are dispersed across 33 taxonomic orders, including crustaceans, chilopods, diplopods, pauropods, Symphyla, mites, arachnids, and insects. However, insects largely dominate, accounting for more than 87% of the species, far in excess of mites (6.4%). Three of the insect orders, namely Coleoptera, Hemiptera and Hymenoptera, overall account for nearly 65 % of the total. The alien fauna seems to be highly diverse with a total of 257 families involved, of which 30 have no native representatives. However, just 11 families contribute more than 30 species, mainly aphids, scales and hymenopteran chalcids. For a number of families, the arrival of alien species has significantly modified the composition of the fauna in Europe. Examples are given. The number of new records of aliens per year has increased exponentially since the 16th century, but a significant acceleration was observed since the second half of the 20th century, with an average of 19.6 alien species newly reported per year in Europe between 2000 and 2008. This acceleration appears to be mainly related to the arrival of phytophagous species, probably with the plant trade, whereas the contribution of detritivores, parasitoids and predators has decreased. Some taxa have not shown any acceleration in the rate of arrivals. Asia has supplied the largest number of alien arthropods occurring in Europe (26.7 %), followed by North America (21.9%) but large differences in the region of origin are apparent between taxa. Once established, most alien species have not spread throughout Europe, at least yet, with 43.6 % of the species only present in one or two countries, and less than 1% present in more than 40 countries. Large differences also exist between European countries in the total number of alien arthropods recorded per country. Italy (700 species) and France (690 species), followed by Great Britain (533 species), host many more species than other countries. The number of alien species per country is significantly correlated with socioeconomic and demographic variables. Copyright Alain Roques. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 12 Alain Roques / BioRisk 4(1): 11–26 (2010) Keywords aliens, arthropods, Europe, globalization, taxonomy, Asia, drivers of biological invasion Introduction Expanding world-wide trade, globalisation of economies and climate change are all factors that contribute to an accelerated international movement and establishment of alien organisms, allowing them to overcome geographic barriers (Hulme et al. 2008, Hulme 2009, Walther et al. 2009, Roques 2010). These alien species have already been shown to impose enormous costs on agriculture, forestry as well as to threaten human health and biodiversity (Williamson 1996, Wittenberg and Cock 2001, Pimentel et al. 2002, 2005, Vilá et al. 2009). Although terrestrial arthropods, and particularly insects, represent a large part of the alien species problem, they appear to have received disproportionately less attention compared to plants, vertebrates, and aquatic organisms, especially regarding their possible ecological impact (Kenis et al. 2009). Most of the works concerning alien terrestrial invertebrates have dealt with case studies of pests having a high economic or sanitary impact, such as gypsy moth (Lymantria dispar (L.)) in North America (Liebhold et al. 1992), Asian long-horned beetles (Anoplophora spp.; Haack et al. 2010), or Asian tiger mosquito (Aedes albopictus (Skuse); Eritja et al. 2005). More synthetic studies have been carried out at guild level (e.g., bark beetles; Brockerhoff et al. 2005) or at ecosystem level, especially for forest insects (Liebhold et al. 1995, Mattson et al. 1996, 2007, Niemelä and Mattson 1996, Langor et al. 2009). However, continental inventories of alien arthropod species, or even of alien insects, are still lacking in most regions, although such studies are needed to assess which taxonomic or bio-ecological groups of alien species are better invaders or more harmful to the economy or environment, and which ecosystems or habitats are at greater risk (Mondor et al., 2007). In Europe, the potential problems caused by alien arthropods have traditionally been considered as less severe than in North America, Australasia or South Africa (Niemelä and Mattson 1996). As a result, unlike other groups of animals and plants, no checklist of alien terrestrial arthropods was available in any of the European countries until the early 2000s. However, in the last 20 years, several exotic pests of economic concern, to name a few, the western corn rootworm (Diabrotica virgifera virgifera LeConte), the red palm weevil (Rhynchophorus ferrugineus (Olivier)), the harlequin labybeetle (Harmonia axyridis (Pallas)), or the chestnut gall maker (Dryocosmus kuriphilus (Yasumatsu)), have invaded Europe, inducing more interest in the issue of alien arthropods. The horse-chestnut leaf miner, Cameraria ohridella Deschka and Dimić, an alien in Europe originating from the Balkans, has also raised much public concern because of its spectacular damage to urban trees in invaded areas of Central and Western Europe (Valade et al. 2009). Thus, checklists of alien arthropods began to be compiled from 2002 onwards, successively covering Austria (Essl and Rabitsch 2002), Germany (Geiter et al. 2002), Taxonomy, time and geographic patterns. Chapter 2 13 the Netherlands (Reemer 2003), the Czech Republic (Šefrová and Laštůvka 2005), Scandinavia (Nobanis 2005), the United Kingdom (Hill et al. 2005, Smith et al. 2007), Italy (Pellizzari et al. 2005), Serbia and Montenegro (Glavendekić et al. 2005), Switzerland (Kenis 2005), Israel (Roll et al. 2007), Albania, Bulgaria and Macedonia (Tomov et al. 2009), and Hungary (Ripka 2010). However, a major advance in the knowledge of alien arthropod species established in Europe was the European project DAISIE (Delivering Alien Invasive Species Inventories for Europe) in 2008. Besides furnishing national and regional lists, this project provided for the first time an overview of the alien fauna of arthropods that has established on the continent. DAISIE identified a total of 1517 alien terrestrial invertebrates, of which 1424 arthropods. However, limited expertise in some taxa during the DAISIE project meant full coverage of all the terrestrial arthropods could not be achieved with the same level of precision. The working group formed on this occasion therefore decided to continue its activity over the next two years, enlarging its taxonomic scope and competencies, in order to provide the most exhaustive list of the alien terrestrial arthropods of Europe as possible, with detailed information about each species. The update of the DAISIE list revealed in this book accounts for 1590 arthropod species alien to Europe, i.e. 166 more species, including both additions and deletions from the former list, and a much better coverage of taxonomic groups other than insects and spiders (i.e., mites, myriapods and crustaceans). In order to allow a comparison of their invasive patterns, the different taxonomic groups are presented separately in 21 chapters which have the same format. Because of the large number of species in some groups, these have been divided into several distinct chapters; i.e., four chapters for Hemiptera and five chapters for Coleoptera. Each chapter successively analyzes the taxonomy of the alien species component compared to that of the native fauna, the temporal trends of introduction, the biogeographic patterns, including both details of the region of origin and the distribution of the species in Europe, the pathways of introduction, the ecosystems and habitats which are invaded, and the economic and ecological impact of the biological invaders. At the end of each chapter, a table summarizes key information regarding all species in the taxa which are alien to Europe; i.e. of ascertained exotic origin or cryptogenic (see Chapter 1 for definitions): family, feeding regime, date and country of first record in Europe, invaded countries, habitats, plant or animal host, and one reference at least (usually that of the first record). In a number of cases, a second table includes a list and similar information for the species considered as alien in Europe; i.e. spreading to new countries within Europe, especially for species of Mediterranean origin recorded in more northern areas and species of continental Europe which have colonized islands. We did not provide such tables systematically. Indeed, it was difficult to ascertain for a lot of these species whether they have been introduced in other parts of Europe through direct or indirect human activity - and thus meet our definition of aliens (see Chapter I) - or they are naturally expanding, e.g. with global warming, or even if their native distribution range was incompletely known before their ‘’discovery’ in these new areas. The geographic range covered in this book is primarily Europe in geographic sense, with the main Mediterranean islands and archipelagos (Balearic Islands, Corsica, Sar- 14 Alain Roques / BioRisk 4(1): 11–26 (2010) dinia, Sicily, Malta, Crete, and the Ionian, North Aegean and South Aegean islands) and those of the North Sea (Aland, Svalbard) which are considered separately from the associated continental countries. Ireland was considered as a single biogeographic entity (i.e., Republic of Ireland plus Northern Ireland). Because of their possible importance as a first step for the invasion of continental Europe, the islands of the Altantic Ocean (Madeira, the Canary Islands, The Azores Archipelgao), were also included in the analysis but they may also correspond to a source of aliens of Macaronesian origin colonizing the European continent. This substantial work allowed us to figure out the relative importance of the different taxa of alien arthropods in a standardized fashion to other groups as well as to compare their respective habitats (Pyšek et al. 2009), and environmental and economic impacts (Vilá et al. 2009). The present chapter presents the most important patterns exhibited by the terrestrial arthropods alien to Europe. 2.2 Taxonomy of arthropods alien to Europe Alien terrestrial arthropods represent the second most numerous group of organisms introduced to Europe (Roques et al. 2009). A total of 1364 species originating from other continents have established so far, to which we add 226 more or less cosmopolitan species of uncertain origin (cryptogenic) for a total of 1590 species. Insects largely dominate this list, accounting for more than 87%, far in excess of mites (6.4%) (Figure 2.1). These alien species are dispersed across 33 taxonomic orders, including two orders of crustaceans, 10 of myriapods (three of chilopods, five of diplopods, one of pauropods and one of Symphyla), four of mites, one of arachnids, and 16 of insects. However, the relative importance of each order is highly variable (Figure 2.2). Three of the insect orders, namely Coleoptera, Hemiptera and Hymenoptera, overall account for nearly 65 % of total alien arthropods, representing 25.0%, 20.0% and 18.7%, respectively. The number of alien Hymenoptera established in Europe is thus much higher than previously considered (Daisie 2009). Diptera (6.2 %), Lepidoptera (6.1 %) Thysanoptera (3.3 %) and Psocoptera (3.1 %) have much lower importance as do Prostigmata mites (4.9 %- see Chapter 7.4) and Aranea (3.0 %), the only non-insect orders to exhibit more than 45 alien species. The other orders are anecdotal. It should be noted that some orders show no alien species whereas there are important components of the native fauna such as Trichoptera. More generally, at the order level, the taxonomic composition of the alien fauna significantly differs from that of the native European arthropod fauna. Calculations done on insects have revealed that establishment patterns differ between orders (Roques et al. 2009). Hemiptera are nearly three times better represented in the alien fauna than in the native fauna (20.0% vs. 8.0%). The alien entomofauna also includes proportionally more thrips (3.3 vs 0.6%), psocids (3.1 vs. 0.3%) and cockroaches (1.1 vs. 0.2%) than the native fauna, but much fewer dipterans (6.2 vs. 21%) and hymenopterans (18.7 vs. 25%). Differences are less pronounced for Coleoptera (25.0 vs. 30.0%) and Lepidoptera (6.1 vs. 10%). Taxonomy, time and geographic patterns. Chapter 2 15 Figure 2.1. Relative importance of the different phyla in the 1590 species of arthropods alien to Europe. Species of ascertained exotic origin and cryptogenic species are presented separately. The number to the right of each bar indicates the total number of alien species observed per phylum. The alien fauna seems to be highly diverse with a total of 257 families involved. However, only 38 of these families contribute 10 and more alien species, while 11 families more than 30 species (Figure 2.3). These 11 families mostly include hemipterans comprising aphids (Aphididae with the highest number of alien species - 102 spp.) and scales (Diaspididae and Pseudococcidae), as well as hymenopteran chalcids used for biological control such as Aphelinidae (63 spp.) and Encyrtidae (55 spp.), mites (Eriophyidae), and thrips (Thripidae). All of these except snout beetles (Curculionidae) and ants (Formicidae) are tiny arthropods. Noticeably, whilst these families dominate the alien fauna of arthropods, they are less intercepted by the phytosanitary quarantine services at European borders. A comparison done by Roques (2010) between interceptions and establishments of alien species in Europe during the period 1995 – 2005 for the alien insects and mites associated with woody plants in Europe has revealed that the major families of invaders were largely undetected (e.g. aphids, midges, scales, leafhoppers and psyllids). In contrast, the groups which were predominantly intercepted (e.g. long-horned and bark-beetles), actually made little contribution to the established alien entomofauna. Similar results were obtained at country level for Austria, the Czech Republic, and Switzerland (Kenis et al. 2007). For a number of families, the arrival of alien species has significantly modified the composition of the fauna presently observed in Europe. First, a total of 30 families had no representatives in Europe before the arrival of aliens. These include seven families of myriapods (Henicopiidae - 5 spp., Haplodesmidae, Rhinicricidae, Oryidae, Siphonotidae, Oniscodesmidae, Pseudospirobolellidae, Spirobolellidae, Trigoniulidae - 1 sp. each), four mite families (Listrophoridae - 4 spp., Myocoptidae, Pyroglyphidae and Varroidae - 1 sp. each), and one spider family (Sicariidae - 2 spp.). For insects, no native species existed for three alien families of psocids (Lepidopsocidae - 5 spp., Psyl- 16 Alain Roques / BioRisk 4(1): 11–26 (2010) Figure 2.2. Relative importance of the different taxonomic orders in the 1590 species of arthropods alien to Europe. Species of ascertained exotic origin and cryptogenic species are summed. The number to the right of each bar indicates the total number of alien species observed per order. lopsocidae - 5 spp., and Psoquillidae - 3 spp.), three lice families (Gliricolidae - 2 spp., Gyropidae and Trimenopidae - 1 sp. each), two Blattodea families (Blaberidae - 10 spp., and Blattidae - 6 spp.), two scale families (Phoenicococcidae and Dactylopiidae 1 sp. each), two beetles families (Ptylodactylidae or little ash beetles - 2 spp. and Acanthonemidae or toe-winged beetles - 1 sp.), one lepidopteran family (Castniidae - 1 sp., the palm moth Paysandisia archon (Burmister)), one Phasmatodea family (Phasmatidae - 4 spp.), one family of Hemiptera Auchenorrhnycha (Acanaloniidae - 1 spp.), and one thrips family (Merothripidae - 1 sp.). Taxonomy, time and geographic patterns. Chapter 2 17 Figure 2.3. Families of arthropods contributing most to the fauna alien to Europe. Only the families with numbers of alien species equal to 10 or more are shown. Corresponding taxonomic orders are indicated by different colors. The number to the right of each bar indicates the total number of alien species observed per family. In some other families, alien species could be over-represented. This is especially true for scales, where aliens now represent nearly half of the total Diaspididae fauna observed in Europe (60 out of 130 species - 44.6 %), a third of the Coccidae fauna (23 out of 70 species - 32.3 %), and a fourth of the Pseudococcidae fauna (37 out of 141 species 18 Alain Roques / BioRisk 4(1): 11–26 (2010) - 25.7 %). Similar high proportions of aliens are observed for psocids (Pachytroctidae - 66.7%, Ectopsocidae - 57%, and Liposcelidae - 26.4 %), hemipterans (Aleyrodidae - 39.1 % and Adelgidae - 36.0 %), hymenopterans (Agaonidae - 40.0 %, Apheliniidae 24.2 %, and Siricidae - 23.8%), and saturnid lepidopterans (30.0 %). Even if the relative proportions are lower, the arrival of a large number of alien species has also largely modified the faunal taxonomic structure in dermestid beetles (21.9 % of aliens), tetranychid mites (15.1 %), drosophilid flies (14.8 %), and encyrtid chalcids (7.2 %). 2.3 Temporal trends of arrival in Europe of alien arthropods Some alien arthropods were introduced to Europe long ago accompanying human movements. For instance, a number of ectoparasites of humans and early-domesticated animals such the head louse (Pediculus capitis De Geer), the crab louse (Phtirus pubis (L.)), the cat flea (Ctenocephalides felis felis (Bouché)), the rat flea (Xenopsylla cheopis (Rothschild)) or the human flea (Pulex irritans L). are probably allochtonous in Europe, having arrived in ancient times with their hosts (Mey 1988; Beaucournu and Launay, 1990). Thus, Pulex irritans was shown to have been present in Europe since the Bronze Age at least, having been found in remains of lake dwellings in the French Jura, dating back to 3100 B.C. (Yvinec et al. 2000). Fragments of insects related to stored products were also found in Roman and Viking graves (e.g., Sitophilus granarius; Levinson and Levinson 1994). However, unlike plants and other animal groups, a clear identification of the archaeozoans* has appeared difficult for arthropods. Therefore, we only qualified as aliens the neozoan* species, i.e. those having likely been introduced after 1500. The introduction of alien arthropods is usually accidental, the release of biological control agents remaining limited, as well as the escape of arthropod ‘pets’ from captivity (see Chapter 3). Thus, the introduction phase is rarely observed and pathways of introduction are poorly known. Consequently, an alien arthropod is usually discovered when it is already established, spreading and causing damage. The precise date of arrival in Europe is not known for most species. Even conspicuous species, such as the Asian long-horned beetle, Anoplophora glabripennis (Motschulsky), have been reported with a delay of at least 3–5 years since establishment (Herard et al. 2006). However, taking into account these caveats, the date of first record in Europe- the single temporal datapoint usually obtainable- may be used as a proxy for the date of first arrival. The date of first record in Europe, relying on published papers, could be obtained for 1421 of the 1590 alien species (89.4%). The number of new records per year appears to have increased exponentially since the 16th century, but a significant acceleration was observed during the second half of the 20th century (Figure 2.4a). As a probable result of globalization, this trend is still increasing with an average of 19.6 alien species newly reported per year in Europe between 2000 and 2008; i.e., a value nearly double the 10.9 species that were observed per year during the period 1950- 1974. In order to understand better this process, we decompose the values according to the feeding regime of the alien species (Figure 2.4b). Fluctuations in the number of Taxonomy, time and geographic patterns. Chapter 2 19 Figure 2.4. Temporal changes in the mean number of new records per year of arthropod species alien to Europe from 1500 to 2008. A Total arthropods (Best fit: y= -0.411- 0.407x + 0.304 x2; r2 = 0.965) B Detail per feeding regime. total arthropods newly arriving per year in Europe appear to be strongly dependent on the increasing arrival of phytophagous species, especially during the last ten years. In contrast, the number of detritivores and parasitoids/ predators has appeared to decrease during this last decade, and contributed much less to the overall increase, whereas these three feeding guilds had contributed more or less equally during the first half of the 20th century. After the period 1950- 2000 when alien parasitoids and predators markedly increased probably in relation with the wave of releases of biological control agents, the explosion of ornamental trade since the 1990s is likely to have triggered the arrival of alien phytophagous arthropods, as has been shown for insects related to woody plants (Roques 2010). Specific analyses per taxa have confirmed these tendencies. Whereas the arrival of mites (see Figure 7.4.2), scales (see Figure 9.3.2.), flies (see Figure 10.2) or lepidopterans (see Figure 11.2), which are mainly phytophagous groups, has revealed a similar acceleration in the number of newly recorded aliens during the last period, no such trend has been observed for the parasitic lice and fleas (see Chapter 13.4), nor for the detritivorous Blattodea (see Chapter 13.3). 2.4 Biogeographic patterns of arthropod species alien to Europe Origin of the species alien to Europe A precise region of origin was ascertained for 1271 species (79.9%) of the 1590 alien arthropod species, while 93 species were only known to be native to tropical or subtropical regions. The remaining 226 cryptogenic invertebrates are mostly cosmo- 20 Alain Roques / BioRisk 4(1): 11–26 (2010) politan species for which there is no agreement regarding their area of origin. This is particularly true for stored products pests and for some ectoparasites on cattle and pets that occur on other continents. A few other cryptogenic species have appeared in Europe without having been detected elsewhere. However, data on their phylogeography, population ecology, parasitoids and dispersal biology strongly suggest that they originate from another continent. The horse-chestnut leaf miner, Cameraria ohridella, is illustrative of the difficulty in identifying the native range of such species. Whereas this leaf miner was previously considered as an extra- European alien, recent genetic studies indicate that it originates from the southern Balkans (Valade et al. 2009). Asia has supplied the major part of the alien arthropods occurring in Europe (26.7 %) followed by North America (21.9%) (Figure 2.5). Analysing specifically insect data per time unit has revealed that the relative contribution of Asia and North America was stable over time (Roques et al. 2009). During the periods 1950–1989 and 1990–2007, 29% and 21% of the established insects were of Asian and North American origin respectively. The contribution of tropical and subtropical areas is surprisingly important. The overall contribution of species from Australasia, Africa, Central and South America in combination with species of undefined tropical areas represents 37% of all alien insects in Europe. While we agree that insect species coming from these areas are not just native to tropical ecosystems, this proportion is nevertheless outstanding. Unlike the temporal trends, the regions of origin do not differ significantly between feeding regimes. Asia is the main region of origin for alien phytophages, parasitoids/ predators and detrivorous species although a bit less important for the latter group (Figure 2.5). Figure 2.5. Region of origin of the 1590 arthropod species alien to Europe. Total arthropods and breakdown per feeding regime are presented. Percentages of the total per category are shown under each region. Taxonomy, time and geographic patterns. Chapter 2 21 However, a comparison of the native range of species from the different orders revealed significant differences (χ2= 388.26; P=0.0000). Most mites (51.5% - see Figure 7.4.3), hymenopterans (32.3 % - see Figure 12.3), and dipterans (30.6 %- see Figure 10.3) have arrived from North America whilst 37.2 % of lepidopterans (see Figure 11.3) and 31.5 % of hemipterans have originated from Asia. Coleoptera have come from various regions, including a significant component from Australasia (9.5%) mostly linked to the introduction of Eucalyptus and Acacia spp. in the Mediterranean regions of Europe. Coleoptera also represent a large proportion of the cosmopolitan stored product pests that are predominantly of tropical or subtropical origin. Patterns of spread Once established, most alien species have not spread throughout Europe, at least yet. We used the presence in a country as a proxy of the invaded range because it appeared impossible to get sufficient data for a quantitative assessment of this invaded range area for most alien species. A total of 694 species (i.e., 43.6 %) have not invaded more than one country/ island additional to the one where they arrived, and 63.6 % are present only in five European countries (Figure 2.6). Less than 1% (12 out of 1590) of the alien arthropods are present in more than 40 countries; among these are the melon and cotton aphid, Aphis gossypii Glover, and several beetles associated with stored products especially seed bruchids (e.g.,. Callosobruchus chinensis (L)). Detritivorous species appeared to have dispersed significantly more (8.5±0.5 countries) than phytophagous species (7.1±0.3) and parasitoids/ predators (5.5±0.3) (Krsukall-Wallis test. F2,1598= 10.97; P=0.0000). Figure 2.6. Geographic spread of the arthropod species alien to Europe expressed as the number of countries colonized by these species and their frequency. 22 Alain Roques / BioRisk 4(1): 11–26 (2010) Figure 2.7. Comparative colonization of continental European countries and islands by dipteran species alien to Europe. Archipelagos: 1 Azores 2 Madeira 3 Canary islands. Large differences also exist between European countries in the total number of alien arthropods recorded per country (Figure 2.7 and 2.8). Italy (700 species) and France (690 species), followed by Great Britain (533 species), host many more species than other countries. The same ranking is obtained when the number of alien species per km2 is considered. Differences in sampling effort may have affected the analyses. However, the number of alien insects is significantly and positively correlated with country surface area (r= 0.3621; P= 0.0384). More westerly countries and islands appear in general relatively more colonized. The number of alien species significantly decreases with the longitude of the countries’ centroids (r= -0.6988; P= 0.0038) whereas latitude does not seem to have a significant influence (r=-0.378; P= 0.168). Islands also host proportionally more alien species than continental countries relative to their size (Kruskall-Wallis test on the number of alien species per km2; F1,53 = 6.20; P=0.0160) but this is independent of the coast length (r= 0.174; P= 0.384). In continental countries, bordering the sea does not influence the number of alien insect spe- Taxonomy, time and geographic patterns. Chapter 2 23 Figure 2.8. Comparison between the number of first records for Europe observed for the alien species in a country (left) and the total number of alien species now present in the country (right). cies (P=0.6404). In addition, the country or island where a species was first recorded in Europe has been identified for 1399 species out of the 1590 alien arthropods (Figure 2.8). 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London: Chapman and Hall. 244 pp. Wittenberg R, Cock MJW (Eds) (2001) Invasive alien species: a toolkit of best prevention and management practices. Wallingford, UK: CABI Publishing. 228 pp. A peer reviewed open access journal BioRisk 4(1): 27–43 (2010) doi: 10.3897/biorisk.4.60 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Pathways and vectors of alien arthropods in Europe Chapter 3 Wolfgang Rabitsch Environment Agency Austria, Dept. Biodiversity & Nature Conservation, Spittelauer Lände 5, 1090 Vienna, Austria. Corresponding author: Wolfgang Rabitsch (wolfgang.rabitsch@umweltbundesamt.at) Academic editor: David Roy | Received 26 April 2010 | Accepted 18 May 2010 | Published 6 July 2010 Citation: Rabitsch W (2010) Pathways and vectors of alien arthropods in Europe. Chapter 3. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 27–43. doi: 10.3897/biorisk.4.60 Abstract This chapter reviews the pathways and vectors of the terrestrial alien arthropod species in Europe according to the DAISIE-database. The majority of species (1341 spp., 86%) were introduced unintentionally, whereas 218 species (14%) were introduced intentionally, almost all of these for biological control purposes. The horticultural/ornamental-pathway is by far the most important (468 spp., 29%), followed by unintentional escapees (e.g., from greenhouses, 204 spp., 13%), stored product pests (201 spp., 12%), stowaways (95 spp., 6%), forest and crop pests (90 spp. and 70 spp., 6% and 4%). For 431 species (27%), the pathway is unknown. The unaided pathway, describing leading-edge dispersal of an alien species to a new region from a donor region where it is also alien, is expected to be common for arthropods in continental Europe, although not precisely documented in the data. Selected examples are given for each pathway. The spatiotemporal signal in the relevance of pathways and vectors and implications for alien species management and policy options are also discussed. Identifying and tackling pathways is considered an important component of any strategy to reduce propagule pressure of the often small and unintentionally translocated, mega-diverse arthropods. This requires coordination and clear responsibilities for all sectors involved in policy development and for all associated stake-holders. Keywords alien species, non-native species, pathways, vectors, Europe Copyright W. Rabitsch. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 28 Wolfgang Rabitsch / BioRisk 4(1): 27–43 (2010) 3.1 Introduction To become an alien species, boundaries of natural distribution ranges must be overcome with the help of man-made structures, goods and services. These activities and purposes are the pathways of invasions. A plethora of vectors, which are the agents of these translocations, is available to break new grounds and reach new areas. Interestingly, there is no common understanding in this separation in the biological invasion literature (e.g. Ruiz and Carlton 2003, Carlton and Ruiz 2005, Nentwig 2007, Hulme et al. 2008). In this overview, however, pathways are understood as the routes (including motivations to use them) and vectors as the physical objects (ships, plants etc) that carry species along. Several attempts to further classify pathways and vectors are available (e.g. Carlton and Ruiz 2005), but here I follow Hulme et al. (2008), who identified six principal pathways for biological invasions (Table 3.1). Only one of these is founded by intentional motivations, that is the deliberate release of organisms, with biological control as the most important example. The others are utilised unintentionally, accidentally and may come from any direction. These are escapes from contained environments and captivity; contaminants of commodities; stowaways, transported as hitch-hikers with vehicles and cargo; corridors, where transport infrastructure enables the spread of a species; and the unaided pathway, where an alien species conquers a nearby region under its own dispersal capacity. Evidently, these different pathways have major implications for risk assessment, regulations, management and control (Hulme et al. 2008, Hulme 2009). Human-mediated translocations differ from natural dispersal by orders of magnitude both quantitatively and qualitatively as can be seen by island colonization rates (e.g. Gillespie and Roderick 2002, Gaston et al. 2003) and genetic consequences (e.g. Wilson et al. 2009). Also, the origin of the source differs as natural colonization usually happens from adjacent populations, whereas translocated individuals may come from all over the world. In the DAISIE-database, three levels of pathways, are distinguished. At the first level, intentional and unintentional ambitions are classified. At the second level, pathways are identified, except that the contaminant, stowaway and corridor pathways are summarized as “transport”. At the third level, these are further specified into broad categories (e.g. biological control, crops, horticultural/ornamental, forestry, stored products). In addition, at the second and third level, the category “unknown” is also used and assigned to 392 and 431 species, respectively (25–27%). This is a similar contingent as for the exotic insects in Japan (24%, Kiritani and Yamamura 2003). Introductions of species are not necessarily restricted to one pathway; many species can be considered “polyvectic” (Carlton and Ruiz 2005), transported by more than one pathway or multiple vectors. Accordingly, some species in the DAISIE-database were assigned to more than one pathway/ vector. Furthermore, it has to be said very clearly that many assignments were only “best guess” or “most likely” assessments, deduced from the preferred habitats, food Chapter 3: Pathways and vectors of alien arthropods in Europe 29 Table 3.1. Pathway terminology and examples of vectors of terrestrial alien arthropod species in Europe. Pathway Motivation Vectors Release Intentional None Escape Unintentional None Contaminant Unintentional Food sources, ornamentals, vegetables, fruits, wood, animals, ... Stowaway Unintentional Any cargo Corridor Unintentional Ships, cars Unaided Unintentional None Examples Biological control Greenhouses Stored product pests, Wood-borers, Leaf-miners, Gall-producers, Endoparasites Ants, Cockroaches Cameraria ohridella Secondary spread from point of entry plants or ecology, because the intimate pathway/vector of many arthropod species often remains ambiguous. In this chapter, pathways and vectors of the terrestrial alien arthropods in Europe are reviewed, with the few alien aquatic insects included, but excluding other freshwater alien arthropods such as crayfish species. There are a multitude of further pathways relevant for the marine and freshwater environments (e.g. ballast water, hull-fouling) and for other organisms such as vascular plants and vertebrates (e.g. seed contamination, hunting, pets) (e.g. García-Berthou et al. 2005, Galil et al. 2009, Genovesi et al. 2009). 3.2. Intentional release With few exceptions, terrestrial arthropods are not intentionally imported. Such exceptions are grasshoppers and crickets as pet food and – more significantly – domesticated honeybees (Apis mellifera) of different provenances (subspecies), which are used for breeding, with the aim of producing higher honey yields (Jensen et al. 2005, Moritz et al. 2005). The same is true for the bumblebee subspecies used for pollination in greenhouses (e.g., Bombus terrestris dalmatinus in the UK, Ings et al. 2006). At the end of the 19th century, two saturniid moths, Samia cynthia and Antheraea yamamai, were introduced from Asia for silk production, but yields was not profitable enough for this to be continued. Both species persist locally in the wild in Europe with most populations being initiated by escapes or releases by amateur lepidopterabreeders. Intentional releases for human food consumption are more prevalent for organisms such as molluscs, fish and aquatic Crustacea (oysters, snails, crayfish, crabs), which are not included in this book. Also, there are no “game insects”, and only a few pets. Further, there are no introductions of arthropods for aesthetic or conservation purposes (but see further below), a major pathway for other animal groups around the globe (e.g. Nentwig 2007). In the DAISIE-database, 218 species (14%) were introduced intentionally, almost all of these for biological control purposes (Table 3.2). 30 Wolfgang Rabitsch / BioRisk 4(1): 27–43 (2010) Table. 3.2. Pathways of the alien arthropod species in Europe, according to the DAISIE-database. Due to double entries the sum differs. Pathway Intentional Released Unintentional Animal husbandry Greenhouse escapees Crops Forestry Horticultural/Ornamental Leisure Stored products Stowaways Unknown Number of species (%) 218 (14%) 175 (11%) 1341 (86%) 42 (2.6%) 204 (13%) 70 (4.3%) 90 (5.6%) 468 (29%) 13 (0.8%) 201 (12%) 95 (5.9%) 431 (27%) 3.2.1. Biological control: Ecology vs Economy The most important pathway for deliberate release of terrestrial alien arthropods is biological control (BC). There has been some controversy about the pros and cons of this technique to control pest organisms (e.g. Howarth 1991, van Lenteren et al. 2006, Babendreier 2007, Murphy and Evans 2009). Whereas non-target effects are considered problematic by conservationists, these are often considered acceptable from an economic point of view. Hence, the underlying basic assumptions and intentions for this controversy are entirely different and comparisons awkward. BC makes use of the “enemy-release” of introduced organisms, which are disburdened from their natural predators or parasites and boom in the new range. Subsequently, mass-reared releases of those enemies from the original area are conducted, aiming at permanent establishment and control of the pest organisms below damaging thresholds. Not particularly from a “pathway point-of-view”, but from a general assessment of non-target effects, it is useful to distinguish between this classical BC and augmentative BC, where control is achieved by periodic releases without permanent establishment intended. Similarly, flightless strains of H. axyridis were released in the Czech Republic in 2003 to control for aphids with the goal of no further unaided spread (Brown et al. 2008). In Europe, there are both success-stories and failures to report from intentional releases, with the former prevailing (e.g. Encarsia formosa used against whiteflies in greenhouses; Trichogramma brassicae, an “alien in Europe” used against European corn borer Ostrinia nubilalis; Aphelinus mali from North America used against the Woolly apple aphid Eriosoma lanigerum). Occasionally, released enemies are aliens from other regions than their targets. In Europe, for example, the San Jose scale Diaspidiotus perniciosus, described from Califor- Chapter 3: Pathways and vectors of alien arthropods in Europe 31 nia, but introduced with infested trees or fruits from Asia, is considered a pest in commercial fruit orchards causing economic losses due to reduced yields. Negative effects are mitigated by application of Neem and other oils, but also by release of the North American parasitoid wasp Encarsia perniciosi, which is used for control in North America. In general, however, the application of BC has been of subordinate relevance in Europe, compared to other regions of the world. The same is true for the application of other technologies where arthropods are released (SIT – Sterile Insect Technique; RIDL – Release of Insects carrying a Dominant Lethal), which may be applied to control alien agricultural pests and mosquitos (Thomas et al. 2000, Alphey et al. 2009). Ex-situ conservation or reintroduction programmes in insects are still rare, but they do occur for some native species in Europe (butterflies in the UK: Oates and Warren 1990; Erebia epiphron in the Czech Republic: Schmitt et al. 2005; Gryllus campestris in the UK and Germany: Witzenberger and Hochkirch 2008). Recently, controversial discussions on assisted colonization have emerged in the context of protecting species from climate change by translocating and releasing them beyond their current range limits (e.g. Hoegh-Guldberg et al. 2008, Ricciardi and Simberloff 2009). 3.3. Unintentional release The unintentional translocation of species is the most common pathway for alien arthropod species invasions into Europe (86% of the species, Table 3.2). 3.3.1. Escapes: Out of the Green Arthropods are infrequently domesticated, reared and used as pets, although examples of tropical species do exist (e.g. tarantulas, walking sticks and leaves, leaf-cutting ants, millipedes). Establishment in the wild in Europe is highly unlikely for such species, even under severe climate change scenarios. However, escapes from captivity do regularly occur, although they are rarely noticed and published. Insects reared as living food for vertebrate pets (e.g. crickets, grasshoppers, mealworms) seem to be of limited significance, whereas pests and insects used for biological control in semi-contained environments, particularly greenhouses, are of much greater importance. Greenhouses are not escape-proof facilities for insects as confirmed by surveys in the areas surrounding such buildings (e.g. Vierbergen 2001, Aukema and Loomans 2005). Well-known examples include the Western Flower Thrips Frankliniella occidentalis, the Cotton Aphid Aphis gossypii, and the Cotton Whitefly Bemisia tabaci, all of which reproduce in the field in southern Europe but are restricted to greenhouses in western, central, or northern Europe. Serving as stepping stones, it is expected that some future invaders in Europe will be recruited out of this pool of species, particularly if climate warms as predicted. In the DAISIE-database, more than 200 arthropod species are listed as living in greenhouses. 32 Wolfgang Rabitsch / BioRisk 4(1): 27–43 (2010) One of the most famous stories of a greenhouse escapee is the Multicoloured Asian lady beetle or Harlequin ladybird Harmonia axyridis, termed the “most-invasive ladybird on Earth” (Roy et al. 2006). This large coccinellid beetle, native to East-Asia, was introduced to North America and Europe for aphid control in greenhouses, but escaped into the wild. It is a highly competitive intra-guild predator reducing and displacing native coccinellid species and other members of the aphid-feeding guild (Roy and Wajnberg 2008). Its subsequent unaided spread across much over Europe within just a few years (Brown et al. 2008) highlights the capacity of invasive alien species to successfully conquer naïve environments. 3.3.2. Contaminant: Going for a ride? The contaminant pathway describes the unintentional transport of species within or on a specific commodity, contrary to stowaways, which are accidentally associated with any commodity. Stored product pests, for example, are translocated with the movements of the products and many species have subsequently achieved a cosmopolitan distribution. In Europe, 201 alien insect species (12%) were introduced as stored product pests, feeding on a variety of food sources (e.g. cereals, rice, seeds, nuts, fruits) with considerable economic damage, including species which are likely to have been introduced by human activities in neolithic or pre-Christian centuries, e.g. Sitophilus granarius and Oryzaephilus surinamensis (Levinson and Levinson 1994). In Europe and temperate regions in general, care of stored products achieves higher protection levels than in subtropical and tropical areas, where up to 10% of weight loss may occur, representing loss of nutritional quality, with associated impacts on human welfare (Rees 2004). Other pest species are strictly associated with the exchange or trade of their host plants (e.g. ampelophagous species feeding exclusively on grapevines - Viteus vitifoliae, Scaphoideus titanus; species feeding exclusively on palms - Rhynchophorus ferrugineus, Diocalandra frumentii; monophagous leaf-miners and gall-producers - Parectopa robiniella, Phyllonorycter robiniella, Dryocosmus kuriphilus) and therefore directly related to these vectors. Other examples include phytophagous species translocated with ornamentals or horticultural host plants (e.g. scales and aphids) and xylophagous bark- and woodinfesting insects, above all beetle larvae, feeding in living trees. One of the best known examples is the Citrus longhorned beetle Anoplophora chinensis, which has repeatedly been reported infesting Bonsais imported from China. Larvae of A. chinensis and more often of the Asian longhorned beetle Anoplophora glabripennis were also intercepted with wood packaging material (see Haack et al. 2010 for a review). Recognizing the relevance of this vector enforced adoption of the International Standard for Phytosanitary Measures No. 15, which sets standards for thermal and chemical treatment of wood packaging material used for international trade. Although now found in lower numbers, living beetles are still being intercepted, indicating some gaps in this procedure. Chapter 3: Pathways and vectors of alien arthropods in Europe 33 Roques (2010) assembled examples of the possible introduction of alien insects during major international events such as the 2004 Olympic Games in Athens, where imported palm trees were widely planted and coincided with the first arrival of the red palm weevil Rhynchophorus ferrugineus. The most striking example of contamination is associated with the introduction of the Potato (Colorado) beetle, Leptinotarsa decemlineata, to Europe. Spanish conquistadors in the 16th century brought the potato plant from South America to Europe, although it was not appraised as a human food source until the mid-17th century. After a severe decline of potato cultivation in Ireland in 1845–1857, caused by the introduced potato blight fungi Phytophthora infestans, emigrants brought the plant to North America, where the beetle exploited the new host plant. Between 1876 and 1922, the beetle was subsequently introduced into Europe on several occasions, not being able to establish in European potato fields until 1922, when it succeeded in France. The beetle has since spread east throughout Europe and Asia, reaching China in the 1980s (Alyokhin 2009). It should also be noted that the Colorado beetle was involved in propaganda to defame Great Britain and the United States of America during World War II and the Cold War. Kenis et al. (2007) found that the majority of alien insects for Austria and Switzerland were contaminants and stowaways, with, in decreasing order, host plants (40% of which on ornamentals and 20% on vegetables and fruits), stored products and wood material as the main sources. Similar results were obtained with interceptions documented by EPPO between 1995 and 2004 (Roques and Auger-Rozenberg 2006). Altogether, introductions of arthropods with ornamental and horticultural plants and plant material, cut flowers, vegetables, and fruits, clearly preponderate in the DAISIE-data (29%, Table 3. 2). It is self-evident that there is a taxonomic bias with the type of commodity. For example, plant-feeding species (e.g. aphids, scales) are closely associated with ornamental plants, whereas wood-boring species (e.g. scolytids, cerambycids) are linked to living and dead wood imports. A rather uncommon invasion history pertains to the inadvertent introduction of the nearctic waterboatman Trichocorixa verticalis into Portugal and Spain. It is likely to have happened with the import and release of Eastern Mosquitofish Gambusia holbrooki for mosquito control (Sala and Boix 2005). Living organisms as well as commodities can be contaminated. For example, many haematophagous alien arthropod species (e.g. Culicidae, Siphonaptera, Phthiraptera, Ixodidae) host parasites and pathogens and serve as reservoir, carriers or biovectors of human and animal infectious diseases. Moreover, phytophagous alien arthropod species (e.g. Hemiptera) may transmit plant pathogens (e.g. phytoplasmas, viruses). Several examples are associated with beekeeping. Both endoparasites (the tracheal mite Acarapis woodi) and ectoparasites (the notorious Varroa-mite Varroa destructor), inquiline scavengers (the Small Hive Beetle Aethina tumida, captured only once in Europe and eradicated in quarantine in Portugal), and bacterial and fungal diseases (chalkbrood, foulbrood, nosemosis) are exchanged throughout the globe through honeybee imports (e.g. Sammataro et al. 2000, Coffey 2007). 34 Wolfgang Rabitsch / BioRisk 4(1): 27–43 (2010) The ultimate agent of Colony Collapse Disorder (CCD) known from North America, Europe and Asia is still under debate (e.g. Ratnieks and Carreck 2010) and it may well be a multi-triggered phenomenon, which causes the complete disappearance of adult worker bees of a colony. Beside environmental causes (e.g. pesticides), several diseases and pathogens are suspected to contribute or elicit CCD, e.g. Nosema ceranae, a microsporidian native to Asia and suspected to have host-switched to the European honeybee (Klee et al. 2007, Higes et al. 2009). 3.3.3. Stowaways: Where do you want to go today? Stowaways are unintentionally introduced organisms that are related to transport infrastructure and vehicles, but independent of the type of commodity. Translocation with ballast water or soil movement are typical examples. In terrestrial environments, any cargo transported by air, water or land has the potential to move species beyond their natural range and habitat boundaries. Several cockroach species, e.g. Blatta orientalis and Periplaneta americana, are typical stowaways, having been translocated worldwide. Kiritani and Yamamura (2003) argued that passenger hand luggage arriving in airplanes to Japan may contain one consignment infested by fruit flies each day. Roughly two thirds of the intercepted pest species at US ports of entry between 1984 and 2000 were associated with baggage, and a further 30% with cargo (McCullough et al. 2006). However, to a certain extent, the separation between the contaminant and the stowaway pathway is ambiguous or not mutually exclusive. Roques et al. (2009) cites the Asian tiger mosquito Aedes albopictus as an example of the stowaway pathway, this species being translocated as eggs and larvae within any small amount of standing water. Water within used tyres or ornamental plants (lucky bamboo Dracaena spp.) is a cause of the trans-continental introduction of A. albopictus to Europe, North America, Africa and Australia (e.g. Reiter 1998). Short-distance dispersal seems to be limited to passive transport by cars and trucks, or movement of infested tyres and plants (Scholte and Schaffner 2007). Establishment in other parts of Europe is very likely within the next decades, supported by climate change (Schaffner et al. 2009). Aedes albopictus is a vector of several viruses (e.g. Dengue, Chikungunya, West Nile) and of increasing relevance for Europe (Scholte and Schaffner 2007, van der Weijden et al. 2007). The movement of used tyres is also likely to be responsible for the most recently introduced mosquito species, Ochlerotatus atropalpus, native to North America and detected in several European countries (France, Italy, Netherlands), where it was subsequently eradicated (Scholte et al. 2009). Many insects are attracted to light and most transport hubs (airports, seaports) are illuminated during night-times, increasing the probability of translocation with vehicles after boarding a vector. For example, it is speculated that the attraction to light facilitates the repeated introduction of adult Diabrotica virgifera with aircrafts from Chapter 3: Pathways and vectors of alien arthropods in Europe 35 North America to Europe, because of regular “first” records of the species in the vicinity of airports. From there the species spreads unaided depending on habitat (maize fields) availability. Ants (Formicidae) are among the most invasive organisms globally, particularly hazardous on oceanic islands (e.g. Holway et al. 2002, Lach and Hooper-Bùi 2010). Entire colonies with gynes and workers may be translocated as stowaways with soil and litter accompanying ornamental plants, with logs or with other commodities offering shelter. The majority of introduced ants in the USA have been detected on plant material (Suarez et al. 2005). Some of the characteristic traits of tramp ants, e.g. preference for disturbed habitats, polygyny, budding, small body size, support successful translocation and subsequent establishment around the globe (e.g. McGlynn 1999). In Europe, the Argentine ant Linepithema humile and the garden ant Lasius neglectus are currently considered to be of prime importance (see Kenis and Branco, chapter 5). Whereas the former was introduced as a stowaway with unknown commodities to Europe (Madeira and mainland Portugal) in the 19th century (Wetterer et al. 2009), the origin (likely Asia Minor), pathway and vector (eventually contaminant of garden soil) and successful secondary spread of the latter are still under debate (Ugelvig et al. 2008). Two more examples of Hymenoptera, initially introduced as stowaways, are the oriental mud dauber Sceliphron curvatum and the Asian hornet Vespa velutina. The former was introduced in the late 1970s via air cargo from Central Asia to Austria and produces conspicuous mud nests in which paralysed spiders are provisioned as food supply for the developing larvae (Schmid-Egger 2004). The latter was only recently detected in France, probably introduced with pieces of pottery from China (Villemant et al. 2006). These two species have subsequently spread rapidly, unaided, and may be of increasing relevance to native sphecids, hornets and honeybees. 3.3.4. Corridors: Like a rolling stone The corridor pathway highlights the role transport infrastructures play in the introduction of alien species; shipping canals are the most important example. Gilbert et al. (2004) have shown that the spread of Cameraria ohridella in Germany was related to the highway routes, Pekar (2002) argues that the spread of the spider Zodarion rubidum was facilitated by the railway system and there is anectodal evidence for repeated northwards transport of the flightless Southern Oak Bush Cricket (Meconema meridionale) and the Speckled Bush-Cricket (Leptophyes punctatissima) with cars along highways from Southern Europe. Although infrastructure networks undoubtedly contribute to the distribution of alien terrestrial arthropod species in Europe, it seems to be of subordinate relevance and is often intermingled with the contaminant/ stowaway pathway. 36 Wolfgang Rabitsch / BioRisk 4(1): 27–43 (2010) 3.3.5. Unaided: One day I’ll fly away The unaided pathway describes leading-edge dispersal, that means situations where spread results in alien species arriving in a new region from a donor region where it is also alien. This holds true for many alien arthropods occurring in the wild in Europe, being introduced once and spreading after successful establishment. Several examples were mentioned in the chapters above, although this is not reflected in the DAISIE-database (Table 3. 2). Unaided spread often follows initial introduction by one of the other pathways into Europe, although long-distance dispersal events may contribute to the distribution patterns and accelerate rates of spread, as shown for the horse chestnut leafminer Cameraria ohridella in Germany and France (Gilbert et al. 2004, 2005). The chestnut gall wasp Dryocosmus kuriphilus was introduced with infested plant material from China to Italy and is now spreading unaided to neighbouring countries, but may also bridge larger distances with transport of infested plant material. Dispersal capacities of arthropods can be impressively high. The conifer seed bug Leptoglossus occidentalis and the Harlequin ladybird Harmonia axyridis spread over much of Europe within just a decade (e.g. Lis et al. 2008, Rabitsch 2008, Brown et al. 2008) presumably on their own wings. In addition, repeated and independent introductions from the area of origin and/or secondary introductions from the alien range over long distances undoubtedly occur, but such events are difficult to prove and require specific techniques (e.g. molecular biology) (e.g. Diabrotica virgifera – Miller et al. 2005, Ciosi et al. 2008). Controversy surrounds the definition of the alien status of species extending their range due to recent anthropogenic climate change. As long as they utilize the beforementioned pathways, e.g. are translocated with vehicles, but then find suitable climate conditions to establish populations, they should be considered alien. If a species extends its range unaided, but only colonizes disturbed or secondary habitats under strong human influence, such species may be considered as alien. Particularly in arthropods, however, it is sometimes difficult or even impossible, to unambiguously identify the boundaries of the natural range of a species. Historic introductions of today’s cosmopolitan species, taxonomic impediment and the lack of recording schemes for most groups cause a high degree of uncertainty in the delimitation of the native range of some species. Host plant distribution, habitats, and molecular techniques may serve as a clue for disentangling factors (e.g. Kavar et al. 2006, Valade et al. 2009). Unaided dispersal is also often assumed for modelling rates of spread of alien species. Liebhold and Tobin (2008) provided examples for the radial rate of spread in alien insects, which span from 1 to 500 km year-1. In Europe, the western flower thrips Frankliniella occidentalis stays ahead with up to 249 km year-1 (Kirk and Terry 2003). However, in many if not most cases, additional pathways including long-distance dispersal or at least a combined stratified dispersal need to be taken into account for more realistic scenarios of spread (e.g. Gilbert et al. 2004 for the horse chestnut leafminer Cameraria ohridella). Chapter 3: Pathways and vectors of alien arthropods in Europe 37 3.4. Future trends and management There is no reason to assume a decrease in people’s movements and restrictions in the transport of goods in the near future. Biological homogenization will tie continents and biodiversity, increasing species richness locally and decreasing it globally; the rate of change will be much more rapid than the hypothesised formation of Neopangaea (Scotese 2001). The ultimate consequences of such a process for the functioning of ecosystems and their services to mankind are far from being well understood. There is a spatiotemporal signal in the relevance of pathways and vectors. Whereas soil was used as ship ballast in earlier days of European colonization (e.g. Vazquez and Simberloff 2001) this was replaced by ballast water in later years. With the construction of bigger and faster ships, even more organisms were translocated rapidly and with the advent of aircrafts this rate was yet further accelerated. Fast transit enables more species to survive transport and subsequently establish successfully in new regions. In addition, continental, land-locked areas became easily accessible (Mack 2003). Asia has recently gained increasing relevance as a country of export globally (Roques 2010) and as a donor region of alien species, particularly for insects associated with woody plants introduced to Europe (Roques et al. 2009). New trends in the ornamental trade by changed consumer behaviour has created new markets. Only two decades ago, bonsais were rare in European households, but have become a recent fashion; sales are increasing in most areas. Generally, the horticultural/ornamental pathway is of paramount significance for the alien arthropods of Europe (Kenis et al. 2007, Table 3. 2) and there is ample scope for enhancing existing plant protection services (e.g. by increasing personnel at points of entry) and providing best-practice guidance to the ornamental trade industry. It has been shown, however, that interception and establishment data of alien insects for Europe differ significantly (Kenis et al. 2007, Roques 2010). This discrepancy may eventually be explained by the changed relevance of pathways and time-lag phenomena (Crooks 2005). In any case, it demonstrates that additional endeavours are necessary to abate undesirable effects on ecology and economy. Import and export of goods follows economic rules and global trade mirrors biological invasion patterns (e.g. Levine and D’Antonio 2003, Taylor and Irwin 2004, Kobelt and Nentwig 2008, Westphal et al. 2008, Roques et al. 2009). Chiron et al. (2009) showed such a pattern for bird introductions on both sides of the “iron curtain” in Europe and it is expected that a similar pattern will be found for arthropods. However, information on introduction dates, number of propagules, etc. are usually lacking for arthropod invasions, so that such analyses are difficult to achieve. Anthropogenic climate change acts upon several levels of biological invasions (e.g. Walther et al. 2009, Thomas and Ohlemüller 2010). It may directly change the realized climatic niche of species, cause habitat shifts (e.g. stepping-stone scenarios) and range shifts in latitude and altitude. Ødegaard and Tømmerås (2000) showed that eight out of 25 alien ground-beetle species used compost heaps as stepping-stones for subsequent establishment in the wild in northern Europe. Global climate change, 38 Wolfgang Rabitsch / BioRisk 4(1): 27–43 (2010) however, may further act indirectly in changing trade and consumer habits, influencing invasion pathways and vectors by creating new opportunities and depleting traditional routes. Species-specific eradication plans are a legally binding obligation in the plant health sector and – to some extent – also in the human and veterinary medical sectors. Regulation and harmonization in Europe, however, lags far behind other regions (Hunt et al. 2008) and this is even worse for species of environmental concern. Thinking of arthropods as a mega-diverse group it is highly likely that numbers and impacts of alien species will increase worldwide. For invasive species management, it is pivotal to tackle pathways, especially in the case of small and unintentionally translocated arthropod species. For example, Skarpass and Økland (2009) proposed measures of how to reduce introduction risk of bark beetles with timber imports. Whereas considerable knowledge has been accumulated for marine pathways, one has to conclude, in agreement with Lockwood et al. (2007), that surprisingly little information is available on the exact magnitude, direction and variation of terrestrial pathways. This is especially true for Europe, where targeted research on invasion pathways should be encouraged. Following identification of the most important pathways, relevant vectors need to be thoroughly tested for their likelihood of interception (e.g. quarantine) or disruption (e.g. import ban or special obligatory and certified treatments) aiming at reducing propagule pressure. There are different options for action to be taken between maximal prevention at border controls and free trade. However, it has to be assumed that “vector management serves as a filter and not as a wall to exotic species” (Carlton and Ruiz 2005: 48). Anoplophora species provide instructive examples of how obligatory management actions are dealt with in practice in Europe. The reasonable goal of complete eradication is hampered by the implementation of national legislations, by costs borne by individual countries, and repeated introductions as a consequence of the single market policy. A united Europe should opt for better coordination, the polluter-paysprinciple, an alien emergency fund, and clear responsibilities. Ultimately, a dedicated independent agency is necessary to deal effectively with biological invasions in Europe (Hulme et al. 2009). 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A peer reviewed open access journal BioRisk 4(1): 45–50 (2010) doi: 10.3897/biorisk.4.66 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Invaded habitats Chapter 4 Carlos Lopez-Vaamonde1, Milka Glavendekić2, Maria Rosa Paiva3 1 INRA UR633, Zoologie Forestière, Centre de recherche d'Orléans, 2163 Avenue de la Pomme de Pin, CS 40001 Ardon, 45075 Orléans Cedex 2, France 2 Department of Landscape architecture and Horticulture, Faculty of Forestry, University of Belgrade, Belgrade, Serbia 3 DCEA, Faculdade de Ciências e Tecnologia, Universidade Nova de Lisboa, 2829-516 Campus de Caparica, Lisbon, Portugal Corresponding authors: Carlos Lopez-Vaamonde (carlos.lopez-vaamonde@orleans.inra.fr), Milka Glavendekić (milka.glavendekic@sfb.rs), Maria Rosa Paiva (mrp@fct.unl.pt) Academic editor: Alain Roques | Received 10 April 2010 | Accepted 20 May 2010 | Published 6 July 2010 Citation: Lopez-Vaamonde et al. (2010) Invaded habitats. Chapter 4. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 45–50. doi: 10.3897/biorisk.4.66 Abstract More than 65% (1040 species) of arthropod species alien to Europe are associated with human-made habitats, especially parks and gardens, human settlements and agricultural lands, whereas woodlands are yet colonized by less than 20% of the alien fauna, which still has a negligible representation in the other natural and semi-natural habitats. Large differences in habitat affinity are observed between alien taxonomic groups. Phytophagous species are predominant among aliens, representing 47.2% of species alien to Europe. Keywords alien, arthropod, habitat, Europe, level of invasion, urban, semi-urban 4.1 Introduction The lack of a general assessment on the level of habitat invasion in Europe has up to now limited the possibilities of evaluating the risks arthropod invaders pose to different habitats. Such an assessment is a fundamental component of early detection and identification of those environments that are more prone to invasion, that will provide a baseline for optimizing actions to prevent, monitor and control invasion (Pyšek et Copyright C. Lopez-Vaamonde. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 46 Carlos Lopez-Vaamonde et al. / BioRisk 4(1): 45–50 (2010) al. 2010). For that reason, here we present a synthesis of the data on habitat preference of terrestrial arthropods alien to Europe compiled from chapters 7-13 of this book, providing an overview of which habitats are most invaded in Europe, and to assess differences among alien taxa in terms of habitat affinity. We compared the numbers of established alien species occurring in 11 European habitats defined according to the European Nature Information System, level 1 (EUNIS) (Davies et al. 2004). This standard classification of European habitats has been chosen as a platform in several different studies on biological invasions in Europe (Chytrý et al. 2008, Daisie 2009, Pyšek et al. 2010). In this classification, a ‘habitat’ is defined as ‘a place where plants or animals normally live, characterized primarily by its physical features (topography, plant or animal physiognomy, soil characteristics, climate, water quality, etc.) and secondarily by the species of plants and animals that live there’ (Davies et al., 2004). Appendix II presents the different habitat types used throughout the taxa chapters. For more convenience, our analysis grouped them into the following broad categories roughly corresponding to the level I of EUNIS: coastal habitats (EUNIS class B); wetlands and riparian habitats (C); mires (D); grasslands (E); heathlands, hedgerows and shrub plantations (F); woodlands (G); cultivated habitats (I1); parks and gardens (we grouped the classes I2 and X11, X22, X23, X24, X25); and urban settlements (J) to which we added a specific code for greenhouses (J100). These broad categories may not precisely reflect the habitat(s) actually colonized by some species, but their use standardizes comparisons between very different taxa such as arthropods, plants and vertebrates. The habitats in the system adopted here differ considerably in the number of alien arthropod species they contain. Aliens show a strong affinity for the habitats intensively disturbed by human activities (Figure 4.1.). Considering all established alien terrestrial arthropods, the highest percentage occurs in parks and gardens (500 out of the 1590 alien species found in Europe- 31.4%) and in human settlements (31.0 %), whilst slightly less occur in cultivated habitats, which host 29.7% of these alien species. Altogether, human-made habitats host 65.4% (1040 species) of the fauna of arthropods alien to Europe, most of these species being likely to occur in several different habitats. In contrast, less than 10% of the alien species have yet colonized natural and semi-natural habitats such as wetlands, riparian habitats, grasslands and heathlands, and less than 20% occur in woodlands and forests (Figure 4.1). These results confirm the analysis of Roques et al. (2009) which relied on a lower number of alien arthropod species. Pyšek et al. (2010) also stated that alien plants are mostly found in human-made, urban or cultivated habitats, unlike vertebrates, which are more evenly distributed among habitats, the most invaded of which are aquatic and riparian habitats, woodland and cultivated land. Some habitats are differentially preferred by certain taxonomic groups (Table 4.1). For instance, many alien species are pests of ornamental plants in parks and gardens. In particular, mites are an important group attacking urban trees, shrubs and flowering plants. More than 40% of alien mites are observed in this habitat. Similarly, alien hemipterans, especially aphids, and lepidopterans have colonized parks and gar- Invaded habitats. Chapter 4 47 Figure 4.1. Main European habitats colonized by the 1590 species of terrestrial arthropods alien to Europe. The number over each bar indicates the absolute number of alien species recorded per habitat. Note that a species may have colonized several habitats. dens effectively, 78.9% and 56.7% of their species being observed there, respectively (Table 4.1). Built-up, industrial and other artificial habitats are invaded to a high degree by spiders. Indeed, more than 90% of the alien spiders are found in buildings. Psocoptera is another well-represented group in this habitat with 81.6% of its alien species in Europe occurring there, as is Phthiraptera (67.7%) and Coleoptera (57.3%), a number of species of the latter group being associated with stored products. By contrast, alien Hymenoptera are mostly present in agricultural lands which are colonized by 65.0% of the alien species in this taxon, probably in relation with the multiple parasitoid releases that have occurred for biological control purposes. Greenhouses constitute another important man-made habitat type, which hosts most alien myriapods (64.7%) and thrips (55.8%). Why do most introduced terrestrial arthropods apparently stay confined to human- modified habitats in their alien range of distribution? Several ecological conditions may be considered: i) disturbed urban and semi-urban areas may have a lower resistance to aliens, especially because of a lower pressure of potential natural enemies and, for phytophagous aliens, less vigorous host plants; ii) some species may prefer human-related habitats in their native range and are thus more likely to be carried into a new area by human transport, than species living in natural environments (Kenis et al. 2007). For instance, exotic ornamental plants are generally used in man-made habitats such as nurseries, parks and gardens and roadside plantings and shelter belts. Most alien phytophagous species introduced alongside these ornamentals remain as yet strictly associated with their original, exotic host (46.4% in Europe; Roques, 2008). They have not so far colonized native trees, and thus they develop only in parks and gardens and in hedgerows where such exotic plants are planted. A striking example 48 Carlos Lopez-Vaamonde et al. / BioRisk 4(1): 45–50 (2010) is that of the horse-chestnut leaf-mining moth Cameraria ohridella, which in its area of origin, the southern Balkans, lives in mountain ravines, whereas in its introduced area of Central and Western Europe, preferentially colonizes urban parks and gardens where its host tree has been extensively planted (Valade et al. 2009). However, there could be a time-lag between the introduction to human habitats and adaptation and spread to natural habitats. Therefore, many alien species currently confined to human-made habitats should be monitored for their potential spread to natural areas (Roques et al. 2009). For instance, species such as the Asian longhorn beetles, Anoplophora spp., (Coleoptera, Cerambycidae) have the potential to live in urban areas, in cultivated lanes (e.g. those planted with poplars) as well as in natural forests where potential host plants occur. However, dispersal from man-made habitats to natural forests appears to be a slow process. For the first twenty-two years since its arrival in North America, A. glabripennis was restricted to trees in urban areas, but in 2008, it was found in natural forests dominated by Acer trees (Haack et al. 2010). Finally, phytophagous species are predominant among the alien terrestrial arthropods, representing 47.2% (751 of 1590) of alien species to Europe, Parasitoids and predators only account for 32.6 % (518 spp.) whilst detritivores represent 20.8% (331 spp.). A few species exhibit several phytophagous guilds, whilst the habits of just 19 species are still unknown. 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Pyšek P, Bacher S, Chytrý M, Jarošík V, Wild J, Celesti-Grapow L, Gassó N, Kenis M, Lambdon P, Nentwig W, Pergl J Roques A, Sádlo J, Solarz W, Vilà M, Hulme PE (2010) Contrasting patterns in the invasions of European terrestrial and freshwater habitats by alien plants, insects and vertebrates. Global Ecology and Biogeography 19: 317–331. Roques A. (2008). The pan-European inventory of alien species established on trees on shrubs, a tool for predicting taxa and ecosystems at risk -final results of the DAISIE project. In Alien invasive species and international trade, 2nd meeting of IUFRO Working Unit 7.03.12, Invaded habitats. Chapter 4 49 May 26-30, 2008, National Conservation Training Center, Shepherdstown, WV, USA. Available at [http://www.forestry.gov.uk/pdf/IUFRO_ Shepherdstown_Roques_Sheperdstown_ end.pdf/FILE/IUFRO_Shepherdstown_ Roques_Sheperdstown_end.pdf ] Roques A, Rabitsch W, Rasplus JY, Lopez-Vaamonde C, Nentwig W, Kenis M (2009) Alien terrestrial invertebrates of Europe. In: Daisie (Ed) Handbook of Alien Species in Europe. Dordrecht: Springer, 63–79. Valade R, Kenis M, Hernandez A, Augustin S, Mari Mena N, Magnoux E, Rougerie R, Lakatos F, Roques A, Lopez-Vaamonde C (2009) Mitochondrial and microsatellite DNA markers reveal a Balkan origin for the highly invasive Horse-Chestnut leaf miner Cameraria ohridella (Lep. Gracillariidae). Molecular Ecology 18: 3458–3470. 3 (8.8) 1 (2.9) 2 (5.9) 2 (5.9) 4 (11.8) 2 (5.9) 3 (8.8) 6 (5.9) 6 (12.8) 2 (2.0) 6 (12.8) 9 (8.8) 6 (12.8) 10 (9.8) 6 (12.8) 6 (12.8) 33 (32.4) 12 (3.0) 6 (6.1) 2 (0.6) 2 (2.1) 5(1.3) 4 (4.1) 1 (0.3) 3 (1.0) 3 (0.8) 4 (4.1) 1 (0.3) 1 (0.3) 24 (6.0) 6 (6.1) 19 (6.0) 8 (2.7) 3 (3.1) 39 (9.8) 4 (4.1) 16 (5.0) 4 (1.3) 13 (13.4) 77 (19.3) 12 (12.2) 61 (19.2) 74 (24.9) 19 (19.6) 2 (0.5) 1 (1.0) 1 (0.3) 2 (0.7) 2 (2.1) 87 (21.9) 18 (18.4) 91 (28.6) 193 25 (25.8) (65.0) I2/X- Parks, gardens 9 (26.5) 42 (41.2) 69 (17.3) 17 (17.3) 251 (78.9) 23 (7.7) 55 (56.7) J- Urban, semi17 (100.0) 8 (23.5) 43 (91.5) 11 (10.8) 228 (57.3) 25 (25.5) 7 (2.2) 31(10.4) 33 (34.0) urban J100 - Greenhouses 22 (64.7) 2 (4.3) 13 (12.7) 12 (3.0) 6 (6.1) 80 (25.2) 63 (21.2) 16 (16.5) Total species 17 34 47 102 398 98 318 297 97 Zygentoma/ Collembolla Thysanoptera Siphonaptera Psocoptera Polyneoptera1 Phthiraptera Lepidoptera Hymenoptera Hemiptera Diptera Coleoptera Acari Aranea Myriapods Crustacea A- Marine habitats B- Coastal habitats C- Riparian habitats D- Mires, bogs, fens E- Grasslands F- Heathlands G- Woodlands 3 (17.6) H- Bare lands I- Cultivated lands - 1 (3.2) 4 (12.9) 7 (18.9) 2 (28.6) 3 (5.8) 1 (3.2) 2 (5.4) 1 (14.3) 2 (3.8) 8 (28.6) 1 (2.7) 12 (19.0) 2 (28.6) 2 (3.8) 1 (2.7) 2 (3.2) 7 (18.9) 1 (1.6) 7 (13.5) 1 (16.7) 7 (14.3) 2 (5.4) 8 (12.7) 1 (14.3) 15 (28.8) 1 (16.7) 21 (71.4) 20 (54.1) 40 (63.5) 5 (71.4) 1 (1.9) 3 (50.0) 31 5 (13.5) 37 49 7 29 (55.8) 3 (50.0) 52 6 Carlos Lopez-Vaamonde et al. / BioRisk 4(1): 45–50 (2010) EUNIS categories 50 Table 4.1. Comparative colonization of European habitats by the different taxonomic groups of terrestrial arthropods alien to Europe. The total number of established alien species observed in each habitat is figured. A species may have colonized several habitats. The percentage of species observed in the habitat with regard to the total number of alien species in the taxonomic group in Europe (last line) is given between brackets. ‘Polyneoptera’ includes Blattodea, Dermaptera, Isoptera, Orthoptera and Phasmatodea (see Chapter 13.3). A peer reviewed open access journal BioRisk 4(1): 51–71 (2010) doi: 10.3897/biorisk.4.42 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Impact of alien terrestrial arthropods in Europe Chapter 5 Marc Kenis1, Manuela Branco2 1 CABI Europe-Switzerland, 1, Rue des Grillons, CH- 2800, Delémont, Switzerland 2 Centro de Estudos Florestais, Instituto Superior de Agronomia, Technical University of Lisbon, Tapada da Ajuda, 1349-017 Lisboa, Portugal. Corresponding authors: Marc Kenis (m.kenis@cabi.org), Manuela Branco (mrbranco@isa.utl.pt) Academic editor: David Roy | Received 31 January 2010 | Accepted 18 May 2010 | Published 6 July 2010 Citation: Kenis M, Branco M (2010) Chapter 5: Impact of alien terrestrial arthropods in Europe. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 51–71. doi: 10.3897/biorisk.4.42 Abstract This chapter reviews the effects of alien terrestrial arthropods on the economy, society and environment in Europe. Many alien insect and mite species cause serious socio-economic hazards as pests of agriculture, horticulture, stored products and forestry. They may also affect human or animal health. Surprisingly, there is relatively little information available on the exact yield and financial losses due to alien agricultural and forestry pests in Europe, particularly at continental scale. Several alien species may have a positive impact on the economy, for example parasitoids and predators introduced for the biological control of important pests. Invasive alien arthropods can also cause environmental hazards. They may affect native biodiversity through various mechanisms, including herbivory, predation, parasitism, competition for resource and space, or as vectors of diseases. They can also affect ecosystem services and processes through cascading effects. However, these ecological impacts are poorly studied, particularly in Europe, where only a handful cases have been reported. Keywords Biological invasions, economic impact, environmental impact, alien arthropods 5.1. Introduction Alien insects and other terrestrial arthropods are among the most numerous invaders worldwide. In Europe alone, the update of the DAISIE database (Roques et al. 2009) which is presented in this book considers that 1590 terrestrial arthropod species of Copyright Marc Kenis, Manuela Branco. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 52 Marc Kenis & Manuela Branco / BioRisk 4(1): 51–71 (2010) non-European origin are established in Europe, including 1390 insects, 47 spiders,102 mites, 34 myriapods and 17 crustaceans. Many others originate from a restricted region in Europe but have invaded other parts of the continent. The establishment and spread of these alien species may have various effects. The best documented impacts are economic, particularly due to agricultural or forest pests (Pimentel et al. 2002a, 2002b). Alien arthropods also impact the environment by affecting populations or communities of native species and by disturb natural ecosystem processes and services (Kenis et al. 2009). They affect human and animal health. Finally, alien organisms have a social impact when they influence human well-being (Binimelis et al. 2007). In this chapter, we review the socio-economic and environmental impacts caused by alien terrestrial arthropods in Europe. Human and animal health impacts will be considered with socio-economic impacts since they represent measurable economic and social costs. Although the social costs of invasions are often difficult to measure in monetary terms, we could not find a single example of an alien arthropod in Europe that primarily affects human well-being without an additional economic burden. The impact of alien species is usually considered to be negative. In some cases, however, the introduction of an alien arthropod may have a positive impact on the economy or the environment, for example when an exotic biological control agent successfully controls a pest, reducing yield losses or preventing the use of pesticides. Positive impacts of alien arthropods will also be considered in this review. The review is partly based on the DAISIE database, a pan-European inventory of alien species commissioned by the European Union (Hulme et al. 2009). When building the list of alien organisms in Europe, experts were asked whether the organism had an economic or environmental impact in a particular country. Although their judgement provides valuable opinions, these have to be taken with caution because they were largely subjective and often unsupported by published references. 5.2. Socio-economic impact The economic impact of alien species has been described as the consequence of an interaction between the invader and economically valuable indigenous species (Williamson 1996). Alien arthropods can affect the economy and society in various ways, through their impact on agriculture, horticulture, forestry, stored products, human and animal health, or various services. Economic impacts can be direct or indirect. Direct economic impacts occur when alien species that affect valuable species or goods cause damage that results in yield losses and increasing production costs. These types of economic impacts are those most often described and can be easily expressed in monetary values (Pimentel et al. 2002a, 2002b). Pest management costs contribute largely to the direct economic impact of alien species. Insect pests imply the yearly application of more than 3000 million kilograms of insecticides globally (Pimentel 2007), a large share of it targeting alien pest Impact of alien terrestrial arthropods in Europe. Chapter 5 53 species. An alien pest may also cause yield losses in its role as vector of other pests and diseases, through interference with indigenous pollinators or as competitors, parasites or predators of beneficial organisms. Indirect socio-economic effects associated with the introduction of an alien pest include, among others, restrictions on trade flow, effects on market access, changes in market values, changes to domestic or foreign consumer demand for a product resulting from quality changes, changes in land use and landscape structure, public health concerns, costs associated with research and educational services, societal effects such as unemployment, effects on tourism, etc. Indirect effects are often difficult to evaluate because many of them cannot easily be expressed in monetary terms (Born et al. 2005). Vilà et al. (2010) estimated from the DAISIE database that 24.2% of the alien invertebrates in Europe have an economic impact. More than a half (51.6%) of the terrestrial arthropods alien to Europe are herbivores and, similarly, about 50% of those with economic impact are phytophagous species. Kenis et al. (2007) found that 40% of the alien insects in Switzerland and Austria had at least one web page describing damage and control methods, suggesting a socio-economic impact. Kenis et al. (2007) also estimated that the rate of native insects reaching pest status in temperate countries is probably much lower than 5%. Alien arthropods are well known for being serious plant pests worldwide. More than half of alien arthropods of economic concern are plant pests, which may directly affect yield losses of a variety of forestry and agricultural crops, such as timber, fruits, vegetables, cereals, ornamentals, etc. Insect pests destroy approximately 14% of all potential food production globally (Pimentel 2007). It is estimated that between 30 and 45% of the insect pests in agriculture and forestry worldwide are of alien origin (Pimentel et al. 2002a, 2002b), despite the fact that they only represent a few percent of the insect fauna. Economic studies on the impact of alien arthropods worldwide are numerous, but less so in Europe. Born et al. (2005) also stated that most economic analysis on the impacts of alien species has been undertaken outside Europe, particularly in North America, South Africa and Oceania. Below, we discuss the most serious economic alien pests of agriculture, protected horticulture, stored products and infrastructures, forestry and urban trees and human and animal health in Europe. Positive impacts of alien arthropods on the economy are discussed separately. 5.2.1. Outdoor agricultural and horticultural pests Many alien arthropods affect European agriculture and horticulture, mainly through yield losses and management costs, but also though quarantine measures, market effects and foreign trade impact. Reliable data on average yield and financial losses due to alien agricultural pests are not frequently published, particularly in Europe. This may be partly due to the lack of controlled, replicated experiments in commercial fields required to document such information. Furthermore, crops are often attacked by several pest species and the contribution of yield or monetary loss due to a single species is difficult to assess. Pimentel (2002) has calculated for the British Isles that, since each 54 Marc Kenis & Manuela Branco / BioRisk 4(1): 51–71 (2010) year arthropods damage or destroy approximately 10% of the crops and 30% of the pests are of exotic origin, alien arthropods cause yield losses of $960 million per year. A similar calculation for the entire European Union would lead to annual economic losses of approximately 10 billion € caused by alien arthropods. This does not include control, eradication or quarantine costs, nor costs linked to foreign trade impact or market effects. The agricultural/horticultural insecticide market represents over one billion € per year in Europe (ECPA 2007), of which probably at least 20 to 30% is to control alien pests. The first major alien agricultural insect pest that hit the European economy was the American vine phylloxera, Viteus vitifoliae, which, in the late 19th century completely destroyed nearly one-third of the French vineyards in the country, i.e. more than 1.000,000 ha, with incalculable economic and social consequences (CABI 2007). The problem was largely solved by replanting European cultivars grafted onto resistant American rootstocks, although some phylloxera biotypes have developed that may overcome the resistance of certain rootstock cultivars. Another major arthropod that invaded the European fields a while ago is the Colorado potato beetle, Leptinotarsa decemlineata. Since its first occurrence in France in 1922, it has spread to most European countries, causing considerable yield losses in potato fields. Nowadays, effective routine control of the beetle has been incorporated into potato cultivation systems and it is difficult to properly assess the economic cost of the beetle alone. In the eastern USA, the cost of controlling infestations averages between US$138 and $368 per hectare but, in this region, infestations are higher than in Europe because of the local development of resistance to the major insecticides (CABI 2007). Leptinotarsa decemlineata has not yet invaded the whole of Europe and some countries are still spending significant amounts of money to prevent its entry. For example, in Finland, pre-entry control measures against the beetle cost an average of EUR 171,000 per year in the period from 1999 to 2004 (Heikkilä and Peltola 2006). A cost-benefit analysis showed that the benefit of these protection measures strongly depends on future scenarios, in particular regarding local climatic conditions and agricultural policies. In the 1990s, the introduction into Europe of the western corn rootworm beetle Diabrotica virgifera virgifera, a serious maize pest in North America, generated much attention. A few years after its introduction, mean yield losses in Serbian Maize fields were estimated to be around 30% (Sivcev and Tomasev 2002). Baufelt and Enzian (Baufeld and Enzian 2005) calculated that the potential pecuniary losses in maize due to D. virgifera virgifera in a selection of European countries was as high as 147 million €/year, based on a conservative average yield loss of 10%. Consequently, most European countries apply costly regulatory control measures to prevent the pest’s establishment in their countries. Nevertheless, in some countries, regulatory control measures may not be economically justified. For example, in UK a cost/benefit analysis showed that, in the absence of a statutory campaign, yield losses of 5% caused by the beetle in maize could have a present value of £0.6 to £2.8 million over 20 years. However, costs Impact of alien terrestrial arthropods in Europe. Chapter 5 55 of a statutory campaign against the pest over the same period could range from £2.5 to £7.1 million (MacLeod 2006). Fruit orchards are particularly prone to alien insect invasions. Many of the most serious pests in European orchards are alien, such as the San José scale, Diaspidiotus perniciosus, the Mediterranean fruit fly, Ceratitis capitata, the oriental fruit moth, Grapholita molesta, the citrus leaf miner, Phyllocnistis citrella, the woolly whitefly, Aleurothrixus floccosus, etc. Some arthropods are harmless by themselves but are vectors of serious diseases, such as the leafhopper Scaphoideus titanus, vector of Flavescence dorée in vineyards. These arthropods, and many other alien agricultural and horticultural pests are described in the factsheets (see Chapter 14). Despite their economic importance, there is little information on the exact costs related to orchard pests. However, when data are available, they are impressive. For example, in Israel, Palestine and Jordan, the annual fruit losses due to C. capitata were estimated to be about U.S. $365 million, an amount which represents more than half of the total fruit revenue of the area (Enkerlin and Mumford 1997). 5.2.2. Pests of protected horticulture Most plant pests that occur in greenhouses and other protected environments are of tropical or sub-tropical origin. Some of them also occur on outdoor crops in Southern Europe. Among the most serious alien pests of protected crops in Europe are the leaf miners Liriomyza huidobrensis and L. trifolii, the whiteflies Bemisia tabaci and Trialeurodes vaporariorum, the aphids Aphis gossypii, Myzus persicae and Macrosiphum euphorbiae, the western flower thrips Frankliniella occidentalis (see factsheets 23, 24, 33, 35, 37 and 78), the citrus mealybug Planococcus citri and the moth Opogona sacchari. Several of these, particularly aphids, whiteflies and thrips, are vectors of important plant viruses. Mediterranean arthropods such as the lepidopteran defoliator Cacoecimorpha pronubana, the leaf mining fly Liriomyza bryoniae and the spotted spider mite Tetranychus urticae have now invaded protected crops throughout Europe (Brødsgaard and Albajes 1999). These alien pests cause enormous economic damage to the greenhouse and protected crops industry, through yield losses, control costs, contingency plans, eradication costs or losses in consignments for export. For example, Roosjen et al. (Roosjen et al. 1998) estimated that the annual cost of F. occidentalis to the Dutch greenhouse could be US$30 million, plus a further US$19 million from the effects of Tomato spotted wilt tospovirus transmitted by the thrips. An intensive eradication programme carried out to control an outbreak of the melon thrips, Thrips palmi in a UK greenhouse in 2000 cost £178,000 (MacLeod et al. 2004). A cost/benefit analysis showed that this eradication programme was four to 19 times cheaper compared with potential losses forecast by modelling the spread and impact of T. palmi in glasshouse crops over ten years. In another example, Rautapaa (1984) comparing all the costs caused by exclusion measures (eradication + quarantine) to maintain Finland free from Liriomyza 56 Marc Kenis & Manuela Branco / BioRisk 4(1): 51–71 (2010) trifolii, with the costs of living with the pest, obtained ratios 1:3 to 1:13 in favour of eradication/quarantine measures. 5.2.3. Stored product and infrastructure pests In Europe, 113 alien insect species are pests of stored products, feeding on products such as grains, seeds, fruits, fabrics, and wood products. Most are Coleoptera (e.g. Anobiidae, Bostrichidae, Chrysomelidae, Cucujidae, Curculionidae, Dermestidae, Mycetophagidae, Nitidulidae, Ptinidae, Silvanidae and Tenebrionidae), Lepidoptera (mainly Pyralidae; Gelechiidae and Tineidae) and Blattodea (cockroaches). Several alien xylophagous beetles and termites may also seriously damage public infrastructures and domestic impairments, furniture and buildings. Alien stored product and infrastructure pests are usually cosmopolitan insects of tropical or sub-tropical origin, being transported worldwide with their food (Rees 2004). Both the quantity and quality of the stored products may be affected by pests. An economic evaluation has been carried out for three species in Germany (Reinhardt et al. 2003). The annual costs arising from the two grain beetles Oryzaephilus surinamensis and Rhyzopertha dominica vary from 11.2 to 35.3 million € and that of the flour moth Ephestia kuehniella from 4.6 to 12.3 million €. Considering that these numbers are only for Germany and for three pest species, it is likely that the costs due to the two dozen economically significant alien stored product arthropod pests in Europe exceed 1 billion € per year. 5.2.4. Forestry and urban tree pests Alien arthropods can have severe economic impacts on forest plantations and urban parks. A total of 438 alien insects are associated with woody plants, representing 28.7% of all European alien species (Roques 2010). So far, European forests have suffered less from invasive arthropods than other continents, and the most important forest pests in Europe are still indigenous species. However, several potentially damaging alien forest pests have recently become established, such as the chestnut gall wasp Dryocosmus kuriphilus, the ambrosia beetle, Megaplatypus mutatus and the two Asian longhorned beetles Anoplophora glabripennis and A. chinensis (see factsheets 6, 7, and 17). Exotic trees tend to suffer more from alien pests than native trees (Day and Leather 1997). Forty-seven percent of the alien pest species affecting forest and urban trees are associated mainly or exclusively with exotic tree and shrub species (Roques 2010). For example, eucalyptus trees are particularly prone to damage by invaders from Australia. Nine alien arthropods are presently found in Europe feeding on eucalyptus, including two woodborers, Phoracantha semipunctata and P. recurva, the eucalyptus snout beetle, Gonipterus scutellatus, three psyllids Ctenarytaina eucaliptii, C. spatulata and Glycaspis brimblecombi, two gall wasps Leptocybe invasa and Ophelimus maskelli and an eriophid mite, Rhombacus eucaliptii. In southern Spain, after the first detection of P. semipunctata in 1981, the average tree mortality in the subsequent two years was estimated to be about 3%, Impact of alien terrestrial arthropods in Europe. Chapter 5 57 equivalent to a loss of 6207 ha, despite the costly control measures applied during this period (Gonzalez Tirado 1986). Gonipterus scutellatus is considered to cause tree growth losses of up to 30% in Galicia (Mansilla et al. 1996). The arrival of alien forest pests may also have indirect effects on land use and land value. For example, in Portugal, in the years following the arrival of P. semipunctata, eucalyptus plantations situated in marginal areas, poorly suitable for the cultivation of this tree species, were abandoned and the land was used for other purposes (M. Branco, unpublished observation). In contrast to what is observed in forests, a large proportion of the arthropod pests attacking ornamental and urban trees in streets, parks and gardens in Europe are alien, partly because many tree species planted in urban areas are exotic. Common non-European pests of urban trees and shrubs include, among others, the lace bug Corythucha ciliata, the scales Pulvinaria regalis and Pseudaulacaspis pentagona, the American false webworm Hyphantria cunea and the arborvitae leaf miner Argyresthia thuiella (See factsheets 41, 45, 52, 64 and 77). The citrus longhorned beetle Anoplophora chinensis was recently introduced from Asia to Italy, where it is now established and spreading, despite an eradication programme. This polyphagous wood borer has already killed thousands of urban trees and shrubs in an area of nearly 200 km2 (Tomiczek and Hoyer-Tomiczek 2007). Ornamental palms and their trade in the Mediterranean region are seriously threatened by several alien insects, in particular the Asian weevil Rhynchophorus ferrugineus and the South American moth Paysandisia archon (EPPO 2008a, 2008b). Several of the most important tree pests in Europe invaded from other parts of the continent. The maritime pine bast scale, Matsucoccus feytaudi, an Iberian species, destroyed thousands of hectares of maritime pine forest in South-eastern France, Corsica and Italy, e.g. (Covassi and Binazzi 1992, Jactel et al. 1998, Riom 1994). Important ornamental tree pests in Central and Western Europe originate from the Balkans, such as the horse-chestnut leaf miner Cameraria ohridella (Tremblay 1984) and possibly the plane leaf miner, Phyllonorycter platani (Schönrogge and Crawley 2000). Many forest pests from continental Europe have invaded the British Isles, where they may cause severe damage to forest plantations, such as the spruce aphid Elatobium abietinum or the larger spruce bark beetle Dendroctonus micans (Day and Leather 1997). Tree pests may have a direct economic effect through decrease of timber value, wood increment loss and tree mortality, treatment costs and costs related to early harvesting and replanting. There are few examples where the costs of alien forest pests have been calculated precisely in Europe. In the British Isles, the estimated cost to losses in forestry products due to alien arthropods is about $2 million per year, that is about 2% of the cost of alien arthropods in the agricultural sector (Pimentel 2002). These numbers may suggest that the direct economic impacts on forest products are much lower than on agricultural crops. The difference might partly be explained by the fact that trees may often sustain pest attacks without substantial growth loss and without tree mortality (Speight and Wainhouse 1989). Furthermore, dead trees may still have economic value as salvage. Still, it should be considered that forests account for only 11% of land cover in the British Isles (Forestry Commission 2006). In other European 58 Marc Kenis & Manuela Branco / BioRisk 4(1): 51–71 (2010) countries where the percentage of forest land cover is higher (e.g. 72% in Finland), the relative direct economic impact of alien forest pests will be much higher. Higher impact values are obtained when control costs are included. For example, Reinhardt et al. (Reinhardt et al. 2003) estimated that the control of the horse-chestnut leaf miner, Cameraria ohridella, in Germany would cost 10.02 to 33.8 millions € per year and the replacement costs for all horse-chestnut trees would be as high as 10.7 billion €. The eradication and control costs against A. chinensis in Northern Italy amounted to 900,000 € in 2005/2006, but are supposed to reach 10 million € in the period 2008–2010 (Ciampitti 2009). Furthermore, forest ecosystems provide a variety of environmental services with high socio-economic value, such as water resources, soil protection, climate amenity, carbon sequestration and leisure. All these may be seriously hampered by tree defoliation and tree mortality caused by alien forest pests. 5.2.5. Arthropods affecting human and animal health Human and animal health can be affected by various groups of alien arthropods, in particular detritivorous and hematophagous species. These generate economic costs related to control strategies, public health measures, health treatments, sick leave, educational programmes, etc. Some detritivores may affect human health by both food poisoning and disease transmission. For example, cockroaches, four of which are listed as alien in the DAISIE database, can carry microbes on their body surface and infest human and animal food. They can also provoke allergic reactions, including asthma (Brenner et al. 1987, Rivault et al. 1993). Hematophagous arthropods, besides being a human nuisance through their biting behaviour, are also able to transmit diseases or to cause allergies and dermatitis to human or domestic animals (Lounibos 2002). Seven alien mosquitoes (Diptera: Culicidae) are found in Europe. The Asian tiger mosquito, Aedes albopictus, and the Asian rock pool mosquito, Aedes japonicus, have already invaded several European countries. They both are natural vector of various viruses and filaria for humans and domestic animals (Mitchell 1995, Schaffner et al. 2009). In summer 2007, in Italy, for the first time in Europe A. albopictus was found to be the vector of an infectious disease, the Chikungunya virus (Enserink 2007). Tropical and sub-tropical mosquito species are often accidentally introduced in Europe and, with global warming, there is a risk that more mosquito species and their associated diseases could become established, particularly in southern Europe. The DAISIE database also mentions six fleas (Syphonaptera), 27 sucking louses (Phthiraptera) and 20 mites that are also able to transmit diseases or to cause allergies and dermatitis to human and animals (Roques et al. 2009). Worth mentioning are the rat flea, Nosopsyllus fasciatus, which is the primary vector for bubonic plague and murine typhus (Beaucornu and Launay 1990) and alien ticks of the genus Hyalomma that represent emerging risks for humans and animals in Europe by transmitting tickborne rickettsial diseases (Parola 2004) (see chapter 7.2.). Finally, although the vast majority of the 48 alien Araneae in Europe are of no medical concern, several species Impact of alien terrestrial arthropods in Europe. Chapter 5 59 of importance to human health are increasingly intercepted at entry ports, and a few are reported as being established, such as two Loxosceles spp. from America and a black widow, Latrodectus hasselti, from Australia (Kobelt and Nentwig 2008). 5.2.6. Arthropods with a positive economic impact Although alien arthropods are mostly associated with negative effects, some alien species may generate substantial economic benefits. For example, many predators and parasitoids introduced as biological control agents to control alien pests have a positive economic impact. The update of the DAISIE database presented in this book lists 217 nonEuropean arthropods acting as biocontrol agents of plant pests, or pests of stored products. Parasitoids include mostly chalcidoid wasps, in particular Aphelinidae (63 spp.) and Encyrtidae (55 spp.) whereas the most numerous introduced predators are Coccinellidae (12 spp.). Most of these species were intentionally introduced to control alien plant pests in outdoor crops or used as augmentative biological control agents in greenhouses. In Europe, the majority of the vegetable greenhouse area is under biological control or IPM (van Lenteren 2007), using a large variety of predators and parasitoids (van Lenteren et al. 1997). Various cost-benefit analyses have shown that, in greenhouses, biological control is the most cost-effective method (van Lenteren 2007). Many natural enemies established in the wild in Europe have a substantial impact on plant pests, such as the aphelinid Aphelinus mali, parasitoid of the woolly aphid Eriosoma lanigerum, and the coccinellid Rodolia cardinalis, predator of the cottony cushion scale Icerya purchasi (Greathead 1976). Some species released locally have been to spread quickly and rapidly become established in the wild. For example, the Australian parasitoid wasp, Psyllaephagus pilosus, which was released locally in southern France in 1997 to control the eucalyptus psyllid Ctenarytaina eucalypti, by 1998 had become established and spread westwards by more than 85 km (Malausa 1998). Interestingly, some of the most efficient natural enemies in Europe were introduced unintentionally, such as Avetianella longoi, an egg parasitoid of the eucalyptus woodborer Phoracantha semipunctata in Italy and Portugal (Farrall et al. 1992, Siscaro 1992), and Closterocerus chamaeleon, an Australian parasitoid of the eucalyptus gall wasp Ophelimus maskelli found in Portugal in 2007 (Branco et al. 2009). Pollinators are other insects whose introductions are often considered beneficial. Species and sub-species of honeybee and bumblebee have been introduced into many parts of the world, including Europe, to improve pollination of cultivated plants, either in outdoor crops or in greenhouses (Ings et al. 2005a, 2005b, Moritz et al. 2005). However, the introduction of exotic pollinators and biological control agents may also have negative effects on the environment (see section 5.3 below). 5.3. Environmental impact Alien arthropods can affect native biodiversity and ecosystem services and processes through various mechanisms (Kenis et al. 2009). Herbivores feeding on native plants 60 Marc Kenis & Manuela Branco / BioRisk 4(1): 51–71 (2010) can have a direct effect on host plant populations. Similarly, predators, parasites and parasitoids may directly affect their indigenous prey or host. Alien species may hybridize with native species, causing disturbances in native genetic resources. They can also affect the native flora and fauna and ecosystems indirectly, through cascading effects, or by carrying diseases, competing for food or space or sharing natural enemies with native species. However, these ecological impacts, their strength and the mechanisms underlying these impacts are poorly studied. Their interaction with the native fauna and flora has been rarely investigated, particularly if their habitat is of little economic concern. Based on the DAISIE database, Vilà et al. (2010) estimated that the percentage of alien terrestrial invertebrates having an ecological impact in Europe was 13.8%. However, in most cases, the notification of environmental impact was based on the fact that an alien arthropod may feed on a native plant or animal species and not on scientific evidence that populations or communities of native species are affected, or ecosystem processes are disturbed. In their extensive literature survey on the ecological effects of alien insects, Kenis et al. (2009) identified 72 alien insects worldwide for which an ecological impact had been investigated, and evidence for impact in the field was found for 54 of them. Among these, only a handful of cases came from Europe and, until now, none of them has had a tremendous impact on the environment, in contrast to what is observed in other continents. Table 1 shows the species for which an ecological effect on native biodiversity or ecosystems has been observed or investigated in Europe, and a selection of species for which an effect is suspected but for which scientific evidence is still lacking. 5.3.1. Impact by herbivores In most continents, herbivores account for the largest number of alien insects of ecological concern. For example, several forest pests of Eurasian origin cause dramatic and irreversible effects on various forest ecosystems in North America (Kenis et al. 2009). In Europe, despite the fact that phytophagous insects largely dominate the alien fauna, hardly any are known to have an ecological impact on native biodiversity and ecosystems. A potential exception is the introduction of a butterfly, the small white, Pieris rapae, in Madeira, which coincided with the extinction of a congeneric species, the Madeiran large white, P. brassicae wollastoni (Wakeham-Dawson et al. 2002). The mechanisms involved in this extinction are unclear. Gardiner (2003) suggests that the introduction of P. rapae brought a different strain of the granulosis virus for which the native butterfly had no resistance, although loss of habitat, pollution from agricultural fertilisers and an exotic parasitoid are also blamed. Another study worth mentioning is that of Schönrogge and Crawley (2000), who investigated the impact of the invasion, in UK, of cynipid gall wasps of the genus Andricus on native gall wasps through the sharing of parasitoids and inquilines. They did not find evidence that the alien species had a long term effect on populations and communities of native species. Péré et al. (2010) observed that horse-chestnut trees Aesculus hippocastanum infested by the invasive leaf miner Cameraria ohridella had a negative effect on neighbouring populations Impact of alien terrestrial arthropods in Europe. Chapter 5 61 and communities of native leaf miners. Although they suspected that the effect is due to shared natural enemies, further studies did not confirm this hypothesis (Péré and Kenis, unpubl. data). Since recently, however, introductions of phytophagous insects in Europe are causing increasing concern for their current or potential impact on the native fauna or flora. The two most serious alien palm pests in Europe, Rhynchophorus ferrugineus and Paysandisia archon, are not only a problem for the trade of ornamental plants. They are also able to develop on, and kill three endemic palm species, Phoenix theophrasti in Crete and P. canariensis in the Canary Islands, in the case of both insects, and Chamaerops humilis in the western Mediterranean region in the case of P. archon (EPPO 2008a, 2008b). The Geranium bronze, Cacyreus marshalli is a South African lycaenid butterfly introduced into southern Europe, where is has developed as a serious pest of cultivated Pelargonium spp. Laboratory tests in Italy showed that it can also develop and kill native Geranium spp. (Quacchia et al. 2008) but further studies are needed to assess better the risk and impact on the wild flora and on native Geranium-consuming lycaenids. The citrus longhorned beetle Anoplophora chinensis is presently still restricted to urban areas in Northern Italy, but it is expected to invade forests, where it could kill a large number of tree and shrub species and modify natural ecosystems. The chestnut gall wasp, Dryocosmus kuriphilus, a Chinese species damaging chestnut in Japan and North America has been recently found in Italy and is rapidly spreading to neighbouring countries, representing a serious threat for the European chestnut, a keystone species in some European forest ecosystems (Quacchia et al. 2008). Other alien phytophagous insects for which the ecological impact should be investigated include, among others: the western conifer seed bug, Leptoglossus occidentalis, which may affect the natural regeneration of conifers (Rabitsch and Heiss 2005); several seed chalcids of the genus Megastigmus that are suspected of displacing native congeneric species (AugerRozenberg and Roques 2008, Fabre et al. 2004); and Metcalfa pruinosa, a planthopper that massively attacks hundreds of different plant species in Southern Europe (Girolami et al. 1996). However, the alien insect that represents the most serious threat to European biodiversity and ecosystems may well be the emerald ash borer, Agrilus planipennis, an Asian wood borer that was detected in North America in 2002. In a few years, it has already killed over 15 million ash trees, Fraxinus spp. (Poland and McCullough 2006). The beetle has recently been detected in the region of Moscow, where it has started to cause similar damage (Baranchikov et al. 2008). Considering its dispersal capacities, there is no doubt that A. planipennis will quickly invade the rest of Europe and poses a serious threat to the three European ash species which are valuable components of various European forest ecosystems. 5.3.2. Impact by ants The alien arthropod which has been most studied for its ecological impact in Europe is undoubtedly the Argentine ant, Linepithema humile, a South American ant species 62 Marc Kenis & Manuela Branco / BioRisk 4(1): 51–71 (2010) that has invaded most continents, becoming one of the most damaging invasive insects on earth (Holway et al. 2002). In Europe, it has been reported in several countries, and has established large wild populations in Spain, Portugal, southern France and Italy. In Spain and Portugal, L. humilis was observed to displace the native ants including myrmecochorous ants, which had a negative effect on seed dispersal of native plants (Carpintero et al. 2005, Gómez and Oliveras 2003, Gómez et al. 2003, Way et al. 1997). Blancafort and Gómez (2005) noted that the invasion of L. humile reduces fruit-set and seed set of the native plant Euphorbia characias. In Madeira, however, it seems that L. humile and another invasive ant, Pheidole megacephala have little impact, even after 150 or more years of residence, and are dominated by the better adapted native ant, Lasius grandis (Wetterer et al. 2006). Way et al. (1997) noted that the displacement of native ants in Portugal was most noticeable on disturbed habitats. Also, L. humile preys on and reduces populations of serious tree pests such as the pine processionnary moth, Thaumetopoea pityocampa, and the eucalyptus wood borer (Way et al. 1992, 1999). Lasius neglectus is another invasive ant in Europe, originating from Asia Minor. It is found in several European countries, but mainly in human-modified habitats, from strictly urban sites to gardens and urban woods. Nevertheless, it can be very aggressive against native ants and some populations in Spain have displaced other surfaceforaging ants as well as other invertebrates, such as Lepidoptera (Espadaler and Bernal 2008). Lasius neglectus also tends arboreal aphids that may have a detrimental impact on trees. In England, Oliver et al. (2008) conducted laboratory studies on competitive interactions between native ants and Technomyrmex albipes, another alien ant that is presently restricted to protected habitats but may become invasive outdoors with future climate warming. 5.3.3. Impact by other predators and parasitoids Biological control agents are usually considered as beneficial because they reduce the impact of pests and the use of pesticides. In some cases, however, they may become pests themselves and threaten non-target species or other beneficial organisms. The best known case in Europe is the harlequin ladybird, Harmonia axyridis, an Asian species used in biological control programmes against aphids on greenhouse and field crops since the 1980s. The first feral populations in Europe were found in Germany in 1999 and, since then, it has spread to at least 15 countries (Brown et al. 2008). In North America, where it was released earlier, it is known to displace native ladybirds through intra-guild predation and competition for food (Koch and Galvan 2008), and it is feared that the same effects will be observed on European ladybird species. Laboratory tests have already shown that European species are vulnerable to predation by H. axyridis (Burgio et al. 2002, Ware and Majerus 2008, Ware et al. 2008), but evidence for displacement in the field needs to be further studied (Adriaens et al. 2008). Two parasitoids released to control plant pests in Europe are known to have affected populations of native parasitoids. The North American aphid parasitoid Lysip- Impact of alien terrestrial arthropods in Europe. Chapter 5 63 hlebus testaceipes, introduced in Mediterranean countries to control Aphis spiraecola, may have displaced two congeneric parasitoids, L. fabarum and L. confusus (Tremblay 1984). Similarly, the introduction of the South American Cales noacki in Italy to control the whitefly Aleurothrixus floccosus, has resulted in the displacement of the indigenous parasitoid Encarsia margaritiventris, parasitoid of the viburnum whitefly Aleurotuba jelineki (Viggiani 1994). However, in a recent paper, Viggiani (2008) stated that, in the two cases, the effects on the native parasitoids were largely local, that none of the affected native parasitoids is now endangered and that this displacement had no effect on pest populations. Alien mosquitoes are not only a threat for human or animal health. They may also affect native mosquito species through competition (Juliano and Lounibos 2005). Following the invasion of the tiger mosquito, Aedes albopictus in Italy, Carrieri et al. (2003) carried out laboratory experiments to investigate potential competitive interactions with the native Culex pipiens. They found that A. albopictus was competitively superior in resource competition but, to date, the displacement of native mosquitoes has not been demonstrated in the field. 5.3.4. Impact by pollinators and impact on pollination In Europe, as in other continents, insect pollinators, particularly bees, are declining, which may have dramatic consequences for the functioning of natural ecosystems and agriculture (Biesmeijer et al. 2006). Although the exact mechanisms leading to bees’ decline is a matter of debate, there is no doubt that the accidental introduction of natural enemies has played a significant role. In particular, the parasitic mite, Varroa destructor, which originates from the Far East and was accidentally introduced into most continents since the 1950s, has largely contributed to the decline of cultivated honeybee, partly because of its association with viruses (Sammataro et al. 2000). This has surely had an indirect ecological effect on plant pollination, although this effect is difficult to quantify. In other parts of the world, it has been shown that V. destructor also has a serious impact on feral honeybee populations (Kraus and Page 1995), but such studies are still lacking in Europe. Honeybees and wild bees may soon be threatened by a new invader, the Asian hornet, Vespa velutina (see factsheet 64). This species was introduced in south-western France some years ago, probably in pieces of pottery imported from China (Villemant et al. 2006). It is known as an important predator of bees in Asia, and it has already been reported preying on domestic honeybees in France. In addition, it may displace the European hornet, Vespa crabro. The current and potential impact of this new alien species should be assessed for the whole of Europe and management measures should be developed. The release in western and Northern Europe of two subspecies of the honeybee Apis mellifera originating from southern and eastern Europe, A. m. ligustica and A. m. carnica, has caused large-scale gene flow and introgression between these sub-species and the native black honeybee, A. m. mellifera (De La Rùa et al. 2002, Jensen et al. 2005, Moritz et al. 2005). In the Canary Islands, Dupont et al. (2003) showed that the 64 Marc Kenis & Manuela Branco / BioRisk 4(1): 51–71 (2010) introduced honeybees depleted nectar of a native plant, which reduced visitation by native pollinators and may have consequences on pollination. 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Impact of alien terrestrial arthropods in Europe. Chapter 5 71 Table 5.1. Examples of alien species with current or potential environmental impact in Europe. A Species for which field studies have been published B Species for which only laboratory studies have been published C Species that may have an environmental impact now or in the near future and for which studies are needed. Details and references are found in the text. A Andricus spp. (Hym.: Cynipidae) Apis mellifera L. subspecies carnica, caucasica and ligustica (Hym.: Apidae) Bombus terrestris (L.) subspecies dalmatinus and sassaricus (Hym.: Apidae) Cales noacki Howard (Hym.: Aphelinidae) Cameraria ohridella Deschka & Dimic (Lep.: Gracillariidae) Lasius neglectus Van Loon, Boomsma & Andrásfalvy (Hym.: Formicidae) Linepithema humile (Mayr) (Hym.: Formicidae) Lysephlebus testaceipes (Cresson) (Hym.: Braconidae) Megastigmus rafni Hoffmeyer (Hym. : Torymidae) Megastigmus schimitscheki Novitzky (Hym.: Torymidae) Pieris rapae (L.) (Lep.: Pieridae) Pheidole megacephala (F.) Hym.: Formicidae) B Aedes albopictus (Skuse) (Dipt.: Culicidae) Cacyreus marshalli Butler (Lep.: Lycaenidae) Harmonia axyridis (Pallas) (Hym.: Coccinellidae) Technomyrmex albipes Smith (Hym.: Formicidae) C Agrilus planipennis Fairmaire (Col.: Buprestidae) Anoplophora chinensis (Forster) (Col. : Cerambycidae) Dryocosmus kuryphilus Yasumatsu (Hym.: Cynipidae) Leptoglossus occidentalis Heidemann (Hem.: Coreidae) Metcalfa pruinosa Say (Hem. : Flatidae) Paysandisia archon (Burmeister) (Lep.: Castniidae) Rhynchophorus ferrugineus (Olivier) (Col.: Curculionidae) Varroa destructor Anderson & Trueman (Acari: Parasitidae) Vespa velutina nigrothorax Lepeletier (Hym.: Vespidae) Impact observed In the field In the lab No Yes Yes Yes Yes Yes Yes Yes Yes Yes Unclear No Yes Yes Yes Yes A peer reviewed open access journal BioRisk 4(1): 73–80 (2010) doi: 10.3897/biorisk.4.67 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Future trends Chapter 6 Jean-Yves Rasplus UMR Centre de Biologie et de Gestion des Populations, CBGP, (INRA/IRD/CIRAD/Montpellier SupAgro), Campus international de Baillarguet, CS 30016, 34988 Montferrier-sur Lez, France Corresponding author: Jean-Yves Rasplus (rasplus@supagro.inra.fr) Academic editor: Alain Roques | Received 15 April 2010 | Accepted 20 May 2010 | Published 6 July 2010 Citation: Rasplus J-Y(2010) Future trends. Chapter 6. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 73–80. doi: 10.3897/biorisk.4.67 Introduction The data supplied in the preceding chapters clearly confirm that the ever-increasing rate of arthropod invasions can be attributed to the upward trend in international freight transport, to passenger travel and, more generally, to globalization. The role that humans play in pest introductions as well as their likely dispersion is obvious and consequently there are strong geographic associations between higher numbers of alien pest occurrences and urban areas as already been noted by Colunga- Garcia et al. (2010) and Pyšek et al. (2010). Another important source of introduced arthropods comes from intentional releases, especially of alien hymenopterans, for the purpose of biological control programs. Invasive alien species threaten forests, agriculture, human and animal health. While economic losses attributed to exotic plant pests are poorly estimated in Europe (but see Vilá et al. 2009), they have been estimated at US $37.1 billion per year in U.S. agricultural and forest ecosystems (Pimentel et al. 2005). Invasive species can also cause irreversible changes to ecosystems, but there is no estimate of the full economic costs of their effects on ecosystems and on the human population that is dependent on them. There is little chance that biological exchanges over borders may decrease in the next decades. Rather, the number of arthropod invasions will continue to grow, threatening economy and ecosystems globally. More and more people or agricultural commodities will cross borders, increasing the likelihood that arthropods will be translocated from Copyright J.-Y. Rasplus. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 74 Jean-Yves Rasplus / BioRisk 4(1): 73–80 (2010) one area of the world to another (Liebhold et al. 2006). In Europe, monitoring, detection of arthropod invasion mostly relies on poorly connected structures hosted by different countries, using non-interoperable tools that imply months if not years to detect the data for characterizing and managing new aliens. Such delays are unacceptable in cases where immediate action has to be taken. Globalization of biological exchanges should be met by globalization of the tools used to predict, detect and manage future bio-invasions. Until now, no integrated biosecurity tool has been developed for arthropods (this is also true for all other bioinvaders). An ideal web-based integrated tool would encompass different interlinked modules to: 1. Identify the most likely future arthropod invaders 2. Provide generic and accurate identification tools 3. Compile biological information on these species 4. Predict where such aliens might potentially thrive, and their future distributions in a warmer climate or under 5. Estimate the full costs of the most likely alien arthropods 6. Finally, quantify and map risks associated with these non-indigenous species and prioritize them Identify the most likely future arthropod invaders Determining which species to target for development of detection tools, distributional area and risk estimation is not an easy task. However, it is increasingly important to identify potential invasive species prior to their introduction and establishment. This may help to reduce the likelihood of alien invasions and better define management scenarios. Only few studies have been published that help to select the most likely future arthropod invaders to Europe within the many thousands of potential bioinvader arthropods. Worner and Gevrey (2006) recently developed an original and efficient method to identify potential invasive insects that should be subject to more detailed risk assessments. They based their study on 1) the assumptions that geographical areas with similar pest assemblages share similar biotic and abiotic conditions, 2) a comprehensive database of the global presence or absence of pests. They used artificial neural network analysis to propose a list of species that are ranked according to the risks they pose. It is important to develop further methods of this kind, to implement databases and make them easily accessible through web interfaces. The development of integrated European projects such as PRATIQUE (Enhancements of Pest Risk Analysis Techniques) is a step towards this goal (Baker et al. 2009). The search for taxa that are particularly invasive worldwide may also benefit from phylogenetic or hierarchical clustering studies. Recent work on the hierarchical patterns in biological invasions has produced results that show both clustering as well as overdispersion of certain life-history traits that are associated with invasion success Future trends. Chapter 6 75 (e.g. reproductive traits) (Lambdon 2008, Procheş et al. 2008). In some cases, traits associated with invasiveness observed in a set of taxa tend to be more similar in closely related taxa, a phenomenon supposed to be linked to the conservation of ecological niches in closely related species. This observation provides promise that analysing these traits in a strict phylogenetic framework may help to predict better the most likely potential invasive species. However, few phylogenetic analyses of invasiveness have been proposed for arthropods. Such analyses may benefit from the development of DNA barcoding applied to multiple genes (see below) that could help in particular to reconstruct phylogenies within species complexes. Another approach, for phytophagous invaders at least, could be to identify and establish ‘sentinel’ host plants in not yet invaded regions, to evaluate the impact of indigenous potential invaders in source regions should they become introduced as exotics at a later date (Britton et al. 2009). This is currently carried out in China for potential pests of European tree species (Roques et al. 2009; Roques 2010). Provide generic and accurate identification tools In the last few years, the application of molecular diagnostic methods have greatly accelerated. At the same time, DNA barcoding based on the mtDNA COI gene as well as nuclear markers, have shown great potential to improve the detection of invasive species. DNA barcoding has been used to detect pests efficiently (Armstrong 2010) and may also enable the flagging of invasive species trapped during biodiversity surveys (deWaard et al. 2009). Consequently, DNA barcoding many provide an efficient new tool in the biosurveillance armoury for detection of alien species. Next generation sequencing technologies (e.g. pyro and single-molecule sequencing) may further help to reduce costs and to increase both speed and quantity of molecular detection of arthropod species. In the near future, it is likely that most identifications of arthropods will proceed through comparison of multiple gene sequences to an online global library whose quality is vastly enhanced by taxonomic knowledge. Consequently, developing a worldwide DNA library of barcodes of the most likely invasive species, including all pests and their natural enemies that could be used in biological control project, is of strategic importance to enhance our ability to detect and manage invasive populations. Such a comprehensive database coupled to real time analysis of trapping may help to detect species even at low densities, long before they become established. Developing such an integrated detection toolkit may clearly improve both biosurveillance and biosecurity in the future. Compile biological information on these species Any introduced arthropod has an area of origin where it could already be a pest and where it may already have been studied and its biology described. Available lists of 76 Jean-Yves Rasplus / BioRisk 4(1): 73–80 (2010) invasive species (NISIC, DAISIE, NOBANIS, etc) do not always provide an up-todate compilation of all available biological information and so may be of limited use for improving future management or predicting spread. To infer better the potential distribution, costs and risks associated with the most likely arthropod invaders, we need to compile all available information on their biology and life-history traits, both in their native and, when possible, in their invaded ranges (Broennimann and Guisan 2008). Predict where such aliens might potentially thrive Predicting which arthropods can invade where is critical for their management, and ultimately in limiting the negative impacts of bioinvaders. Niche-based models are widely used to predict potential distributions of invasive insects, mites or other arthopods. These methods use observations either from the invaded or the native range of an invasive species to predict the potential range in the area of introduction. However, despite its increasing use, environmental niche modelling is based on fundamental assumptions that are easily violated and lead to incorrect prediction of the full extent of biological invasions. For example, the alien species may not occupy all suitable habitats when its ecological requirements have changed during the invasion process. Furthermore, predictions are sensitive not only to occurrence and environmental data, but also to the methods used to calibrate the models. These approaches have also been criticised for their lack of consideration of species interactions (natural enemies), dispersal, availability and synchrony with the host plant or host. However, unless we can accurately parameterize the relationship between a species and its environment, no single model predicting the invasive range is likely to represent reality. This task may prove to be not feasible for most arthropods, for which knowledge of their distribution and interactions is as yet fragmentary if not rudimentary. Consequently, multiple modelling methods are required to provide better prediction and error estimates for arthropod distributional areas, especially when based on poor observation datasets. Moreover, identification of consensus areas of distributional estimate consistency using these different methods may help to produce more reliable estimates of species’ potential distributions (Roura-Pascual et al. 2009). A recent study also showed that using predictions based on both abiotic variables (usually climate) and biotic ones (for insect or host assemblage) may be more accurate than predictions based on climatic factors alone (Watts and Worner 2008). Consequently, in an effort to improve the management of invasive arthropods to Europe, we need to 1) develop a comprehenive database of life-history traits and worldwide occurrences of invasive arthropods; 2) build or implement a system providing the most accurate projections based on this database; 3) develop free access tools that implement all these methods; 4) allocate research investment to such a task that will strongly improve both predictive methodology and knowledge of the most likely invasive arthropods and their natural enemies. Future trends. Chapter 6 77 Estimate the full costs of the most likely alien arthropods Until now few general models of the economic costs of biological invasion have been developed. The goal of such models is to develop effective management programs, that seek both to estimate current or future impacts of alien invasive species, and to prevent, control, or mitigate their biological invasion. Estimates of the full costs of biological invasions (i.e., beyond direct damages or control costs) are still rare, since the costs of such complex problems are hard to calculate. Vilá et al. (2009) provided a first continent-wide assessment of impacts on ecosystem services by all major alien taxa, including invertebrates, in terrestrial, freshwater, and marine environments. They tried to compare how alien species from the different taxonomic groups affect “supporting”, “provisioning”, “regulating”, and “cultural” services and interfere with human wellbeing. However, many of these components are difficult if not impossible to quantify, such as the impacts of alien invasive species on biodiversity, ecosystem functions, human health and other indirect costs, for instance the impacts themselves of control measures. Furthermore, estimating the costs of an invasive arthropod that threatens biodiversity rather than agricultural production is particularly challenging. Precise economic costs associated with the most ecologically damaging alien species are simply not available. Consequently, we need to develop analysis of the ecological impact of introduced arthropods, especially those that are intentionally introduced for biological control purposes (Kenis et al. 2009). This is particularly important if we want in the near future to decrease our intake of pesticides and promote biological control. Economic applications are also essential to provide more accurate and comprehensive assessments of the benefits and costs of control alternatives that can increase the effectiveness and efficiency of publicly funded programs. There is also a need for the development of better databases and modelling approaches to estimate better damages from invasive species and their control costs. Further research should also be conducted to narrow the uncertainty of the estimates. Work in these areas should help improve invasive species policy and achieve a more effective use of resources. Future cost estimates should be computed, within a real-time estimation procedure, using updated infestation measuresand regional input-output economic data. Quantify and map risks associated with these non-indigenous species In the case of invasive species, risk can be defined as the probability that an invader will become established in an area along with some evaluation of the economic consequences of this event. Traditionally, quantifying risks associated with arthropod invasive species require studies on 1) the process of introduction, dispersion and the pathways used; and 2) the economic consequences of spread in recently contaminated areas (Yemshanov et al. 2009). However - as emphasized above - biology, life history and full costs of most potential invasive arthropods are still poorly known and most risk assessment studies rely on expert judgment or rudimentary analytical approaches. 78 Jean-Yves Rasplus / BioRisk 4(1): 73–80 (2010) Here again the need of integrated tools is overwhelming to produce efficient risk assessment for policy-makers. Toward a global european tool Already 1590 alien arthropod species have been introduced and established in Europe and increased efforts are needed to minimize the risk of introductions and spread of additional species in the future. Europe is poorly structured to detect rapidly, efficiently manage and control invasive arthropod species. In face of this global problem, European countries mostly have responded through nation-specific strategies and disconnected or weakly integrated projects. This disappointing situation must be changed. Faced with increasing economic pressure and despite already large grants in the past, the European Community has to invest more on invasive species prevention, detection and management. One of the key elements is the need to establish a European early warning system and rapid response framework (Genovesi 2009). In the present situation where ornamental trade is a dominant pathway for invasion by phytophagous arthropods, a more thorough survey of parks, gardens and nurseries may function as such an early warning system. This could also be accompanied by the installation of more sophisticated quarantine and control measures at invasion ‘hubs’ for the ornamental plant trade (e.g. in the Netherlands) (Roques 2010). 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A peer reviewed open access journal BioRisk 4(1): 81–96 (2010) RESEARCH ARTICLE doi: 10.3897/biorisk.4.54 BioRisk www.pensoftonline.net/biorisk Alien terrestrial crustaceans (Isopods and Amphipods) Chapter 7.1 Pierre-Olivier Cochard1, Ferenc Vilisics2, Emmanuel Sechet3 1 113 Grande rue Saint-Michel, 31400 Toulouse, France 2 Szent István University, Faculty of Veterinary Sciences, Institute for Biology, H-1077, Budapest, Rottenbiller str. 50., Hungary 3 20 rue de la Résistance, 49125 Cheffes, France Corresponding authors: Pierre-Olivier Cochard (pierre-olivier.cochard@wanadoo.fr), Ferenc Vilisics (vilisics. ferenc@gmail.com), Emmanuel Sechet (e-sechet@wanadoo.fr) Academic editor: Alain Roques | Received 28 January 2009 | Accepted 20 May 2010 | Published 6 July 2010 Citation: Cochard P-O et al. (2010) Alien terrestrial crustaceans (Isopods and Amphipods). Chapter 7.1. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 81–96. doi: 10.3897/biorisk.4.54 Abstract A total of 17 terrestrial crustacean species aliens to Europe of which 13 isopods (woodlice) and 4 amphipods (lawn shrimps) have established on the continent. In addition, 21 species native to Europe were introduced in a European region to which they are not native. The establishment of alien crustacean species in Europe slowly increased during the 20th century without any marked changes during the recent decades. Almost all species alien to Europe originate from sub-tropical or tropical areas. Most of the initial introductions were recorded in greenhouses, botanical gardens and urban parks, probably associated with passive transport of soil, plants or compost. Alien woodlice are still confined to urban habitats. Natural habitats have only been colonized by three amphipod species in the family Talitridae. Keywords Woodlice, lawnshrimps, Europe, alien 7.1.1. Introduction The orders in the arthropod subphylum Crustacea are mainly composed of aquaticliving species, at least during part of their life-cycle. Most alien terrestrial crustaceans belong to the order Isopoda, suborder Oniscidea, commonly named woodlice. But Copyright P-O. Cochard et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 82 Pierre-Olivier Cochard et al. / BioRisk 4(1): 81–96 (2010) several species recorded in Europe belong to the order Amphipoda, and are commonly known as “lawn shrimps” or “landhoppers”. In 2004, the total number of valid Isopod species worldwide was 3637 (Schmalfuss 2003). Woodlice are adapted to various terrestrial environments from sea shores to deserts and have established on all continents. As decomposers of organic plant material, isopods play an important role in ecosystems (Holthuis et al. 1987, Zimmer 2002). Most European species prefer humid and moist micro-habitats (Vandel 1960) like soil, leaf litter, mosses and decaying wood. Several species are known for their myrmecophylic nature. Amphipods are generally marine or limnicolous, and only a few species can live permanently on land (mainly in the family Talitridae). Some live near the sea, on beaches where they hide under logs and dead algae and vegetation. The true terrestrial amphipods live on the surface of mulch and moist ground (Fasulo 2008). Many of the habitat features of terrestrial amphipods are similar to those of isopods. These little animals are most commonly noticed by their strong, rapid jumps upon being disturbed. 7.1.2.Taxonomy of alien terrestrial crustaceans Thirty-eight species belonging to ten different families were recorded during this study. The four most commonly represented families (all belonging to Isopoda) are Trichoniscidae (seven species), Porcellionidae (five species), Philosciidae and Armadillidiidae, both with five species (Figure 7.1.1.). Two main categories were considered: • • Aliens to Europe, including 17 crustacean species originating from other continents (Table 7.1.1). Aliens in Europe, represented by 21 crustacean species native to a region of Europe but introduced in another European region to which they are not native. Several other species considered as cryptogenic or cosmopolitan are probably also aliens in some parts of Europe. However, in most cases it was not possible to distinguish their alien range from the native one. Below only those species we classify as aliens in Europe: Armadillidiidae: Armadillidium assimile Budde-Lund, 1879, Armadillidium kossuthi Arcangeli, 1929, Armadillidium nasatum Budde-Lund, 1885, Armadillidium vulgare (Latreille, 1804); - Oniscidae: Oniscus asellus Linnaeus, 1758; - Philosciidae: Chaetophiloscia cellaria (Dollfus, 1884); - Platyarthridae: Platyarthrus schoblii Budde-Lund, 1885; - Porcellionidae: Porcellio dilatatus Brandt, 1833, Porcellio laevis Latreille, 1804, Porcellio scaber Latreille, 1804, Porcellionides pruinosus (Brandt, 1833), Proporcellio vulcanius Verhoeff, 1908; - Schiziidae: Paraschizidium coeculum (Silvestri, 1897); Alien terrestrial crustaceans (Isopods and Amphipods). Chapter 7.1 83 Figure 7.1.1. Taxonomic overview of the Isopoda and Amphipoda species alien to and Alien in Europe. - Styloniscidae: Cordioniscus stebbingi (Patience, 1907); Trachelipidae: Agabiformius lentus (Budde-Lund, 1885); Trichoniscidae: Androniscus dentiger Verhoeff, 1908, Buddelundiella cataractae Verhoeff, 1930, Haplophthalmus danicus Budde-Lund, 1880, Metatrichoniscoides leydigi (Weber, 1880), Trichoniscus provisorius Racovitza, 1908, Trichoniscus pusillus Brandt, 1833. Some of the species above have proved to be very successful colonizers and are currently considered as part of the native fauna in parts of Europe, e.g. in Hungary. However, their synanthropic nature and their extremely wide distribution range suggest a long colonisation history as it is the case for Armadillidium vulgare. In the remainder of this chapter, we will focus mainly on the species alien to Europe. 7.1.3.Temporal trends of introduction in Europe of alien terrestrial crustaceans The total number of crustaceans alien to Europe has slowly increased during the 20th and the early 21st centuries, but without any acceleration in the rate of arrival. Two alien species were first discovered in Europe in the 19th century, about nine species in the first half of the 20th century and only five species since then. The majority of these alien species have been found in several other countries after their discovery in Europe. However, the number of occupied countries over time has grown steadily rather than exhibiting exponential growth. A similar pattern is apparent for woodlice species alien to Europe. However, because of sparcer information on this group, the date for the first introduction is roughly known for only approximately 50% of species. To our knowledge, at least six species of 84 Pierre-Olivier Cochard et al. / BioRisk 4(1): 81–96 (2010) woodlice classified as aliens of Europe were noticed in the first half of the 20th century and only five more species since then. Thus, unlike many other invertebrate phyla, the temporal trend in alien crustaceans (both intra-European and alien) has shown no marked changes during recent decades. As “silent invaders” (Hornung et al. 2007) no terrestrial crustaceans are classified as pests in Europe; they are elusive animals. We suspect frequently a large gap between the date of introduction and “discovery” of alien woodlice species. For example, during an intense eight year survey of the isopod fauna in a large region representing 15% of Hungary, three new alien species for this country were found (Farkas 2007). To conclude, the atypically gradual trend in the number of alien terrestrial Crustacea in Europe could be an artefact of incomplete knowledge. Because of both the increasing worldwide trade in ornamental plants and the general ecology of terrestrial crustaceans (i.e. often hidden in soils), it is more realistic to expect a future exponential increase in the number of alien species (especially intra-European aliens). 7.1.4. Biogeographic patterns of the alien Crustaceans 7.1.4.1. Origin of the alien species Species alien to Europe almost all originate from sub-tropical or tropical areas (Table 7.1.1.). Only one species – Protracheoniscus major (Dollfus, 1903)- is likely to be native from Central Asia. For several species, their ranges are poorly known (they are also often introduced in other tropical areas). However, several species do have a precise origin. The most widely distributed alien woodlouse in Europe is the tropical American Trichorhina tomentosa (Budde-Lund, 1893), while the most widely distributed amphipod is Talitroides alluaudi Chrevreux, 1901. It should be noted that a least six of the seventeen alien species were originally described from Europe (Great Britain, France and Germany) after their introduction. The crustaceans alien in Europe generally originate from the Mediterranean basin (seven species), from western and south-western Europe (five species). 7.1.4.2 Distribution of the alien species in Europe Within Europe, Crustaceans of alien origin have mainly been recorded in western countries, where they appeared first. The four countries with most species are Great Britain (11 species), the Netherlands (10 species) and Germany (nine species) (Figure 7.1.2). Comparatively few alien species have been recorded in central and eastern Europe to date (e.g. only four species in Hungary). In this part of Europe, the CentralAsian P. major is one of the most widespread alien crustaceans. The high number of aliens in western European countries may be linked to the high number of scientists and the intensity of soil research (Hornung 2009). Alien terrestrial crustaceans (Isopods and Amphipods). Chapter 7.1 85 Figure 7.1.2. Colonization of continental European countries and main European islands by myriapod species alien to Europe. Archipelago: 1 Azores 2 Madeira 3 Canary islands. There are only very few records of alien crustaceans on European islands. Trichoniscus pusillus has been reported from the Azores and Madeira, T. provisorius and A. assimile from the Azores but these species are native of Continental Europe. To our knowledge, the only alien aliens recorded on islands are talitrids, Arcitalitrus dorrieni (Hunt, 1925) in Scilly and Guernsey, Talitroides topitotum (Burt, 1934) in the Azores and Madeira, and T. alluaudi in the Azores and the Canaries. All these species occur outdoors and are therefore considered as naturalised. The rarity of alien terrestrial crustaceans on European islands is likely to be due to the primarily introduction route being major greenhouses in large metropolitan cities (see below). Crustaceans classified as aliens of Europe are typically species which have expanded their range approximately northwards and eastwards. The eastern and central countries have a higher number of these species than more westerly countries of Europe. For example, Germany and the Czech Republic, taken together, have nine species of alien woodlice of European origin, about 45% of the total in this category. 86 Pierre-Olivier Cochard et al. / BioRisk 4(1): 81–96 (2010) b a Figure 7.1.3. Alien terrestrial crustaceans. a Trichorhina tomentosa (Isopoda, woodlice) (credit: Vassily Zakhartchenko) b Arcitalitrus dorrieni (Amphipoda, lawn shrimp) (Credit: John I. Spicer). A striking example of successful colonization and establishment of such species is given by A. nasatum. This woodlouse is believed to be native to Italy, southern France and Spain (Vandel 1962). Since the start of the 20th century, it has been introduced into greenhouses in a number of additional countries of Northern and Central Europe (e.g. Denmark, Finland, Germany, Hungary, Poland, Slovakia, Sweden), making this species one of the most widely distributed alien woodlice of Europe. Moreover, numerous reports highlight the successful establishment of outdoor populations in several western and central European countries (e.g. the Netherlands, Czech Republic, Romania, Slovenia) (Berg et al. 2008, Giurginca 2006, Navrátil 2007, Vilisics and Lapanje 2005). Some of the aliens of Europe have also invaded other continents and can be considered as very successful invaders. The most notable ones are A. vulgare, P. scaber and P. pruinosus. Armadillidium vulgare and P. pruinosus are probably native from Mediterranean regions. In northern temperate parts of Europe, these species are restricted to synanthropic habitats (e.g. gardens, cellars, compost heaps). P. pruinosus is one of the woodlice that has been spread most by man across the world (Vandel 1962) and can now be considered as “synanthropically cosmopolitan” (Schmalfuss 2003). A consequence of the dominance of Mediterranean origin for species classified as aliens of Europe is their decreasing number towards the north of the continent (Vilisics et al. 2007). In the northernmost countries of Europe (e.g. Finland (Vilisics and Terhivuo 2009)) only the most tolerant habitat-generalists, as well as intra-European aliens, are able to become successfully established. Alien terrestrial crustaceans (Isopods and Amphipods). Chapter 7.1 87 7.1.5. Pathways of introduction of alien terrestrial Crustaceans Because a great majority of the first isopod introductions were recorded in greenhouses, botanical gardens or urban parks, it is clear that many were associated with passive transport of soil, plants or compost. With few visible effects in such biotopes, terrestrial crustaceans colonize and spread as undetected “silent invaders” (Hornung et al. 2007). Thus, most introductions were unintentional. The one known exception is the spreading of T. tomentosa, commonly sold as pet food, triggered by trading activity in Europe. This probably explains why, among all the alien crustaceans, T. tomentosa is the most widespread species in Europe. Another interesting case is the Mediterranean species P. schoblii. This myrmecophylous woodlouse is a commensal of the ant Lasius neglectus Van Loon, Boomsma & Andrásfalvy, 1990 and was first recorded in Hungary in 2001, a few years after the introduction of the ant. P. schoblii was probably introduced at the same time as its ant host (Tartally et al. 2004). It has since been found regularly (Hornung et al. 2005, Tartally et al. 2004, Vilisics 2007, Vilisics et al. 2007) and is now considered established, as is L. neglectus. 7.1.6 Ecosystems and habitats invaded in Europe by alien terrestrial Crustaceans To our knowledge, the only alien crustaceans invading natural habitats are three talitrid species. Arcitalitrus dorrieni has invaded leaf litter understoreys of deciduous woodlands in Great Britain and Ireland (Cowling et al. 2003, Vader 1972). Talitroides alluaudi is known outdoors in the Canary Islands, and T. topitotum in the Madeira Islands, both species in the Azores (Vader 1972). All other species are generally limited to highly artificial habitats and artificial ecosystems: mostly greenhouses, urban parks and houses (especially cellars). The proportion of introduced isopods can be very high in urban areas. A study in Budapest revealed that 35% of the total species (n = 28) were introduced (Vilisics and Hornung 2009). The major settlements of Hungary were characterised as “hotspot for non-native species” (Hornung et al. 2008). This could certainly be applied to many major cities in other European countries. For the tropical species, especially those recorded only once or twice in Europe, they may not be considered as established (Table 7.1.1.) since their survival is completely dependent on warm greenhouses. Among all alien woodlice, none have spread to more natural habitats. However, the situation is different for intra-European woodlice native to southern or Mediterranean Europe. These established aliens can successfully expand by dispersal from very disturbed areas (where they were originally introduced) to more semi-natural habitats in rural-suburban zones (Vilisics and Hornung 2009). With global warming and the large-scale disturbance of biomes in Europe, that trend could increase, especially for the species with large ecological spectra. 88 Pierre-Olivier Cochard et al. / BioRisk 4(1): 81–96 (2010) 7.1.7. Ecological and economic impact of alien terrestrial Crustaceans Alien crustaceans in Europe are not known to carry diseases or to have an impact on native species and natural habitats. Further, they have no economical impact. Based on existing literature, the occurrence of alien woodlice is strictly bound to the urban environment (e.g. greenhouses, botanical and private gardens); alien terrestrial isopods do not yet seem able to survive or to expand to more natural ecosystems. The case of the alien amphipod A. dorrieni is quite different. Terrestrial amphipods are known to have many effects on the soil and leaf litter (Friend and Richardson 1986). Arcitalitrus dorrieni has invaded deciduous and coniferous woodlands in western parts of Great Britain. In Ireland, a study showed that 24.7% of annual litter fall in a coniferous woodland was ingested by this species. It is suggested that “this introduced species plays a more important role than native macrofaunal species in nutrient turnover in this particular woodland habitat” (O’Hanlon and Bolger 1999). It is possible that other, as yet undetected, ecological impacts are likely. Terrestrial crustaceans can represent a large percentage of biomass and abundance in the soil macrofauna (Gongalsky et al. 2005). Thus any successful invasion by a terrestrial alien crustacean could induce some disturbance if it established in relatively natural habitats. For example, in a forested area of Florida, a study on the introduced European woodlouse A. vulgare showed that this species’ activity “had a strong effect on the chemistry of the mineral layer” (Frouz et al. 2008) and concluded that in some cases it may significantly alter soil conditions”. Woodlice classified as aliens of Europe are usually associated with synanthropic habitats and often gain dominance in urban environments (e.g. urban parks, villages, private gardens). The successful colonisation of human- influenced biotopes may lead to the uniformity of local Isopod assemblages. With the decrease of native species in the urban isopod fauna, an ongoing process of biotic homogenisation is prevalent in cities across Europe (Szlávecz et al. 2008, Vilisics and Hornung 2009). Acknowledgements The authors would like to thank Matty Berg (Associate Professor, Vrije Universiteit Amsterdam, the Netherlands); Samuel Danflous (Entomologist, France); Elisabeth Hornung (Associate professor, Szent István University, Hungary); Spyros Sfenthourakis (Associate professor, University of Patras, Greece); Stefano Taiti (Dr, Istituto per lo Studio degli Ecosistemi, Italy); Ivan H. Tuf (PhD, Palacký University, Czech Republic); and Wim Vader (Professor, Tromsø Museum, Norway). 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Memoranda Societatis pro Fauna et Flora Fennica 85: 9–15. Wouters K, Tavernier JM, Meurisse L (2000) Distribution and bibliography of the terrestrial Isopoda (Crustacea) of Belgium. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique 70: 195–205. Zimmer M (2002) Nutrition in terrestrial isopods (Crustacea: Isopoda): an evolutionary-ecological approach. Biological Reviews 77: 455–493. Native range 1st record in Europe Invaded countries Habitat References Central Asia? 1903, PL/ UA AT, CZ, DE, J EE, HU, PL, RO, SK, UA Dudich (1926),Dudich (1933), Dyduch (1903), Dominiak (1970), Flasarová (1986), Flasarová (1988), Flasarová (1995), Forró and Farkas (1998), Frankenberger (1959), Ilosvay (1985), Schmölzer (1974), Semenkevitsh (1931), Strouhal (1929), Strouhal (1951), Verhoeff (1930) Detritivorous Pacific islands 1930, DE DE, FR, GB, J HU, NL Detritivorous Congo 2003, NL NL J Detritivorous Tropical regions 2003, NL GB, NL J Berg et al. (2008), Grüner (1966), Holthuis (1947), Holthuis (1956), Kesselyák (1930a), Kesselyák (1930b), Kontschán (2004), Schmalfuss (2003), Soesbergen (2003), Vandel (1962), Verhoeff (1937) Berg et al. (2008), Schmalfuss (2003), Soesbergen (2003), Soesbergen (2005) Berg et al. (2008), Gregory (2009), Schmalfuss (2003), Soesbergen (2003) Detritivorous East Africa 1928, DE DE, NL J Detritivorous Brazil (Southeast) Asia ?, DE DE J Berg et al. (2008), Ferrara and Taiti (1982), Holthuis (1945), Schmalfuss (2003), Verhoeff (1928) Schmalfuss (2003) 1947, GB GB J Harding and Sutton (1985), Holthuis (1947) Isopoda Armadillidae Reductoniscus costulatus Kesselyák, 1930 Synarmadillo pallidus Arcangeli, 1950 Venezillo parvus (Budde-Lund, 1885) Isopoda Philosciidae Anchiphiloscia balssi (Verhoeffff, 28) Benthana olfersii (Brandt, 1833) Burmoniscus meeusei (Holthuis, 1947) Detritivorous Pierre-Olivier Cochard et al. / BioRisk 4(1): 81–96 (2010) Order Species Regime Family Isopoda Agnaridae Protracheoniscus major Detritivorous (Dollfus, 1903) 94 Table 7.1.1. List and main characteristics of the Crustacean species alien to Europe. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Only selected references are given. Last update 16/10/2009. Order Family Species Regime Burmoniscus orientalis Detritivorous Green, Ferrara & Taiti, 1990 Native range Asia 1st record Invaded in Europe countries 2005, AT AT Habitat J References Uteseny (2009) Platyarthridae Trichorhina tomentosa Detritivorous (Budde-Lund, 1893) America (Tropical) 1896, FR AT, CH, BE, J CH, CZ, DE, FR, GB, HU, IE, NL, NO, PL1 Berg et al. (2008), Dollfus (1896a), Foster (1911), Foster and Pack-Beresford (1913), Harding and Sutton (1985), Holthuis (1945), Jedryckowsky (1979), Korsós et al. (2002), Meinertz (1934), Olsen (1995), Pack-Beresford and Foster (1911), Polk (1959), Schmalfuss (2003), Verhoeff (1937), Wouters et al. (2000) Styloniscidae Styloniscus spinosus (Patience, 1907) Detritivorous Madagascar, Mauritius 1907, GB GB Edney (1953), Harding and Sutton (1985), Patience (1907), Schmalfuss (2003) Trachelipodidae Nagurus cristatus (Dollfus, 1889) Detritivorous Pantropical 1956, NL DE, GB, NL, J RO Detritivorous Tropical regions 1985 GB GB, IE J Detritivorous USA (East) ? 1908,GB DE, GB J Isopoda J Isopoda Nagurus nanus Budde-Lund, 1908 Allspach (1992), Berg et al. (2008), Harding and Sutton (1985), Holthuis (1956), Oliver and Meechan (1993), Radu (1960), Schmalfuss (2003) Foster (1911), Foster and Pack-Beresford (1913), Harding and Sutton (1985), Schmalfuss (2003), Sutton (1980) Isopoda Trichoniscidae Miktoniscus linearis (Patience, 1908) Alien terrestrial crustaceans (Isopods and Amphipods). Chapter 7.1 Isopoda Kesselyák (1930a), Patience (1908), Schmalfuss (2003), Vandel (1962) 95 Regime Detritivorous Native range Australia (East) 1st record in Europe 1925, GB Tropical 1912, GB regions? Tropical 1896, FR regions, Seychelles Isl.? Talitroides topitotum (Burt, 1934) Indo-Pacific Detritivorous 1942, DE Habitat GB, IE, NL G1, J GB, NL J BE, CH, CZ, G1, J DE, DK, ESCAN, FI, FR, GB, HU, NL, PL, PTAZO, SE DE, GB, NL, G,J PT-AZO, PT-MAD References Cowling et al. (2003), Cowling et al. (2004a), Cowling et al. (2004b), Hunt (1925), Moore and Spicer (1986), Peart and Lowry (2006), Spicer and Tabel (1996) Calman (1912), Friend and Richardson 1986, Vader (1972) Chevreux (1896), Dudich (1926), Friend and Richardson (1986), Hunt (1925), Vader (1972) Friend and Richardson (1986), Stock and Biernbaum (1994), Vader (1972) 1 Trichorhina tomentosa is on sale as reptile food in many European pet shops. After this table was established, Gregory (2009) mentioned the presence of two more alien species in Great Britain, Styloniscus mauritiensis (Barnard, 1936) (Styloniscidae) from Hawaii and Mauritius and Setaphora patiencei (Bagnall, 1908) (Philosciidae) from The Réunion and Mauritius islands. Pierre-Olivier Cochard et al. / BioRisk 4(1): 81–96 (2010) Brevitalitrus hortulanus Detritivorous Calman, 1912 Talitroides alluaudi Detritivorous (Chevreux, 1896) Invaded countries 96 Order Species Family Amphipoda Talitridae Arcitalitrus dorrieni (Hunt, 1925) A peer reviewed open access journal BioRisk 4(1): 97–130 (2010) doi: 10.3897/biorisk.4.51 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Myriapods (Myriapoda) Chapter 7.2 Pavel Stoev1, Marzio Zapparoli2, Sergei Golovatch3, Henrik Enghoff4, Nesrine Akkari5, Anthony Barber6 1 National Museum of Natural History, Tsar Osvoboditel Blvd. 1, 1000 Sofia, Bulgaria 2 Università degli Studi della Tuscia, Dipartimento di Protezione delle Piante, via S. Camillo de Lellis s.n.c., I-01100 Viterbo, Italy 3 Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky prospekt 33, Moscow 119071 Russia 4 Natural History Museum of Denmark (Zoological Museum), University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen, Denmark 5 Research Unit of Biodiversity and Biology of Populations, Institut Supérieur des Sciences Biologiques Appliquées de Tunis, 9 Avenue Dr. Zouheir Essafi, La Rabta, 1007 Tunis, Tunisia 6 Rathgar, Exeter Road, Ivybridge, Devon, PL21 0BD, UK Corresponding author: Pavel Stoev (pavel.e.stoev@gmail.com) Academic editor: Alain Roques | Received 19 January 2010 | Accepted 21 May 2010 | Published 6 July 2010 Citation: Stoev P et al. (2010) Myriapods (Myriapoda). Chapter 7.2. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 97–130. doi: 10.3897/biorisk.4.51 Abstract Alien myriapods in Europe have never been subject to a comprehensive review. Currently, 40 species belonging to 23 families and 11 orders can be regarded as alien to Europe, which accounts approximately for about 1.8% of all species known on the continent. Millipedes (Class Diplopoda) are represented by 20 alien species, followed by centipedes (Class Chilopoda) with 16, symphylans with 3 and pauropods with only 1. In addition there are numerous cases of continental species introduced to the Atlantic and Mediterranean islands or others of southern origin transported and established in North European cities. The earliest record of an alien myriapod dates back to 1836, although the introduction of some species into Europe could have begun already in historical times with an increase in trade between ancient Greeks and Romans with cities in the Near East and North Africa. In post-medieval times this process should have intensified with the trade between Europe and some tropical countries, especially after the discoveries of the Americas and Australia. The largest number of alien myriapods (25, excl. intercepted) has been recorded from Great Britain, followed by Germany with 12, France with 11 and Denmark with 10 species. In general, northern and economically more developed countries with high levels of imports and numerous busy sea ports are richer in alien species. The various alien myriapods have different origins, but most of them show tropical or subtropical links (28 species, 70%). Eight of them (20%) are widespread in the Tropical and Subtropical belts, eleven (circa 28%) are of Asian origin, seven show links with South and Central America, and one each originates from North America, North Africa, Australasia, and islands in Copyright P. Stoev et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 98 Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010) the Indian Ocean. Ten myriapods are of unknown origin (cryptogenic). Only 12 species (ca. 30%) of all have established in the wild in Europe. At the present time alien myriapods do not cause serious threats to the European economy and there is insufficient data on their impact on native fauna and flora. Keywords Diplopoda, Chilopoda, Symphyla, Pauropoda, Europe, alien, invasions, intercepted species, biogeographical patterns 7.2.1. Introduction Myriapods are terrestrial wingless arthropods with elongated bodies composed of more or less similar segments, most of which bear one or two pairs of legs. Four classes are recognised: Pauropoda, Symphyla, Chilopoda and Diplopoda. Approximately 15 000 species from nearly 160 families are currently known in the world. The Diplopoda is by far the most diverse group, comprising roughly 11 000 species (Adis and Harvey 2000). A total of 2,245 myriapod species or subspecies from 15 orders and 70 families are currently known in Europe (http://www.faunaeur.org/statistics.php), of which 1,529 are Diplopoda, 481 Chilopoda, 41 Symphyla and 125 Pauropoda. All members of the class Diplopoda (millipedes) have two pairs of legs per diplosegment for most segments. Several morphotypes have been recognised, i.e. juloid, polydesmoid, polyxenoid, platydesmoid and glomeroid (Kime and Golovatch 2000), of which the former two are especially rich in species both in Europe and worldwide. Most of the species are cylindrical or flattened dorsally, often with prominent lateral projections, generally medium- to large-sized (up to 8–9 cm in the genera Pachyiulus and Eurygyrus). Some species of the order Glomerida, or pill millipedes, are oniscomorph, capable of rolling up into a tight ball. Members of the order Polyxenida, or “dwarf millipedes”, are minute in size and with peculiar hairs along the body arranged in groups and tufts like small pin-cushions or brushes. The number of legs varies between species, often (especially in juloids) even individually, the record being 375 pairs in the North American siphonophoridan species Illacme plenipes Cook & Loomis, 1928 (Marek and Bond 2006). Species of the class Chilopoda (centipedes) have an elongated flattened trunk and bear one pair of legs per segment, with a total number ranging between 15 and 191 pairs. Centipede body length varies from a few millimeters in some species of genus Lithobius (Monotarsobius) to approximately 30 cm in the Neotropical species Scolopendra gigantea (Minelli and Golovatch 2001). All centipedes have a pair of poison claws, or forcipules, which represent modified first body appendages. About 3,500 valid species and subspecies from 5 orders and 22 families are known in the world (Minelli 2006, Edgecombe and Giribet 2007). The other two myriapod classes – Symphyla and Pauropoda – consist of very small species, with body length of 2–8 and 0.5–1.9 mm respectively, both still remaining very poorly studied. The number of described symphylans and pauropods in the world is roughly estimated to about 200 and 700, respectively (Adis and Harvey 2000). Myriapods (Myriapoda). Chapter 7.2 99 Most millipedes, as well as all pauropods and symphylans, are phytophages, detritivores or saphrophages. A few millipedes can be regarded as omnivores, e.g. Blaniulus guttulatus (Fabricius, 1798), Uroblaniulus canadensis (Newport, 1844), or even predators, like Apfelbeckia insculpta (L. Koch, 1867), Callipus foetidissimus (Savi, 1819), and Abacion magnum (Loomis, 1843), which have been observed feeding on earthworms, flies and spiders (Hoffman and Payne 1969, Golovatch 2009). Other species feed on their own exuvia or fecal pellets (Minelli and Golovatch 2001). Centipedes are mostly predatory, feeding on different available prey items in the soil (earthworms, enchytraeids, snails, slugs, small insects – both larvae and adults – and other arthropods). More details on the biology and ecology of millipedes, centipedes and the two other, smaller myriapod classes can be found in Hopkin and Read (1992), Lewis (1981), and Verhoeff (1933, 1934). Little information is as yet available on the non-indigenous myriapods in Europe (DAISIE 2009, Roques et al. 2009). The most recent overview of alien organisms in Europe (see DAISIE 2009, p. 225) lists two centipedes (Lamyctes emarginatus, Lamyctes caeculus) and three millipedes (Oxidus gracilis, Eurygyrus ochraceus, Sechellobolus dictyonotus = Paraspirobolus lucifugus) as alien to Europe. Some papers have been, however, published on the ecology, morphology and post-embryonic development of several alien centipedes (Andersson 1984, 2006, Bocher and Enghoff 1984, 1975a, Negrea 1989) and millipedes (Enghoff 1975b, 1978, 1987, Golovatch et al. 2000, et al. 2002). Lists of alien species have been published for a few countries only, such as Germany (Kinzelbach et al. 2001), Austria (Gruber 2002, Gruber and Christian 2002), the Czech Republic (Šefrová and Laštůvka 2005), Switzerland (Wittenberg 2005), Italy (Zapparoli and Minelli 2005) and Great Britain (Barber 2009a, b). Increasing attention has been paid in the last decades to species which have accidentally arrived in Europe (see Barber 2009a, BBC News 2005, Christian and Szeptycki 2004, Gregory and Jones 1999, Lewis 1988, Lewis and Rundle 1988 for centipedes and Andersson and Enghoff 2007, Enghoff 2008a and Read 2008 for millipedes). 7.2.2.Taxonomy of the myriapod species alien to Europe Altogether, 40 species belonging to 23 families and 11 orders can be regarded as alien to Europe (Table 7.2.1). This accounts approximately for about 1.8% of all myriapods known on the continent. Millipedes are represented by 20 alien species, followed by centipedes with 16, symphylans with 3 and pauropods with only 1. The relative proportion of alien species is highest in Symphyla (7.3%) and Chilopoda (3.3%), and the lowest in Pauropoda (0.8%) and Diplopoda (1.3%). The centipede family Henicopidae is the richest in alien species (5 species), followed by Scutigerellidae, Mecistocephalidae, Scolopendridae, Paradoxosomatidae and Pyrgodesmidae, each with three species. The remaining 17 families are represented by only one or two species each (Figure 7.2.1). 100 Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010) Figure 7.2.1. Relative importance of each family in the alien (right side) and native (left side) myriapod fauna in Europe. Number near the bar indicates the number of species. Families are listed in a decreasing order based on the number of alien or, in alternative, native species. Striking is the absence of alien species in Europe of the species-rich order Spirostreptida since spirostreptidans, for instance Hypocambala anguina (Attems, 1900) and Glyphiulus granulatus Gervais, 1847, are quite widespread in the tropical countries and show a clear tendency to anthropochorism (Jeekel 1963, Shelley 1998). Several myriapods have been intercepted at their arrival in Europe from consignments from other countries but have never managed to establish themselves. Barber (2009a) provided a list of centipede species captured and registered by the Central Science Laboratory (now FERA) in the UK when imported with exotic plants, fruits and luggage (Table 7.2.2). Two of them, Lithobius forficatus and L. peregrinus, are European natives which have long been introduced to Australia and New Zealand, so their interception in Great Britain is a clear case of re-introduction. A similar list for intercepted millipedes examined by the Central Science Laboratories between 1975 and 2006 (S. Reid pers. comm.) is more substantial with some 85 entries over this period of time (Table 7.2.2). Of these 36 were records of Oxidus gracilis from a wide range of different parts of the world (W & S. Europe, Canary Islands, Israel, N., C. and S. America, Australia, China, Japan, Malaysia, Singapore, India, Nepal, N., W. and S. Africa and Madagascar). Other types found included members of the Polydesmida (dalodesmids, parodoxomatids, polydesmids), Spirosteptida (from Australia, New Zealand and Africa), Julidae and Blaniulidae. Amongst species from the latter two families were the NW European Blaniulus guttulatus and Cylindroiulus Myriapods (Myriapoda). Chapter 7.2 101 londinensis (both from Australia) and Ommatoiulus moreletii (originating in the Iberian Peninsula, introduced to Australia in 1953 and now a pest species there; in this list reported from both that country (tree fern) and South Africa (melon fruit)). Man-aided introductions of species from one part of Europe to another have played a prominent role. One of the most common synanthropic centipedes in North Europe is the Mediterranean “house centipede” Scutigera coleoptrata (Linnaeus, 1758). It has been introduced to a number of North European cities, e.g., Copenhagen, Edinburgh, Aberdeen, Leiden, etc., where it survives only in buildings. The earliest record in the British Isles of this species is perhaps that by Gibson-Carmichael (1883) who recorded it from a paperworks near Aberdeen. It could have been established there already for 25 years and arrived in bundles of rags from South Europe (Barber 2009a); at the present time it is sporadically reported from inside buildings in various parts of Britain and seems to be common in houses in St. Peter Port (Guernsey) and St. Helier (Jersey) in the Channel Islands from where it has also been reported from outdoor sites. Other cases of south or central European species being introduced to northern countries that perhaps still survive only in buildings, hothouses, gardens or similar man-made habitats are: Tuoba poseidonis (Verhoeff, 1901) in Finland, Dicellophilus carniolensis (C.L. Koch, 1847), Lithobius lucifugus L. Koch, 1862, Lithobius peregrinus Latzel, 1880, Haplopodoiulus spathifer (Brölemann, 1897) and Cylindroiulus salicivorus Verhoeff, 1908 in Great Britain, Cylindroiulus vulnerarius (Berlese, 1888) in Sweden, Pachyiulus varius (Fabricius, 1781) in Norway, etc. (Barber 1995, Barber and Eason 1986, Barber and Keay 1988, Bergersen et al. 2006, Lee 2006, Read 2008). Even within the same geographic area some indigenous species occur at localities that are not part of their primary distribution area, most probably as a consequence of accidental anthropogenic introductions. Examples are the records from Italy of Lithobius infossus Silvestri, 1894 near Padua (Minelli 1991), of L. peregrinus Latzel, 1880 in northeastern and central Italy (Minelli 1991, Zapparoli 1989, Zapparoli 2006), of Pleurolithobius patriarchalis (Berlese, 1894) in the Egadi, Pontine and Campania islands (Zapparoli and Minelli 1993), and of Scolopendra cingulata near Milan (Manfredi 1930). Island invasions by continental species is another phenomenon worth mentioning. Eason in a study on the Icelandic fauna, concluded that most centipede and millipede species probably arrived by human transport (Eason 1970). Examples of recent introductions to Iceland are Geophilus truncorum Bergsøe & Meinert, 1866, Polydesmus inconstans Latzel, 1884, and Brachydesmus superus Latzel, 1884, which “… have only been found on Heimaey, one of the Vestman Islands, which supports a town and where casual introduction by human transport is likely: they have probably been introduced quite recently and the two millipedes seem still to be confined to the outskirts of the town”. Regarding the other two possibly allochthonous species, Lithobius forficatus (Linnaeus, 1758), and Lithobius erythrocephalus C.L. Koch, 1847, Eason wrote, “these two species may be confined to the south owing to the relatively warm and humid southern climate, but their restricted distribution might also be explained by their having been introduced by Norse settlers ....”. The first Norse set- 102 Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010) tlements on Iceland were established in the ninth century A.D., so this must have happened after that time. According to Enghoff (2008b), of the 21 species of centipedes recorded in Madeira 17 are introduced and 2 are probably introduced. High rates of introduction are also known for the Azores and Canary Islands (Borges and Enghoff 2005, Zapparoli and Oromi 2004) (Table 7.2.3). All symphylans on the Canary Islands have been considered as possibly introduced. Likewise, only two of 21 millipede species are considered native on the Azores (Enghoff and Borges 2005). The geophilomorph centipede Nyctunguis persimilis Attems, 1932 was originally described from Turkey and has not been found there since in spite of the active work of the second author who has published several papers on the Turkish centipede fauna during the last 20 years. Taking into account that the species was recently found in synanthropic habitats in the outskirts of Vienna (Christian 1996) and that all other congeners occur in the Nearctic region, it is very likely that the type locality (the surroundings of Ankara, Turkey) is erroneous and the material was actually mislabeled (Zapparoli 1999). Mecistocephalus maxillaris (Gervais, 1837), one of the first alien centipedes to be recorded in Europe, is another poorly known species. It was described from the gardens of the Muséum National d’Histoire Naturelle, Paris, and subsequently recorded from numerous places around the world. However, most of the records were certainly based on misidentifications with the morphologically similar M. guildingii or M. punctifrons actually being involved (Bonato et al. 2009). According to Bonato et al. (2009), most of the records in Europe e.g., those from Germany, Great Britain, France (not the type specimen but material taken subsequently from a greenhouse in the Paris Museum, cf. Brolemann 1930) and Portugal (Madeira), are referable to M. guildingii, while those from the Netherlands and Denmark require further clarification. The actual taxonomic status and native range of Ghilaroviella cf. valiachmedovi remains uncertain. The same applies to the millipede Chondrodesmus cf. riparius which shows some differences from the original description by Carl (1914) and its identity cannot be clarified without a comprehensive review of the entire genus (Enghoff 2008a). 7.2.3. Temporal trends in the introduction of alien myriapod species to Europe Introductions of alien myriapods into Europe probably began several centuries ago, even though a precise arrival date is hard to determine. Only 10 out of 40 species were recorded for the first time in Europe in the 19th century while most of the records date from the 20th (26 species) and 21st centuries (4 records). Gervais was virtually the first person to record alien myriapods in Europe (Gervais 1836, 1837). He described the tropical millipede Iulus lucifugus (now Paraspirobolus lucifugus) and the geophilomorph centipede Mecistocephalus maxillaris from greenhouses of the Paris Museum. The means of arrival of both species remains obscure but Myriapods (Myriapoda). Chapter 7.2 103 must be linked to the establishment of the greenhouses and the planting of tropical flowers, perhaps already by the end of the 18th century. P. lucifugus has been subsequently recorded in intervals of around 60–70 years from greenhouses near Hamburg (Latzel 1895), Hortus Botanicus Amsterdam (Jeekel 1977), a greenhouse in Copenhagen (Enghoff 1975b), and more recently from the Tropical Biome at the Eden project (Lee 2006). This can hardly be regarded as reflecting the actual course of colonization but rather the date of investigation and the level of effort involved in each case. The only alien millipede that has invaded some natural ecosystems in Europe and acclimatized is the East Asian species Oxidus gracilis. Perhaps the earliest records of this species in Europe are those of Tömösváry (1879) from the Margaret Island in Danube, Hungary, and of Latzel (1884) from greenhouses in Zeist, Utrecht, and Amsterdam in the Netherlands. Subsequently it was also found in Edinburgh in 1898 and in Kew Gardens in Great Britain (Evans 1900, Pocock 1902). In Finland the species was first recorded in 1900, but since the sample already contained several specimens the species must have arrived there at least two years earlier (Palmén 1949). The mechanism of dispersal of the species within Europe is certainly related to the trading and growing of tropical plants in the greenhouses as in some places this process must have happened more than once. According to Palmén (1949), the population of O. gracilis in the greenhouses of Hanko, South Finland went extinct during the period 1939–1943 when they were not kept warm. In 1946 a single female was found in a greenhouse with plants imported from Belgium, in 1947 its numbers increased considerably and the next year it was already very abundant in the whole greenhouse complex. Golovatch (2008) suggested that the intense trade ties between the ancient town of Khersonesos in the Crimea and the town of Pergam (= Bergama), a major centre of red ceramics production of the time in present-day Turkey, as possible pathways for the introduction of Eurygyrus ochraceus in the Ukraine. He also pointed out that the Bulgarian population near Varna may owe its origin to the very active commerce in Roman times between Bergama and the colonies in Moesia (= currently northern Bulgaria and southern Romania), including Odessos (= Varna). The area and trade connections were already quite developed by the mid-4th century B.C. or even earlier, under ancient Greeks, so this introduction must have happened around that time. Members of the genus Lamyctes are represented in Europe only by parthenogenetic populations. Males of L. emarginatus are known only from Macaronesia, New Zealand, Tasmania and Hawaii (see also Attems (1935) and Zapparoli (2002) for the record of a single male from Greece), while males of L. coeculus are only known from a greenhouse in Italy and from Cuba (Enghoff 1975a). Taking into account that the entire family Henicopidae is predominantly distributed in the Southern Hemisphere, and presuming that the regions where males are being found are the native areas of the species, L. emarginatus could have been introduced to Europe from one of the above regions, most likely from Australia or New Zealand. The earliest confirmed record is from Denmark in 1868 (see Meinert 1868). Lamyctes coeculus was first found in a greenhouse in Italy at the end of 19th century (Brölemann 1889), but its presence in the area would have been older. It has been recently found in Great Britain (Barber 2009b). 104 Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010) The earliest records of Cylindroiulus truncorum in Europe date from the 1920’s and, according to Schubart (1925), the Central European populations are probably of relatively recent origin. In Finland it was first reported in 1945 and in the following three years its numbers increased considerably. It is completely lacking in older collections (Palmén 1949). One of the recent introductions is the large Neotropical millipede Chondrodesmus cf. riparius which was first recorded in 2000 in a flowerpot in the telephone office of Umeå University, northern Sweden. It was found again elsewhere in Sweden in 2006 and, later, in January 2007, it was also recorded in a flowerpot with a palm (Phoenix robbelini) in an office in Copenhagen and in a flowerpot in Bonn (Enghoff 2008a). There are further records of the species from flowerpots in Germany and also a recent one in Norway (Göran Andersson in litt.), so it seems that the species is dispersing well with palm pots. The study of the invertebrate fauna of Kew Gardens, Great Britain began already at the beginning of 20th century with papers by Pocock (1902, 1906) and continues today (Blower and Rundle 1980, 1986, Read 2008). Some of the species recorded by Pocock such as Scolopendra morsitans, Trigoniulus corallinus and Asiomorpha coarctata have not been re-found since then and most likely could not become established in Kew Gardens. At the same time Paraspirobolus lucifugus, Amphitomeus attemsi, Cylindrodesmus hirsutus, Rhinotus purpureus and Pseudospirobolellus avernus, species not previously known from Britain have been recorded recently in the Tropical Biome at the Eden project in Cornwall (Read 2008, Barber 2009b, Barber et al. 2010). 7.2.4. Biogeographic patterns of the myriapod species alien to Europe Records of exotic species are not evenly distributed in Europe but this is mainly due to the different levels of investigation of this area. The highest number of species (25) has been recorded from Great Britain, followed by Germany with 12, France with 11 and Denmark with 10 alien myriapods (Figure 7.2.2). In general, northern and economically more developed countries with high levels of imports and numerous busy sea ports are richer in alien species. These countries also, in general, have poorer native faunas meaning that a small number of aliens can constitute a large percentage of the fauna. Several species are hitherto known in Europe from a single country only, e.g. Prosopodesmus panporus, Pseudospirobolellus avernus, Tygarrup javanicus and Cryptops doriae, which implies recent introductions or poor dispersal abilities. Others, such as Eurygyrus ochraceus, Paraspirobolus lucifugus and Lamyctes coeculus, have a larger but yet fairly restricted distribution limited to two or more countries. The most widespread species are the parthenogenetic centipede Lamyctes emarginatus, whose range in Europe spreads from the Urals to Iceland [outdoor species], and the bisexual millipede Oxidus gracilis, reported from 33 countries, including several Mediterranean islands. Myriapods (Myriapoda). Chapter 7.2 105 The various alien myriapods have different origins, but most of them show tropical or subtropical links (28 species, 70%). Eight of them (20%) are widespread in the Tropical and Subtropical belts, very often introduced by human agency to islands and synanthropic areas on continents. Their native range cannot so far be determined with certainty (Figure 7.2.3). Eleven (circa 28%) alien myriapods are of Asian origin, the majority (10 species) having their native range in East or Southeast Asia, and only one from West Asia, namely Anatolia. Cylindroiulus truncorum is perhaps the only North African myriapod introduced to Europe just as Brachyiulus pusillus (Leach, 1814) so far is the only European julid introduced to North Africa (Akkari et al. 2009). The only species that seems to be an Australasian native (Australia and New Zealand) is Lamyctes emarginatus. Among henicopids, Rhodobius lagoi and Ghilaroviella cf. valiachmedovi are of particular interest being members of monotypic genera and the only representatives in Europe of the subfamily Anopsobiidae which comprises chiefly species with Gondwanan distribution patterns. Besides Rhodobius, four other monotypic genera represent the subfamily in the Northern Hemisphere, occurring in Vietnam, Japan, Kazakhstan, and Tajikistan (Edgecombe 2003, Farzalieva et al. 2004). Of Central or South American origin are seven species (circa 18%), and one each from North America and islands in Indian Ocean. The sole record of the pantropical geophilomorph centipede Orphnaeus brevilabiatus in Europe comes from Bohuslän, a Swedish province in the northern part of the W coast, where the animal was collected in the 19th century (Andersson et al. 2005). Ten centipedes and millipedes have been considered as cryptogenic (= species of unknown origin which cannot be ascribed as being native or alien). Some of them such as the geophilid Arenophilus peregrinus and the schendylid Nyctunguis persimilis, which have only been reported from the Isles of Scilly, Great Britain and Austria respectively (Barber 2008, Christian 1996) whereas all the other species of these genera live in North America, are of likely Nearctic origins. Another suspected introduction of uncertain origin is Nothogeophilus turki which has hitherto been known only from Scilly and the Isle of Wight, Great Britain (Lewis et al. 1988) and represents a monotypic genus. However, we cannot completely exclude the possibility that some cryptogenic species suspected to be alien are actually native to Europe. Support for this notion we find in the scolopendromorph centipede Theatops erythrocephalus C.L. Koch, 1847, which occurs in various natural habitats (including caves) in the Pyrenees and the western part of the Balkans (with a gap between these geographic areas), while all its other four congeners occur in North America (Minelli 2006). Unknown also is the origin of the symphylid Hanseniella oligomacrochaeta described from a hothouse in the Botanical Garden in Berlin; according to Scheller (2002), all species in the genus Hanseniella have tropical-subtropical distributions. The haplodesmid Prosopodesmus panporus is only known from the Royal Botanic Gardens in Kew, England, while its other described congener, P. jacobsoni Silvestri, 1910, is pantropical (Golovatch et al. 2009). Likewise, it is uncertain whether Napocodesmus endogeus, a millipede described solely from females collected in the garden of Cluj University, is a European native or not. According to Tabacaru et al. (2003), the generic allocation 106 Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010) Figure 7.2.2. Colonization of continental European countries and main European islands by myriapod species alien to Europe. Archipelago: 1 Azores 2 Madeira 3 Canary islands. of the second species described in the genus, N. florentzae Tabacaru, 1975, hitherto known from Romania and Moldova, is not certain and since there are no other records of N. endogeus in nature it might be an introduced species. 7.2.5. Pathways for the introduction of alien myriapod species in Europe All of the alien myriapods have most probably been accidentally introduced to Europe with plant material in relation to human activities and trade between Europe and other continents such as Asia, Australasia and the Americas. This process must have begun with an increase in trade between ancient Greek and Romans with cities in Asia Minor and North Africa and should have intensified in post-medieval times with the trade between Europe and some East Asiatic countries (e.g. Japan, China) and the geographic discoveries of the Americas and, later, of Australia. This process is still going on with Myriapods (Myriapoda). Chapter 7.2 107 Figure 7.2.3. Geographic origin of the myriapod species alien to Europe (in percent). the trade of tropical flowers and other plants and their cultivation in houses and greenhouses or with the importing of goods from tropical countries. Even large species could be transported this way, as is the recent case of the discovery of the largest centipede Scolopendra gigantea, found in 2005 in a house in London, which is thought to have arrived with a cargo of electrical goods or fruit (BBC News 2005). Pocock (1906) suggested the possible countries whence a variety of alien species found in Kew Gardens were introduced with their host plants: India (Scolopendra morsitans, Mecistocephalus guildingii), Sri Lanka (Chondromorpha kelaarti), Barbados (Anadenobolus monilicornis), Saint Vincent Island (A. vincenti). The distribution of the alien diplopods in Europe shows that all the species living here in greenhouses are much more widespread compared to e.g. the restricted outdoor species Eurygyrus ochraceus. It is also likely that the obligate thelytokous parthenogenesis (= sexual reproduction giving rise to females only) shown in continental Europe by several of the exotic millipedes and at least one of the centipedes has facilitated their survival during transport and their establishment on the continent. However, bisexual populations are known from the Azores and the Canary Islands for Lamyctes emarginatus (Enghoff 1975a). Species from other centipede orders, such as the mecistocephalid Tygarrup javanicus also presumably reproduce by parthenogenesis since so far only females have been found in the hothouse at the Eden project, in Great Britain (Barber 2009b). The number of exotic diplopods in Europe is far smaller (3–4 times) than that of European species introduced to other continents. Apparently, this could mean that the arrival and, especially, becoming resident in Europe is much more difficult than the converse process. The asymmetry has probably nothing to do with quarantine controls at European borders. Instead, it may be due to specific ecological and biological patterns exhibited by the successful invaders. Many of the alien millipedes and centipedes which have successfully invaded Europe be- 108 Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010) Figure 7.2.4. Scolopendra gigantea Linnaeus, 1758 [Chilopoda: Scolopendromorpha: Scolopendridae] caught in 2005 in apartment in London, perhaps arrived with a cargo of electric goods or fruit. Source: BBC News: http://news.bbc.co.uk/go/em/fr/-/1/hi/england/london/4201634.stm Figure 7.2.5. Tygarrup javanicus Attems, 1929 [Chilopoda: Geophilomorpha: Mecistocephalidae]. United Kingdom: Eden Project, Cornwall. Credit: Anthony Barber. long to genera moderately rich to rich in species, such as Poratia, Chondrodesmus, Lamyctes, Cryptops, etc. A pertinent question arises as to why often only one species succeeds in establishing populations on foreign continents, sometimes becoming quite widespread to even cosmopolitan, whereas its rather numerous congeners fail to do so. Specific adaptive ecological patterns may be an issue, but, as noticed Myriapods (Myriapoda). Chapter 7.2 109 Figure 7.2.6. Rhinotus purpureus (Pocock, 1894) [Diplopoda: Polyzoniida: Siphonotidae]. Japan: MinamiDaito. Credit: Zoltán Korsós. Figure 7.2.7. Eurygyrus ochraceus C.L. Koch, 1847 [Diplopoda: Callipodida: Schizopetalidae]. Ukraine: Crimea. Credit: Kiril Makarov. above, obligate or opportunist parthenogenesis is probably a major trait favoring dispersal at least because a single founder juvenile or female is sufficient to arrive at destination and found a population. It has to be noted that the successful myriapod invaders tend to be among the smallest species, thus being more easily transported, better fitted to find a suitable microhabitat, and sometimes requiring a shorter time and even a smaller number of developmental stages to reach maturity (Golovatch 2009). 110 Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010) Figure 7.2.8. Chondrodesmus cf. riparius Carl, 1914 [Diplopoda: Polydesmida: Chelodesmidae]. Denmark: Copenhagen. Credit: Gert Brovad. Figure 7.2.9. Oxidus gracilis (C.L. Koch, 1847) [Diplopoda: Polydesmida: Paradoxosomatidae]. Italy: Porto Badino (Borgo Hermada – Terracina). Credit: Massimiliano Di Giovanni. Myriapods (Myriapoda). Chapter 7.2 111 Figure 7.2.10. Paraspirobolus lucifugus (Gervais, 1836) [Diplopoda: Spirobolida: Spirobolellidae]. Japan: Okinawa. Credit: Zoltán Korsós. Another possible pathway of the introduction of exotic myriapods to Europe is their intentional import as ‘pets’, and their further escape from pet keepers. Large Scolopendra spp., as well as some large and colorful millipedes of the orders Spirobolida, Spirostreptida and Sphaerotheriida are quite popular pet animals subjected to trade in pet shops. Although there are many guides and internet resources available for keeping and caring for exotic species, there is no reliable information about the importance of the ‘pet’ trade for the introduction of alien myriapods to Europe. However, the establishment of pet myriapods in the wild is in most cases very unlikely. 7.2.6.The most invaded ecosystems and habitats Man-made artificial environments (pastures and cultivated lands, greenhouses, urban and suburban areas) constitute the main habitat types hosting alien myriapods (Table 7.2.1). Species of tropical and subtropical origin are likely to be restricted to greenhouses or equivalent artificially warmed habitats. Some of them, in the summer season in the southern countries perhaps could survive also outdoors in close proximity to the hothouses. However, 11 species have been reported from natural habitats in Europe, where they most likely were able to establish viable populations. So far the alien species of symphylans and pauropods are unknown in natural areas, which is not the case with several species of the other two myriapod classes. The millipede Oxidus gracilis, which is bisexual everywhere and is naturalized in several areas in Europe and in the Caucasus, has been found in forests close to suburban and urban areas (Tömösváry 1879), in woodlands of Robinia pseudoacacia in the Kanev Nature Reserve, Ukraine 112 Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010) Figure 7.2.11. Trigoniulus corallinus (Gervais, 1847) [Diplopoda: Spirobolida: Trigoniulidae]. Taiwan. Credit: Zoltán Korsós. (Chornyi and Golovatch 1993) and records from caves also exist (Strasser 1974, Vicente and Enghoff 1999). On the Canary Islands the species is quite widespread invading various, mostly dry and warm, habitats (Arndt et al. 2008). According to Palmén (1949), O. gracilis dies when subjected for 2 hours to a temperature of minus 4°C. This means that in North Europe the species can survive only in hothouse conditions. Cylindroiulus truncorum mainly inhabits synanthropic habitats: greenhouses, gardens, parks, woodpiles, school grounds, cemeteries, spoil heaps, horticultural nurseries (Kime 2004, Korsós and Enghoff 1990). Eurygyrus ochraceus occurs in the Crimea only in a patch of semi-natural xerophytic vegetation ca. 1 km long and 100–300 m wide along a watershed. It was reported to be rather common, although not too abundant on the site and is definitely an anthropochore (Golovatch 2008). Lamyctes emarginatus shows remarkable plasticity regarding the surrounding environment, although in the British Isles there is preponderance of rural records in comparison with (sub)urban ones. In artificial habitats it has been reported from gardens, roads, roadside verges, hedges, embankments, crops of Zea mays and Medicago sativa, even in human rubbish (Eason 1964, Minelli and Iovane 1987, Barber and Keay 1988). In natural habitats it lives in various woods (deciduous or mixed coniferous/ deciduous) and has also been recorded from open and coastal areas (Barber and Keay 1988, Zerm 1997, Zapparoli 2006). According to Andersson (2006), it predominates in open and disturbed areas with sparse vegetation. A great many of these localities Myriapods (Myriapoda). Chapter 7.2 113 are associated with lake shores, river gravels or river banks. L. emarginatus shows clear preferences for temporarily flooded sites, no matter for how long the inundation lasts. Its appearance as a pioneer species on mine sites may indicate that the species shows preference to disturbed habitats (Zerm 1997). In close proximity to water pools the species abundance can reach 95% of all centipedes (Minoranskii 1977). Two of the (presumed) alien geophilomorphs, Arenophilus peregrinus and Nothogeophilus turki, have been recorded in coastal areas, where they occur under stones and in soil close to rocky sea cliffs with sparse vegetation although A. peregrinus has been found inland in Cornwall in woodland and one of the Isle of Wight records for Nothogeophilus turki was from an area of demolished buildings with copious rubbish on the ground although no more than 5 m from the tidal river (A.N. Keay pers. comm.). Considerable fluctuation in the abundance of some alien species have been observed by Barber (2009b) in the tropical hothouse of the Eden Project. P. lucifugus which was not found in 2003/4, was rather restricted in its occurrence in 2005, had become abundant throughout by 2009. Likewise, C. doriae which has been relatively uncommon and limited in occurrence in 2005 was the dominant species there in 2009. Conversely, T. javanicus, which had been abundant in 2005, was difficult to find in 2009 (Barber 2009b). 7.2.7. Ecological and economic impact Alien myriapods are unlikely to pose major threats to native biodiversity and ecosystems. The number of species established in the wild being very limited (12 species, ca 30%) for the moment (Table 7.2.1). Diplopods are detrivorous animals, consuming 10–15% of the leaf litter in temperate forest and as thus contribute significantly to soil formation processes through the fragmentation of leaves which stimulates microbial activity. They may thus indirectly influence the fluxes of nutrients (Hopkin and Read 1992). Nevertheless, some alien diplopods could be harmful to cultivated plants, especially in the artificial habitats where temperature and humidity conditions allow species establishment and expansion. Invasive soil invertebrates may also have an impact on the structure and function of natural ecosystems. They can change soil carbon, nitrogen and phosphorus pools and can considerably affect the distribution and function of roots and micro-organisms (Arndt and Perner 2008). In addition, mass occurrences and swarming, which have been observed in several countries in Europe, may have negative ecological and economic impact although the causes still remain obscure (Sahli 1996, Voigtländer 2005). An example of a plant-damaging alien myriapod is Oxidus gracilis, which is regarded as a pest in several European countries. This species is very common in greenhouses where its density may exceed 2500 ind./ m2. It is known for attacking vegetable and fruit crops such as sugar beet, potatoes, strawberries, cucumbers, orchard fruits, roots of wheat, and flowers in outdoor cultivated areas. Furthermore, several thousand O. gracilis were once found after rain in a house in Lenoir City, Tennessee, USA, with most of the city infested during the same outbreak (Hopkin and Read 1992). As a curiosity, one might also mention the report 114 Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010) by the classical writer Theophrastus, according to whom an army of millipedes once overran Rhoeteum in the present province of Çanakkale (northwestern Turkey) and drove its human inhabitants into the sea (Sharples 1994, Enghoff and Kebapći 2008). Several plants can withstand the attacks of symphylans but they may cause severe damage to growing crops both in fields and greenhouses (Scheller 2002). Arndt and Perner (2008) recently carried out a study on the impact of invasive ground-dwelling predatory species, including alien centipedes, in the native laurel forest habitat in the Canary Islands. They found that centipedes in laurel forests seem to be much more variable than carnivorous ground beetles since the 14 recorded species include representatives of three orders with very different characters. They tentatively recognised four functional groups of centipedes: a micro-cephalic schendylid type, (ii) a geophilid type with medium head size and extreme body length, (iii) a scolopendromorph type, and (iv) a macro-cephalic lithobiomorph type. These groups suggest patterns of invasion similar to the coleopteran predators: autochthonous and introduced species of the same size class and group are mutually exclusive (Arndt 2006). The potential role of tropical giant millipedes and centipedes (Scolopendra spp.) kept as pets has been little analyzed as a source of health problems in relation to their defensive fluids or their bites which can cause pathological reactions if exposed to skin, mouth/throat or eyes (Rein 2002). Acknowledgements We thank Helen Read (Farnham Common, UK), John Lewis (Taunton, UK), Greg Edgecombe (London, UK) and Zoltán Korsós (Budapest, Hungary) for their helpful comments and shared literature as a result of which the manuscript was able to be significantly improved. John Lewis and Göran Andersson shared unpublished information on the presence of alien myriapods in UK and Norway, respectively. Darren Mann (Oxford, UK) provided a copy of Pocock’s report on Kew species and the paper by Clarke. 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Zerm M (1997) Distribution and phenology of Lamyctes fulvicornis and other lithobiomorph centipedes in the floodplain of the Lower Oder Valley, Germany (Chilopoda, Henicopidae: Lithobiidae). Entomologica Scandinavica, Supplement 51: 125–132. Table 7.2.1. List and main characteristics of the myriapod species alien to Europe. Status: A Alien to Europe C cryptogenic species. Country codes abbreviations refer to ISO 3166 (see Appendix I). Habitat abbreviations refer to EUNIS (see Appendix II). Only selected references are given. Last update 10/03/2010. Hanseniella oligomacrochaeta Scheller, 2002 Hanseniella orientalis (Hansen, 1903) Chilopoda Geophilomorpha Mecistocephalidae Mecistocephalus guildingii Newport, 1843 Status Native range A Southeast Asia (India, Sri Lanka) A C A A Mecistocephalus maxillaris (Gervais, 1837) C Tygarrup javanicus Atems, 1929 A 1st record in Europe Invaded countries 1958, FR FR, NO Habitat References Andersson et al. (2005) Tropical, subtropical 1903, DK DK, FR, GB, (North America up to MC, NO Mexico, South America, sub-Saharan Africa, Sri Lanka, Galapagos Islands, and possibly New Zealand) Unknown. Tropical, 2000, DE DE subtropical? Tropical, subtropical (South 2000, DE DE and southeastern Asia, Central and South America, islands in the Pacific) J100 Andersson et al. (2005), Scheller (2002), Shear and Peck (1992) J100 Scheller (2002) J100 Scheller (2002) Amphi-Atlantic (coasts of Tropical America, African coast from Gambia to Liberia, Atlantic islands) Unknown, tropical? 1895, DE DE, FR, GB, PTMAD J100 Bonato et al. (2009), Pocock (1906) 1837, FR DK, FR, NL J100 Southeast Asia (Java, Vietnam, Cambodia), The Seychelles 1975, GB AT, GB J100 Andersson et al. (2005), Bonato et al. (2009), Jeekel (1977) Barber (2009b), Christian (1996), Lewis and Rundle (1988) 123 J100 Myriapods (Myriapoda). Chapter 7.2 Class Family Species Order Pauropoda Tetramerocerata Pauropodidae Allopauropus pseudomillotianus Remy & Balland, 1958 Symphyla Symphylomorpha Scutigerellidae Hanseniella caldaria (Hansen, 1903) Family Geophilidae Oryidae Native range 1st record Invaded in Europe countries Unknown, genus present in 1986, GB GB North America Habitat References Arenophilus peregrinus Jones, 1989 C Nothogeophilus turki Lewis, Jones & Keay, 1988 Orphnaeus brevilabiatus (Newport, 1845) C Unknown 1985, GB GB A 19th century, SE Nyctunguis persimilis Attems, 1932 C Tropical, subtropical (Australia, Central and South America, SubSaharan Africa, Madagascar, East Asia, Arabian Peninsula, Hawaii) Unknown. Genus present in North America 1996, AT AT I2? Christian (1996), Christian and Szeptycki (2004), Gruber and Christian (2002) Southeast Asia, Papua New Guinea, The Seychelles Central and South America Tropical, subtropical. North and South America, Atlantic Ocean Islands, Europe, Africa, Arabian Peninsula, Southeast Asia, Indian Ocean Islands, Australia, New Zealand, Pacific Islands East and South Asia 2007, GB GB J100 Barber (2009a), Lewis (2007) 2005, GB GB 1902, GB GB J1 J100 BBC News (2005) Akkari et al. (2008), Pocock (1906) 1902, GB GB J100 Minelli (2006), Pocock (1906) Chilopoda Scolopendromorpha Cryptopidae Cryptops doriae Pocock, 1891 Scolopendridae Status A Scolopendra gigantea Linnaeus, 1758 Scolopendra morsitans Linnaeus, 1758 A A Scolopendra subspinipes Leach, 1815 A SE B3, I2 Barber (2009a), Gregory and Jones (1999), Jones (1989) B3 Barber (2009a), Lewis et al. (1988) Un- Andersson et al. known, (2005) J100? Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010) Schendylidae Species 124 Class Order Class Family Species Order Chilopoda Lithobiomorpha Henicopidae Ghilaroviella cf. valiachmedovi Zalesskaja, 1975 Status A C Lamyctes (Lamyctes) coeculus (Brölemann, 1889) A Lamyctes (Lamyctes) emarginatus (Newport, 1844) A Rhodobius lagoi Silvestri, 1933 C 1st record in Europe Invaded countries Unknown. G. 2004, AT AT valiachmedovi occurs in Central Asia (Tajikistan) Southeast Asia (Java), 1988, ES- ES-CAN Sakhalin Island, CAN Guadeloupe, The Seychelles Habitat I2 References Christian and Szeptycki (2004) H3, H5 Eason and Enghoff (1992), Hollington and Edgecombe (2004) DK, ES-CAN, FI, J100, J Barber (2009a), FR, GB, IT, SE Enghoff (1975a), Zapparoli and Minelli (2005) 1889, IT Tropical, subtropical. Known from Australia, Central and South America, Sub-Saharan Africa, Madagascar Australasia (Australia+ New 1868, DK AT, BE, BG, CZ, DE, DK, ESZealand) is the possible CAN, FI, FR, GB, areas of origin. Known GL, GR, HU, IT, also from North and South LU, NL, NO, PL, America, Africa, Asia PT, PT-AZO, PTMinor, Greenland, Iceland, MAD, RO, RU, New Caledonia, islands in SE, SK, UA the Pacific Unknown, possibly tropical, subtropical. Subfamily Anapsobiinae distributed in South America, South Africa, Australia, Japan, Vietnam, Kazakhstan and Tajikistan 1933, GR-SEG GR-SEG (Rhodes) B1, D, E, F4, F9, G1, G3, J1, J2, J3, J4, J5, J6, I, I1, I2, X6, X7, X23 I? Barber and Keay (1988), Bocher and Enghoff (1984), Meinert (1868), Minelli and Iovane (1987), Negrea (1989), Palmén (1948, 1952), Zapparoli and Minelli (2005) Silvestri (1933), Zapparoli (2002) Myriapods (Myriapoda). Chapter 7.2 Lamyctes (Metalamyctes) albipes (Pocock, 1895) Native range 125 Species Status Native range A Nearctic (USA), Bermuda Islands Diplopoda Polyzoniida Siphonotidae Rhinotus purpureus (Pocock, 1894) A Diplopoda Callipodida Schizopetalidae Eurygyrus ochraceus C.L. Koch, 1847 Diplopoda Polydesmida Chelodesmidae Haplodesmidae Invaded countries Habitat References G Attems (1935), Condé (1961), Vicente and Enghoff (1999) Tropical, subtropical (South 1986, GB GB and Central America, islands in Indian and Pacific oceans) J100 Barber (2010), Read (2008) A Asia (Turkey) 1925, BG BG, UA E1, I2 Golovatch (2008), Stoev (2007), Verhoeff (1926) Chondrodesmus cf. riparius Carl, 1914 A South America 2000, SE DE, DK, NO, SE Cylindrodesmus hirsutus Pocock, 1889 A Prosopodesmus panporus Blower & Rundle, 1980 A Tropical, subtropical (South 1950AT, DE, FR, GB, 1985 HU, SK America, Southeast Asia, Papua New Guinea, islands in Indian and Pacific oceans) Unknown, other species in 1975, GB GB the genus pantropical 1961, PT- ES-CAN, PTMAD MAD J J100 J100 Andersson and Enghoff (2007), Enghoff (2008a) Golovatch and Stoev (2010), Golovatch et al. (2001), Golovatch et al. (2009), Read (2008) Blower and Rundle (1980), Golovatch et al. (2009), Read (2008) Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010) Polyxenus fasciculatus Say, 1821 1st record in Europe 126 Class Family Order Diplopoda Polyxenida Polyxenidae Class Order Family Oniscodesmidae Species Status Native range Amphitomeus attemsi (Schubart, 1934) A South America (Venezuela or Colombia) A Southeast Asia 1st record Invaded Habitat References in Europe countries 1930, DE AT, CH, DE, DK, J100 Barber and Eason GB, HU, NL, (1986), Enghoff PL, SK (1987), Enghoff (2009), Golovatch et al. (2002), Gruber (2002), Korsós et al. (2002) 1906, GB GB J100 Pocock (1906) A India, Sri Lanka 1902, GB GB A Asia (East or Southeast) 1879, HU Pyrgodesmidae Cynedesmus formicola (Cook, 1896) C Unknown, genus native of Central America AT, BE, BG, BY, J, J100, CH, CZ, DE, G DK, ES, ES-BAL, ES-CAN, FI, FR, GB, HU, IE, IS, IT, LT, LU, LV, MC, MD, MK, MT, NL, NO, PL, PT-MAD, PTAZO, RO, RU, SE, SI, SK, UA 1896, ES- ES-CAN, HU, J100 CAN PT-MAD Poratia digitata (Porat, 1889) A Tropical and subtropical (Southern North and Central America) 1889, SE AT, CH, DE, DK, FR, GB, NL, NO, SE J100 J100 Pocock (1906) Blower (1985), Enghoff (2009), Enghoff et al. (2004), Evans (1900), Hoffman (1999), Pocock (1902), Read (2008), Šefrová and Laštůvka (2005), Stoev (2004) Myriapods (Myriapoda). Chapter 7.2 Paradoxosomatidae Asiomorpha coarctata (De Saussure, 1860) Chondromorpha kelaarti (Humbert, 1865) Oxidus gracilis (C.L. Koch, 1847) Attems (1935), Korsós et al. (2002), Vicente and Enghoff (1999) Blower and Rundle (1986), Golovatch and Sierwald (2001), Gruber (2002), Latzel (1895) 127 Family Species Poratia obliterata (Kraus, 1960) Trichopolydesmidae Napocodesmus endogeus Ceuca, 1974 A C Native range 1st record Invaded in Europe countries late DE, FR, HU 1990s, DE Tropical (South and Central America: Peru, Colombia, Brazil, Costa Rica) Unknown, only female/s 1969, RO RO known; the second tentative congener occurs in Romania and Moldova Habitat References J100 Adis et al. (2000), Golovatch and Sierwald (2001), Korsós et al. (2002) Ceuca (1974), Tabacaru et al. (2003) I2? Cylindroiulus truncorum (Silvestri, 1896) A North Africa (Algeria, Tunisia) 1925, DE AT, BE, CH, DE, J, J100, I2 DK, ES-CAN, FI, FR, GB, HU, LT, LU, NL, NO, PL, PT, PT-MAD, RO, SE, UA Enghoff (2009), Kime (2004), Korsós and Enghoff (1990), Read (2008), Schubart (1925) Pseudospirobolellus avernus (Butler, 1876) A 2009, GB GB J100 Barber et al. (2010), Enghoff (2001) Anadenobolus monilicornis (Porat, 1876) Anadenobolus vincenti (Pocock, 1894) A Tropical (Southeast Asia, islands in Indian and Pacific oceans, and Caribbean Sea) Caribbean region 1906, GB GB J100 Spirobolellidae Paraspirobolus lucifugus (Gervais, 1836) Trigoniulidae Trigoniulus corallinus (Gervais, 1847) Hoffman (1999), Pocock (1906) Hoffman (1999), Pocock (1906) Enghoff (1975b), Jeekel (2001), Latzel (1895), Lee (2006), Read (2008) Pocock (1906), Shelley and Lehtinen (1999) Diplopoda Spirobolida Pseudospirobolellidae Rhinocricidae A A Saint Vincent Island, Lesser 1900, GB GB Antilles Tropical. Area of origin 1836, FR DE, DK, GB, NL most likely The Seychelles and/or Mauritius J100 A Southeast Asia J100 1902, GB GB J100 Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010) Diplopoda Julida Julidae Status 128 Class Order Myriapods (Myriapoda). Chapter 7.2 129 Table 7.2.2 List of myriapod species intercepted in Great Britain (Barber 2009a, Clarke 1938, John Lewis, pers. comm., Sharon Reid (FERA), pers. comm.) Species Native Range Class Chilopoda Order Craterostigmomorpha Craterostigmus sp. New Zealand & Tasmania Order Geophilomorpha ?Zelanion (= Steneuryton) sp. Australia, New Zealand, Hawaii Order Scolopendromorpha Scolopendra cingulata Mediterranean Latreille, 1829 region Scolopendra dalmatica C.L. Balkan Koch, 1847 peninsula Scolopendra subspinipes Asia, Africa, subspinipes Leach, 1815 C. & S. America Order Lithobiomorpha Lithobius forficatus Europe (Linneaus, 1758) Lithobius peregrinus Latzel, Europe, 1880 Caucasus Class Diplopoda Order Polydesmida Polydesmida gen. spp. ?Gasterogramma plomleyi Tasmania Mesibov, 2003 ?Mestosoma sp. Akamptogonus novarae ? Australia (Humbert & Saussure, 1869) Habrodesmus falx Cook, West Africa 1896 Habrodesmus sp. ?Oxidus gracilis ?East Asia Oxidus gracilis East Asia Found in/ Country of dispatch/ Year of Interception Dicksonia (Australia or New Zealand, 2008) Dicksonia (Australia, 2005) With luggage (Spain, 2003), potatoes (Greece, 1975), Turkey (2004), Palestine (pre-1992) Found in fruit & vegetable warehouse on Isle of Wight (1983) Trachycarpus wagnerianus (South Korea, 2006), bananas (Jamaica, 1938) Dicksonia (Australia, 2004) Dicksonia (New Zealand, 2004) Dracaena fragans (Belgium, 1979) Dicksonia (Australia, 2004) Bromeliad (Ecuador, 1982) Dicksonia (New Zealand, 2004) Tete leaves (Nigeria, 1981) Orchid (Malawi, 1982) Zelkova (Netherlands, 1995) Aroid (USA,1980), Chaemaerops (Morocco, 2001), Cryptomeria (Japan, 1979), Dracaena (Belgium, 1979), Ficus (West Africa, 1979), Hibiscus (Canary Is.), Lirope (USA, 1999), Orchid (Belize, 1980; Madagascar, 1995; Malaysia,1984; India, 2000), Palm (Canary Is., 1998), Pentas (Canary Is., 2010), Phoenix (USA, 1995), Rhododendron (soil, Nepal, 1981), Sanseviera (USA, 1980), Scindapus (soil, Nepal, 1981), Selaginella (Singapore, 1999; Brazil, 1995), Serissa (China, 1999, 2004), Trachycarpus (Netherlands, 2008), Washingtonia (Italy, 2009), Weeping fig (USA, 1984), Yucca (?Netherlands, 1980), Zamia seed (USA, 1982), Zelkova (China, 1995), unknown (Chile, 1998; South Africa, 2001) 130 Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010) Species Polydesmidae Native Range Polydesmus sp. Order Spirostreptida Spirostreptida Spirostreptus sp. Plusioglyphiulus sp. Order Julida Blaniulidae Blaniulus guttulatus (Fabricus, 1798) Blaniulus sp. Cylindroiulus londinensis (Leach, 1814) Cylindroiulus sp. Ommatoiulus moreletii (Lucas, 1860) Ommatoiulus oxypygus (Brandt, 1841) Ophyiulus targionii Silvestri, 1898 Found in/ Country of dispatch/ Year of Interception Dicksonia (Australia, 2005; New Zealand, 2004), Orchid (Malaysia, 1983), Wild Plant (South Africa, 1983) Miscanthus (Dominica, 2000), Orchid (Australia, 1985) Cyathea (New Zealand, 2005), Dicksonia (Australia, 2004–2008), Dracaena (Rwanda, 1980) Fig (Ivory Coast, 1983) Orchids & Rhododendrons (Borneo, 1979) Europe Echinodorus (Singapore, 2008), Orchid (Brazil, 2003) Orchid (Australia, 1985) Europe Unknown (South Africa, 1999) Phoenix dactylifera (Italy, 2004) Iberian peninsula Italy Dicksonia (New Zealand, 2004) Dicksonia (Australia, 2006), melon fruit (South Africa, 1983) Vitis sp. (Italy, 1979) Italy Unknown (New Zealand, 1982) Table 7.2.3. Relative importance of the non-native species in the myriapod fauna of the Macaronesian islands. The numbers of introduced species correspond to the total non-native species of both exotic and continental European origin (cf., Arndt et al. 2008, Baéz and Oromí 2004, Borges, 2008a,b, Borges and Enghoff 2005, Enghoff 2008b, Enghoff and Borges 2005, Zapparoli and Oromi 2004), some numbers updated according to recent records. * 7 certainly native, 6 probably native, 20 possibly native, ** all probably introduced; *** all possibly native. Diplopoda Chilopoda Symphyla Pauropoda Canary Isl. Azores Isl. Madeira Is. Selvages Isl. Native Introduced Native Introduced Native Introduced Native Introduced 83 24 2 19 40 18 2 0 33* 2** 8 3 2 17+2? 0 2 0 6** 3 0 1 2 no no records records 14*** 0 1 0 10 0 no no records records A peer reviewed open access journal BioRisk 4(1): 131–147 (2010) doi: 10.3897/biorisk.4.48 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Spiders (Araneae) Chapter 7.3 Wolfgang Nentwig, Manuel Kobelt Community Ecology, Institute of Ecology and Evolution, University of Bern, Baltzerstrasse 6, CH-3012 Bern, Switzerland Corresponding author: Wolfgang Nentwig (wolfgang.nentwig@iee.unibe.ch) Academic editor: Alain Roques | Received 27 January 2010 | Accepted 20 May 2010 | Published 6 July 2010 Citation: Nentwig W, Kobelt M (2010) Spiders (Araneae). Chapter 7.3. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 131–147. doi: 10.3897/biorisk.4.48 Abstract A total of 47 spider species are alien to Europe; this corresponds to 1.3 % of the native spider fauna. They belong to (in order of decreasing abundance) Theridiidae (10 species), Pholcidae (7 species), Sparassidae, Salticidae, Linyphiidae, Oonopidae (4–5 species each) and 11 further families. There is a remarkable increase of new records in the last years and the arrival of one new species for Europe per year has been predicted for the next decades. One third of alien spiders have an Asian origin, one fifth comes from North America and Africa each. 45 % of species may originate from temperate habitats and 55 % from tropical habitats. In the past banana or other fruit shipments were an important pathway of introduction; today potted plants and probably container shipments in general are more important. Most alien spiders established in and around human buildings, only few species established in natural sites. No environmental impact of alien species is known so far, but some alien species are theoretically dangerous to humans. Keywords Buildings, urban area, greenhouse, pathways, venomous spiders, Europe, alien 7.3.1 Introduction Spiders are among the most diverse orders in arthropods with a world-wide distribution in all terrestrial habitats and more than 40,000 species, grouped in 109 families (Platnick 2008). The European spider fauna comprises nearly 3600 species of which 47 (= 1.3 %) are alien to Europe, i.e. their area of origin is outside Europe. An ad- Copyright W. Nentwig, M. Kobelt. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 132 Wolfgang Nentwig & Manuel Kobelt / BioRisk 4(1): 131–147 (2010) ditional number of at least 50–100 species are alien within Europe, i.e. they originate, e.g., from the Mediterranean or from eastern parts of Europe and spread gradually into other parts of Europe. Such aliens within Europe are not considered here. Small scale spread, e.g., into an adjacent country, is also not considered here. All spiders are predators and usually prey on arthropods, mainly insects. Since many insects are regarded as pests, spiders are often seen as beneficial. Spiders have unique features such as abdominal silk glands which are used in many ways (e.g., construction of retreat, cocoon, web or dragline) and venom glands to poison their prey (only two families deviate from this). Spiders developed many different ways to catch their prey. Roughly half of them build silken webs to subdue prey and they evolved a large variety of web types. Funnel webs are usually soil-born and closely connected to the retreat of the spider (such as Agelenidae and Amaurobiidae), sheet webs are more often found within the vegetation (examples are Linyphiidae and Theridiidae) and orb webs often bridge the open space between structures (Araneidae and Tetragnathidae). Spiders which do not build a web live as sit-and-wait predators (e.g., Clubionidae, Gnaphosidae, Lycosidae, Sparassidae, and Thomisidae) or actively hunt for prey (such as Salticidae). For this compilation of alien spiders to Europe the DAISIE database (www.europe-aliens.org) was used. In addition a variety of further sources (cited below) was consulted. When speaking about alien species two main problems occur. (1) It may be unclear whether a species is native to Europe or not, e.g., because it is native in an area close to the European borders. This concerns primarily Mediterranean and North or East Palearctic species. We choose a very conservative attitude and did not consider such species. It may also be difficult to decide whether a Holarctic species originates in the Nearctic or in the Palearctic part of the Arctic. We tried to follow the most probable decision. (2) We included only established alien species. In some cases it may be difficult to decide on this because sometimes the discovery of an alien species is communicated but no follow-up reports on its establishment are available. Again, we tried to achieve the most probable point of view. For example, all the reports on tropical Ctenidae or Theraphosidae arriving with banana shipments in Europe never lead to an established population of these spiders and were therefore not included into our chapter. 7.3.2 Taxonomy of alien species The 47 spider species alien to Europe belong to 17 families (Table 7.3.1) with Theridiidae (10 species) and Pholcidae (7 species) being the most species-rich families. Sparassidae comprise five species; Salticidae, Linyphiidae and Oonopidae comprise four species each. Eleven families are represented with only one or two species each. The most astonishing aspect of the composition of the alien spider fauna is that it neither reflects the structure of the global spider community nor the structure of the European spider fauna (Fig. 7.3.1). Spiders (Araneae). Chapter 7.3 133 Globally frequent families (such as Araneidae, Corinnidae, Lycosidae, Theraphosidae, and Zodariidae) are not represented at all among the alien species in Europe. This may be due to some specialisations or restrictions of most species in these families: Araneidae and Corinnidae are usually not associated with human infrastructure and have a rather low probability of becoming transported to foreign areas (see below). Most Theraphosidae (“tarantulas”) depend on their specific microclimate and are among the largest spiders, thus easy to detect and avoid. Lycosidae were also not imported to Europe and the reason for this remains unknown. Other families are overrepresented among the alien community: Sicariidae, Oonopidae, Sparassidae, Pholcidae, and Theridiidae. Their common feature is a preadaptation to human infrastructure, especially buildings. Many species from these families initially live on bark and rocks and/or in arid habitats, thus, they tolerate the dry climate in houses and in urban areas. They can easily sit at the vertical sides of containers (Sparassidae), hide at the underside of pallets or in cracks and cavities (Pholcidae, Theridiidae) or are simply so tiny that they fit everywhere in (Oonopidae). The composition of the spider fauna in Europe will become strongly influenced by alien newcomers if the trend of the last decades continues. Eresidae, Prodidomidae, Scytodidae, and Oonopidae were so far rare families in Europe. Sparassidae and Pholcidae comprise only a few species and the alien add-on may lead to a situation where some families are dominated by alien species. Sicariidae did not even occur previously in Europe. 7.3.3 Temporal trends In the past, there was hardly any systematic check for alien spiders in imported goods. In contrast to herbivores where damage to plants may be of economic importance, alien spiders were only occasionally recorded. Exceptions may be border controls of banana shipments and similar goods because such transports enabled large and dangerous animals to enter Europe. In general, information on arrival data of alien spiders is scarce and when using the date of a scientific publication as a proxy, this information may be considerably fuzzy because some publications compile data of a long period; e.g., for 26 years in Van Keer (2007). 12 first species records were collected in the 19th century, 24 records came from the th 20 century and already 11 records were perceived in the first years of the 21st century. This in itself indicates a steep increase in recording alien species. Of course, it should not be overlooked that the public awareness of alien species and the number of experts increased in the last decades considerably. Both accelerate the probability of detecting new spider introductions. Kobelt and Nentwig (2008) analysed the arrival of 87 alien spider species with known arrival date (alien to Europe and alien within Europe) and concluded that the known number of alien spider introductions still represents an underestimation. They predict a continuous trend of more alien species and give the figure of at least one additional alien spider species annually arriving in Europe in the near future. 134 Wolfgang Nentwig & Manuel Kobelt / BioRisk 4(1): 131–147 (2010) 7.3.4 Biogeographic patterns One third of all alien spiders have an Asian origin. This may include Eastern Palearctic and Indo-Malayan, thus temperate and tropical areas. About one fifth of the species derive from North America and Africa each, and South America and Australia contribute only four species each. In a few cases the origin is not known or subjected to expert guess (Fig. 7.3.2). Such cosmopolitan species are not truly cosmopolitan because they have of course a defined area of origin, but due to early spread among many or all continents and due to lacking phylogeographical information, it is sometimes still impossible to solve such a puzzle. These results suggest that the closer a continent is (Palaearctic) and the more traffic and goods exchange exists (Asia, North America), the more alien species are also imported. An analysis between temperate and tropical origins indicates that about 45 % of species may originate from temperate habitats and 55 % from tropical habitats. Uncertainty, however, is high because for many species nothing or not very much is known about the natural environment in which they live in their area of origin. 7.3.5 Main pathways to Europe Kobelt and Nentwig (2008) analysed the origin of alien spider species in Europe and the intensity of trade between Europe and the native area of these alien spiders in a continent by continent comparison. By including trade volume, area size, and geographical distance, they clearly could demonstrate that trade volume, size of the area of origin, and the geographical distance to Europe are good indicators for the number of alien species transported to Europe. The volume per time curves of agricultural products and mining products fit the increase of alien spiders less well than the curve for manufactures, and therefore it is concluded that the first have a lower number of alien stowaways whereas manufactures have the highest potential to transport alien species (Fig. 7.3.3). More in detail, spiders can survive shipment in or at containers or construction materials for periods long enough to reach most other continents. The rare collection notes on spiders which had been recorded during or after this voyage suggest that spiders frequently occur in container (e.g., with stones, wood, other products), in or at wooden boxes, at wooden pallets, and within shipments of logs or wood products. Consequently, many alien spiders are detected in a harbour, in buildings at or close to a harbour, and in or at warehouses (Van Keer 2007). Up to the 1980s, many alien spiders were detected in banana or other fruit shipments (Forsyth 1962, Reed and Newland 2002). This does not only represent a pathway from a tropical area of origin to Europe, it also enables the spider to travel within Europe. With increasing technical standards to supply the fruits with optimal transport conditions (usually low temperature, oxygen reduction to 1–5 % and a carbon dioxide increase to 1–10 %, see also Hallman (2007)), spiders have less chances to survive this (but see Craemer 2006). Spiders (Araneae). Chapter 7.3 135 Figure 7.3.1 Taxonomic overview of the spider species alien to Europe compared to the native European fauna. Right- Relative importance of the spider families in the alien fauna expressed as the percentage of species in the family compared to the total number of alien spiders in Europe. Families are presented in a decreasing order based on the number of alien species. The number over each bar indicates the total number of alien species observed per family. Left- Relative importance of each family in the native European fauna of spiders and in the world fauna expressed as the percentage of species in the family compared to the total number of spiders in the corresponding area. The number over each bar indicates the total number of species observed per family in Europe and in the world, respectively. Transported plants represent a very important pathway for spiders. This hardly concerns cut flowers but potted plants and plants for planting. There are numerous anecdotes that plants bought in supermarket, in a plant shop or at a plant fair contained a spider or a spider cocoon. Since a considerable amount of such potted plants is produced in China and transported through Italy to different European countries, this indicated the importance of plants as pathway from Asia to Europe. For the further spread of alien spiders within Europe, it is assumed that transport vehicles such as trucks or trains play an important role. The spread of Zodarion rubrum, formerly only known from the French Pyrenees, followed in the last 100 years the main railway connections within Europe. This allowed the small spider to hitchhike over large distances (Pekár 2002). Hänggi and Bolzern (2006) discuss this phenomenon and give evidence for additional species. Spread by vehicles also may explain the fact that quite often the first record of an alien spider had been made at roadsides or in drains along roadsides (Van Keer 2007). 136 Wolfgang Nentwig & Manuel Kobelt / BioRisk 4(1): 131–147 (2010) Figure 7.3.2 Geographic origin of the 47 spider species alien to Europe. In a country-wise comparison within Europe, France, Belgium, The Netherlands, Germany and Switzerland possess the highest numbers of alien spider species (Fig. 7.3.4). These countries are also the ones with the highest level of imports (Fig. 7.3.5). On the other side, the Balkan countries have much lower numbers of alien spiders and Norway, the Baltic States, Belarus, and Russia have the lowest numbers of alien spiders. There is a good correlation between this type of economic activity and the number of alien species, thus, on the country level a comparable picture to the continental level of Kobelt and Nentwig (Kobelt and Nentwig 2008) is obtained. 7.3.6 Most invaded ecosystems and habitats Nearly half of all alien spider species occur only in buildings and/or urban areas. This may be species which inhabit walls of buildings or need the specific microclimatic conditions of houses. One third of all alien species live in greenhouses, botanical gardens, in zoo buildings, or in comparably warm buildings. They rely on the specific temperature conditions but nevertheless are able to establish permanent populations (Holzapfel 1932, Van Keer 2007). In the summer season, in southern countries and under the conditions of climate change some species can colonise the vicinity of buildings and have the potential of further spread. Only five among 47 alien spiders so far were able to establish in natural habitats. They usually are small-sized species, belonging to families which are common in Europe (Dictynidae, Linyphiidae, Tetragnathidae), and they build sheet webs or small orb webs. They originate from North America, Japan and the temperate part of Australia or New Zealand. These parameters probably indicate the conditions which an alien spider should fulfil to be able to survive in natural habitats in Europe. Spiders (Araneae). Chapter 7.3 137 Figure 7.3.3 Increase in global trade (left scale) and the cumulative number of alien spider species introductions (right scale) during the last 50 years. Only cases with known year of introduction are included - from Kobelt and Nentwig (2008). An interesting reason for the obvious high establishment success of alien spiders in human buildings may be found in the rarity of native species at such conditions. This could mean that alien species have much better chances to establish in habitats with no competition by native species. 7.3.7 Ecological and economic impact A family-wise comparison of body sizes of alien and European spider species showed that alien Theridiidae imported to Europe were significantly larger than the native species, Pholcidae and Salticidae showed a trend into the same direction. Kobelt and Nentwig (2008) argue that this reflects the physical transport conditions, especially of temperature and humidity inside a standard ship container (Diepenbrock and Schieder 2006, Naber et al. 2006). These are important stress factors which primarily affect small specimen and can be more easily compensated by large spiders (Pulz 1987). So, even if spiders of all body sizes and from all continents would have more or less equal possibilities to be shipped around the globe, larger species have better chances to survive transportation than smaller ones do. 138 Wolfgang Nentwig & Manuel Kobelt / BioRisk 4(1): 131–147 (2010) Figure 7.3.4 Number of alien spider species for each European country. If alien species could successfully invade European spider assemblages in natural habitats, it could be argued that due to their slightly larger body size they could compete with native species and suppress or even replace them. This would change the dominance structure in natural spider communities within a few years. So far, however, most alien species do not occur in natural spider communities and / or remained rare. Therefore, in Europe no influence of alien spider species on native spiders had been observed so far. This is in agreement with a two-year-analysis of spider communities in California were the occurrence of alien spider species did not negatively affect native species. The most productive habitats contained both the highest proportion of alien and the greatest number of native spiders. No negative associations between native and alien spiders could be detected and, thus, Burger et al. (2001) concluded that the alien spiders do not impact native ground-dwelling spiders. The most frequently occurring alien spider in Europe is probably the North American linyphiid Mermessus trilobatus, first detected in southern Germany in the 1980s and spreading since then. Only in the last years it had been detected that it obviously easily Spiders (Araneae). Chapter 7.3 139 Figure 7.3.5 Relationship between the number of alien spider species and the value of imported goods in European countries (economic data for 2005). establishes in many natural spider communities, especially in grassland and ruderal habitats (Schmidt et al. 2008). With an average body length of 1.6–2.1 mm (Nentwig et al. 2003), M. trilobatus belongs to the smaller linyphiids and it is unlikely that it outcompetes a native species. Competition experiments indeed proved that the invasion success of M. trilobatus is not facilitated by strong competitiveness. Actually it is unknown if other traits (e.g., higher reproduction effort, better dispersal abilities, or nutritional aspects) give some competitive advantage over native species (Eichenberger et al. 2009). So far, the integration success of M. trilobatus into native spider communities seems to confirm the assumption of Burger et al. 2001 on the resilience of native spider communities. An economic impact of spiders may be expected from those spider species which are venomous to humans. Among the alien spiders listed here (Table 7.3.1) species which may be considered as theoretically dangerous to humans comprise the sicariids Loxosceles laeta and L. rufescens and the Australian black widow Latrodectus hasselti (Forster 1984). We are, however, not aware of any report from Europe referring to bites from these species. This is in line with the general assumption that the frequency of spider bites is overestimated (Vetter et al. 2003). Additionally it may be possible that these alien species did not reach relevant densities or that they even did not establish in the long term. Spiders are also known to pollute the faces of buildings and the interior of rooms by their silk spinning activity. Spider webs often stay for long, collect dust and dirt, and are the reason for additional cleaning procedures which cause costs for hygienic reasons. There are only very few reports on this and they only refer to the Mediterranean dictynid spider Dictyna civica spreading since more than 50 years in Central Europe (Billaudelle 1957, Hertel 1968) which occasionally colonises the outside surface of buildings in high densities. Also many native species live inside buildings and cause 140 Wolfgang Nentwig & Manuel Kobelt / BioRisk 4(1): 131–147 (2010) a b c d e f Figure 7.3.6. Alien spiders. a Cicurina japonica female (Dictynidae) b Ostearius melanopygius female (Linyphiidae) c Crossopriza lyoni female with eggsac (Pholcidae) d Spermophora senoculata male (Pholcidae) e Plexipus paykulli female (Salticidae) f Loxosceles rufescens female (Sicariidae). Reprinted with kind permission of Jǿrgen Lissner (© Jǿrgen Lissner, http://www.jorgenlissner.dk). regular cleaning activities due to their web spinning activity but no report concerns additional cleaning costs. 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Biodiversity and Conservation DOI 10.1007/s10531–008–9412–6 Slawson C (2000) Staffordshire biodiversity action plan, Araneae (Arachnida) – spiders. 2nd edn. http://www.sbap.org.uk/focal/checklst/araneae.htm Terhivuo J (1993) Novelties to the Finnish spider fauna (Araneae) and notes on species having bitten man. Memoranda Societatis Fauna Flora Fennica 69: 53–56. Valesova-Zdarkova E (1966) Synanthrope Spinnen in der Tschechoslowakei. Senckenbergiana Biologica 47: 73–75. Van Keer K (2007) Exotic spiders (Araneae): Verified reports from Belgium of exported species (1976–2006) and some notes on apparent neozoan invasive species. Nieuwsbrief Belgische Arachnologische Vereniging 22: 45–54. Vetter RS, Cushing PE, Crawford RL, Royce LA (2003) Diagnoses of brown recluse spider bites (loxoscelism) greatly outnumber actual verifications of the spider in four western American states. Toxicon 42: 413–418. Wunderlich J, Hänggi A (2005) Cicurina japonica (Araneae: Dictynidae) eine nach Mitteleuropa eingeschleppte Kräuselspinnenart. Arachnologische Mitteilungen 29: 20–24. 144 Wolfgang Nentwig & Manuel Kobelt / BioRisk 4(1): 131–147 (2010) Table 7.3.1 List and main characteristics of the spider species alien to Europe. Area of origin: since the area of origin is quite often not well known, this refers to the most probable origin. “cosmopolitan” means that the area of origin is outside Europe but not known, “cosmopolitan” in brackets gives an alternative explanation, South America refers to the tropical part of America. Country codes abbreviations refer to ISO 3166 (see appendix I). Only selected references are given. Last update 30.09.2008. Family Species Amaurobiidae Amaurobius similis (Blackwall 1861) Clubionidae Clubiona facilis O. P.-Cambridge 1910 Dictynidae Cicurina japonica (Simon 1886) Area of origin First Invaded countries record in Europe Habitats Refs AD, BE, CH, DK, J1 DE, ES, FR, GB, IE, MD, NL, NO, PL, RO, SE, UA Fauna Europaea (2005), Harvey (2002), Sacher (1983), Jonsson pers. comm. (2005), Scharff pers. comm. (2005) Australia 1932, GB GB U Fauna Europaea (2005), Platnick (2008) Asia 1990, DE DE, CH, DK E, F, G, Blick and Hänggi (2003), H, I Wunderlich and Hänggi (2005) North America (cosmopolitan) 1915, DK Dysderidae Dysdera aculeata Kroneberg 1875 Eresidae Seothyra perelegans Simon 1906 Gnaphosidae Sosticus loricatus (L. Koch 1866) Asia 1988 HR HR U Deeleman-Reinhold and Deeleman (1988) Africa 1906 FR U Fauna Europaea (2005) Asia Fauna Europaea (2005), Sacher (1983), Terhivuo (1993), Pekar pers. comm. (2005) Zelotes puritanus Chamberlin 1922 North America 1879, SK AT, BG, BY, CS, J1 CZ, DE, EE, FI, FR, GR, HU, IT, LV, LT, MK, PL, RO, RU, SK 1966, CZ AT, CH, CR, CZ , J1 DE, LI, NO, RU, SE, SK Linyphiidae Erigone autumnalis Emerton 1882 Mermessus denticulatus (Banks, 1898) (=Eperigone eschatologica) Mermessus trilobatus (Emerton 1882) Ostearius melanopygius (O. P.-Cambridge 1879) North America North America FR 1990, CH, IT CH 1995, BE BE, CH, DE, ES, NL 1980, North America DE Australia 1906, GB AT, BE, CH, DE, IT, PL AT, BE, BG, CH, CZ, DE, DK, ES, FR, FI, GB, IT, NL, PT, PL, RO, SE, SK Fauna Europaea (2005), Komposch (2002), Pekar pers. comm. (2005) E, F, G, H, I J1, J2.43 Blick and Hänggi (2003), Fauna Europaea (2005) Blick (2004), Blick and Hänggi (2003), Fauna Europaea (2005) E, F, G, H, I E, F, G, H, I Blick and Hänggi (2003), Fauna Europaea (2005) Blick and Hänggi (2003), Fauna Europaea (2005), Komposch (2002), Ruzicka (1995), Pekar pers. comm. (2005), Scharff pers. comm. (2005) Spiders (Araneae). Chapter 7.3 Family Species Area of origin Oonopidae Diblemma donisthor- Asia pei O. P.-Cambridge 1908 Ischnothyreus lymAsia phaseus Simon 1893 Ischnothyreus velox Asia Jackson 1908 Triaeris stenaspis Simon 1891 North America Pholcidae Artema atlanta Wal- Africa ckenaer 1837 Crossopriza lyoni (Blackwall 1867) Micropholcus fauroti (Simon 1887) First Invaded countries record in Europe 1914, GB Habitats Refs GB J1 Platnick (2008), Saaristo (2003) 2005, FR FR U Fauna Europaea (2005) 2003, DE DE, GB, NL J2.43 2001 BE BE, GB, GR J1 Blick (2004), Fauna Europaea (2005), Saaristo (2003) 1896, FR BE, FI, FR, IE, SK J1, J100 Blick (2004), Fauna Europaea (2005), Holzapfel (1932), Koponen (1997), Van Keer (2007), Pekar pers. comm. (2005) Africa 2004, BE Africa 2001, BE Asia 1859, CZ Pholcus phalangioides Asia (Fuesslin 1775) 1857, SK Smeringopus pallidus Africa (Blackwall 1858) Spermophora senocu- Africa lata (Dugès 1836) 2004, NL Pholcus opilionoides (Schrank 1781) 145 1976, SK Blick (2004), Blick and Hänggi (2003), Fauna Europaea (2005), Lee (2005), Platnick (2008), Van Keer (2007) BE E, F, G, Blick (2004), Van Keer H, I, J1 (2007) BE, CH J1 Blick (2004), Blick and Hänggi (2003), Platnick (2008), Van Keer (2007) AD, AT, BG, CH, J1 Fauna Europaea (2005), CS, CZ , DE, ES, Sacher (1983), Pekar pers. FR, GR, HR, HU, comm. (2005) IT, LI, LU, MD, MK, MT, PL, PT, RO, RU, SK, UA AT, BE, BG, BY, J1 Fauna Europaea (2005), CH, CS, CZ , DE, Holzapfel (1932), KomDK, ES, FI , FR, posch (2002), Sacher GB, GR, HU, IE, (1983), Terhivuo (1993), IS, IT, LI, LT, LU, Valesova-Zdarkova MD, MK, MT, (1966), Jonsson pers. NO, NL, PL, PT, comm. (2005), Pekar RO, RU, SE, SK, pers. comm. (2005), UA Scharff pers. comm. (2005) NL J1, Blick (2004) J2.43 BE, BG, CH, CS, J1, J100 Blick (2004), Fauna ES, FR, GR, HR, Europaea (2005), PlatIT, MK, MT, PT, nick (2008), Pekar pers. SI, SK, UA comm. (2005) 146 Wolfgang Nentwig & Manuel Kobelt / BioRisk 4(1): 131–147 (2010) Family Species Area of origin Prodidomidae Zimiris doriai Simon Asia 1882 Salticidae Hasarius adansoni Africa (Audouin 1826) Menemerus bivittatus (Dufour 1831) Panysinus nicholsoni (O. P.-Cambridge 1899) Plexippus paykulli (Audouin 1826) Scytodidae Scytodes venusta (Thorell 1890) Sicariidae Loxosceles laeta (Nicolet 1849) Loxosceles rufescens (Dufour 1820) Africa 2005, DE DE Habitats J1 1901, FR BE, CH, CZ, DE, J2.43 DK, ES, FR, GR, IE, IT, MT, NL, PL Refs Jäger (2005) Blick and Hänggi (2003), Bosmans and Vanuytven (2002), Fauna Europaea (2005), Hänggi (2003), Holzapfel (1932), Pekar pers. comm. (2005), Scharff pers. comm. (2005) Fauna Europaea (2005), Montardi (2006) Fauna Europaea (2005) 1831, ES CZ, ES, FR, GB, IT, PT 2005, FR FR J1 Asia 1819, FR ES, FR, GB, GR, IT, MT J1 Fauna Europaea (2005), Montardi (2006) Asia 2004, NL NL J1 Blick (2004), Fauna Europaea (2005), Platnick (2008), Pekar pers. comm. (2005) South America North America (cosmopolitan) 1963, FI J1 Asia Sparassidae Barylestis scutatus Africa (Pocock 1903) Barylestis variatus Africa (Pocock 1899) Heteropoda venatoria Asia (Linnaeus 1767) Olios sanctivincentii (Simon 1897) Tychicus longipes (Walckenaer 1837) First Invaded countries record in Europe FI, IT J1 1820, ES ES, FR, GR, HR, IT, NL, MT, PT J1, J2.43 Fauna Europaea (2005), Huhta (1972) Blick (2004), Fauna Europaea (2005) 1961, IE IE J1 Forsyth (1962) 1961, IE GB, IE J1 Forsyth (1962), Slawson (2000) Blick and Hänggi (2003), Fauna Europaea (2005), Hänggi (2003), Ruzicka (1995), Valesova-Zdarkova (1966), Ruzicka pers. comm. (2005), Scharff pers. comm. (2005) Forsyth (1962), Slawson (2000) Platnick (2008) 1960, CZ CH, CZ, DE, DK, J2.43 ES, NL, NO, PL Asia 1961, IE GB, IE Asia 1837, NL NL J1 J2.43 Spiders (Araneae). Chapter 7.3 Family Species Tetragnathidae Tetragatha shoshone (Levi 1981) Theridiidae Achaearanea tabulata Levi 1980 Achaearanea acoreensis (Berland 1932) Achaearanea tepidariorum (C.L. Koch 1841) Area of origin First Invaded countries record in Europe North America 1992, DE South America North America South America (cosmopolitan) 1991, AT AT, CH, DE, PL, RU, BG, UA 2002, BE BE 147 Habitats Refs AT, CZ, DE, HU, E, F, G, Fauna Europaea (2005) MK, RO, SK H, I J1 J1, J2.43 J1 1867, AT AT, BE, BG, CH, CZ, DE, DK, ES, FI, FR, GB, GR, HU, HR, IE, IS, IT, LV, LI, MK, MT, NL, NO, PL, PT, RO, RU, SK, SE, UA Achaearanea verucu- Australia 1885, BE BE, GB J1, lata (Urquhart J2.43 1885) Chrysso spiniventris Asia 1949, NL NL J2.43 (O. P.-Cambridge 1869) Coleosoma floridaAsia 1981, AT, CH, DE, FI, J1, num Banks 1900 GB GB, NL J2.43 Latrodectus hasselti Thorell 1870 Australia 2001, BE BE, DK Nesticodes rufipes (Lucas 1846) Steatoda grossa (C.L. Koch 1838) South America Cosmopolitan Steatoda triangulosa (Walckenaer 1802) Cosmopolitan Thomisidae Bassaniana versicolor Keyserling 1880 North 1932, FR FR America J2.43 1996, AT AT, BE, CZ, ES, J2.43 MT, PT 1850, SE AT, BE, BG, BY, J1 CS, CZ, DE, DK, EE, ES, FI, FR, GB, GR, HU, IE, IT, LT, LV, MD, MK, MT, NL, PL, PT, RO, RU, SE, SI, SK , UA 1852, AT AD, AT, BE, BG, J1 CH, CS, CZ, DE, ES, FR, GB, GR, HR, HU, LV, MK, MT, NL, PT, RO, RU, SI, SK, UA U Blick and Hänggi (2003), Fauna Europaea (2005) Van Keer (2007) Fauna Europaea (2005), Komposch (2002), Sacher (1983), Valesova-Zdarkova (1966), Koponen pers. comm. (2005), Pekar pers. comm. (2005), Scharff pers. comm. (2005) Blick (2004), Platnick (2008), Van Keer (2007) Blick (2004) Blick (2004), Blick and Hänggi (2003), Fauna Europaea (2005), Hänggi (2003), Harvey (2002), Komposch (2002) Blick (2004), Platnick (2008), Scharff pers. comm. (2005) Blick (2004), Komposch (2002), Van Keer (2007) Fauna Europaea (2005), Komposch (2002), Sacher (1983), Valesova-Zdarkova (1966), Jonsson pers. comm. (2005), Pekar pers. comm. (2005), Scharff pers. comm. (2005) Fauna Europaea (2005), Harvey (2002), Komposch (2002), ValesovaZdarkova (1966), Scharff pers. comm. (2005) Fauna Europaea (2005) A peer reviewed open access journal BioRisk 4(1): 149–192 (2010) doi: 10.3897/biorisk.4.58 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Mites and ticks (Acari) Chapter 7.4 Maria Navajas1, Alain Migeon1, Agustin Estrada-Peña2, Anne-Catherine Mailleux3, Pablo Servigne4, Radmila Petanović5 1 Institut National de la Recherche Agronomique, UMR CBGP (INRA/IRD/Cirad/Montpellier SupAgro), Campus International de Baillarguet, CS 30016, F-34988 Montferrier sur Lez, cedex, France 2 Faculty of Veterinary Medicine, Department of Parasitology, Miguel Servet 177, 50013-Zaragoza, Spain 3 Université catholique de Louvain, Unité d’écologie et de biogéographie, local B165.10, Croix du Sud, 4-5 (Bâtiment Carnoy), B-1348 Louvain-La-Neuve, Belgium 4 Service d’Ecologie Sociale, Université libre de Bruxelles, CP231, Avenue F. D. Roosevelt, 50, B-1050 Brussels, Belgium 5 Department of Entomology and Agricultural Zoology, Faculty of Agriculture University of Belgrade, Nemanjina 6, Belgrade-Zemun,11080 Serbia Corresponding author: Maria Navajas (navajas@supagro.inra.fr) Academic editor: David Roy | Received 4 February 2010 | Accepted 21 May 2010 | Published 6 July 2010 Citation: Navajas M et al. (2010) Mites and ticks (Acari). Chapter 7.4. In: Roques A et al. (Eds) Arthropod invasions in Europe. BioRisk 4(1): 149–192. doi: 10.3897/biorisk.4.58 Abstract The inventory of the alien Acari of Europe includes 96 species alien to Europe and 5 cryptogenic species. Among the alien species, 87 are mites and 9 tick species. Besides ticks which are obligate ectoparasites, 14 mite species belong to the parasitic/predator regime. Among these species, some invaded Europe with rodents (8 spp.) and others are parasitic to birds (2 spp). The remaining 77 mite species are all phytophagous and among these 40% belong to the Eriophyidae (37 spp.) and 29% to the Tetranychidae (27 spp.) families. These two families include the most significant agricultural pest. The rate of introductions has exponentially increased within the 20th century, the amplification of plant trade and agricultural commodities movements being the major invasion pathways. Most of the alien mite species (52%) are from North America, Asia (25%), and Central and South America (10%). Half of the ticks (4 spp.) alien to Europe originated from Africa. Most of the mite species are inconspicuous and data regarding invasive species and distribution range is only partially available. More research is needed for a better understanding of the ecological and economic effects of introduced Acari. Keywords Europe, alien, mite, tick, Acari, Eriophyidae, Tetranychidae, biological control, Tetranychus evansi, Oligonychus perseae, Polyphagotarsonemus latus, Brevipalpus californicus, Aceria sheldoni, Aculops pelekassi, Dermatophagoides evansi, Varroa destructor Copyright M. Navajas et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 150 Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) 7.4.1. Introduction The subclass Acari, which includes mites and ticks, forms an important part of the class Arachnida, with a worldwide distribution and with over 55,000 (Krantz and Walter 2009) species described to date. An estimate of up to half a million to a million more species await discovery (Krantz and Walter 2009). Mites and ticks are a very diverse group ranging in size from about 0.08 mm up to 1 centimetre long. Acari differ from others Arachnida by the fusion of the abdominal segments as in Araneae (spiders) and from spiders by the presence of a gnathosoma containing mouthparts, the fusion of the posterior part of the prosoma (the podosoma, bearing legs) and fusion of an opisthosoma into an idiosoma (Evans et al. 1996). Most species are free living and have different trophic modes, including phytophagous, predators feeding on a variety of small invertebrates, fungivores and detritivores. Some species have developed complex parasitic relationships with both vertebrate and invertebrate animals. A number of acarine groups are injurious to crops and to livestock, both because of their feeding activities and because of their capacity as vectors for a variety of disease organisms to their plant or animal host. While the Oribatida is an important group (more than 6,000 species) having a key role in soil equilibrium, data regarding invasive species and distribution range remain largely unavailable. Ticks are very peculiar acarines, since they are obligate ectoparasites. In this sense they form a very homogenous group, with the order Ixodida composed of only three families. In this chapter, the two groups of Acari, mites and ticks, will be treated separately. The ticks will be presented through the description of a few significant case studies. By contrast, mites being much diversified in their biology and habitat use, and being truly ubiquitous, will be presented systematically. Mites have successfully colonized nearly every known terrestrial, marine, and freshwater habitat. The most studied and observed invaders are found among the phytophagous mites of the families Tetranychidae and Eriophyidae, which include important agricultural pests. There is a growing awareness of the economic relevance of eriophyids as crop pests, including their importance as vectors of plant viruses, their role as alternative food for predators of plant pests, and their potential as weed control agents (Sabelis and Bruin 1996). A description on spider mite biology and their control is presented in the extensive review by Helle and Sabelis (1985). In addition to plantfeeding mites, a second group includes the alien parasitic mites. Among them, some invaded Europe with rodents such as muskrats (six alien species of mites), and brown rats (two aliens), while others are bird parasites (two species). Dermatophagoides evansi (Pyroglyphidae) is not associated with rodents and it has probably been accidentally introduced by humans (Bigliocchi and Maroli 1995,Hughes 1976,Thind and Clarke 2001). A single species in the family Varroidae, Varroa destructor, is alien to Europe (De Rycke et al. 2002, Griffiths and Bowman 1981). Ticks are important parasites of livestock, wild animals, and humans. After their parasitic phase, they spend most of their life cycle outside their hosts, where prevailing climate conditions may constrain their ability to colonize a given territory. While Mites and ticks (Acari). Chapter 7.4: 151 some tick species are highly restricted to particular combinations of climatic variables, or have defined host species, others may occur in widely variable climate conditions and have catholic feeding habits. Some species of ticks can be considered as invasive species, since the uncontrolled movements of domestic animals may introduce alien species into Europe or disperse some species outside their native distribution ranges. The introduction via large-bodied host vectors (such as passerine birds) and the uncontrolled importation of reptiles, are important means for colonizing newly available areas. Furthermore, one species of tick, Rhipicephalus sanguineus, is spreading in parts of Europe out of its current range because of the movements of domestic dogs. 7.4.2 Taxonomy of the mite species alien to Europe A total of 101 mite species have been considered as alien to Europe, including 96 species shown to have originated from other continents and 5 cryptogenic species (Table 7.4.1). These species involve 16 different families of mites (Figure 7.4.1). In addition, Table 7.4.2 provides some examples of mite species alien in Europe; i.e., European species introduced from one part of Europe to another where they are not native. Alien mites belong to two super orders, Acariformes (Actinotrichida) and Parasitiformes (Anactinotrichida). Most of these species belong to two orders of Acariformes, Prostigmata and Astigmata. Prostigmata includes the three most important superfamilies: * Tetranychoidea comprises two main families containing alien mites. The Tetranychidae family, or spider mites, includes 1,250 described species (http://www1. montpellier.inra.fr/CBGP/spmweb/). Among them, 100 can be considered as pests and 10 as major pests of agricultural crops. All stages are phytophagous and feed on parenchyma cells. No viruses associated with spider mites have been observed. The most widely distributed species is the highly polyphagous and ubiquitous Tetranychus urticae (two spotted spider mite), found on nearly 1,000 plant species. In Europe, alien spider mites are generally more specialized and occur on a single genus or family of plants. Due to their minute size (200 to 900 μm) typical of many species of Acari, spider mites remain undetected until major plant damage occurs. The members of another family, Tenuipalpidae, or false spider mites, are important obligate phytophagous mites. They are elongate, dorsoventrally flattened and usually have a reddish colour. * Eriophyoidea includes three families: – Eriophyidae, to which belong ca. 88% of all known Eriophyoidea in the fauna of Europe (Fauna Europaea 2009). These are vermiform, four legged mites. The family includes important economic pests of broadleaved plants. All known mite vectors of plant pathogens and nearly all gall-forming species belong to this family. About half are vagrants. Most of the species in the genera Aceria and Eriophyes cause specific galls on the leaves, green twig, flower buds, vegetative buds, or fruit of the hosts (Oldfield 1996). Others, especially Epitrimerus, Phyllocoptes, Aculops and Aculus cause discolouration and other non-distortive damage to their hosts. 152 Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) Figure 7.4.1. Relative importance of the mite families in the alien and native fauna in Europe. Families are presented in a decreasing order based on the number of alien species. Species alien to Europe include cryptogenic species. Only the most important families of native species (> 50 spp.) have been considered. The number over each bar indicates the number of species observed per family. – Phytoptidae, which are obligate phytophagous and gall mites, with a high degree of specificity. They are also vermiform and four-legged. The family Phytoptidae is well represented on conifers (half of the described phytoptid species) and monocots. Phytoptidae is less represented than Eriophyidae or Diptilomiopidae on dicotyledons. Four alien species out of a total of 56 species have been reported in the fauna of Europe. – Diptilomiopidae, which are predominantly leaf vagrants, only inhabiting leaves of dicotyledons, and rarely causing notable damage to their hosts (Keifer 1975). Two monotypic genera are known from only two families of monocotyledons (Poaceae and Palmae) occurring in the tropics. Rhyncaphytoptus species are mainly represented on several families of deciduous trees in the Holarctic region. Two alien species have been reported, out of the total 61 in the European fauna. * Tarsonemoidea represented by the family Tarsonemidae includes economically important mites. Most of them are mycophagous. Some species are phytophagous, whereas others are parasites of bark beetle eggs, or predators of tetranychid eggs. The most redoubtable pest species in the family is the broad mite, Polyphagotarsonemus latus (=Hemitarsonemus latus), which was described in 1890 and has recently been redefined and considered as being a species complex (Gerson 1992). The order Astigmata is less represented in the alien fauna. A few species belong to the super-family Sarcoptoidea, and especially to families Listrophoridae and Myocopidae. Members of Listrophoridae are usually small, elongate mites and are skin or hair parasites of mammals. The palpae and/or legs I-II are often highly modified for grasping hairs. Four species of Listrophoridae mites have invaded Europe, grasped to the fur of muskrats: Listrophorus americanus, L. dozieri, L. faini and L. validus (Šefrová and Laštůvka 2005). Myocopids, or hair mites, live on skin of marsupial and rodents (Bauer and Whitaker 1981, Šefrová and Laštůvka 2005, Mites and ticks (Acari). Chapter 7.4: 153 Whitaker 2007). Myocoptes ondatrae is an ectoparasite that has invaded Europe by grasping the fur of muskrats (Bauer and Whitaker 1981, Šefrová and Laštůvka 2005, Whitaker 2007). Other species belong to the super-family Acaroidea and families Epidermoptidae and Pyroglyphidae. Epidermoptidae are skin parasites of birds. Epidermoptes bilobatus causes avian scabies. Pyroglyphidae are external parasites living on bird feathers or are nidicolous. Dermatophagoides evansi feeds on human detritus, and lives in house dust as well as within bird nests (Piotrowski 1990, Razowski 1997). Among the super-order Parasitiformes (Anactinotrichida), aliens belong to orders Ixodida and Mesostigmata. Ixodida is represented by the species in the family Ixodidae, which is treated in a separate section at the end of the chapter. Alien Mesostigmata belong to superfamilies Ascoidea and Dermanyssoidea. The first superfamily is represented by a single family with aliens, Phytoseiidae, which are predators of spider mites. In Europe, species such as Phytoseiulus persimilis, Amblyseius (Neoseiulus) californicus and Iphesius (Amblyseius) degenerans are used as biological control agents against phytophagous pests (Bartlett 1992, Croft et al. 1998, Easterbrook 1996, EPPO 2002, Garcia Mari and Gonzalez-Zamora 1999, Helle and Sabelis 1985, McMurtry and Croft 1997). Three families of Dermanyssoidea contain alien species. Varroidae mites are ectoparasites of honeybees. Varroa destructor is at present the most important parasite of Apis mellifera (L.). Varroa feeds on the haemolymph of adult, larval and pupal bees. Laelapidae mites live in soil, are nidicoles or parasitize small mammals and insects. Ondatralaelaps multispinosus is an ectoparasite of muskrats (Šefrová and Laštůvka 2005). Laelaps echidninus is a common worldwide ectoparasite of spiny rats, wild brown rats and is occasionally found on the house mouse and cotton rat (Wharton and Hansell 1957). Macronyssidae mites are haematophagous, have a large dorsal shield, prominent chelicerae and inconspicuous body setae (Easterbrook et al. 2008). Ornithonyssus bacoti is a parasite of rats, living in rat nests and their surroundings (Cole et al. 2005, Easterbrook et al. 2008, Fan and Petit 1998, Whitaker 2007). Ornithonyssus bursa is a natural parasite of common birds including pigeons, starlings, sparrows, Indian mynahs, poultry, and some wild birds, such as the robin (Berggren 2005). 7.4.3 Temporal trends of introduction in Europe of alien mite species The rate of arrival of alien mites in Europe is increasing exponentially (Figure 7.4.2). An average of 2.1 alien species was newly recorded per year in Europe during 2000– 2007 whereas only half this number was recorded during the period 1950–1974 (1 species/year). However, large differences were found between families. The first records for Europe of all alien Tetranychidae are extensively documented in this chapter. There are no records reported before 1950; however, only few taxonomists were specialized on the family before this date. Since the second half of the 20th century, tetranychid species have been reported at an average rate of one new species every two years, with an acceleration of reports (one species per year) since 2000. 154 Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) Figure 7.4.2. Temporal changes in the mean number of records per year of mite species alien to Europe from 1800 to 2009. The number over each bar indicates the absolute number of species newly recorded per time period. Most of these mites represent agricultural pests, and therefore have been widely studied which explains the overrepresentation of crop pest species as Tetranychidae aliens. The mean number of records of Eriophyoidae species alien to Europe increased rapidly during the third quarter of the 20th century. Only one species Aceria alpestris, which is alien in Europe, was recorded within the period 1850–1899. This species was described from the host plant Rhododendron ferrugineum L. from Tirol (Austria). The species was later recorded in mainland Italy, Czech Republic, Slovenia and Serbia, but it is not clear if it was associated with cultivated Rhododendron. Species recorded intensively between 1900–1924 (although described from Germany in 1857) are categorized as cryptogenic (Eriophyes pyri, the pear blister mite) or alien in Europe, like Aculus hippocastani (recorded in 1907, but probably introduced in Europe from the 17th century when its host plant Aesculus hippocastanum L. was intensively cultivated), and Aceria loewi (probably introduced in the 16th century when lilac started to be cultivated in France). Aculops allotrichus, which is alien to Europe, was recorded in 1912 but was probably, introduced with its host Robinia pseudoacacia L. which was for the first time introduced into France at the beginning of 17th century. Aceria erinea and A. tristriata were suspected to have an Asian origin and have been designated as aliens. They were recorded on 1903, but probably were present on its host, Persian walnut, in the Balkans and South Europe much earlier. Only one species in the Eriophyoidae was recorded between 1925–1949, e.g. Aceria petanovicae, the lilac rust mite. Being for long time known under the name of Aculops massalongoi the species is alien in Europe. Mites and ticks (Acari). Chapter 7.4: 155 Six alien species to Europe were recorded between 1950–1974. Two pests of citrus, Aceria sheldoni (citrus bud mite) and Aculops pelekassi (citrus rust mite) and the azalea mite Phyllocoptes azaleae, are suspected to have been introduced from Asia. Characteristic symptoms of deformed lemon fruits caused by A. sheldoni were drawn by Battista Ferrari in Italy in 1664 (Ragusa 2002). Three pests have been reported from North American maple trees (Acer negundo L., A. saccharinum L. and A. rubrum L.), i.e. Shevtchenkella brevisetosa, Vasates quadripedes and Rhyncaphytoptus negundivagrans. The 25 species recorded during the period 1975–1999 almost all have a North American origin (only Epitrimerus cupressi is designated as cryptogenic, because of the Mediterranean origin of its host Cupressus sempervirens L.). During the period from 2000 to 2007, one species alien to Europe, Rhyncaphytoptus bagdasariani, has been recorded as being introduced from Asia and the serious pest Aceria fuchsiae (a species on the European quarantine list) was introduced from South America. As for other phytophagous mites, the most probable explanation for the acceleration in the pace of introductions of alien eriophyids is intensification of international trade. Most of these alien species inhabit ornamental trees and shrubs, flowers and potted ornamental plants. Some alien parasitic mites have invaded Europe with rodents such as muskrats and brown rats. The muskrat (Ondatra zibethicus L.) is an invasive rodent native to North America. It was introduced around 1905, by humans as a fur resource in several parts of Europe, as well as in Asia and South America. Six species of mites, native from North America (Bauer and Whitaker 1981, Whitaker 2007), have invaded Europe grasping its fur (Glavendekić et al. 2005, Šefrová and Laštůvka 2005). The first report of muskrat mites was recorded in 1955, and a second in 2000, both in Czech Republic. Two other parasitic species, Laelaps echidninus and Ornithonyssus bacoti, are also alien ectoparasites of rodents that have invaded Europe and were identified in the 1950’s (Šefrová and Laštůvka 2005), but the exact pathway of introduction is not known. One possible vector is the wild brown rat, Rattus norvegicus (Berkenhout). Thought to have originated in northern China, this rodent spread in Europe in the middle ages and is now the dominant rat in the continent. Birds are vectors of a second group of alien parasitic mites, that include Epidermoptes bilobatus and Ornithonyssus bursa, both identified in the 1950’s, in the Czech Republic (Šefrová and Laštůvka 2005). The exact route of introduction is not known with confidence, but a possible vector is the chicken (Gallus gallus domesticus L.). In the 20thcentury, with the intensifications of poultry production, concerns have been raised about the increasing risk of transfer of diseases and mites (from chickens to native bird species). Whereas the exact date of arrival of alien mites is generally unknown, deliberately released biological control agents are the exception to this rule. Among them, three phytoseiids are mainly used as predatory species against pests (McMurtry and Croft 1997). Phytoseiulus persimilis was introduced for the first time in the 1970’s in Bulgaria and Czech Republic (EPPO 2002, Šefrová and Laštůvka 2005). Neoseiulus californicus was introduced for the first time in 1991 in Great Britain (EPPO 2002). It was also introduced at the same period in the Czech Republic (EPPO 2002, Šefrová and Laštůvka 156 Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) 2005). The third introduced mite is Iphiseius degenerans. It is native from the Mediterranean region and was introduced for the first time in 1993 in Czech Republic (EPPO 2002, Šefrová and Laštůvka 2005). Nowadays, these three biological agents have been introduced in most European countries. 7.4.4 Biogeographic patterns of the mite species alien to Europe 7.4.4.1 Origin of the mite species alien to Europe Figure 7.4.3. presents the region of origin of the 101 alien species of mites. Most of the alien mite species (52%) came from North America, then from Asia (25%), and Central and South America (10%). The origin of phytophagous alien mites can usually be inferred from the origin of the host plant. These mites are dispersed over long distances mainly by the introduction of plant material and spread further by plant cultivation in newly colonized regions. Aerial distribution is possible and most frequent, but mainly over short distances (Margolies 1993, Margolies 1995). In the case of highly polyphagous species such as several Tetranychidae, their ubiquity and highly diverse host uses might be misleading and the origin can be difficult to ascertain. Twelve out of 27 alien Tetranychidae originated in North America, nine in Asia and only five in Central and South America. Temperate regions provide the majority of the alien species (16 vs. 11 for tropical areas). The majority of eriophyoid species are mono- or oligophagous and are distributed within the host range. North America appears to be the dominant source of the alien eriophyoid fauna with half of the species originating from this continent. Around 26% of species originate from Asia, and less than 10% from South America. A few species are designated as cryptogenic or with questionable origin. For example, Rhyncaphytoptus negundivagrans, although described from Hungary, probably originated from North America with its host plant, Acer negundo. Whereas the camellia rust mite, Cosetacus cameliae (described from California) was probably introduced to Europe from the USA, it probably has an Asian origin considering that Camelia japonica L. comes from subtropical and tropical regions of Southeast Asia. The pouch gall mite of plum leaves, Eriophyes emarginatae, first discovered in the USA, has also been recorded in Serbia and Japan. This mite is very closely related to the European E. padi (Nalepa) (Petanović 1997) and may even be the same species, with synonymous names (Keifer 1975). Epitrimerus cupressi was described from North America, but according to the origin of its host plant Cupressus sempervirens, which is from the Mediterranean region, the mite probably has an European origin too. The gall mite Phytoptus hedericola (Phytoptidae) is native from South Africa (Glavendekić et al. 2005), and Trisetacus chamaecypari (Phytoptidae) from North America (Ostojá-starzewski and Halstead 2006, Smith et al. 2007). Among the false spider mites (Tenuipalpidae), Brevipalpus californicus, B. obovatus and Tenuipalpus pacificus originated from Central and South America, and Florida Mites and ticks (Acari). Chapter 7.4: 157 Figure 7.4.3. Origin of the mite species alien to Europe. (USA) (Denmark 1968, Manson 1967). Six alien species of rodents bear parasitic mites originating from North America, and belong to the families Listophoridae (four species), Laelapidae (one species), and Myocoptidae (one species). In their native country, they are all ectoparasites of murskrats. There are also some bird parasites: one species of Epidermoptidae, Epidermoptes bilobatus, is an ectoparasite native from South Asia, and Ornithonyssus bursa is probably native from Trinidad. A single Varroa species, V. destructor, is alien to Europe (Griffiths and Bowman 1981). Its native range is South East Asia, where it was originally confined on its original host, the Asian honeybee, Apis cerana F. This mite came to be a parasite of the European honeybee, Apis mellifera, in the mid-twentieth century. Importation of commercial A. mellifera colonies into areas with A. cerana brought the previously allopatric bee species into contact and allowed V. destructor to switch to the new host 7.4.4.2 Distribution in Europe of the alien mite species Alien mite species are not evenly distributed throughout Europe. Large differences in the number of aliens are noticed between countries (Figure 7.4.4) but it may reflect differences in sampling efforts and in the number of local taxonomic specialists. Among the Tetranychidae, 19 alien species are found around the Mediterranean Basin and 12 in the rest of Europe. With relatively warm winters, the Mediterranean region provides suitable climatic living conditions for many species of temperate climates, but also for the establishment of many species of tropical or sub-tropical origin. Except for Panonychus citri and the cryptic species Tetranychus ludeni, which can be found in glasshouses in Europe, all tropical alien spider mites are restricted to the area around the Mediterranean Sea. 158 Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) Figure 7.4.4. Comparative colonization of continental European countries and islands by mite species alien to Europe. Archipelago: 1 Azores 2 Madeira 3 Canary Islands. Most alien Eriophyids have a very restricted distribution. More than 40% of the species have been observed in only one country (17 species), more than 40% (21 species) in 2–5 countries, and approximately 20% (7 species) in 6–11 countries. Eight European countries have no recorded occurrence of alien eriophyoids to date. Only one species, the pear blister mite Eriophyes pyri (which has cryptogenic status), has been recorded from 32 European countries. Besides E. pyri, the more widely distributed eriophyoid species are: Aceria erinea, A. loewi, A. sheldoni, Aculops pelekassi and Eriophyes canestrini. The gall mite Phytoptus hedericola (Phytoptidae) entered Europe in 2002 and has been observed in Serbia (Glavendekić et al. 2005). Trisetacus chamaecypari (Phytoptidae) entered Europe in 2002 (Ostojá-starzewski and Halstead 2006, Smith et al. 2007). The status of Typhloctonus squamiger (Phytoseiidae), a poorly known phytophagous mite found on trees in Italy since 1991 (Rigamonti and Lozzia 1999), is questionable. The distribution of biological agents belonging to the Phytoseiidae family is wellknown. Phytoseiulus persimilis is now present in nearly all of Europe (Table 7.4.1) Mites and ticks (Acari). Chapter 7.4: 159 (EPPO 2002). Neoseiulus californicus has been found in the same countries except Austria, Hungary, Morocco, Slovakia, Sweden and Turkey. The third introduced phytoseid mite, Iphesius degenerans, is also present in several countries (Table 7.4.1). The broad mite Polyphagotarsonemus latus (Tarsonemidae) is now cosmopolitan. In Europe, it was reported for the first time in 1961 and since then the mite has invaded almost all countries (Table 7.4.1) (CAB-International 1986, Fan and Petit 1998, Natarajan 1988, Parker and Gerson 1994); it is potentially now in all parts of Europe. Three species of false spider mites (Tenuipalpidae) are major invaders in Europe. Brevipalpus californicus, found in 316 orchid and tree species of 67 genera and 33 families, was first recorded in 1960 and is mainly observed in citrus trees around the Mediterranean basin (Denmark 1968, Manson 1967). The privet mite, Brevipalpus obovatus is found in 451 herb, ornamental and shrub species (19 genera, 55 families) (Manson 1967) has been recorded from Austria, Cyprus, France, Germany, Israel, Netherlands, Serbia and Spain (Manson 1967). Tenuipalpus pacificus (the Phalaenopsis mite) is found in greenhouses of Phalaenopsis orchids in Germany, Great Britain, Netherlands and Serbia (Denmark 1968, Manson 1967). The introduced range of Varroa destructor is practically worldwide. It was first reported in Eastern Europe in the mid- 1960s and it has spread rapidly all over the continent. Two different genotypes, characterized by mitochondrial DNA sequences, have spread as independent clonal populations (Solignac et al. 2005), the Korean and the Japanese haplotypes, the latter having been found, besides Asia, in the Americas only. 7.4.5. Pathways of introduction in Europe of alien mite species Although colonisation routes are poorly documented for the Tetranychidae, it is known that many species travel with their host plant. Small organisms like tetranychids are easily transported with plant material (leaves and in bark crevices). Only five species feed mainly on herbaceous plants (Tetranychus evansi, T. macfarleni, T. sinhai, Schizotetranychus parasemus, and Petrobia lupini), whereas all other alien species in the family feed on perennial shrubs. As for tetranychids, the horticultural and ornamental trade is probably the most important factor for accidental introductions of almost all species of alien Eriophyoidae. Just a few species of Eriophyoids are on European quarantine lists, as plants are rarely inspected for presence of these mites. Infested plant material is not regularly intercepted at borders even in the case of important pests such as the grape rust mite Calepitrimerus vitis (Nalepa) or the blackberry fruit mite Acalitus essigi (Hassan), which are frequently disseminated with plant seedlings. During recent decades more than 50% of aliens were imported with ornamental plants. Among eriophyids, which are obligate plant parasites, only one trophic group which is associated with weeds, can be subject to intentional introduction. Although these mites were recently nominated as potential agents for classical biological control of weeds (few species are imported for this purpose), they have not yet been used for this purpose in Europe. Four species of 160 Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) alien eriophyoids which were probably introduced along with their host plants may have the potential as biological control agents of serious alien weed pests. In particular, Aceria ambrosiae can be used against the allergenous weed Ambrosia artemisifolia L. that was imported into Europe from North America. As for other phytophagous species, the broad mite Polyphagotarsonemus latus (Tarsonemidae) has mainly been dispersed by human activities, but also by wind or insect transfer. Movement by insects should not be neglected: this concerns almost only females that get attached to the legs of aphids and the whiteflies Bemisia argentifolii (Bellows and Perring), Bemisia tabaci (Gennadius) and Trialeurodes vaporariorum (Westwood) (Homoptera: Aleyrodidae) (Fan and Petit 1998, Natarajan 1988, Parker and Gerson 1994). Although including important crop pest species, the dispersal potential of false spider mites (Brevipalpus spp.), Tenuipalpidae, remains unclear (Childers et al. 2003a, 2003b). Intentional introductions of mites represent a low proportion of alien arrivals. Only three phytoseeid predators were introduced purposely for biological control and have established. Some of these biological control agents were released in the field but others were first released in glasshouses, and then escaped and became established outdoors. International travel and commerce has facilitated the dispersal of Varroa destructor. Once established in a new region, the mite spreads using drifting, robbing, and swarming behaviour of the host. Human mediated varroa dispersion also occurs via apicultural practices. 7.4.6. Ecosystems and habitats invaded in Europe by alien mite species Alien mites established in Europe predominantly live in agrosystems or anthropogenic environments (ca. 92%; Figure 7.4.5). This is especially verified in Tetranychidae and Eriophyidae. Among eriophyoids, some are present in man-made habitats, parks and gardens (22 species), agricultural lands (13 species), and greenhouses (10 species); very few species inhabit woodland and forest, costal, alpine or sub alpine habitats. Most alien species in this superfamily are leaf vagrants (13 species). Twelve species cause leaf galls, erinea* and leaf rolling, 11 cause leaf and/or fruit russeting or other type of discolouration, six live predominantly in buds causing bud galls, three species cause stunting of whole plants and/or plant organs and two cause flower and/or fruit deformations. Among the leaf gall makers, the most important horticultural pests are distributed in many European countries, such as E. pyri, A. erinea, A. tristriata or, such as A. fuchsiae which is on quarantine lists. Among the rust mites, only a few are important horticultural pests like A. theae, A. pelekassi and C. carinatus. Most species are pests of ornamental trees, shrubs or flowering plants, having an important aesthetic impact on plants in parks and streets in most European towns and cities (i.e. A. gleditsiae, A.ligustri, A. petanovicae, S. strobicus, P. chrysanthemi), an exception being A. sawatch- Mites and ticks (Acari). Chapter 7.4: 161 Figure 7.4.5. Main European habitats colonized by the established alien species of mites. The number over each bar indicates the absolute number of alien dipterans recorded per habitat. Note that a species may have colonized several habitats. ensae which inhabits weeds. Two Eriophyoids which cause plant stunting, A. paradianthi and T. califraxini, are important pests of ornamental plants and one species, A. ambrosiae, is a potential biocontrol agent against the alien weed Ambrosia artemisifolia. Two species which cause flower and/or fruit deformations, A. alpestris and A. sheldoni, are respectively pests of Rhododendron and citrus trees. The gall mite Phytoptus hedericola lives on ivy (Hedera helix L.) and Trisetacus laricis switched from American larch to European larch (Larix decidua Mill.). The broad mite Polyphagotarsonemus latus (Tarsonemidae) has a very short life cycle of a few days, damaging crops abruptly. Being highly polyphagous, the species has been reported on 57 plant families (Gerson 1992) both in open field crops and in greenhouses. This is an important pest of crops and ornamental plants such as azaleas, castor bean, chillies, citrus fruits, cotton, cucumber, mango, papaya, pepper, potato, sweet potato, tea, tomato and winged bean (Gerson 1992, Glavendekić et al. 2005, Heungens 1986, Raemaekers 2001). Nevertheless, in Europe this mite is found mainly in greenhouses because the mite cannot survive winter conditions outdoors. False spider mites (Brevipalpus spp.; Tenuipalpidae) present a risk of invasion in greenhouses. Brevipalpus obovatus (the privet mite) is found on ornamentals and shrubs like citrus and azaleas and could become of great importance in glasshouses for ornamentals (Childers et al. 2003a, 2003b). Tenuipalpus pacificus (the Phalaenopsis mite) is one of the rare monophagous mites in the family, but it is a very destructive pest of orchids under greenhouses, mainly because it has several generations per year and has a two-month life cycle (Denmark 1968, Manson 1967). 162 Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) A Pyroglyphidae mite, Dermatophagoides evansi, is a cosmopolitan free-living species, often encountered in synanthropic situations and has probably been accidentally introduced by humans (Bigliocchi and Maroli 1995, Hughes 1976). 7.4.7. Ecological and economic impact of alien mite species Seven species of alien Tetranychidae are important pests. On citrus, four alien species are found: Panonychus citri, Eotetranychus lewisi (also on grapes) Eutetranychus banksi and E. orientalis, the last presently spreading to Southern Portugal and Spain from Huelva to Murcia and Alicante. Oligonychus perseae is found on avocado and produces very severe damage in southern Spain (Malaga, Granada and Huelva) and in the Canary Islands. Stigmaeopsis celarius is found on bamboos and causes important visual damage to these ornamental plants. Tetranychus evansi is found on solanaceous crops and can reach very high density as observed in France, Spain and Canary Islands. All these mites are present in the Mediterranean Basin, which appears to be the region most threatened by alien species. Only two of these species can be found outside the Mediterranean area: Panonychus citri, especially in glass-houses, and Stigmaeopsis celarius. In humid citrus-growing regions of the world, eriophyoid mites are considered to be the major mite pests (Jeppson et al. 1975, McCoy 1996). Two alien species, Aceria sheldoni and Aculops pelekassi, distributed worldwide, are among the most important pests infesting citrus. The pear blister mite, Eriophyes pyri, widely distributed in Europe, probably does little harm to the tree, but in severe infestations, the tree leaves may become disfigured, and most importantly the mite may damage fruits (Easterbrook et al. 2008). Besides fruit orchards, species in the superfamily inhabiting wild trees in natural forests are: Aceria tristriata and A. erinea which appear to be the most common and most injurious eriophyoids found on Juglans regia L. (Castagnoli and Oldfield 1996). Among the five species of eriophyoid mites reported from commercially important beverage crops in different parts of the world, wherever tea is grown, the purple tea mite Calacarus carinatus and the pink tea mite Acaphylla theae are economically important in Southeast Asian countries, and in India (Channabasavanna 1996). Both species are aliens to Europe, reported from mainland Italy (A. theae) and from Hungary, Poland and Spain (C. carinatus). Records concerning host plant range in the case of C. carinatus are, besides tea, Viburnum opulus L. and Capsicum annuum L. (Amrine and Stasny 1994). Bearing in mind that congeneric Calacarus citrifolii has an extremely wide host range (Oldfield 1996), this might be also the case for C. carinatus, which would convey on the latter serious pest status in Europe. Economic impact of alien pest species of eriophyoids on ornamentals has been observed for Aculops gleditsiae on honey locust, Aceria petanovicie on lilac, Aculops ligustri on privet hedges, Aculops allotrichus on black locust, Reckella celtis on Celtis australis L., Shevtchenkella brevisetosa on Acer negundo, Vasates quadripes on silver maple, Phytoptus hederae on English ivy, and Setoptus strobicus on Pinus strobus L. (Petanović 2004). Flower and foliage aesthetic impact has been observed indoors (business centers, restaurants, shopping centers, hotels, etc.) for a few alien eriophyoids, Cecidophyopsis hendersoni causing a powdery Mites and ticks (Acari). Chapter 7.4: a b c d e f g h i j 163 Figure 7.4.6. Alien mites and their damage. a Curling and rusting of black locust leaves caused by Aculops allotrichus b Chlorotic and misshapen leaves of Acer negundo caused by Shevtchenkella brevisetosa (left) and uninfested leaves (right) c Leaf rusting of lilac leaves caused by Aceria petanovicae d Aceria petanovicae, dorsal view-SEM photograph e Rusting of Pinus strobus needles caused by Setoptus strobacus f Setoptus strobacus eggs, juveniles and adults between needles of Pinus strobus g Leaf distortion and unopened damaged flower buds of chrysanthemum caused by Paraphytoptus chrysanthemi h Deformed flower heads of chrysanthemum caused by Paraphytoptus chrysanthemi i Colony of Cecidophyopsis hendersoni on Yucca leaf j Panonychus citri. (a–i Credit: Radmila Petanović; j Credit: Alain Migeon). appearance on Yucca leaves, Cosetacus cameliae causing bud rust and abortion on flower buds of Camelia plants, and Paraphytoptus chrysanthemi causing deformed buds, hairy leaves and rust on Chrysanthemum (Petanović 2004). The broad mite Polyphagotarsonemus latus (Tarsonemidae) and the false spider mites (Brevipalpus spp.) (Tenuipalpidae) are major pests of great agronomical impor- 164 Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) tance because of their broad host range, worldwide distribution and economic impact (CAB-International 1986, Fan and Petit 1998, Gerson 1992, Heungens 1986, Natarajan 1988, Parker and Gerson 1994, Raemaekers 2001). The most important threat for Brevipalpus spp. is the spread of citrus viruses (Childers et al. 2003b). Among parasitic mites, the hair mites (muskrat mites) are currently considered non-pathogenic for humans although they are sometimes found in the fur of other mammals. Laelaps echidninus (Laelapidae) is a common worldwide ectoparasite of the spiny rats (hystricognath rodents), wild brown rat and is occasionally found on the house mouse, cotton rat and other rodents. It is a bloodsucking mite and the natural vector of Hepatozoon muris Balf. (Protozoa, Adeleidae), a haemogregarine parasite pathogenic for white rats (Smith et al. 2007) but which should not be overlooked as a possible vector of disease to humans (Wharton and Hansell 1957). Ornithonyssus bacoti (Macronyssidae) is a parasite of rats and inhabits the area in and around the rat’s nesting area. This mite is the only one of the common rat mites which frequently deserts domestic rats to bite man or his domestic and laboratory animals (Cole et al. 2005). It is also a bloodsucking mite and its bite is painful and causes skin irritation, itching and skin dermatitis in humans (James 2005). Ornithonyssus bacoti, is a known vector of the murine filarial nematode Litomosoides carinii Travasaos. In addition, it is susceptible to the transmission of endemic typhus, Rickettsia typhi (Wolbach and Todd) 1943 (= R. mooseri Monteiro) to humans (Berggren 2005, Bowman et al. 2003). Epidermoptes bilobatus (Epidermoptidae) is a bird parasite causing avian scabies. This endoparasite burrows into the skin causing inflammation and itchiness. The skin thickens with brownish-yellow scabs, which may become secondarily infected with a fungus. It is difficult to control and can cause death. Culling infested birds is usually required (Department of the Environment and Heritage 2006). Ornithonyssus bursa (Macronyssidae) is an haematophagous natural parasite of common birds including pigeons, starlings, sparrows, Indian mynahs, poultry, robin (Berggren 2005). These pest mites and parasites are and will remain a long term problem for poultry housing (Gjelstrup and Møller 1985). Although none of these two species of mites are truly parasitic on humans and pets, they readily bite humans and are liable to cause allergies and dermatitis in human (Denmark and Cromroy 2008, James 2005). Dermatophagoides evansi (Pyroglyphidae), and a species alien in Europe, Glycyphagus domesticus (Glycyphagidae), have been accidentally introduced by humans and often encountered in synanthropic situations (Bigliocchi and Maroli 1995, Hughes 1976, Thind and Clarke 2001). Glycyphagus domesticus also occurs in bird, bat and mammal nests. It is associated with moist and humid conditions that promote the growth of mould on which they feed (Thind and Clarke 2001). Dermatophagoides evansi (Pyroglyphidae) feeds on detritus and is also found in house dust, birds’ nests and poultry houses (Piotrowski 1990, Razowski 1997). Dermatophagoides evansi represents a source of airborne allergens in indoor house dust (Eriksson 1990, Musken et al. 2000) that may cause sensitization, dermatitis, rhinopharyngitis and asthma especially among farmers. The honeybee ectoparasite Varroa destructor causes serious losses through feeding injury in apiaries in Europe but also almost worldwide. While the populations of the Mites and ticks (Acari). Chapter 7.4: 165 a b c d Figure 7.4.7. Ixodidae ticks on tortoises and snakes. a Hyalomma aegyptium on tortoise b Amblyomma exornatum semi-engorged on Python head c Amblyomma sp. on snake head (Credits: Nicasio Brotons) d Female of Varroa destructor on abdomen of Apis mellifera (Credit: Alain Migeon). parasite reach only a small size within colonies of A. cerana and do not damage the colony, infested A. mellifera colonies die. The problems with varroa control are typical of those encountered in curbing arthropod pest population. Varroas are becoming resistant to the acaricides used by beekeepers to control them. The recent discovery in several parts of the world (notably the United States of America (Harbo and Harris 2005) and Europe (Le Conte et al. 2007)) of honeybee bee colonies able to tolerate heavy infestations of V. destructor opens the door to lasting solutions for controlling the parasite. A positive impact is recognized for the three mite species deliberately introduced to Europe for biological control of house flies and tetranychid mites. Phytoseiulus persimilis and N. californicus are two well-known biological control agents used against spiders mites such as Tetranychus urticae Koch (Garcia Mari and Gonzalez-Zamora 1999, Helle and Sabelis 1985) and Phytonemus pallidus (Banks) (James 2005). The third introduced mite, Iphiseius degenerans, targets numerous species of thrips (van Houten and van Stratum 1993, van Houten and van Stratum 1995), e.g. Thrips tabaci Lindeman and Frankliniella occidentalis (Pergande) (Albajes et al. 1999, Bartlett 1992, McMurtry and Croft 1997, Sengonca et al. 2004). 7.4.8. Alien tick species: case studies It is difficult to ascertain if a tick may have permanent populations outside of its native range or, to the contrary, they are just isolated records. In some cases, a few examples 166 Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) of a given species have been reported for a small area or found over non-resident hosts. This may result from the introduction of a few specimens, commonly immature stages. The most important means of introduction and expansion of ticks (provided that suitable climate and host is available) is by means of engorged females, because of their huge potential to lay thousands of eggs. The movements of domestic ungulates have introduced some tick species, that may be considered to produce permanent and viable populations out of their native range. An example is the introduction of Hyalomma dromedarii into the Canary Islands, by the importation of dromedaries (Camelus bactrianus L.). The native range of this tick is northern Africa where C. bactrianus is the main adult host, and H. dromedarii is abundant in wide areas of Mauritania and Morocco. The current population of dromedaries in the Canary Islands was introduced from Morocco at the end of 18th Century, and it seems that this tick came into these islands using dromedary hosts. H. dromedarii may use a wide range of hosts in immature stages, thus increasing risk of spread and permanent establishment (Apanaskevich and Horak 2008, Apanaskevich et al. 2008). It is difficult, however, to assess the reliability of records of Hyalomma anatolicum excavatum. A recent review of the original two subspecies (H. a. anatolicum and H. a. excavatum), concluded that they should be considered as separate species, although the matter is hard to decide as both taxa have a well defined allopatric range (Apanaskevich 2003). H. excavatum is restricted to central and eastern Asia and H. anatolicum colonizes wide areas of northern Africa. The records of H. excavatum from Bulgaria, Albania, Greece, and Italy should be cautiously treated, as they may probably represent H. anatolicum imported from northern Africa with domestic ungulates, as is the case for Hyalomma detritum. The formerly recognized species H. detritum, restricted to northern Africa, is now considered to be a synonym of the European H. scupense, which occurs not only in scattered localities of mainland Europe but is present in wide areas of northern Africa. Similarly, caution should be also applied for the single record of Hyalomma truncatum in the Canary islands. This tick is currently known to be restricted to parts of Asia, while a close species, H. rufipes, is common in sub-saharan Africa. While the adults of H. rufipes feed on a variety of hosts, including domestic ungulates, the immature stages commonly attach to diverse passerine birds. Most of these birds perform long distance travel in their migratory flights from Africa to Europe, and they have been found carrying hundreds of immature ticks (Hoogstraal 1956). However, as mentioned above, it is difficult for a population of nymphs to produce a viable and permanent population of resident ticks. To our knowledge, H. rufipes has been recorded only in Cyprus and Macedonia (Apanaskevich and Horak 2008), and we still do not know if these are permanent populations or only accidental records on their passerine hosts on migration to lower latitudes from sub-saharan Africa. The scenario for the tortoise tick, Hyalomma aegyptium, is however different. Its presence outside northern Africa has been reported in countries such as Romania, Spain, Italy, Greece, Bulgaria, Croatia, and even farther north in Belgium (Siroky Pet al. 2007). The tick has permanent populations in areas of southern Russia (Robbins et al. 1998). There have been also introductions of this tick by tortoises imported form Mites and ticks (Acari). Chapter 7.4: 167 northern Africa or eastern Europe, where this tick is common. The only record of a permanent population of H. aegyptium as a consequence of an accidental importation recorded for eastern Spain (Brotóns and Estrada-Peña 2004). Since the ticks attach to portions of the neck and legs of the host body, it may be difficult to find feeding stages even after careful observation of the hosts. In the reported case of introduction of several specimens of Testudo graeca infested by ticks, the hosts were kept in a large private garden with a Mediterranean-type climate and vegetation. After some years of recurrent tick parasitism in the tortoises without new importations and repeated treatments, it was realized that the tick had permanent populations in the garden, and the hosts became infested according to the seasonal activity of the ticks. An interesting case of tick introduction into mainland Europa are ticks commonly found on snakes, like Amblyomma latum and A. exornatum (both formerly in the genus Aponomma). These ticks feed for a long period on the host, and owing to their small size and preference to feed under host scales, they are commonly unrecognized while importing a host out of its native range. Amblyomma latum is a very common parasite of Python spp., which is becoming increasingly popular as a pet in Europe. The only known case of an importation of A. exornatum was noticed on specimens of Varanus niloticus that arrived into Spain (Estrada-Peña (Unpubl.)). These imported ticks founded a permanent population in the terrarium where the lizards live, under suitable conditions of high relative humidity and controlled temperature. A very peculiar case of tick introduction is an alien in Europe, the brown dog tick, Rhipicephalus sanguineus. While feeding on domestic dogs, this tick is endophilic and is normally restricted to the Mediterranean region, being abundant in kennels, human constructions and private gardens where dogs remain unprotected against tick bites. Because of its endophilic behaviour, this tick may survive independently of prevailing environmental conditions, since human habitations buffer harsh climate. Therefore, unprotected pets travelling may harbor feeding ticks, and introduce them to uninfested areas which might be far from their native range. 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Shetchenko VG, Rupais AA (1964) Four Leggd Mites (Acarina:Eriophyidae)- pests of parks in Latvia. Fauna Latviskoi SSR,IV, Riga: 203–239. Sibomana G, Geerts S, De Vries T (1986) L’établissement de Rhipicephalus sanguineus (Latreille, 1806) à l’intérieur des maisons en Belgique. Annales de la Société belge de médecine tropicale 66 : 79–81. Siroky P, Petrzelkova KJ, Kamler M, Mihalca A, Modry D (2007) Hyalomma aegyptium as dominant tick in tortoises of the genus Testudo in Balkan countries, with notes on its host preferences. Experimental and applied Acarology 40: 279–290. Smith TG (1996) The genus Hepatozoon (Apicomplexa: Adeleina). Journal of Parasitology 82: 565–585. Smith RM, Baker RHA, Malumphy CP, Hockland S, Hammon RP, Ostojá-Starzewski JC, Collins DW (2007) Recent non-native invertebrate plant pest establishments in Great Britain: origins, pathways, and trends. Agricultural and Forest Entomology. 9, 307–326. 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Mites and ticks (Acari). Chapter 7.4: 177 Van Houten YM, van Stratum P (1993) Biological control of western flower thrips in greenhouse sweet peppers using nondiapausing predatory mites. Experimental and Applied Acarology 4: 229–234. Van Houten YM, van Stratum P (1995) Control of western flower thrips on sweet pepper inwinterwith Amblyseius cucumeris (Oudemans) and A. degenerans Berlese. NATO ASI series A, Life Sciences 276: 245–248. Vappula NA (1965) Pest of cultivated plants in Finland. Acta Entomologica Fennica 19: 1–239. Vierbergen G (1989) Panonychus citri - na vele malen geïmporteed, tenslotte geïntroduceerd. Verslagen en Medelingen Plantenziektenkundige Dients Wageningen 167: 52–53. Vierbergen G (1990) The spider mites of the Netherlands and their economic significance (Acarina: Tetranychidae). Proceedings of the Section Experimental & Applied Entomology of the Netherlands Entomological Society 1: 158–164. Vierbergen G (1994) Entomology. 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Family Species Diptilomiopidae Rhinophytoptus bagdasariani Shev. et Pog.,1985 Status Regime Native range 1st record in Europe A Phytophagous Asia South West 2002, RS RS I2, X11 Rhyncaphytoptus negundivagrans Farkas,1966 Epidermoptidae Epidermoptes bilobatus Rivolta, 1876 Eriophyidae Acaphylla theae (Watt & Mann, 1903) Acaricalus hederae Keifer,1939 Aceria ambrosiae Wilson, 1959 C Phytophagous North America? 1960, HU HU, RS I2, X11 A parasitic/predator Asia- Tropical 1948, CZ CZ I, J Gallus Šefrová and Laštůvka (2005) A Phytophagous Asia 1983, IT IT, ES I2 Camellia A Phytophagous 1997, RS RS I2, X11 Hedera helix A Phytophagous North America North America 1999, RS RS J( J1–J4) Aceria byersi Keifer,1961 A Phytophagous 1981, RS RS X24, X25 Aceria caliberberis Keifer, 1952 A Phytophagous 1998, RS RS I2, X11 Aceria erinea (Nalepa, 1891) A Phytophagous Ambrosia psilostachya, Ambrosia artemisifolia Cucumis sativus Berberis californica, Mahonia dyctiota Juglans regia Fauna Europaea (2009), Pérez Otero et al. (2003) Petanović and Stanković (1999) Petanović (1999) Asia South West 1903, BG BE, BG, CZ, GB, LU, ME, RO, RS Habitat I1, I2, X11, X13 Hosts Ulmus, Quercus macranthera, Salix caprea Acer negundo References Petanović (2004) Petanović (in prep.), Ripka (2007) Petanović (1988), Petanović (1997) Petanović (1998) Petanović (1988) Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) North America Asia South West Invaded countries 178 Table 7.4.1. List and characteristics of the mite species alien to Europe. Status: A: Alien to Europe; C: cryptogenic species. Country codes abbreviations refer to ISO 3166 (see Appendix I). Habitat abbreviations refer to EUNIS (see Appendix II). Family Species Aceria ligustri (Keifer,1943) Status Regime Native range 1st record Invaded in Europe countries North 1995, RS BE, HU, PL, America RS Phytophagous A Phytophagous A Phytophagous Aceria petanovicae Nalepa, 1925 A Phytophagous Aceria sawatchense Keifer, 1965 A Phytophagous Aceria sheldoni (Ewing, 1937) A Phytophagous Asia ? Aceria tristriata (Nalepa, 1890) Aculops allotrichus (Nalepa, 1894) Aculops fuchsiae Keifer,1972 A Phytophagous A Phytophagous A Phytophagous Asia South West North America South America A Phytophagous Aceria neocynarae (Keifer,1939) Aceria paradianthi (Keifer,1952) Aculops gleditsiae (Keifer, 1959). Hosts References I2,FB, X11 Ligustrum ovalifolium , Ligustrum sp. Cynara scolimus Dianthus sp. Petanović (1997), Petanović (1998), Soika and Labanowski (1998), Witters et al. (2003) Fauna Europaea (2009), González Núñez et al. (2002) Anagnou-Veroniki et al. (2008), Fauna Europaea (2009) Fauna Europaea (2009), Fauna Italia, Petanović and Stanković (1999), Ripka (2007) Petanović et al. (1983) North America North America 1998, ES GR, IT-SIC, PT, ES 1987, GR IT, PL, GR Mediterranean East North America 1939, IT FI, GB, HU, IT, RS I2, X11 Syringa 1981, RS RS J (J1–J4) Polygonum douglasii ssp. johnstoni, Polygonum lapatifolium Citrus North America 17th, IT I J100 I, X13 Mijušković and Tomašević (1975) X13 Juglans Petanović (1996), Trotter (1903) 2003, FR DE, FR, GB I1,I2 Fuschia 1993 RS HU, IT, RS X11 Gleditsia triacanthos Deutsche Dahlien, Fuchsien, Gladiolen und Kübelpflanzen, Ostojá-Strazewski (2007) Fauna Italia, Petanović (1993), Petanović (1997), Ripka (2007), Ripka and De Lillo (1997) 179 ES, GR, IT, IT-SAR, ITSIC, ME, PT 1903, RS BG, CZ, GB, LU, ME, RS 1912, RO BG, CZ, RO Mites and ticks (Acari). Chapter 7.4: A Habitat Status Regime A Phytophagous A Phytophagous A Phytophagous A Phytophagous A Phytophagous Calacarus carinatus (Green, 1890) A Phytophagous Asia 1983, IT ES, HU, IT, PL Cecidophyes malifoliae Parrot, 1906 A Phytophagous North America 1991, RS RS Cecidophyopsis hendersoni (Keifer,1954) A Phytophagous North America 1991, RS RS, PL Coptophylla lamimani (Keifer, 1939) A Phytophagous North America 1981, RS IT, RS, ME Cosetacus camelliae Keifer,1945 A Phytophagous North America 1990, ME ES, ME Aculops rhodensis (Keifer,1957) Aculus ligustri Keifer, 1938 Anthocoptes punctidorsa Keifer, 1943 Anthocoptes transitionalis Hodgkiss,1913 North America North America Habitat I, X13 X11, X13 X11, X13 1991, IT IT I2, FB 1989, RS RS X13 I2 X13 J100, J1 I2, FB, X13 I2, J100 Hosts References Citrus Mijušković and Tomašević (1975) Salix alba, Salix elegnos Ligustrum ovalifolium , Ligustrum sp. Ulmus laevis, U. pumila Acer rubrum, A. monspessulanum Camellia, Capsicum, Viburnum Malus x domestica, Aremonia agrimonoides Yucca glauca, Yucca gloriosa Fauna Italia Corylus avellana, Corylus colurna Camelia japonica Fauna Italia, Petanović and Stanković (1999), Ripka (2007) Rigamonti and Lozzia (1999) Glavendekić et al. (2005), Petanović (1997) Fauna Europaea (2009) Petanović and Stanković (1999) Glavendekić et al. (2005), Labanowski (1999), Petanović (2004) Petanović (1988), Rigamonti and Lozzia (1999) Estación Fitopatolóxica do Areeiro (1998), Petanović (1997), Petanović and Stanković (1999) Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) Native range 1st record Invaded in Europe countries Asia 1958, GR ES, GR, IT, IT-SAR, IT-SIC, ME, MT North 1997, HU HU, IT America North 1993, IT HU, IT, RS America 180 Family Species Aculops pelekassi (Keifer, 1959) Family Species Epitrimerus cupressi Keifer,1939 Eriophyes emarginatae Keifer,1939 Status Regime Native range 1st record Invaded in Europe countries California? 1986, ME FR, ME Habitat Phytophagous A Phytophagous North America 1978, RS RS Eriophyes pyri (Pagenstecher, 1857) C Phytophagous Cryptogenic Paraphytoptus chrysanthemi Keifer,1940 Phyllocoptes amaranthi (Corti, 1917) A Phytophagous North America 1903, ME AT, BA, BE, BG, CH, CY, CZ, DE, DK, ES, FI, FR, GB, GR, GR-CRE, HR, HU, IE, LT, LV, MD, MK, MT, NL, NO, PL, PT, RO, RU, SE, SI, YU 1997, RS RS X25,J100 A Phytophagous South America 1981, RS RS J (J1–J4) A Phytophagous Asia- East A Phytophagous Armenia 1952, CZ BG, CZ, DE, IT, NL 1995, RS MK, RS Phyllocoptes azaleae Nalepa, 1904 Reckella celtis Bagdasarian,1975 I2 I, X13,G1 I G G1, X13 References Cupressus sempervirens Prunus emarginata, P. americana, P.domestica Pear, apple, plum Guttierez et al. (1986), Petanović (1993) Petanović (1997), Petanović and Dobrivojević (1987) Chrysanthemum morifolium Amaranthus muricatus, A. retroflexus Rhododendron Petanović (1997), Petanović and Stanković (1999) Celtis caucasiaca, Celtis australis Bebić (1955), Fauna Europaea (2009), Hadžistević (1955), Trotter (1903) Mites and ticks (Acari). Chapter 7.4: C Hosts Petanović et al. (1983) Fauna Europaea (2009), Šefrová and Laštůvka (2005) Petanović et al. (1997) 181 Status Regime Native range 1st record Invaded in Europe countries North 1999, RS HU, PL, RS America Habitat Phytophagous Shevtchenkella erigerivagrans (Davis, 1964) A Phytophagous North America 1989, RS RS Tegolophus califraxini (Keifer, 1938) Vasates quadripedes Shimer 1869 A Phytophagous 1988, IT HU, IT A Phytophagous North America North America 1957, LV HU, LV, RS, PL A parasitic/predator Africa 2004, ES ES E A Parasitic/predator Africa 2004, ES ES E A parasitic/predator A parasitic/predator North America Africa Ixodidae Amblyomma latum Koch, 1844 Amblyomma exornatum Koch, 1844 Dermacentor variabilis (Say, 1821) Hyalomma aegyptium (L., 1758) ?, DK DK 1911, DE AL, BE, BG, CY, DE, ES, FR, GB, GR, GR-CRE, IT, PT, RO, RU X11, X24 Acer negundo. A. negundo var. californicum, A.campestre J (J1–J4) Erigeron strigosus , Taraxacum officinale, Artemisia absinthium I2, X10–X13, Fraxinus X20 angustifolia I2,FB Acer saccharinum, A.pseudoplatanus, A. rubrum G I Reptile, python Reptile, phyton Dog (transmit Lyme disease) Tortoises ( transmit Borellia) References Petanović (in prep.) Petanović and Stanković (1999) Fauna Italia, Ripka (2007), Ripka and De Lillo (1997) Petanović and Stanković (1999), Ripka (2007), Shetchenko and Rupais (1964), Soika and Labanowski (1999) Brotóns and Estrada-Peña (2004) Estrada-Peña (Unpubl.) Brotóns and Estrada-Peña (2004), Feider (1965), Neumann (1911), Robbins et al. (1998), Schulze (1927), Siroky Pet al. (2007) Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) A Hosts 182 Family Species Shevtchenkella brevisetosa (Hodgkiss,1913) Status Hyalomma truncatum Koch 1844 Rhipicephalus rossicus Yakimov & Kolyakimova, 1911 Laelapidae Laelaps echidninus Berlese, 1887 A parasitic/predator Native range 1st record Invaded in Europe countries Cryptogenic 1929, CY CY Habitat Hosts A parasitic/predator Africa 1929 A parasitic/predator Cryptogenic 1940 A parasitic/predator Cryptogenic A parasitic/predator Cryptogenic A parasitic/predator Asia- Tropical 1955, CZ CZ G spiny rat A parasitic/predator North America 1955, CZ CZ C Muskrat A parasitic/predator North America 1955, CZ CZ C, I muskrat F4, F5, F6, F7 Cattle BG, ES-CAN F4, F5, F6, F7 Camels AL, BG, CY, ES-CAN, GR, GRCRE, IT 1956 ES- ES-CAN CAN 1965, RO RO F4, F5, F6, F7 Cattle References Apanaskevich (2003), Schulze and Schlottke (1929) Drenski (1955), Schulze and Schlottke (1929) Apanaskevich (2003), Battelli et al. (1977), Drenski (1955), Rosicky et al. (1960) F4, F5, F6, F7 Cattle Hoogstraal (1956) F4, F5, F6, F7 Domestic animals, hedgehogs, occasionally humans (transmit Crimean congo haemorragic fever) Feider (1965) Šefrová and Laštůvka (2005), Smith et al. (2007), Wharton and Hansell (1957) Šefrová and Laštůvka (2005) Bauer and Whitaker (1981), Šefrová and Laštůvka (2005), Whitaker (2007) 183 Ondatralaelaps multispinosus (Banks, 1909) Listrophoridae Listrophorus americanus Radford, 1944 Regime Mites and ticks (Acari). Chapter 7.4: Family Species Hyalomma anatolicum Koch 1844 Hyalomma dromedarii Koch 1844 Hyalomma excavatum Pomerantsev 1946 Status Regime Native range 1st record Invaded in Europe countries North 2004, CZ CZ America Habitat Hosts parasitic/predator Listrophorus faini Dubinina, 1972 A parasitic/predator North America Listrophorus validus Banks, 1910 A parasitic/predator North America A parasitic/predator Asia- Tropical 1952, CZ CZ G, I, J tropical rat, rat, mices, little rodents A parasitic/predator C&S America 1948, CZ CZ, DK G, I, J birds, mammals A parasitic/predator North America 2004, CZ CZ C, 1 Muskrat Bauer and Whitaker (1981), Šefrová and Laštůvka (2005),Whitaker (2007) A Phytophagous South Africa 2002, RS RS I2, X11 Hedera helix Glavendekić et al. (2005) A s 2005, RS RS A Phytophagous North America North America Ornithonyssus bursa (Berlese) Myocopidae Myocoptes ondatrae Lukoschus & Rouwet, 1968 Phytoptidae Phytoptus hedericola Keifer, 1943 Setoptus strobicus Keifer,1966 Sierraphytoptus alnivagrans Keifer, 1939 muskrat 2004, CZ CZ C, I muskrat 2004, CZ CZ C, I muskrat 2007, RS RS G3F, X25, X11 Pinus strobus G1 Bauer and Whitaker (1981), Šefrová and Laštůvka (2005),Whitaker (2007) Bauer and Whitaker (1981), Šefrová and Laštůvka (2005),Whitaker (2007) Bauer and Whitaker (1981), Šefrová and Laštůvka (2005),Whitaker (2007) Bowman et al. (2003), Cole et al. (2005), Easterbrook et al. (2008), James (2005), Šefrová and Laštůvka (2005), Whitaker (2007) Berggren (2005), Denmark and Cromroy (2008), Gjelstrup and Møller (1985), James (2005) Petanović (in prep.) Alnus glutinosa Petanović (in prep.) Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) A Macronyssidae Ornithonyssus bacoti (Hirst, 1913) C, I References 184 Family Species Listrophorus dozieri Redford, 1994 Family Species Trisetacus chamaecypari Smith, 1977 Status Regime Native range 1st record Invaded in Europe countries North 2002 GB America Phytophagous A parasitic/predator South America Amblyseius (Neoseiulus) californicus (McGregor 1954) A parasitic/predator Typhloctonus squamiger Wainstein 1960 A A Phytoseiidae Phytoseiulus persimilis Athias-Henriot 1957 References I2 Chamecyparis, Ostojá-starzewski and lawsonianna, Halstead (2006), Smith et al. C. nootkaensis, (2007) Cupressus macrocarpa, Juniperus virginiana 1974,CZ BG, CZ, BE, DE, ES, GB, IT I Predator of Tetranychus North America 1991, GB BG, CZ, GB, IT I Predator of Tetranychus Phytophagous Cryptogenic 1991, IT IT I Acer platanoides, Prunus serratulata parasitic/predator North America Unknown NL, NO, PL, IT J house dust Bartlett (1992), Croft et al. (1998), Easterbrook (1996), EPPO (2002), Garcia Mari and Gonzalez-Zamora (1999), Helle and Sabelis (1985), McMurtry and Croft (1997), Šefrová and Laštůvka (2005) Croft et al. (1998), Easterbrook (1996), EPPO (2002), Garcia Mari and Gonzalez-Zamora (1999), Helle and Sabelis (1985), McMurtry and Croft (1997), Šefrová and Laštůvka (2005) Rigamonti and Lozzia (1999) Bigliocchi and Maroli (1995), Eriksson (1990), Hughes (1976), Musken et al. (2000), Piotrowski (1990), Razowski (1997), Thind and Clarke (2001) 185 Pyroglyphidae Dermatophagoides evansi Fain, Hughes et Johnston, 1967 Hosts Mites and ticks (Acari). Chapter 7.4: A Habitat Status Regime Native range 1st record in Europe Invaded countries Hosts References polyphagous: crops, vegetables, fruits and leaves CAB-International (1986), Fan and Petit (1998), Gerson (1992), Heungens (1986), Natarajan (1988), Parker and Gerson (1994), Raemaekers (2001) A Phytophagous Sri Lanka IT, 1965 DK, ES, GB, IT, IT-SAR, IT-SIC, NL, RO, RS, BE, DE I A Phytophagous North America I2, J100 Citrus, Camellia sinensis CAB-International (1986), Childers et al. (2003a), Childers et al. (2003b) A Phytophagous I2, J100 Phytophagous Brevipalpus obovatus Donnadieu, 1875 A Phytophagous North America Citrus, ornamentals Polyphagous, Citrus, Gardenia, Hibiscus, Ilex, Ligustrum; Ficus, Phoenix, Prunus Citrus, Camellia, Coffea, Mentha, Solanum Childers et al. (2003a) A North America Tropical IT, 1998 CY, FR, GRCRE, GR, IT, IT-SAR, ITSIC, PT, IL Unknown BG, FR, GR, RO IT, 1998 ES, GR, IT, NL Brevipalpus russulus (Boisduval 1867) A Phytophagous C&S America Tenuipalpidae Brevipalpus californicus (Banks, 1904) Brevipalpus lewisi (McGregor 1949) Brevipalpus phoenicis (Geijskes 1939) IT, 1986 AT, FR, DE, IL, NL, SP, RS, BE, BA, BG, HR, CY, GR, IT, PT, RO, UA 1867, FR BE, DE, FR, GB, GR, NL, PT, UA I2, J100 I2 J100 Cactaceae Childers et al. (2003a), Childers et al. (2003b) CAB-International (1986), Childers et al. (2003a), Childers et al. (2003b), Glavendekić et al. (2005), Manson (1967) Denmark (1978) Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) Habitat 186 Family Species Tarsonemidae Polyphagotarsonemus latus (Banks, 1904 Family Species Tenuipalpus caudatus (Dugès 1834) Tenuipalpus pacificus Baker 1945 A Phytophagous A Phytophagous A Phytophagous A Phytophagous A Phytophagous A Phytophagous A Phytophagous A Phytophagous A Phytophagous A Phytophagous A Phytophagous A Phytophagous Native range 1st record Invaded in Europe countries Tropical Unknown FR, GR, IT, PT C&S Unknown DE, GB, NL, America RO, RS C&S America North America North America Habitat I2, J100 J100 Hosts References Citrus Manson (1967) Orchids: Phalaenopsis, etc.. Denmark (1968), Glavendekić et al. (2005), Manson (1967) 1990, PT- PT-MAD MAD 2004, RS AL, MK, RS I Citrus, Carica Carmona (1992) I Populus Glavendekić et al. (2005) 1970, HU FR, HU I2 Coniferous Bozai (1970), Migeon (2003) AsiaTemperate 1974, HU HU G Picea Bozai (1974) C&S America Asia-Tropical 2001, ES ES, PT I Citrus Garcia et al. (2003) 2001, ES ES I Citrus Garcia et al. (2003) North America? AsiaTemperate 1972, IT IT-SAR, ITSIC, IT, PT 1985, IT IT, NL I2 I2 North America AsiaTemperate 1964, PL PL I2 1990, NL NL I2 Quercus robur, Rigamonti and Lozzia (1999) Castanea Azalea, Rota and Biraghi (1987) Rhododendron, Camelia Larix Boczek (1964), Doboz et al. (1995) Juniperus Vierbergen (1990) chinensis 187 Oligonychus laricis Reeves, 1963 Oligonychus perditus Pritchard & Baker, 1955 Regime Mites and ticks (Acari). Chapter 7.4: Tetranychidae Eotetranychus lewisi (McGregor, 1943) Eotetranychus weldoni (Ewing, 1913) Eurytetranychus admes Pritchard & Baker, 1955 Eurytetranychus furcisetus Wainstein, 1956 Eutetranychus banksi (McGregor, 1914) Eutetranychus orientalis (Klein, 1936) Oligonychus bicolor (Banks, 1894) Oligonychus ilicis (McGregor, 1917) Status Status Regime Native range 1st record Invaded in Europe countries North 2004, ES ES America Habitat Phytophagous A Phytophagous A Phytophagous Panonychus citri (McGregor, 1916) A Phytophagous Asia 1950, FR AL, BG, ES, ES-CAN, FI, FR, GB, GRCRE, GR, HR, HU, IT, IT-SAR, IT-SIC, MK, NL, NO, PL, PT, RO, SI, UA, YU Petrobia (Tetranychina) lupini (McGregor, 1950) Schizotetranychus bambusae Reck, 1941 Schizotetranychus parasemus Pritchard & Baker, 1955 A Phytophagous North America 1968, GR GR I A Phytophagous 2001, FR FR I2 A Phytophagous AsiaTemperate North America 1964, PL PL I North America C&S America References I1 Persea americana 1984, PL PL G 1988, FR- FR-COR COR I2 Quercus robur Kropczynska (1984), Doboz et al. (1995) polyphagous: Bolland et al. (1998) Quercus, Juglans, Eucalyptus Citrus Balevski (1967), Bernini et al. (1995), Bowman and Bartlett (1978), Bozai (1970), Ciampolini and Rota (1972), Ciglar and Barić (1998), Delrio et al. (1979), Emmanouel and Papadoulis (1987), Fauna Europaea (2009), Garcia Mari and de Rivero (1981), Jeppson et al. (1975), Mijušković (1953), Pande et al. (1989), Petanović (1980), Rambier (1958), Vacante (1983), Vappula (1965), Vierbergen (1989) Lupinus., Hatzinikolis (1970), Fragaria, Papaioannou-Souliotis et al. Poaceae (1993) Bambusaceae Auger and Migeon (2007), Migeon et al. (2004) Cynodon, Boczek and Kropczynska Poaceae (1964) I1, I2 Alcázar et al. (2005) Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) A Hosts 188 Family Species Oligonychus perseae Tuttle, Baker & Abbatiello, 1976 Oligonychus pritchardi (McGregor, 1950) Oligonychus punicae (Hirst, 1926) Family Species Stigmaeopsis celarius Banks, 1917 Status Regime Native range 1st record Invaded in Europe countries Asia1985, FR BE, FR, GB, Temperate NL Habitat Tetranychus canadensis (McGregor, 1950) A Phytophagous North America 1954, HU HU, PL Tetranychus evansi Baker & Pritchard, 1960 A Phytophagous C&S America 1991, PT ES, ES-BAL, ES-CAN, FR, IT, PT, PTMAD, PT Tetranychus kanzawai Kishida, 1927 A Phytophagous Asia-Tropical 1966, GR BE, GR Tetranychus macfarlanei Baker & Pritchard, 1960 Tetranychus mcdanieli McGregor, 1931 A Phytophagous Asia-Tropical 1989, ES- ES, ES-CAN CAN I A Phytophagous North America 1981, FR FR I A Phytophagous Tropical 1989, ES- ES-CAN CAN I A Phytophagous North America 1964, PL PL I A Phytophagous North America 1986, GR GR I Tetranychus neocaledonicus André, 1933 Tetranychus sinhai Baker, 1962 Tetranychus tumidellus Pritchard & Baker, 1955 I, X11, X22, Bambusaceae X23, X24, X25 I2 I, J100, X J100 Auger and Migeon (2007), Bolland et al. (1998), OstojaStarzewski (2000), Witters et al. (2003) Boczek and Kropczynska (1964), Hetenyi (1954) Polyphagous: Rosaceae, Carya, Corylus Solanaceae Castagnoli et al. (2006), Ferragut and Escudero (1999), Ferragut et al. (1997), Ferreira and Carmona (1995), Migeon (2005), Migeon (2007) Saxifragaceae: Hance et al. 1998, Hydrangea Hatzinikolis (1968), Hatzinikolis (1986) Musa, Pande et al. (1989) Ipomoea, etc Vitis, Acer, Lonicera, Fragaria, Ulmus, etc. Polyphagous: Citrus, Fabaceae Helianthus, Agropyron, Prunus Sambucus, Passiflora, Solanum Rambier (1982) Ferragut and Santonja (1989) Boczek (1964) Hatzinikolis (1986) 189 Phytophagous References Mites and ticks (Acari). Chapter 7.4: A Hosts Regime A Phytophagous A parasitic/predator Native range 1st record Invaded in Europe countries C&S 1981, GR GR, GR-CRE America Asia 1964 RS AL, BG, CZ, DE, DK, EE, ES, FI, FR, GB, GR, HU, IE, IT, ITSAR, IT-SIC, MT, PL, PT, RO, RS, RU, SI, SK Habitat I2, X J Hosts References Plumeria, Lonicera, exotic Fabaceae Hatzinikolis (1986) bee parasite Colin (1982), De Rycke et al. (2002), Griffiths and Bowman (1981), Morse and Goncalves (1979), Ruttner (1983), Ruttner and Marx (1984) Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) Varroidae Varroa destructor Anderson & Trueman, 2000 Status 190 Family Species Tetranychus yusti McGregor, 1955 Table 7.4.2. List and characteristics of the mite species alien in Europe. Country codes abbreviations refer to ISO 3166 (see Appendix I). Habitat abbreviations refer to EUNIS (see Appendix II). Family Species Argasidae Argas reflexus (Fabricius, 1794) Aculus hippocastani (Fockeu, 1890) Eriophyes canestrinii (Nalepa, 1891) Glycyphagidae Glycyphagus domesticus (De Geer, 1778) Ixodidae Hyalomma scupense Delpy 1946 parasitic/ predator Native range Europe Phytophagous Alps 1st record Invaded countries in Europe 19th , DE AT, BE, BG, CH, CZ, DE, DK, ES, FR, GB, GR, IT, PL, RO, RU, UA 1952, CZ AT, CZ, IT, RS Phytophagous Mediterranean 1901, RO AT, BG, CZ, CY, East DE, FR, HU, IT, LT, LV, PL,GB Phytophagous Mediterranean 1907, CZ BG, CZ, IT, RO, East FR Phytophagous Mediterranean 1998, RS AT, BG, CZ, DE, region HU, IS, PL Habitat* Hosts References J Rat F2 Rododendron Petanović and Stanković (1999) ferrugineum Syringa Fauna Europaea (2009) I2, X11 G1,G4, X11 X 11, X24 Aesculus Dautel and Kahl (1999) Fauna Italia Buxus Petanović (1998) sempervirens detrivorous Europe Unknown DK, FÖ, IT, NO, PL, SE J1, J2 Houes dust Bigliocchi and Maroli (1995), Hughes (1976), Musken et al. (2000), Piotrowski (1990), Razowski (1997), Thind and Clarke (2001) parasitic/ predator Europe Unknown AL, BG, ES, ESCAN, FR, GR, HR, IT, IT-SAR, IT-SIC, MK, RU, RS, YU J Cattle Morel et al. (1977) Mites and ticks (Acari). Chapter 7.4: Eriophyidae Aceria alpestris (Nalepa,1892) Aceria loewi (Nalepa, 1890) Regime 191 1st record Invaded countries Habitat* in Europe Mediterranean Unknown BE, CH, CZ, DE, J region DK, GB, IE, NL, NO, PL Phytophagous Alps parasitic/ predator 1912 BA,DE, GB, HR, SI Mediterranean 1993, CZ CZ, GB, GR, IT,PT Hosts References Dogs Černý (1985), Fauna Europaea (2009), Garben et al. (1980), Sibomana et al. (1986) I2 Larix Fauna Europaea (2009) I Predator of Tetranychus Albajes et al. (1999), Bartlett (1992), EPPO (2002), Šefrová and Laštůvka (2005), Sengonca et al. (2004), van Houten and van Stratum (1993), van Houten and van Stratum (1995) Maria Navajas et al. / BioRisk 4(1): 149–192 (2010) Phytoptidae Trisetacus laricis \(Tubeuf 1897) Phytoseiidae Amblyseius (Iphesius) degenerans (Berlese 1889) Native range 192 Family Regime Species Rhipicephalus sanguineus parasitic/ (Latreille 1806) predator A peer reviewed open access journal BioRisk 4(1): 193–218 (2010) RESEARCH ARTICLE doi: 10.3897/biorisk.4.56 BioRisk www.pensoftonline.net/biorisk Longhorn beetles (Coleoptera, Cerambycidae) Chapter 8.1 Christian Cocquempot1, Åke Lindelöw2 1 INRA UMR Centre de Biologie et de Gestion des Populations, CBGP, (INRA/IRD/CIRAD/Montpellier SupAgro), Campus international de Baillarguet, CS 30016, 34988 Montférrier-sur-Lez, France 2 Swedish university of agricultural sciences, Department of ecology. P.O. Box 7044, S-750 07 Uppsala, Sweden Corresponding authors: Christian Cocquempot (cocquemp@supagro.inra.fr), Åke Lindelöw (Ake.Linde- low@ekol.slu.se) Academic editor: David Roy | Received 28 December 2009 | Accepted 21 May 2010 | Published 6 July 2010 Citation: Cocquempot C, Lindelöw Å (2010) Longhorn beetles (Coleoptera, Cerambycidae). Chapter 8.1. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 193–218. doi: 10.3897/biorisk.4.56 Abstract A total of 19 alien longhorn beetle species have established in Europe where they presently account for ca. 2.8 % of the total cerambycid fauna. Most species belong to the subfamilies Cerambycinae and Laminae which are prevalent in the native fauna as well. The alien species mainly established during the period 1975–1999, arriving predominantly from Asia. France, Spain and Italy are by far the most invaded countries. All species have been introduced accidentally. Wood-derived products such as wood- packaging material and palettes, plants for planting, and bonsais constitute invasive pathways of increasing importance. However, only few species have yet colonized natural habitats outside parks and gardens. Present ecological and economical impacts, and future trends are discussed. Keywords Cerambycidae, Europe, Introductions, Establishments, Biogeographical origins, Pathways, Impacts 8.1.1 Introduction The coleopteran family Cerambycidae (longhorn beetles) is currently classified in the superfamily Chrysomeloidea, along with the families Vesperidae and Distenidae (Hunt et al. 2007, Szeoke and Hegyi 2002). Cerambycidae is a large family comprising about Copyright C. Cocquempot, Å. Lindelöw. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 194 Christian Cocquempot & Åke Lindelöw / BioRisk 4(1): 193–218 (2010) 40000 described species worldwide. Longhorn beetles are all phytophagous. Larvae may be found in conifer, deciduous and fruit trees, in bushes and herbaceous plants. They are mainly xylophagous borers of living, decaying or dead wood. Some species also bore small twigs, roots or fruit endocarps. They usually have a long period of larval development, some species being capable of developing in woody material a long time after the death of the tree. They are thus very susceptible to transport with wood products, facilitating their introduction and establishment. The oldest known introduction of a longhorn beetle from one continent to another was probably that of the house borer, Hylotrupes bajulus (L., 1758), which was first described by Linnaeus from both Europe and ‘America septentrionali’ (von Linnaeus 1758). Since a study by Duffy in 1953 (Duffy 1953a) for Great Britain, there has been no further large synthesis of the alien cerambycid species introduced to Europe. Since 1999, the development of research interests in the Asian longhorn beetles, Anoplophora spp., in North America has raised awareness of the risks presented by cerambycid importation and provided a baseline for subsequent studies (Haack et al. 2000, Haack et al. 2010). There is an urgent need for a comprehensive literature review of the alien cerambycids that have successfully established in Europe. The exponential growth in the volume of international trade in both horticulture and forestry has allowed an increasing number of wood products and ornamental plants potentially containing cerambycids to arrive in Europe. More than 250 species have been introduced to Europe or moved within Europe since the middle of the 18th century (Cocquempot 2007) but most of them never established. We have identified 19 species alien to Europe that have established in Europe but have not yet been eradicated. 8.1.2 Taxonomy of the Cerambycid species alien to Europe Taxonomy in Cerambycidae sensu lato is not well established (e.g., Hunt et al. 2007, Lawrence and Newton 1995, Napp 1994, Özdikmen 2008, Sýkorová 2008) but a general consensus exists about the presence in Europe of 7 subfamilies, namely Cerambycinae, Lamiinae, Lepturinae, Necydalinae, Prioninae, Spondylidinae, and Vesperinae (the latter being sometimes considered as a valid family). A total of 677 native species are known to occur in Europe (Althoff and Danilevsky 1997, Fauna Europaea), being largely dominated by 3 subfamilies (Lamiinae- 343 spp.; Cerambycinae- 158 spp.; Lepturinae- 130 spp.) which account for 93.2% of the total. The 19 alien species established in Europe belong to only 3 of these subfamilies, Cerambycinae, Laminae and Prioninae (Table 8.1.1). The alien species are mostly represented by the subfamily Cerambycinae, followed by Lamiinae but the relative proportion of aliens compared to the total cerambycid fauna is still limited (<6%) in these two subfamilies. By contrast, the proportion of aliens is much more important in Prioninae with 2 species adding to 10 native ones (Fig. 8.1.1.). In addition, Parandrinae, a subfamily which is not represented in the native European entomofauna, is represented by Parandra brunnea, a North American species introduced in Germany (Nüssler 1961). Longhorn beetles (Coleoptera, Cerambycidae). Chapter 8.1 195 Figure 8.1.1. Relative importance of the subfamilies of Cerambycidae in the alien and native entomofauna in Europe. Subfamilies are presented in a decreasing order based on the number of alien species. Species alien to Europe include cryptogenic species. The number over each bar indicates the number of species observed per family. Two more alien species have been introduced and established in Israel, Batocera rufomaculata (DeGeer, 1775) (Bytinski-Salz 1956, Chikatunov et al. 1999, Sama et al. 2010) and Xystrocera globosa (Olivier, 1795) (Chikatunov et al. 2006, Sama et al. 2010), but they have not yet spread to Europe and were not considered in Table 8.1.1. Table 8.1.2 gives a list of species of European origin introduced through human activity in another part of Europe (aliens in Europe). These species are mostly of Mediterranean origin introduced in more northern areas and species from Continental Europe introduced to the Atlantic islands. 8.1.3 Major biological characteristics of the cerambycid species alien to Europe Lepturinae but also Prioninae and Parandrinae share some biological characteristics that reduce their probability of introduction. Larvae in these subfamilies develop in decaying wood and are rarely imported with wood products or living plants. Interceptions have shown that they are mainly introduced through accidental importation in industrial packages or in stocks of perishable vegetables. Only a few species of Lepturinae (Tribe Rhagiini, and some Lepturinii) developing on recently felled trees are likely to be successfully introduced through the wood trade. The importation of living potted plants is also a potential new pathway for Prioninae. Cerambycinae and Lamiinae seem more predisposed to introduction. Most species develope in living plants and several Cerambycinae undertake their entire life-cycle in dead wood, e.g. the cosmopolitan tribe Hesperophanini and the species Hylotrupes bajulus and Gracilia minuta. Thus, Cerambycinae and Lamiinae can easily survive 196 Christian Cocquempot & Åke Lindelöw / BioRisk 4(1): 193–218 (2010) throughout the importation process of living plants including bonsai (e.g. Anoplophora chinensis (Cocquempot 2007, EPPO 2006, van Rossem et al. 1981, Schmidt and Schmidt 1990)), recently felled logs and other non-aged wood products (e.g. Anoplophora glabripennis (Cocquempot et al. 2003, Haack et al. 2000), Monochamus spp. (Cocquempot 2007, Cocquempot (Unpubl.), Duffy 1953a), Chlorophorus annularis (Cocquempot 2007) and Phoracantha spp. (Cocquempot and Debreuil 2006)). Species in the genera Hesperophanes, Trichoferus, and Stromatium can emerge from wood products even several years after importation (Duffy 1953a). Once a population is introduced, the capability for natural dispersal constitutes an important factor for establishment success. Although our knowledge about the dispersal behaviour of alien longhorn beetles is still rather limited and mostly concerns only a few species of recent invaders such as Anoplophora glabripennis (Smith et aol. 2001) and A. chinensis (Adachi 1990, Komazaki and Sakagami 1989), this variable is important when designing an eradication attempt (MacLeod et al. 2002). 8.1.4 Temporal trends of introduction in Europe of alien Cerambycids Figure 8.1.2 presents the temporal changes in the records of Cerambycid species alien to Europe from 1492 to 2007. Cerambycids have tracked trade routes since the beginning of overseas communications. The first species to have moved are those which live in dry wood and undergo a long stage of larval development. These species have become cosmopolitan (e.g. Hylotrupes bajulus) or nearly so (e.g. Stromatium spp.). With the increased speed of international transport from 1850 to 1925, species with shorter life cycles were able to reach Europe alive and become established, e.g. Neoclytus acuminatus (Reineck 1919, Sama 2002, Tassi 1969). Later, only two species were introduced from North America to Europe via the US effort to supply extra furniture and increase military material after the 1st World War (i.e., Parandra brunnea, Neoclytus acuminatus). Subsequently, 50 years passed until a second wave of introduction arrived alongside with the rapid development of international exchange of goods and transportation after the 2nd World War. During the recent period, two further species have been detected in the wild - Anoplophora chinensis in 2000 in Italy (Colombo and Limonta 2001) and A. glabripennis in 2001 in Austria (Dauber and Mitter 2001). The number of interceptions of Cerambycids is still increasing throughout Europe. However, more effective control at borders is like to have reduced establishments following interception or introductions. The importation of exotic plants also offers opportunities for introduction but also constraints the establishment of some alien species. For example, Phoracantha spp. could not have been introduced without the importation and mass cultivation of its host plants, Eucalyptus spp. in the Mediterranean basin. In south-eastern France, an Australian cerambycid, Bardistus cibarius (Newman, 1841) could survive only on its original host plant, an introduced grass tree (Xanthorrhoea sp., Xanthorrhoeaceae); the beetle population disappeared immediately after the infested host plants were removed (Cocquempot 2007). The case of Batocera Longhorn beetles (Coleoptera, Cerambycidae). Chapter 8.1 197 Figure 8.1.2. Temporal changes in the mean number of new records per year of Cerambycid species alien to Europe from 1492 to 2007. The number over each bar indicates the absolute number of species newly recorded per time period. rufomaculata (DeGeer, 1775) found in Munster’s Zoo (Germany) is similar (Cocquempot 2007) although this tropical species has established in Israel since at least 1948 (Bahillo de la Puebla and Iturrondobeitia-Bilbao 1995, Plavilstshtikov 1934, Sama et al. 2010). The combination of importation of longhorn beetle species with their specific host plant or groups of plants followed by establishment is rare. However the establishment of A. chinensis is an exception. Other species are frequent intercepted at border controls, e.g. Mimectatina meridiana (Matsushita, 1933) with Cycas fruits from Japan (Cocquempot 2007) or Trichoferus campestris (Faldermann 1835) with Salix timber from China (Cocquempot 2007). The degree of polyphagy is also an important factor in the likelihood of establishment. Polyphagous species appear to have a higher potential to establish than oligophagous and monophagous species. The large number of hosts utilised by Anoplophora spp. (Cocquempot et al. 2003, Hérard and Roques 2009, Maspero et al. 2007a) is a main factor in the difficulty in eradicating this species for example. These difficulties appear much less important for oligophagous species such as Callidiellum rufipenne (Bahillo and Iturrondobeitia-Bilbao 1995, Campadelli and Sama 1988, Plavilstshtikov 1934) 198 Christian Cocquempot & Åke Lindelöw / BioRisk 4(1): 193–218 (2010) or Phoracanthine species. It is also the case for the North American wood borer Saperda candida (Fabricius, 1787), which was introduced in Germany in 2008 but apparently did not established yet (EPPO 2008, Nolte Krieger 2008). By contrast, Monochamus species have a regime close to polyphagy, including a large number of conifer species, and may spread throughout Europe. There is no example of establishment in Europe of a strictly monophagous exotic long-horned beetle. Species with a limited host range do not seem to be capable of going beyond the interception or introduction stage, e.g. Bardistus cibarius (Cocquempot 2007). 8.1.5 Biogeographic patterns of the cerambycid species alien to Europe Alien species established in Europe mostly originated from Asia, followed by Africa (Figure 8.1.3). The region of origin appears to depend on the major trade routes developed by each country. Some North African species have colonized Mediterranean countries such as Spain, France, and Malta for example. Other African species have often been intercepted but only Phryneta leprosa has established in Malta where the climate is favourable for development (Mifsud and Dandria 2002). Long-established trade routes between Iberian countries and South American countries have resulted in some historic, isolated establishments in the Spanish and Portuguese Atlantic Islands but with a limited risk of further expansion (Lemos-Perreira 1978, Méquignon 1935). With the numerous interceptions in the U.K (Duffy 1953a) together with the colonial trade routes with African and Asiatic countries, it is surprising that only Trinophylum cribratum has established to date (Gilmour 1948); the incompatible climate may negate the development of tropical and subtropical species. Two species native to North America, Parandra brunnea and Neoclytus acuminatus, also colonized Europe at the beginning of the last century. The first species is well established but restricted to Dresden (Germany) (Nüssler 1961). The second is widely established in the Mediterranean area but its populations appear to be declining (Brustel et al. 2002). Beside these two species, there have been no further establishments originating from North America; the pathway of transported material is mainly in the reverse direction, from Europe to America. Some Australian species have reached Europe but only those using Eucalyptus (Phoracantha spp.) have successfully established (Cocquempot and Sama 2004) and only in areas newly planted with these fast-growing tree species. The large differences in species composition between the floras of Australia and Europe probably accounts for the failure of Australasian longhorn beetles such as in Bardistus cibarius on Xanthorrhoea sp. (Cocquempot 2007) to establish. Recent increases in commercial traffic from Asia (especially China) to Europe has accounted for the introduction of a number of new species of cerambycids. Striking examples are Callidiellum rufipenne which has recently established in Spain (Bahillo de la Puebla and Iturrondobeitia-Bilbao 1995) and Italy (Campadelli and Sama 1988), Anoplophora glabripennis and A. chinensis which can be considered as established or Longhorn beetles (Coleoptera, Cerambycidae). Chapter 8.1 199 Figure 8.1.3. Origin of the Cerambycidae species alien to Europe not eradicated in several countries (Hérard and Roques 2009, Maspero et al. 2007a), Psacothea hilaris (Pascoe, 1857) under eradication in Italy (Cocquempot 2007, Jucker et al. 2006), and Monochamus alternatus Hope, 1842 intercepted a number of times in Germany (Cocquempot 2007) and France (Cocquempot Unpubl.) but not yet established. A final case, Xylotrechus stebbingi, is less clear. It is believed that an initial introduction from its native area of central Asia to Asia Minor was followed by a step-wise expansion into southern Europe and North Africa (Cocquempot and Debreuil 2006, Sama 2002, Šefrová and Laštůvka 2005). Alien cerambycid species are not evenly distributed throughout Europe. Large differences in the number of aliens are apparent between countries, France, Italy and Spain being by far the most invaded (Figure 8.1.4). 8.1.6 Main pathways of introduction to Europe of alien cerambycid species All alien longhorn beetles established in Europe have been introduced accidentally; there are no examples of a successful, deliberate introduction. The principal pathways of arrival have been identified and presented by Frank 2002 and each relates to the import of immature stages that subsequently emerge as adults. There are relatively few records of living adults imported with vegetables or fruits although Eucalyptus beetles, Phoracantha recurva, were found in a cluster of bananas (Bosmans 2006). The longest established pathway is timber importation for house construction (Hylotrupes bajulus) or building furniture (e.g. Trichoferus spp., Stromatium spp. and Chlorophorus annularis arriving with bamboo- made objects (Cocquempot 2007)). Species 200 Christian Cocquempot & Åke Lindelöw / BioRisk 4(1): 193–218 (2010) Figure 8.1.4. Comparative colonization of continental European countries and islands by Cerambycidae species alien to Europe. Archipelago: 1 Azores 2 Madeira 3 Canary islands. introduced through this pathway have traditionally required a long life cycle but more rapid travel now enables the introduction of species with a one year life cycle. The second pathway is via the importation of timber for pulp (e.g., for Phoracantha spp.). A third, more recent, pathway concerns wood packages, palettes and other wood-derived products (e.g., for Anoplophora glabripennis) (Hérard and Roques 2009). The final pathway is the importation of plants for planting in nurseries, including the bonsai industry, which has resulted in the arrival of species such as Anopolophra chinensis (Cocquempot 2007, EPPO 2006, van Rossem et al. 1981, Schembri and Sama 1986), Callidiellum rufipenne and Bardistus cibarius. All pathways are still prevalent but they vary in importance. Most recent interceptions (from the end of the 20th Century) have related to wood-manufactured products (e.g. Chlorophorus annularis and Trichoferus campestris). Importation of Eucalyptus wood for pulp has also resulted in the introduction of a second species of Phoracantha, P. recurva (Miquel 2008). If such importations continues a number of Longhorn beetles (Coleoptera, Cerambycidae). Chapter 8.1 201 additional species of this genus, which are mainly related to Eucalyptus (Wang 1995), are expected to arrive. Since their first usage, wood packaging and palettes have constituted an important introduction pathway. The source material spends sufficient time as logs without sanitary controls to be colonized by longhorn beetles. When the wood is turned into packages or palettes, infestation occurs mainly as unnoticed early stages (eggs or firstinstar larva). Development continues in the woody material during importation and emergence of adults occurs often unnoticed in warehouses, weeks or months after arrival. This is the case for A. glabripennis, P. hilaris and M. alternatus which may already complete their entire lifecycle before the source wood is processed or destroyed. Wood package is often produced using low quality timber often colonized by longhorn beetle species, which is increasing its potential as a vector. Other, less significant, introduction pathways have also been identified, yet they typically only transported one or a few individuals which fail to establish. The introduction route is unknown for other species such as Acanthoderes jaspideus (Méquignon 1935), Oxymerus aculeatus (Alluaud 1935), Deroplia albida, and Phryneta leprosa (Mifsud and Dandria 2002) but they may be related to the uncontrolled importation of wild plants. Natural range expansion cannot be ruled out for a few species which have a nearby native range, e.g. Lucasianus levaillantii (Mayet 1905, Pellegrin and Cocquempot 2001) and Xylotrechus stebbingi (Šefrová and Laštůvka 2005) originating from North Africa and the Middle East, respectively. 8.1.7 Ecosystems and habitats invaded in Europe by alien cerambycid species Although all natural or artificial terrestrial ecosystems and anthropogenic areas which contain trees, bushes or wood products are potentially occupied by alien longhorn beetles, establishment in Europe is concentrated in man-made habitats to date, especially in parks and gardens (Figure 8.1.5). To date, only the two clytine beetles, Neoclytus acuminatus and Xylotrechus stebbingi, have colonized natural habitats. X. stebbingi is very common on Eucalyptus cut wood in Crete (Sama 2002) for example and may be related to the polyphagous nature of these two species. Other polyphagous species such as Anoplophora spp. also have the potential to live in urban areas, in cultivated lanes (e.g. planted with poplars) as well as in natural forests where potential host plants occur. However, dispersal from man-made habitats to natural forests appears to be a slow process. For the first twenty-two years since its arrival in North America, A. glabripennis has been restricted to trees in urban areas until 2008 when it was found in natural forests dominated by Acer trees (Haack et al. 2010). Although such a process has not yet been observed in Europe, there is a strong risk that Anoplophora spp. will spread to naturally-forested landscapes, if the ongoing eradication attempts in Austria, Germany, France and Italy are unsuccessful. The expansion of oligophagous species is inevitably more dependant on the presence of suitable host plants. Those using largely- planted trees can spread more easily. 202 Christian Cocquempot & Åke Lindelöw / BioRisk 4(1): 193–218 (2010) Figure 8.1.5. Main European habitats colonized by the established alien longhorn beetles. The number over each bar indicates the absolute number of alien longhorn beetles recorded per habitat. Note that a species may have colonized several habitats. Thus, Phoracantha spp. that live only in eucalypt trees have colonized ornamental tree plantations in urban areas as well as old plantations such as those found on the Mediterranean islands and in neighbouring countries, and industrial plantations created for paper pulp. Other established species mostly have a distribution restricted to Mediterranean and Atlantic islands. In these areas, anthropogenic ecosystems are mainly colonized. A species of considerable concern with conifer forests is Monochamus alternatus, which could potentially become established in coniferous plantations and forests and subsequently transfer the pine wood nematode (Bursaphelenchus xylophilus Steiner & Buhrer, 1934). 8.1.8 Ecological and economic impact of alien cerambycid species Although there is concern about the potential ecological impact of the invasive longhorn beetles N. acuminatus and X. stebbingi, there is no measure of their impact on trees or any estimation of possible competitive displacement of the native fauna. The ecological impact of Anoplophora species may also be important if they establish in European forests. Anoplophora could compete with other arthropods occupying the same niche, but they also create niches for other arthropods that live in tunnels in decaying wood or compete with other saproxylic beetles. The joint introduction Longhorn beetles (Coleoptera, Cerambycidae). Chapter 8.1 203 and establishment of the Citrus longhorn beetle, A. chinensis, and its parasitoid, Aprostocetus anoplophorae Delvare, 2004, exemplifies the potential risk of adaptation of imported parasitoids which themselves might not specialise on the native fauna (Delvare et al. 2004). Although the ecological niche occupied by an alien species may be vacant there remains a risk of secondary infection resulting from their damage. For example, secondary infestation by the pine wood nematode vectored by Monochamus spp. (Evans et al. 2008, Kawai Miho et al. 2006) may cause serious impacts to coniferous trees in all landscapes. M. alternatus has only been intercepted in Germany and France (Cocquempot 2007, Cocquempot (Unpubl.)); yet the pine wood nematode which it vectors was recorded from Portugal in 1999 (Mota et al. 1999). After having been contained for several years in a limited area, the nematode has spread throughout Portugal, as well as being eradicated following incursions into Spain in 2008 and Madeira in 2009. A novel association with the native species, M. galloprovincialis (Villiers 1967) has also been reported. The expansion as well as new introductions of the pine wood nematode could potentially have a substantial level of economic impact in all areas of coniferous cultivation in Europe. Other economic impacts are mainly associated with ornamental trees in urban areas, cultivated trees such as poplars and eucalypts and nurseries, including these for bonsai production. Studies of Anoplophora glabripennis in North America and A. chinensis in China indicate the possible scale of economic damage following establishment of these species in a new country or in a plantation, especially of poplar or Citrus trees (Cocquempot et al. 2003, Haack et al. 2010, MacLeod et al. 2002). As a control measure, ornamental trees colonized by invasive longhorns must be eliminated without consideration of their aesthetic value. Eradication measures entail high costs to be borne by local communities or private owners. Special attention is paid to A. chinensis necessitating complete removal of trees, including the rootstock (Haack et al. 2010). Poplars or eucalypt plantations can be highly affected as has already been the case in China (A. glabripennis on poplars) and in Spain (Phoracantha spp.), where infested trees become unsuitable for pulp and wood exploitation. The Citrus longhorn beetle is also considered as an important risk for all Citrus fruit production in the Mediterranean area and its islands. The nursery industry is already concerned. There are several examples of introductions or establishments of potentially invasive species such as Callidiellum rufipenne and Anoplophora chinensis, with the imports of nursery plants. Nurseries can themselves be vectors of aliens when they dispatch their products. The eradication process established for quarantine species aims to limit introductions although only a few eradications have been officially reported in Europe, e.g. as for Anoplophora chinensis in France (Hérard et al. 2006, Hérard and Roques 2009). Phytosanitary interceptions at borders are likely to have prevented a number of introductions and further establishments (e.g., Monochamus alternatus, Trichoferus campestris in France, Anoplophora glabripennis and A. chinenis in several countries) (Cocquem- 204 Christian Cocquempot & Åke Lindelöw / BioRisk 4(1): 193–218 (2010) a b c f e g d Figure 8.1.6. Adults of some alien longhorn beetle species. a Phoracantha semipunctata b Phoracantha recurva c Mimectatina meridiana (Credit: Christian Cocquempot) d Xylotrechus stebbingi (Credit: Vítěslav Maňák) e Bardistes cibarius (Credit: Christian Cocquempot) f Psacothea hilaris g Parandra brunnea (a, b, e, f, g: Credit: Henri-Pierre Aberlenc). pot 2007) whilst at the same time, several non-quarantine species not submitted to importation controls have become established (e.g., Xylotrechus stebbingi, Phoracantha semipunctata, Neoclytus acuminatus). This illustrates the importance of quarantine species lists, which should be preventive and not only curative to be most effective. Human-mediated dispersal should also be tightly controlled during the eradication process. Without due respect for control obligations, eradication can fail. For example, the long delay by Italian authorities in applying control measures and strong management measures against Anoplophora chinensis (EPPO 2009, Jucker et al. 2007) or inadvertent movement of untreated wood material for A. glabripennis in New-York (Haack et al. 1997) are examples of ineffective eradication efficacy. Longhorn beetles (Coleoptera, Cerambycidae). Chapter 8.1 205 8.1.9 Expected trends The combination of increasing volumes of trade, the increased speed of import of potential vectors, the diversity of sources and sites for introduction is likely to result in increasing invasion risk (Cocquempot 2007). All recently established species alien to Europe have been intercepted too late after their introduction and have been outside official institutional controls. These factors make it increasingly difficult for rapid eradication after initial arrival. Effective monitoring of each point of possible entry is unfeasible when the key pathways identified here have different vectors and locations of arrival (e.g. airports, harbours, stations, lorry parks), and there are major difference in the quality of phytosanitary controls between European countries, particularly following the enlargement of the EU. The risk depends on volume and diversity of vector material imported, and subsequently there is greatest risk in countries such as the UK, France, Spain, Italy, Netherlands, Belgium and Germany. The case of Anopolophora glabripennis in North America and Europe clearly demonstrates the possibility of spread in our continent; such detailed assessment is required for all potentially invasive longhorn beetles (MacLeod et al. 2002). According to Worner (2002), progress in the knowledge of invasion processes and associated preventive measures have not been followed by actions since the late 1980’s. Preventive methods are still routinely applied, e.g. the application of ISPM 15 (International Standard for Phytosanitary Measures No.15), which set standards for heat treatment and fumigation of wood product materials used in international trade is likely to limit the arrival of longhorn beetles related to these materials although a few have been found to survive (Haack et al. 2010). However, this method is not uniformly applied to all imported living trees, shrubs plants for planting or bonsais. Thus, a high number of imported bonsais or other nursery trees infested with Anoplophora chinensis are still discovered (Hérard and Roques 2009). Although importation controls could be improved, they will never offer full protection. Further, controls which reduce the risk of introduction are mainly restricted to quarantine species. 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Status Regime Native range A phytophagous Brazil 1st record in Invaded Europe countries 1880, PT-AZO PT-AZO A phytophagous phytophagous Brazil 1977, PT China South- 2000, IT Central IT, NL Habitat Hosts Anoplophora glabripennis (Motschulsky, 1853) A phytophagous China South- 2001, AT Central AT, DE, FR, IT Callidiellum rufipenne (Motschulsky, 1860) A phytophagous Eastern Asia, 1906, FR Japan ES, FR, IT Chlorophorus annularis (Fabricius, 1787) Cyrthognathus forficatus (Fabricius, 1792) Derolus mauritanicus Buquet, 1840 A phytophagous phytophagous phytophagous AsiaTemperate Africa 1991, ES ES Borges et al. 2005, Méquignon 1935, Serrano 1982 I2 Moraceae, Lemos-Perreira 1978 , Vives Apocynaceae 1995 FB, FA, Acer, Betula, Cocquempot 2007, Colombo I2, G Carpinus and Limonta Citrus, Corylus, 2001, 2009a, EPPO 2009b, Rosa and Evans et al. 2008, Hérard et deciduous al. 2006 shrubs (polyphagous) FB, FA, I Acer, Aesculus, Carter et al. 2009, Betula, Cocquempot 2007, Carpinus, Cocquempot et al. 2003, Fagus, Populus, Dauber and Mitter 2001, Salix EPPO 2004, Hérard et al. 2006, 2009 FA, FB, Cupressaceae Bahillo and Iturrondobeitia G1, G5, J4 (Cupressus 1995, Campadelli and Sama macrocarpa) 1988, Cocquempot 2007 G Bamboo Vives 1995 1872, MT MT U Northern Africa 1884, FR ES ?, FR ? Nerium E7, F5, F8, FB, I2, oleander X11 Acrocinus longimanus (Linnaeus, 1758) Anoplophora chinensis (Förster, 1848) (=A. malasiaca Thompson, 1865) A A A PT, PT-MAD I2 Acacia, Albizzia Unknown References Bertolini 1872 Brustel et al. 2002, Fauvel 1884, Mendizábal 1944, Verdugo 2004 Christian Cocquempot & Åke Lindelöw / BioRisk 4(1): 193–218 (2010) Family Species Acanthoderes jaspidea Germar, 1824 214 Table 8.1.1. List and characteristics of the Cerambycidae species alien to Europe. Status: A: Alien to Europe; C: cryptogenic species. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Family Species Deroplia albida (Brullé, 1838) Lucasianus levaillantii (Lucas, 1846) Status A A Regime Native range phytophagous phytophagous Canary Islands Northern Africa 1st record in Invaded Europe countries 1988, ES ES 1905, FR ES, FR, PT Habitat Hosts E7, F6, Pelargonium FB, G5 FA, G, FB Cupressus References Vives 1995 A phytophagous SouthCentral U.S.A. 1908, IT CH, CZ, DE, FR, HR, HU, IT, ME, PTMAD, RS, SI FB, G, G1, Ulmus, G5, I2, Fraxinus, X11 Juglans Oxymerus aculeatus lebasi Dupont, 1838 Parandra brunnea (Fabricius, 1789) C phytophagous phytophagous Unknown Unknown ES-CAN U Calophyllum North America 1916, DE DE G, J1 Phoracantha recurva Newman, 1840 A phytophagous Australia 1992, IT ES, GR, IL, IT, G1 IT-SAR, ITSIC, MT, PT Tilia, Populus, Grämer 1961, Nüssler 1961 deciduous trees Eucalyptus Bercedo and Bahillo 1998, Bercedo and Bahillo 1999, Černý 2002, Cocquempot 2007, Cocquempot and Sama 2004, Friedman et al. 2008, Mazzeo and Siscaro 2007, Mifsud 2002, Miquel 2008, Orousset 2000, Palmeri and Campolo 2006, Pérez Moreno 2001, Ruiz and Barranco 1998, Sama and Bocchini 2003, Sama et al. 2010, Wang 1995 A 215 Neoclytus acuminatus (Fabricius, 1775) Longhorn beetles (Coleoptera, Cerambycidae). Chapter 8.1 Brustel et al. 2002, Cocquempot et al. 2007, Mayet 1905, Pellegrin and Cocquempot 2001, Plaza Lama 1990, Vives 1995 Bijaoui 1980, Brustel et al. 2002, Cocquempot 2007, Heyrovský 1951, Ilić 2005, Picard 1937, Pil and Stojanović 2005, Reineck 1919, Sama 1984, Tassi 1969, Villiers 1979, Winkler 1932, Wittenberg 2005 Alluaud 1935 Status Regime Native range phytophagous Australia Phryneta leprosa (Fabricius, 1775) A phytophagous Taeniotes cayennensis Thomson, 1859 Trinophylum cribratum (Bates, 1878) A phytophagous phytophagous South Tropical Africa Central America India Xylotrechus stebbingi Gahan, 1906 A A phytophagous Central Asia 1997, FR FR, MT G Morus nigra 1858, PT PT-AZO U Tropical trees Unknown GB I2 1990, IT References Berger 1992, Brustel et al. 2002, Cadahia 1980, Cavalcaselle 1983, Černý 2002, Cocquempot 1993, Cocquempot 2007, Cocquempot and Sama 2004, Mifsud and Booth 1997, Orousset 1984, Orousset 1991, Sama et al. 2010, Teunissen 2002, Vives 1995, Wang 1995 Mifsud and Dandria 2002, Vincent 2007 Sama 2006a Deciduous Duffy 1953b, Gilmour 1948 trees, Larix, Pinus (polyphagous) CH, CY, FB, G, G1, Alnus, Ficus, Cocquempot 2007, DE, FR, GR, G5, I2, Morus, Populus Cocquempot and Debreuil GR-CRE, X11 2006, Dioli and Vigano 1990, GR-NEG, GRKöhler 2000, Sama 2006b, SEG, IL, IT, Sama et al. 2010, Šefrová and IT-SAR Laštůvka 2005, Tomiczek and Hoyer-Tomiczek 2008, Wittenberg 2005 Christian Cocquempot & Åke Lindelöw / BioRisk 4(1): 193–218 (2010) A 1st record in Invaded Habitat Hosts Europe countries FB, G, G1, Eucalyptus 1948, IL CY, FR, FRG5, I2, COR, ES, ES-CAN, GR, X11 IL, IT, IT-SAR, IT-SIC, MT, PT, PT-MAD 216 Family Species Phoracantha semipunctata (Fabricius, 1775) Table 8.1.2. List and characteristics of the Cerambycidae species alien in Europe. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Regime phytophagous Native range Invaded countries Continental PT-AZO, PTEurope MAD Continental PT-AZO Europe Balkans MT phytophagous Balkans MT phytophagous Continental Europe Southern Europe PT-MAD phytophagous phytophagous phytophagous Icosium tomentosum atticum Ganglbauer, 1881 Monochamus galloprovincialis (Olivier, 1795) Monochamus sartor (Fabricius, 1787) Monochamus sutor (Linnaeus, 1758) phytophagous Morimus asper funereus Mulsant, 1863 phytophagous phytophagous phytophagous phytophagous AT, CH, , ESCAN, IE, LV, LT, PT-AZO, PT-MAD Southeastern FR Europe Southwestern NL Europe Northern BE, , NL Europe, Alps Central and BE, PT Northern Europe Southeastern CZ, MT Europe Habitat G3 Hosts References I2 Pinus, Picea, Abies, Larix Fauvel 1897, Picard 1937, Serrano 1982 Salix, Populus, Alnus Borges et al. 2005 G Prunus Sama and Cocquempot 1986 G Pyrus, Malus Fauvel 1897, Schembri and Sama 1986 Picard 1937, Wollaston 1854 E5, G, Deciduous trees G1, G5 (polyphagous) F3, G, G5 Deciduous trees (polyphagous Borges et al. 2005, Bytinski-Salz 1956, Lucht 1987, Speight 1988, Wollaston 1863 G3 Cupressaceae G3 Pinus Cocquempot et al. 2007, Pellegrin 1990 De Fluiter 1950 G3 Picea Fauvel 1884, Wiel 1956, Lucht 1987 G3 Picea, Pinus Speight 1988, Weyers 1876 G Deciduous trees (polyphagous Schembri and Sama 1986, Šefrová and Laštůvka 2005 Longhorn beetles (Coleoptera, Cerambycidae). Chapter 8.1 Family species Arhopalus rusticus (Linnaeus, 1758) Aromia moschata (Linné, 1758) Cerambyx carinatus Küster, 1846 Cerambyx nodulosus Germar, 1817 Clytus arietis (Linné, 1758) Gracilia minuta (Fabricius, 1781) 217 Regime phytophagous Deciduous and conifer trees (polyphagous) G Deciduous and fruit trees, preferably on Quercus Quercus, Castanea phytophagous Continental Europe Poecilium lividum (Rossi, 1794) phytophagous Southeastern BE, CH, CZ, Europe DE, LU, NL G,J1 Rhagium inquisitor (Linné, 1758) phytophagous Continental Europe G3 Stictoleptura rubra (Linné, 1758) Stromatium unicolor (Olivier, 1795) phytophagous Trichoferus fasciculatus (Faldermann, 1837) Trichoferus griseus (Fabricius, 1792) Xylotrechus arvicola (Olivier, 1795) phytophagous phytophagous phytophagous phytophagous IE Central MT Europe, Alps Central PT-AZO Europe Southeastern PT-MAD Europe Southeastern CH, PT-MAD Europe Southeastern CZ Europe Southeastern SP-CAN Europe Hosts F3 Phymatodes testaceus (Linné, 1758) Rosalia alpina (Linné, 1758) phytophagous PT-AZO Habitat Conifers (Pinus, Picea, Abies, Larix); deciduous trees (Betula, Fagus, Quercus) G, I2, J1 Fagus, and other deciduous trees G3 Conifers (Pinus, Picea, Abies, Larix) G Deciduous trees (mostly) and conifers (polyphagous) G Deciduous trees (polyphagous) G Ficus, Pistacia, Rosa G Deciduous trees (polyphagous) References Adlbauer 2006, Borges et al. 2005, Duffy 1953a, Heyrovský 1930, Korcynski 1985, Lucht 1987, Sliwinski 1958, Speight 1988, Weidner 1973, Weyers 1875 Fauvel 1897, Picard 1937, Wollaston 1854 Lucht 1987, Heyrovský and Sláma 1992, Horion 1974, Šefrová and Laštůvka 2005, Wittenberg 2005 Speight 1988 Horion 1974, Schembri and Sama 1986 Borges et al. 2005 Fauvel 1897, Picard 1937 Allenspach 1973, Picard 1937 Šefrová and Laštůvka 2005 Demelt 1974 Christian Cocquempot & Åke Lindelöw / BioRisk 4(1): 193–218 (2010) Native range Invaded countries Southwestern AT, BE, CH, Europe CZ, DE, GB, IE, LU, LV, PL, PT-AZO 218 Family species Nathrius brevipennis (Mulsant, 1839) A peer reviewed open access journal BioRisk 4(1): 219–266 (2010) RESEARCH ARTICLE doi: 10.3897/biorisk.4.64 BioRisk www.pensoftonline.net/biorisk Weevils and Bark Beetles (Coleoptera, Curculionoidea) Chapter 8.2 Daniel Sauvard1, Manuela Branco2, Ferenc Lakatos3, Massimo Faccoli4, Lawrence R. Kirkendall5 1 INRA, UR633 Zoologie Forestière, 2163 Avenue de la Pomme de Pin, CS 40001 ARDON, 45075 Orléans Cedex 2, France 2 Centro de Estudos Florestais, Instituto Superior de Agronomia, Technical University of Lisbon, Tapada da Ajuda, 1349-017, Lisboa, Portugal 3 University of West-Hungary, Institute of Silviculture and Forest Protection, Bajcsy-Zs. u. 4., H-9400 Sopron, Hungary 4 Department of Environmental Agronomy and Crop Sciences, Viale dell’Università 16, 35020 Legnaro (PD), Italy 5 University of Bergen, Biology Institute, Postbox 7803, N-5020, Bergen, Norway Corresponding author: Daniel Sauvard (Daniel.Sauvard@orleans.inra.fr) Academic editor: Alain Roques | Received 16 March 2010 | Accepted 25 May 2010 | Published 6 July 2010 Citation: Sauvard D et al. (2010) Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 219–266. doi: 10.3897/biorisk.4.64 Abstract We record 201 alien curculionoids established in Europe, of which 72 originate from outside Europe. Aliens to Europe belong to five families, but four-fifths of them are from the Curculionidae. Many families and subfamilies, including some species-rich ones, have few representatives among alien curculionoids, whereas some others are over-represented; these latter, Dryophthoridae, Cossoninae and specially Scolytinae, all contain many xylophagous species. The number of new records of alien species increases continuously, with an acceleration during the last decades. Aliens to Europe originate from all parts of the world, but mainly Asia; few alien curculionoids originate from Africa. Italy and France host the largest number of alien to Europe. The number of aliens per country decreases eastwards, but is mainly correlated with importations frequency and, secondarily, with climate. All alien curculionoids have been introduced accidentally via international shipping. Wood and seed borers are specially liable to human-mediated dispersal due to their protected habitat. Alien curculionoids mainly attack stems, and half of them are xylophagous. The majority of alien curculionoids live in human-modified habitats, but many species live in forests and other natural or semi-natural habitats. Several species are pests, among which grain feeders as Sitophilus spp. are the most damaging. Copyright D. Sauvard. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 220 Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Keywords Europe, Coleoptera, Curculionoidea, Curculionidae, alien species, invasive species, xylophagy, seed feeder 8.2.1. Introduction The superfamily Curculionoidea encompasses the weevils and the bark and ambrosia beetles; here we will use „weevils“ to refer to the entire superfamily. It is the most species-rich beetle clade, with more than 60,000 described species (Oberprieler et al. 2007). Four fifths of all weevils are in the family Curculionidae. Curculionoids are distributed worldwide, everywhere vegetation is found. This is a rather homogeneous group, its members being generally easily recognizable despite various aspects. Adults are primarily characterized by the head being produced into a rostrum (snout) to which the antennae and mouthparts are attached. The rostrum is highly variable in size and shape, varying from as long as the body to very short or absent. Larvae, generally white and C-shaped, are catepillar-like (eruciform), soft-bodied, with legs being either vestigial or (usually) absent, except in some species of the primitive family Nemonychidae. Except for a few rare species, adults and larvae of Curculionoidea are phytophagous. Larvae are mainly endophytic or subterranean. Weevils feed on a large variety of plants, attacking all parts. Many species are important pests for agriculture or forestry. The Macaronesian islands1 pose a special problem. While many of their weevils are only found on single islands or groups of islands and are thus clearly endemic, other species are shared between island groups, or between Macaronesian islands and the continental Europe or North Africa. For example, a number of scolytines specialized to Euphorbia are shared between the Canary Islands and Madeira, or between the Canary Islands and the Mediterranean and North Africa (Table 8.2.1). Given the difficulties involved with dispersal by these tiny insects over vast expanses of salt water, we have chosen to interpret the distributions of non-endemic species as resulting from recent human transport. We are well aware that rare instances of natural dispersal do occur, at least on evolutionary time scales: after all, such natural dispersal has resulted in many instances of well documented species radiations (Emerson 2008, Juan et al. 2000). Because of the inherent uncertainty in distinguishing between recent anthropogenic spread and older natural dispersal, we classify nonendemic species of these archipelagos as presumed aliens (they are indicated in tables 8.2.1 & 8.2.2). Without contradictory data, we consider: 1) species known from Europe and found on a Macaronesian island as presumed alien in Europe; 2) species known from Africa (and not from Europe) and found in Macaronesia as presumed alien to Europe; 3) species from the Canary Islands which also occur further north on Madeira or the Azores as presumed alien 1 We include in our coverage the Macaronesian islands associated with European countries (Madeira, the Azores, the Canary Islands); we exclude the Cape Verde Islands. Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 221 from the Canary Islands and presumed alien to Europe. Presumed alien are often considered below separately than others, due to the uncertainty attached to their status and the geographical and biogeographical differences between Macaronesia and Europe. We consider that 201 alien curculionoids currently live in Europe, of which 72 species originate outside of Europe (aliens to Europe, Table 8.2.1; 20 presumed alien are included) and 129 species originate from other parts of Europe (aliens in Europe, Table 8.2.2; 60 presumed alien are included)2. Except where otherwise noted, our discussion of exotic curculionoids only pertains to alien to Europe. 8.2.2.Taxonomy and biology The systematics of the superfamily Curculionoidea have long been controversial, in part due to the enormous number of taxa involved, in part due to extensive parallel evolution arising from the similar ecologies of unrelated clades (Alonso-Zarazaga and Lyal 1999, Oberprieler et al. 2007). We follow here the current classification of Fauna Europaea (Alonso-Zarazaga 2004), which notably considers the traditional Platypodidae and Scolytidae families as subfamilies of Curculionidae. About 5,000 native curculionoids live in Europe, distributed among 13 families. Comparatively, the alien entomofauna is very limited with only 72 established species recorded at this time (Fig. 8.2.1). These alien species belongs to five families, all of which have native representatives. Anthribidae. Principally present in tropical areas, these largely fungus-feeding curculionoids generally live primarily in fungus-infested wood. There is only one alien species in Europe, Araecerus coffeae, which is a seed feeder, an exceptional biology in this family. Apionidae. Characterized in part by their non-geniculate antennae and endophytous larvae, these tiny curculionoids are represented in Europe by three alien species, all living on alien ornamental Alcea (Malvaceae). Dryophthoridae. This family contains large weevils mainly living on woody monocotyledons. Alien dryophthorids consist of woody monocotyledons borers and seed feeders. They are particularly numerous compared with the world fauna (Fig. 8.2.1) and especially with respect to the few native species in Europe (8 aliens vs 6 natives, according to Fauna Europaea (Alonso-Zarazaga 2004)). This situation could be explained first by the few woody monocotyledons in Europe-native flora in contrast with the several woody monocotyledons introduced in Europe for ornamental or agricultural purpose. The human-mediated transport of seeds, and consequently seed feeders, is probably a further explanation. 2 Other aliens have been recorded, but have not been taken into account here because their establishment have not been confirmed. We have also excluded some possible presumed aliens due to the uncertainty about their distribution. 222 Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Figure 8.2.1. Taxonomic overview of Curculionoidea species alien to Europe compared to the native European fauna and to the world fauna. Right- Relative importance of the Curculionoidae families and subfamilies in the alien entomofauna is expressed as percentage of species in the family/ subfamily compared to the total number of alien Curculionidea in Europe. Subfamilies of Curculionidae and other families of Curculionidea are presented in a decreasing order based on the number of alien species. The number over each bar indicates the total number of alien species observed per family/ subfamily. LeftRelative importance of each family/ subfamily in the native European fauna of Curculionidea and in the world fauna expressed as percentage of species in the family/ subfamily compared to the total number of Curculionidea in the corresponding area. The number over each bar indicates the total number of species observed per family/ subfamily in Europe and in the world, respectively Erirhinidae. Curculionoids of this small family mainly live on herbaceous monocotyledons, often aquatic ones. With two alien species, they are relatively well represented in Europe. Curculionidae. This huge family encompasses more than 80% of weevils and notably includes the bark beetles and pinhole borers (Scolytinae and Platypodinae). Curculionids have a large variety of habits, but are all phytophagous. The European species are distributed in 16 subfamilies. The alien species belong to 10 subfamilies, all having native representatives. Many subfamilies, including the world’s largest (Entiminae, Curculioninae and Molytinae), are under-represented among alien curculionoids compared with their world importance in the superfamily (Fig. 8.2.1). On the other hand, the subfamily Cossoninae, which mainly contains wood-boring weevils, are over-represented, but the most remarkable result is the over-representation of Scolytinae. Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 223 Scolytinae are small, cylindrical wood borers, without a rostrum or with only a very reduced one; they include some of the most important forest pests in the world. The majority are phloeophagous, breeding in the inner bark. Most others are xylomycetophagous, feeding on symbiotic fungi which they cultivate in tunnels in the wood (ambrosia beetles). The scolytines represent about 10% of world curculionoids but almost half of curculionoids alien to Europe. Alien bark beetles represent more than 12% of all bark beetle species in Europe. The over-representation of Scolytinae is related to the frequency with which they are transported in wooden packing material, pallets, and timber (Haack 2001, 2006, Brockerhoff et al. 2006). All stages of these beetles can survive long voyages well, since both adults and larvae are in tunnels under bark or in wood and not directly exposed to temperature extremes or dessication. The importance of a stable, protected microenvironment is illustrated by the high prevalence of ambrosia beetles in the Scolytinae plus Platypodinae (35%) among successful aliens to Europe (Table 8.2.1), compared with the prevalence of ambrosia beetles in these groups in temperate climates generally (below 20%: Kirkendall 1993). The establishment of ambrosia beetles in Europe is further facilitated by polyphagy (11/12 spp.) and inbreeding (10/12 spp.), as is generally believed to be the case for ambrosia beetles globally (Kirkendall 1993, Haack 2001). The curculionoids alien in Europe are more representatives of Europe-native fauna. Scolytines (25% of aliens in Europe) are also over-represented compared with their importance among European curculionoids (5%), but not cossonines (3% of aliens in Europe). On the other hand, Entiminae (26% of alien in Europe, mostly Otiorhynchus and Sitona) are under-represented compared with the European fauna, but less so than among aliens to Europe. 8.2.3.Temporal trends Of the five families considered in this chapter, the first information concerning an alien species in Europe was probably the description by Ratzeburg in 1837 of Xyleborus pfeilii based on specimens from southern Germany8. The curculionid Pentarthrum huttoni was introduced to Great Britain from New Zealand in 1854, and has subsequently become naturalized in many European countries (Table 8.2.1). Only three other introduced species were recorded in the second half of 19th century. With the beginning of the 20th century, alien species began to be discovered more frequently, though this was limited to sporadic introductions (about 2 species per decade) confined to southern Europe – which perhaps provided more favourable climatic conditions – and along the main routes of international trade. Since the 1920s the rate of new introductions has slightly increased (Fig. 8.2.2), with a mean of nearly three species every decade, but remaining stable until middle of 1970s. Despite the European laws regulating the trade of plant material, the number of records of new exotic species introduced to Europe has increased rapidly since 1975 and especially since 2000, reaching worrying levels with an average of more than one 224 Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Figure 8.2.2. Temporal trend in establishment of Curculionoidea species alien to Europe from 1492 to 2010. Presumed alien species are excluded. The number besides each bar indicates the absolute number of new records during the time period. For the introduction year of each species see Table 8.2.1. species per year (16 new species from 2000 to 2009: Table 8.2.1), and a peak of five new species per year in 2004 (8 species in 2003–2004). It is too early to say if the relatively low number of establishments observed since 2005 will be confirmed or is only due to stochastic variations. However, if the trend towards increasing rates of introduction continues unabated, in a few decades the mean number of alien species becoming established in Europe could reach several per year. The temporal trend of alien curculionoids establishment is very similar to that observed in Europe for all alien terrestrial invertebrates (Roques et al. 2009, but see also Smith et al. 2007 for contradictory (more limited) data). On the other hand, this trend varies among weevils. Aliens from Asia follow the general trend (half of them have been recorded after 1975, a third after 2000), but the increasing of establishment rate is faster for those from North and South America (two-thirds of them have been recorded after 2000) while it is slower for those from others continents (half of them have been recorded before 1950, and none after 2000). Regarding feeding habits, all aliens follow the general trend except those with spermatophagous larvae, which show no trend. This particularity of the formers seems related to the oldness and intensity of human-mediated seed transport. Unfortunately, for many alien species spread over large parts of Europe, data on the place and time of introduction are lacking, and generally the data on time of arrival of exotic species are very weak. Often, introduced species – especially those which are not pests – are first noticed only many years after arrival, or following subsequent and repeated introductions. As prompt communication of new findings is extremely important for the application of specific monitoring and eradication programs, the poor quality of these data is a major obstacle to aliens management. Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 225 8.2.4. Biogeographic patterns Origin of alien species All presumed aliens probably come from Africa (among which 35% from the subregion Macaronesia). These species are not included in further discussion due to uncertainty of their status and specially because their arrival modes have probably been different from other aliens due to proximity of the source region. A probable region of origin could be specified for 51 of the 52 curculionoid species alien to Europe. There is one species, Sitophilus zeamais (Dryophthoridae), whose region of origin is uncertain (cryptogenic). Cryptogenic species are thus rare in this group compared to all alien terrestrial invertebrates (14%: Roques et al. 2009). Sitophilus zeamais is associated with maize crops, Zea mays, and feeds on maize grain stores, and it is likely that this species is American. More than one-third (40%) of the exotic curculionoid species originate from Asia. Central and South America represents the second most important region of origin, with 19% of the species coming from this area. North America and Australasia both represent 14% of the contributing regions. Africa is a minor region of origin (6%), and the remaining species (6%) arrived from tropical or subtropical areas but the region of origin could not be precisely identified (Figure 8.2.3). This distribution is rather similar to that for all alien terrestrial invertebrates (Roques et al. 2009). The main differences are the under-representation of African aliens (6% vs. 12%) and the overrepresentation of South American (19% vs. 11%) and Australasian (14% vs. 7%) ones. A rather surprising result is that species originated from areas with tropical or subtropical climates all around the world represent about half of alien curculionoids. Thirteen out of the twenty-one alien species originating from Asia are from the family Curculionidae, twelve species belonging to the subfamily Scolytinae and one species to the subfamily Cyclominae. Other families consist of Dryophthoridae (4 spp.), Apionidae (3 spp.) and Anthribidae (1 sp.). Scolytines originate from very different parts of this large continent. For example Cyclorhipidion bodoanus is native to Siberia and temperate northeast Asia, Phloeosinus rudis to Japan, and the three species of the genus Xylosandrus to Southeast Asia. In contrast, all the weevils of the Dryophthoridae family originate from tropical Asia. This group includes the banana root weevil Cosmopolites sordidus, the coconut weevil Diocalandra frumenti, the palm weevil Rhynchophorus ferrugineus and the rice weevil Sitophilus oryzae. The introduced apionids, Alocentron curvirostre, Aspidapion validum and Rhopalapion longirostre, all feed on flowers and seeds of Alcea rosea and other Malvaceae species (Bolu and Legalov 2008); these all originate from central Asia. Finally, the anthribid Araecerus coffeae originates from India. The ten curculionoid species coming from Central and South America consist of curculionids (8 spp.) and dryophthorids (2 spp.). Curculionids originating from this region are as highly diverse taxonomically (they are distributed in six subfamilies) as in feeding habits. The native ranges of many species largely extend through the continent 226 Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Figure 8.2.3. Origin of Curculionoidea species alien to Europe. Presumed alien species are excluded. (including sometimes part of North America), though those of others are more narrow as for Rhyephenes humeralis (central Chile and neighboughring area of Argentina) and Paradiaphorus crenatus (Brazil). Seven alien curculionoids are known to originate from North America. They include five species of the family Curculionidae and two of Erirhinidae. Many curculionids introduced from North America are xylophagous sensu lato7, feeding on several broadleaved or coniferous hosts. The exceptions are the ash seed weevil Lignyodes bischoffi and Caulophilus oryzae, originally from the southeastern USA, which feeds on seeds. In contrast, the two Erirhinidae species feed externally on weed roots and ferns, respectively. Seven curculionoid species come from Australasia, all curculionids: four cossonines, two molytines and one cyclomine. Three woodboring weevils (Pentarthrum huttoni, Euophryum confine and E. rufum, all from Cossoninae), feeding on decaying wood, originate from New Zealand. The four other species were unintentionally introduced from Australia. All feed inside plant material (xylophagous or herbiphagous), except the Eucalyptus snout beetle, Gonipterus scutellatus, a defoliator of Eucalyptus trees originated from Southern Australia. Only three curculionoid species are known to originate from Africa, a curculionine and two scolytines. The palm flower weevil, Neoderelomus piriformis, probably originates from North Africa; it feeds on but also pollinates flowers of palms like Phoenix canariensis. The scolytines both originate from Canary Islands; Dactylotrypes longicollis breeds in Phoenix canariensis seeds, while Liparthrum mandibulare is a highly polyphagous phloeophage. Three cosmopolitan curculionoid species originate from undetermined areas of the tropical and subtropical parts of the world: the tamarind seed borer, Sitophilus linearis (Dryophthoridae), and the palm seed borers Coccotrypes carpophagus and C. dactyliperda (Scolytinae). As seed-feeders, they are readily distributed through commerce, which probably explain the uncertainty about their origin. Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 227 Concerning the curculionoids alien in Europe, nine-tens of these (114 spp. among 129, Table 8.2.2) are introduced from mainland Europe to islands (mainly the Canary Islands, the Azores, the British Isles and Madeira). They are often widespread continental species which have been introduced to islands by human transport. Other cases are mainly species of southern and western regions which were introduced into northern Europe (as Otiorhynchus corruptor), especially to Denmark and Sweden. However, some species have moved westwards (as Otiorhynchus pinastri and Phloeotribus caucasicus) and even southwards (Ips duplicatus). Distribution of alien species in Europe As for the other arthropod groups, alien curculionoid species are unevenly distributed throughout Europe, which may partly reflect differences in sampling intensity (Fig. 8.2.4, Table 8.2.1). In continental Europe, mainland Italy and France host the largest number of species alien to Europe, with 28 and 26 introduced curculionoid species, respectively. These countries are followed by continental Spain (17 spp.), Austria (15 spp.), and Germany, Switzerland and United Kingdom3 (13 spp.). This distribution is similar as that of all alien terrestrial invertebrates (Roques et al. 2009). The number of aliens per country significantly decreases eastwards (y=12 - 0.29*longitude, R2=0.21, F1,31=8.08, p=0.008), but it is mainly correlated with human variables, country population (y=-1.5 + 3.7ln(population), population in million inhabitants, R2=0.39, F1,31=19.6, p=1*10-4) and country importation values (y=-32 + 3.5ln(value), value 2003–2007 in million USD: The World Factbook 2009, R2=0.53, F1,29=32.4, p=4*10-6)4. The best model integrates importations and latitude (y=-19 + 3.6ln(value) - 0.28*latitude, value in million USD, R2=0.60, F2,28=20.6, p=3*10-6), indicating that alien establishment is favored by human trade and warm climate. The abundance of aliens in mainland Italy and France is not fully explained by the model (predicted values 17 and 16 alien species, respectively); it is likely related to a combination of the diversity of habitats and plants present with the favorable climate and the importance in international shipping. Islands have a rather rich alien curculionoid fauna, especially Macaronesia: 29 (of which 14 presumed), 18 (8 presumed) and 10 (2 presumed) species in the Canary Islands, Madeira and the Azores, respectively. These islands are followed by Sicily (10 spp.), Corsica (8 spp.) and Malta (6 spp.). As it has been found for other alien terrestrial invertebrates (Roques et al. 2009), the number of alien curculionoids per km2 in European islands is higher than in continental countries (on average 2.8 vs 0.17 3 4 Concerning species alien to Europe, United Kingdom characteristics are closer to those of continental countries than to those of other islands, so we consider it as part of continental Europe. This is likely related to its large size and population. Computations were performed without small countries where no alien curculionoid is recorded, because this absence is probably due to lack of data. Israel was also excluded due to its special location. 228 Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Figure 8.2.4. Comparative colonization of continental European countries and islands by Curculionoidea species alien to Europe. Archipelagos: 1 the Azores 2 Madeira 3 the Canary Islands. alien/1000km2, R2=0.10, F1,58=6.56, p=0.013). Aliens density is specially high in Madeira and Malta (23 and 19 alien/1000km2, respectively), perhaps because these tiny islands are stopping places on trade routes. Islands show no global trend of alien distribution. However, cold nordic islands (Greenland, Iceland, Svalbard) host few aliens, and in Macaronesia alien number (specially presumed alien number) decreases when distance to continent increases. Near half of alien curculionoid species (33 spp.) have been observed in only one country, most of them (31 spp.) in a peninsular region or on islands: Italy, Iberia, Macaronesian islands, Malta or the British Isles. Aliens introduced to such areas are less likely to move to nearby countries in comparison with aliens in other mainland regions, but Austria and Russia also host each an own alien species. As examples, Syagrius intrudens from Australia is encountered only in Great Britain, Naupactus leucoloma, from South America, is found only in the Azores, and Paradiaphorus crenatus, from Brazil, is known only from the Canary Islands. After the Canary Islands, Italy hosts the Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 229 highest number of alien species unique to one country, eight in total, of which six are from subfamilies Scolytinae and Platypodinae. Also, the recent arrival of these species, most of them having first been discovered later than 2000, may in part explain their currently restricted distribution. Ten alien species (14%) are limited to two countries. In almost all cases, the species are found in neighbour countries, as with the scolytine Dryocoetes himalayensis in France and Switzerland, and Macrorhyncolus littoralis in Great Britain and Ireland. One alien species, Scyphophorus acupunctatus, occurs in two distinct regions, Sicily and France, suggesting the possiblity of multiple introductions (this suggestion is supported by the previous interceptions of this species in different european countries: EPPO 2008). At the other extreme, the rice weevil Sitophilus oryzae has been found in 34 European countries, and two other seed feeders, Sitophilus zeamais and Rhopalapion longirostre, occur in 23 and 21 countries. Their feeding habits in association with frequently transported seeds or stored products presumably explain this broad distribution. Another eleven species are found in 10 or more countries. These include several longestablished species: Xyleborus pfeilii8, the wood-borer Pentarthrum huttoni, the palm seed borer Coccotrypes dactyliperda and the parthenogenetic weevil Asynonychus godmani. However, the relatively recently introduced (1993) palm weevil Rhynchophorus ferrugineus is also widely distributed, occurring in most of the Mediterranean region, which attests their high dispersal capabilities (natural and human-mediated). Overall, alien weevil species are more widespread in Europe than other alien terrestrial invertebrates, with 40% of species distributed in more than two countries vs. only 22% (Roques et al. 2009). 8.2.5. Main pathways and factors contributing to successful invasions There are two components to successful invasion, dispersal and establishment. Dispersal to new continents by phytophagous arthropods is now almost entirely due to human transport, the magnitude of which has inceased exponentially in recent decades. Plant feeding arthropods are carried in and on live plants and fruits, in wood, and as stowaways in shipments and baggage. Deliberate introductions of arthropods are less frequent, and most involve exotic organisms imported for biological control. Establishment of new arrivals depends on availability of appropriate habitats near sites of introduction, ability to compete with similar species already present, and on a reasonable tolerance for the local climate. All exotic species of Curculionoidea have been introduced accidentally in Europe, vs. only 90% for all alien terrestrial invertebrates (Roques et al. 2009). The lack of intentional introductions of weevils could be related to their poor potential for biological control. One exotic weevil species (Stenopelmus rufinasus) has been used successfully for biological control of the American water fern Azolla filicoides in South Africa and to a less extent in the British Isles, but its first introduction in Europe was accidental (Sheppard et al. 2006, Baars and Caffery 2008). 230 Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) As is the case for other regions in the world, many of Europe’s alien curculionoids have presumably arrived via the shipping of wooden materials: pallets, crating, and barked or unbarked timber (Brockerhoff et al. 2006, Haack 2001, 2006). Bark and wood boring species make up half of all alien weevils (50%); these have almost certainly been introduced with wood transport and solid wood packaging materials. Logs with bark are ideal for transporting bark beetles and other weevils. However, even debarked logs can contain live wood borers such as ambrosia beetles. Although some wood-boring beetles have more restrictive requirements (e.g. high humidity and decayed wood: Euophryum confine, E. rufum, Pentarthrum huttoni), even these can often survive a few days or even weeks of transport. The east Asian ambrosia beetle X. germanus provides a typical example for entry by wood-borers. It was introduced to the USA (1932), where it was discovered in imported wine stocks in greenhouses; the species spread rapidly and has become an important nursery pest in warmer parts of eastern North America (Ranger et al. 2010). In Europe, it was first recorded after World War II, in Germany, where the species probably had been introduced with wood imported from Japan to southern Germany early in the 20th century; the present distribution area includes twelve European countries (Table 8.2.1). Seed feeders (20%) are introduced with the seeds, which are also an excellent way for transporting insects. Several of these species are associated with agricultural products (e.g. Caulophilus oryzae, Sitophilus oryzae and S. zeamais), however most species feed on ornamental or forest seeds (e.g. Rhopalapion longirostre on Alcea, Lignyodes bischoffi on ash seeds, Dactylotrypes longicollis on palm seeds). Other alien species (30%) live on or inside leaves and nonwoody stems, or in the soil. The formers can be introduced with their host plants or with host plant products (e.g. Gonipterus scutellatus with eucalyptus, Listroderes costirostris with plants such as tobacco); weevils living around roots (e.g. Asynonychus godmani) are transported with living plants. These feeding habits (plus root boring, which doesn’t exist among aliens to Europe) are more frequent among presumed aliens to Europe and among aliens in Europe (52%); both cases result from a rather short distance transport, which likely allows survival of less protected insects (among wood boring scolytines, phloeophagous species are similarly much more frequent than xylomycetophagous species among presumed aliens to Europe and among aliens in Europe, contrary to what is observed among other aliens to Europe). Currently, most introductions are due to international trade, but the increasing movement of fruits and plants by travelers, which is much more difficult to check, may contribute to the future diffusion of new alien species. Newly arrived phytophages must find suitable hosts. The likelihood of success is greatly enhanced if the species is not too host specific, or if its preferred hosts are abundant. Not surprisingly, the majority of established exotic weevils in Europe are polyphagous, and the hosts of others are often widespread and abundant plants (Table 8.2.1). Parthenogenesis and inbreeding further increase the chances for successful colonization. When an exotic species is first introduced to a new area, it faces a varie- Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 231 ty of problems associated with low density which reduce the likelihood of successful establishment and slow the rate of invasion (Tobin et al. 2007, Liebhold and Tobin 2008, Contarini et al. 2009). New populations create problems for mate finding; parthenogenetic females do not mate, and inbreeding females mate with brothers while in the natal nest, before dispersal (Jordal et al. 2001); in both cases, there is no problem of mate location and new populations can be established by single females. Very small populations (such as those in recent colonizations) may suffer from high levels of inbreeding depression (Charlesworth and Charlesworth 1987); however, regular inbreeding species such as the invasive scolytines have presumably purged their genomes of the deleterious alleles responsible for inbreeding depression (Charlesworth and Charlesworth 1987, Jordal et al. 2001, Peer and Taborsky 2005). Only a few invasive curculionoid species are parthenogenetic: Asynonychus godmani, Lissorhoptrus oryzophilus, Listroderes costirostris (Morrone 1993) and Naupactus leucoloma, whose males are unknown outside its native range (Lanteri and Marvaldi 1995). However, over half of the alien scolytines inbreed (59%, presumed aliens excluded), compared with less than a third of scolytines native to Europe and about a fourth of Scolytinae species worldwide (Kirkendall 1993). 8.2.6. Most invaded ecosystems and habitats All alien curculionoid species are phytophagous, as are nearly all curculionoids worldwide. Most of the species have a cryptic way of life, at least during larval stage, feeding inside plant tissues such as stems or seeds, or living in the soil; only 9% are leaf/stem browsers. Stems and trunks is the major feeding niche of most alien curculionoids (65%). Most of these are bark beetles, ambrosia beetles or other wood borers (50%); herbiphagous (15%) comprise the remaining. Seeds are the second most important feeding niche (18%), followed by leaves (9%; some species could also attack non woody stems) and roots (6%). Last species, Neoderelomus piriformis, feeds on flowers, and acts as pollinator in palm trees. By contrast, of the curculionoids alien in Europe, only 33% are wood borers, among which most are phloeophagous (28%). A third (30%) attack roots, especially root browsers as Otiorhynchus and Sitona (26%), the remaining (4%) being root borers. Herbiphagous (18%), spermatophagous (15%) and leaf/stem browsers (4%) comprise the remaining. Near half of the alien curculionoid species established in Europe colonize urban and peri-urban habitats, primarily parks and gardens (27%) and around buildings (11%). Woodlands is also a frequent habitat for the alien curculionoids (27%), beyond natural heathlands (16%), cultivated agricultural lands (9%) and greenhouses (5%). Only three species occur in wetland habitats, one in coastal and two in inland surface water (Fig. 8.2.5). The importance of natural heathlands is in fact mainly limited to specific areas, most of the species recorded in these habitats being presumed aliens attacking euphorbias in Macaronesian xerophytic heathlands. 232 Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Figure 8.2.5. Main European habitats colonized by Curculionoidea species alien to Europe. The number besides each bar indicates the absolute number of alien curculionoids recorded per habitat. Note that a species may have colonized several habitats. This pattern differs from the average value observed for all arthropods, where only a fourth of the species is recorded in natural or semi-natural habitats, and where agricultural lands and greenhouses contain more alien species than woodlands. That could be obviously related to the high frequency of xylophagous sensu lato7 habits in alien curculionoids. Both deciduous trees, such as Populus sp. and Fraxinus sp, and conifers in the genera Picea and Pinus are colonized by several alien curculionoid species utilizing trees. Eucalyptus plantations are also affected by a defoliating curculionid, Gonipterus scutellatus, both host and weevil originating in Australia. In urban and suburban areas such as gardens and parks, other trees species, mainly exotics and in particular palm trees, are also affected by alien curculionoids. 8.2.7. Ecological and economic impact Ecological impacts of alien insects are poorly known in general (Kenis et al. 2009), and the impacts of Curculionoidea species alien to Europe seem not to have been documented at all. Their economic impact is better known, reflecting the economic importance of many of these alien species. A third of the Curculionoidea species alien to Europe (26 species) have a known economic impact, a much higher proportion than for native weevils, even though the latter contain numerous pests. Nevertheless, this high proportion may partly be an artefact, since pests have a higher probability of being detected. The most damaging species are the four attacking stored products. The rice weevil Sitophilus oryzae and the maize weevil S. zeamais are among the main pests of stored grains worldwide, destroying significant amounts and incurring high pest management Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 233 Figure 8.2.6. Examples of alien curculionoids: Gonipterus scutellatus. Adult damage on Eucalytus sp. (Credit: Alain Roques). costs5 (Balachowsky 1963, Pimentel 1991). Larvae develop in cereal seeds and adults feed on these seeds as well as on a wide variety of stored products, products derived from cereal grains and even dried vegetables. Damages is exascerbated by incompletely dried stored products (Balachowsky 1963). In addition to their direct damage, these species facilitate attacks of grains by other pests. Caulophilus oryzae, a less widespread species, sporadically causes the same kind of damages, while Araecerus coffeae attacks grains but mainly less common products such as stored coffee and cocoa beans. Five species attack native or introduced cultivated plants. Listroderes costirostris attacks a wide range of vegetables and weeds; adults can also damage foliage of fruit trees. The recently established whitefringed weevil, Naupactus leucoloma, is also highly polyphagous; its soil-inhabiting larvae are a serious pest of many agricultural crops. The banana root weevil, Cosmopolites sordidus, and Paradiaphorus crenatus are important pests of tropical cultures (banana and pineapple, respectively). Their economic impact is currently limited in Europe due to the limited distribution of their hosts in this area and a rather low aggressiveness in its climate, but it could increase later in the future according to the global warming. The last species is the rice water weevil, Lissorhoptrus oryzophilus. Recently introduced in Europe, it is a major pest of rice, but also attacks indigenous Carex. Eight species damage different ornamental plants and trees, mainly introduced tropical or subtropical species. The palm weevil Rhynchophorus ferrugineus is a dangerous pest of palms which has rapidly colonized the Mediterannean basin. On the Canary Islands, palms are also attacked by the lesser coconut weevil Diocalandra frumenti. Even if damage are mainly esthetic, they are worrying because this insect princi5 Damages are also due to the grain weevil S. granarius, probably alien too, but not taken into account here because it has been established in Europe at least since Antiquity. 234 Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Figure 8.2.7. Examples of alien curculionoids: Rhynchophorus ferrugineus. Female, larvae and damage (Credit: Juan Antonio Ávalos, Universidad Politécnica de Valencia). pally attacks Phoenix canariensis, an endemic palm which is emblematic of the Canary Islands where it is widely used for landscaping and is a major element of coastal landscape. Asynonychus godmani attacks roots of a large variety of ornamental shrubs and fruit trees, native or introduced. Others species are monophagous or oligophagous on introduced hosts: the tamarind seed borer Sitophilus linearis on Tamarindus indica, Demyrsus meleoides on cycadophyts, Scyphophorus acupunctatus on Agavaceae species, Phloeotribus liminaris on Prunus serotina, Phloeosinus rudis on Cupressaceae species. Five species have an impact on forests or related habitats. Three attack live exotic or native trees. The Eucalyptus snout beetle Gonipterus scutellatus is an important pest of Eucalyptus everywhere it has been introduced (see factsheet 14.12). This defoliator causes severe damage and wood loss, particularly on E. globulus, the major cultivated Eucalyptus species in southern Europe. Rhyephenes humeralis attack another introduced tree, Pinus radiata, but causes less damage. Megaplatypus mutatus is one of the few platypodine beetles which breeds in live trees; it is highly polyphagous, but in Europe it has thus far only been found to damage Populus plantations in Italy (Alfaro et al. 2007). The two other species depreciate wood stock. Gnathotrichus materiarius is a common pest of a large variety of conifer wood, and Xylosandrus germanus sporadically attacks mainly broadleaf wood. Pentarthrum huttoni and the two Euophryum species live in rotting wood, so their economic impact is generally low, though they do attack wood of historically signifi- Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 235 Figure 8.2.8. Examples of alien curculionoids: Scolytinae. Top left: Gnathotrichus materiarius: gallery in wood (Credit: Louis-Michel Nageleisen). Top right: Cyclorhipidion bodoanus: femelle (Credit: LouisMichel Nageleisen). Bottom: Xylosandrus germanus (Blandford 1894): female (Credit: Daniel Adam), adults and gallery holes on wood (Credit: Louis-Michel Nageleisen). cant artefacts or buildings. Finally, as opposed to all previous species, the introduced frond-feeding weevil Stenopelmus rufinasus has a positive impact due to its hability to control the invasive red water fern Azolla filiculoides. 8.2.8. Conclusion The superfamily Curculionoidea is well represented among alien species now established in Europe. Alien weevils show specific characteristics comparing both native and world ones, which seem result from a selection of species having high capabilities to human-mediated dispersal and establishment in a new habitat. Thus, they have often cryptic habits, as seed boring or wood and plant boring, leading to over-representation of bark and ambrosia beetles and other xylophagous sensu lato7 species; alien weevils are consequently more numerous in natural areas than other terrestrial invertebrate aliens. 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Apionidae Alocentron (Alocentron) curvirostre (Gyllenhal 1833) Aspidapion (Aspidapion) validum (Germar 1817) 6 7 Status Feeding habits Native range 1st record in Europe Invaded countries A phytophagous (spe) AsiaTropical A phytophagous (spe) phytophagous (spe) Asia1904, BG AT, BG, CH, CZ, HU, Temperate IT-SIC, MD, PL, RO, RS, SI, SK Asia1960, BG AT, BG, CH, CZ, DE, Temperate FR, HR, HU, IT, MD, PL, PT, RO, SK, UA A 1951, DE AT, BG, DE, FR, GB, IL, IT, MT, PL Habitat J1 Hosts stored products (pp: Coffea, Camellia sinensis, stored products) I2, FA, Alcea rosae (op: FB Malvaceae) I2, FA, Alcea rosae (op: FB Malvaceae) References Essl and Rabitsch (2002), Mphuru (1974), Obretenchev et al. (1990), Sebelin (1951) Essl and Rabitsch (2002), Joakimow (1904), Wittenberg (2005) Abbazzi et al. (1994), Angelov (1960), Essl and Rabitsch (2002), Wittenberg (2005) Platypodines and scolytines adults generally feed as larvae, as do adults of many other species with spermatophagous or xylophagous sensu lato7 larvae. Otherwise adults generally feed externally on leaf and stem regardless of the larval habits. Adults are often more polyphagous than larvae, except platypodines and scolytines. We use the term xylophagous sensu lato to gather species with phloeophagous, xylomycetophagous and xylophagous larvae. Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Family / subfamily Species Anthribidae Araecerus coffeae (Fabricius 1801) 246 Table 8.2.1. Characteristics of the Curculionoidea species alien to Europe. Asterisks indicate presumed aliens. Feeding habits and hosts are those of larvae, which are generally the more damaging stage6. Country codes abbreviations refer to ISO 3166, with extensions (see Appendix I); main Atlantic and Mediterranean islands are treated separately as special „countries“. N/A data non available. Status: A alien to Europe C cryptogenic. Feeding habits: abbreviations between brackets specify the feeding habits; her herbiphagous (larvae bore and feed inside non woody tissue of plant stems or leaves; stem includes branches, twigs, collar, bulb and rootstock) lbw leaf/stem browser (larvae externally feed on leaves or stems, as most caterpillars; early stages could be miner) phl phloeophagous (larvae bore and feed inside tree inner bark) rbo root borer (larvae bore and feed inside roots) rbw root browser (subterranean larvae externally feed on roots; early stages could be root miner) spe spermatophagous (larvae bore and feed inside reproductive organs, generally seeds) xmp xylomycetophagous (larvae live in galleries bored by females inside wood and mainly feed on wood-decaying symbiotic fungi) xyl xylophagous (larvae bore and feed inside wood, including woody materials such as palm stems)7. Native range: the field contains standardized range; if useful, native range could be specified between brackets. 1st record in Europe: date and countries of first known specimen, or first publication. Habitat: habitats in invaded countries; abbreviations refer to EUNIS (see Appendix II). Hosts: recorded hosts in invaded countries, and, between brackets, host breath in native range; host breath in native range is given as monophagous, oligophagous or polyphagous (abbreviated as mp, op and pp), depending if the species normally attacks hosts in one genera, one family or more; hpp: highly polyphagous. Curculionidae Cossoninae Amaurorhinus (Amaurorhinus) monizianus (Wollaston 1860)* Caulophilus oryzae (Gyllenhal 1838) A phytophagous Africa N/A (ES-CAN) A phytophagous (spe) phytophagous (xyl) phytophagous (xyl) phytophagous (xyl) phytophagous (xyl) North America Euophryum confine (Broun 1880) A Euophryum rufum (Broun 1880) A Macrorhyncolus littoralis (Broun 1880) A Pentarthrum huttoni Wollaston 1854 A PT-AZO, PT-MAD 1982, ES-CAN, GB, PT-MAD PT-MAD Abbazzi et al. (1994), Ehret (1983), Essl and Rabitsch (2002), Kozłowski and Knutelski (2003), Markovich (1909), Mazur (2002), Perrin (1984), Perrin (1995), Wittenberg (2005) N/A (Suaeda, Salsola) Base de dados da biodiversidade dos Açores, Oromí and García (1995) J1 grain, stored products (pp: grain, Persea seed) decaying wood (pp: decaying wood) Izquierdo et al. (2004), Morris (2002), O‘Brien and Wibmer (1982) Essl and Rabitsch (2002), Hill et al. (2005), Menet (1998) decaying wood (pp: decaying wood) Hill et al. (2005), O‘Connor (1977) driftwood (pp: decaying wood) Morris (2002), Telfer (2007), Welch (1990) decaying wood (pp: decaying wood) Abbazzi and Osella (1992), Bruge (1994), Buck (1948), Dieckmann (1983), Halmschlager et al. (2007), Hoffmann (1954), Rasmussen (1976), Stachowiak and Wanat (2001), Strejček (1993), Wittenberg (2005) B2 Australasia 1854, GB AT, BE, CH, DE, DK, J1 ES, FR, GB, IE, IT, NL, PL, RU, SK 247 B Australasia 1937, GB AD, AT, CZ, ES, J1, I2 ES-BAL, FR, GB, HU, PT, SE Australasia 1934, GB CH, DK, ES, GB, J1, I2 IE, SE Australasia 1987, GB GB, IE References Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 Family / subfamily Status Feeding Native 1st record Invaded countries Habitat Hosts Species habits range in Europe Rhopalapion longirostre A phytoAsiaAlcea rosae (op: 1875, RO AT, BG, CH, CY, CZ, I2 Malvaceae) (Olivier 1807) phagous Temperate ES, FR, FR-COR, DE, (spe) GR, GR-NEG, HR, HU, IT, MD, NL, PL, RO, RS, SK, UA 1st record Invaded countries in Europe N/A ES-CAN C&S America 2003, ES North America Habitat Hosts References N/A N/A (N/A) Machado and Oromí (2000) ES G1, G5, X11 Pinus radiata (pp: broadleaf trees) Alonso-Zarazaga and Goldarazena (2005) 2001, PL AT, PL G, I2 Fraxinus (op: Fraxinus, Syringa) Essl and Rabitsch (2002), Freude et al. (1983), Gosik et al. (2001) Africa (North) 1992, IT, IT-SIC ES, ES-CAN, FR, IL, IT, IT-SIC, PT-MAD I2 Phoenix canariensis (Phoenix) A phytophagous Africa (North) N/A ES-CAN N/A N/A (Fabaceae) Abbazzi and Osella (1992), Alonso-Zarazaga and Lyal (1999), Friedman (2006), Machado and Oromí (2000), Piry and Gompel (2002) Machado and Oromí (2000) A phytophagous Africa (North) N/A ES-CAN N/A N/A (Fabaceae) Machado and Oromí (2000) A phytophagous (lbw) phytophagous (lbw) Asia1946, SE Temperate DE, DK, FI, FR, GB, LT, LV, NL, SE I2 Atriplex (op: Chenopodiaceae) Meregalli (2004) Australasia 1975, IT ES, ES-CAN, FR, FR-COR, IT, PT I2, G2 Eucalyptus (mp: Eucalyptus) Abbazzi and Osella (1992), Arzone (1976), Carrillo (1999), Machado and Oromí (2000), Mansilla (1992), Mansilla and Pérez Otero (1996), Neid (2003), Paiva (1996), Rabasse and Perrin (1979), Sampò (1976) A Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Tychius (Tychius) antoinei Hustache 1932* Tychius (Tychius) depauperatus Wollaston 1864* Cyclominae Asperogronops inaequalis (Boheman 1842) Gonipterus scutellatus Gyllenhal 1833 Native range Africa (North) 248 Family / subfamily Status Feeding Species habits Pentatemnus arenarius A phytoWollaston 1861* phagous Cryptorhynchinae A phytoRhyephenes humeralis (Guérin-Méneville phagous 1830) (phl) Curculioninae Lignyodes (Lignyodes) A phytobischoffi (Blatchley phagous 1916) (spe) Neoderelomus piriformis A phyto(Hoffmann 1938) phagous (spe) Family / subfamily Species Listroderes costirostris Schoenherr 1826 A Feeding Native habits range phytoC&S phagous America (lbw) 1st record Invaded countries Habitat Hosts References in Europe 1950, ES-BAL, ES-CAN, FR, I, J100 N/A (hpp: vegetables, Balachowsky (1963), Friedman ES-CAN IL, PT weeds) (2009), Germain et al. (2008a), Machado and Oromí (2000), Moncoutier (1982) I N/A (pp: Rosa, ornamentals, fruit trees) I, G N/A (hpp: Fabaceae, Borges et al. (2005) vegetables, Zea mays) Africa N/A (PT-MAD) ES-CAN N/A Foeniculum (N/A) García (2003), Machado and Oromí (2000) phytophagous (lbw) phytophagous (lbw) Africa (North) N/A ES-CAN N/A N/A (N/A) Machado and Oromí (2000) Africa (North) N/A ES-CAN N/A N/A (N/A) Machado and Oromí (2000) A phytophagous Africa (North) N/A ES-CAN N/A N/A (N/A) Machado and Oromí (2000) A phytophagous (xyl) Australasia 1974, IT IT I2 Cycadales (op: Cycadales) Covassi (1974) phytophagous (rbw) C&S America 1908, IT Naupactus leucoloma Boheman 1840 A C&S America 2003, PT-AZO Sitona (Sitona) latipennis Gyllenhal 1834* Hyperinae Donus (Donus) fallax (Capiomont 1868)* A phytophagous (rbw) phytophagous (rbw) Donus (Antidonus) isabellinus (Boheman 1834)* Lixinae Pycnodactylopsis (Louwia) tomentosa (Fåhraeus 1842)* Molytinae Demyrsus meleoides Pascoe 1872 A A Hoffmann (1950), Machado and Oromí (2000), Solari and Solari (1908), Stüben (2003) 249 DK, ES, ES-BAL, ES-CAN, FR, IT, IT-SAR, IT-SIC, MT, PT, PT-AZO, PT-MAD, SE PT-AZO A Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 Entiminae Asynonychus godmani Crotch 1867 Status Status A Feeding Native 1st record Invaded countries habits range in Europe phytoAfrica N/A PT-MAD phagous (ES-CAN) Habitat Hosts References N/A N/A (N/A) Oromí and García (1995) phytophagous (her) Australasia 1998, GB GB J100 Pteridopsida (op: Pteridopsida) Hackett (1998), Hill et al. (2005) A phytophagous (xmp) C&S America 2000, IT IT G1, I2 Populus (pp: broadleaf trees) Tremblay et al. (2000) A phytophagous (xmp) Asia 2008, IT IT G Faccoli et al. (2009) Aphanarthrum affine Wollaston 1860* A Africa 1860, ES-CAN ES-CAN F8 Aphanarthrum bicinctum Wollaston 1860* Aphanarthrum bicolor Wollaston 1860* A phytophagous (her) phytophagous (her) phytophagous (her) phytophagous (her) phytophagous (her) Aesculus hippocastanum, Prunus persica (pp: broadleaf trees) Euphorbia (mp: Euphorbia) Africa 1860, ES-CAN ES-CAN F8 Euphorbia (mp: Euphorbia) Israelson (1972) Africa 1972, PT- PT-MAD (ES-CAN) MAD F8 Euphorbia (mp: Euphorbia) Israelson (1972) Africa ES-CAN F8 Euphorbia (mp: Euphorbia) Israelson (1980) Africa 1972, PT- PT-MAD (ES-CAN) MAD F8 Euphorbia (mp: Euphorbia) Israelson (1972) Platypodinae Megaplatypus mutatus (Chapuis 1865) Scolytinae Ambrosiodmus rubricollis Eichhoff 1875 A Aphanarthrum mairei Peyerimhoff 1923* A Aphanarthrum piscatorium Wollaston 1860* A 1928, ES-CAN Israelson (1972) Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) A 250 Family / subfamily Species Styphloderes (Parastyphloderes) lindbergi Roudier 1963* Syagrius intrudens Waterhouse 1903 Family / subfamily Species Cisurgus wollastonii (Eichhoff 1878)* Status A 1st record Invaded countries in Europe 1860, ES-CAN ES-CAN Hosts References F8 Euphorbia (mp: Euphorbia) Schedl (1946) ES-CAN, PT-AZO, PT-MAD I2 Bright (1987), Kirkendall per. obs. ES-CAN, FR, FR-COR, HU, IT, IT-SAR, IT-SIC, MT, PT-MAD ES-CAN I2 Phoenix, Washingtonia, Arecaceae, Dracaena (pp: Arecaceae, woody seeds) Phoenix, Chamaerops umilis, Arecaceae (pp: Arecaceae, woody seeds) Euphorbia (mp: Euphorbia) Coccotrypes carpophagus (Hornung 1842) A Coccotrypes dactyliperda (Fabricius 1801) A phytophagous (spe) Tropical, 1884, IT subtropical Coleobothrus alluaudi (Peyerimhoff 1923)* A Africa 1928, ES-CAN Cyclorhipidion bodoanus (Reitter 1913) A phytophagous (her) phytophagous (xmp) Asia 1960, FR BE, CH, DE, FR, IT, NL Dactylotrypes longicollis (Wollaston 1864) A phytophagous (spe) Africa 1949, (ES-CAN) FR-COR Dryocoetes himalayensis Strohmeyer 1908 A Asia2004, FR CH, FR Temperate Gnathotrichus materiarius (Fitch 1858) A phytophagous (phl) phytophagous (xmp) North America N/A Habitat F8 G1 ES, FR, FR-COR, HR, I2 IT, IT-SIC, PT-MAD G 1933, FR BE, CH, CZ, DE, ES, G FI, FR, IT, NL, SE Kirkendall and Faccoli (2010), Schedl (1963), Schedl et al. (1959), Targioni Tozzetti (1884) Israelson (1980) Quercus (op: Fagaceae) Audisio et al. (2008), Bouget and Noblecourt (2005), Kirkendall and Faccoli (2010), Schott (2004), Schott and Callot (1994) Phoenix canariensis, Balachowsky (1949), Lombardero Arecaceae, Dracaena and Novoa (1994), Sampò and draco (op: Arecaceae, Olmi (1975), Whitehead et al. Dracaenaceae) (2000) N/A (pp: Juglans Knížek (2004) regia, Pyrus lanata) Picea, Pinus (pp: conifers) Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 Feeding Native habits range phytoAfrica phagous (her) Tropical, phytophagous subtropical (spe) Balachowsky (1949), Faccoli (1998), Kirkendall and Faccoli (2010), Valkama et al. (1997), Wittenberg (2005) 251 Hypothenemus eruditus Westwood 1836 A Liparthrum artemisiae Wollaston 1854* A Liparthrum bituberculatum Wollaston 1854* Liparthrum curtum Wollaston 1854* A Liparthrum inarmatum Wollaston 1860* A Liparthrum mandibulare Wollaston 1854 A Monarthrum mali (Fitch 1855) A A A A Feeding habits phytophagous (phl) phytophagous (phl) Native range Asia 1st record Invaded countries in Europe 1991, MT MT I2 Ficus (pp) Mifsud and Knížek (2009) N/A PT-AZO G1 N/A (hpp) Base de dados da biodiversidade dos Açores 1924, IT-SIC N/A (hpp) N/A ES, ES-CAN, FR, J1 FR-COR, IL, IT, IT-SIC, MT, PT-AZO, PT-MAD PT-MAD F5 Balachowsky (1949), Machado and Oromí (2000), Noblecourt (2004), Pfeffer (1995), Ragusa (1924), Roll et al. (2007) Schedl (1963) N/A ES-CAN, PT-MAD G1 Laurus (mp: Laurus) Israelson (1990) N/A PT-AZO, PT-MAD G1 Israelson (1990) N/A ES-CAN, PT-MAD F8 Castanea, Ficus (pp: Euphorbiaceae, Moraceae, Fabaceae, Fagaceae) Euphorbia (mp: Euphorbia) Africa N/A (ES-CAN) ES, GB, PT-MAD G1 North America IT G C&S America (+ North Am.) phytoC&S phagous America (phl, spe) (+ North Am.) phytoAfrica phagous (ES-CAN) (phl) phytoAfrica phagous (North) (phl) phytoAfrica phagous (ES-CAN) (phl) phytophagous (her) phytophagous (phl) phytophagous (xmp) Africa 2007, IT Habitat Hosts Artemisia (mp: Artemisia) References Israelson (1990) Alnus, Betula, Israelson (1990), Lombardero Castanea, Euphorbia, and Novoa (1993) Erica, Quercus, Rubus (hpp) N/A (pp: broadleaf Kirkendall et al. (2008) trees) Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Status 252 Family / subfamily Species Hypocryphalus scabricollis (Eichhoff 1878) Hypothenemus crudiae (Panzer 1791) Status Phloeotribus liminaris (Harris 1852) A Polygraphus proximus Blandford 1894 A Xyleborinus attenuatus Wood & Bright 1992 A Xyleborus affinis Eichhoff 1868 A phytophagous (xmp) Xyleborus atratus Eichhoff 1875 A Xyleborus pfeilii (Ratzeburg 1837)8 A Xylosandrus crassiusculus (Motschulsky 1866) A phytophagous (xmp) phytophagous (xmp) phytophagous (xmp) 1st record Invaded countries in Europe 1940, FR FR, NL phytophagous (phl) phytophagous (phl) phytophagous (xmp) North America 2004, IT Asia 2000, RU RU G3 Abies (mp: Abies) Asia 1987, AT, AT, CH, CZ, DE, ES, CZ HU, NL, PL, RU, SE, SK, UA G1 C&S America (+ North Am.) Asia 2006, AT AT I Alnus, Betula, Salix, Tilia, Quercus, Corylus, broadleaf trees (pp: broadleaf trees) Dracaena (pp: broadleaf trees) 2007, IT G Asia Asia IT IT Habitat Hosts FA, G5 Thuja, Chamaecyparis, Juniperus chinensis, Cupressaceae (op: Cupressaceae) I2 Prunus serotina (mp: Prunus) 1837, DE AT, BG, CH, CZ, DE, G ES, FR, HR, HU, IT, PL, SI, SK, UA 2003, IT IT G2, J100 N/A: Quercus? (pp: broadleaf trees) References Balachowsky (1949), Moraal (2009) Pennacchio et al. (2004) Chilahsayeva (2008), Mandelshtam and Popovichev (2000) Essl and Rabitsch (2002), Kirkendall and Faccoli (2010) Holzer (2007) Faccoli (2008) Alnus, Betula, Populus Kirkendall and Faccoli (2010), (pp: broadleaf trees) Ratzeburg (1837) Ceratonia siliqua (pp: broadleaf trees, Pinus) Pennacchio et al. (2003) Xyleborus pfeilii was until recently treated as native to Europe, but is now thought to be introduced (Kirkendall and Faccoli 2010). 253 8 A Feeding Native habits range phytoAsia phagous (phl) Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 Family / subfamily Species Phloeosinus rudis Blandford 1894 Status Xylosandrus morigerus (Blandford 1894) A phytophagous (xmp) Asia A phytophagous (xyl) phytophagous (xyl) phytophagous (xyl) phytophagous (xyl) AsiaTropical 2004, ES-CAN ES-CAN, PT-AZO, PT-MAD I N/A (op: Musa, Ensete) Machado and Oromí (2000) AsiaTropical 1998, ES-CAN ES-CAN I2 Phoenix, Arecaceae (op: Arecaceae) C&S America 2004, ES-CAN ES-CAN I1 N/A (Ananas) Gonzales et al. (2002), Machado and Oromí (2000), Salomone Suárez et al. (2000) Machado and Oromí (2000) AsiaTropical 1993, ES CY, ES, ES-CAN, X24, I2 Arecaceae (op: FR, FR-COR, GR, Arecaceae) GR-CRE, GR-SEG, IL, IT, IT-SAR, IT-SIC C&S America 2006, IT-SIC FR, IT-SIC Diocalandra frumenti (Fabricius 1801) A Paradiaphorus crenatus (Billberg 1820) A Rhynchophorus ferrugineus (Olivier 1790) A Scyphophorus acupunctatus Gyllenhal 1838 A phytophagous (her) 1st record Invaded countries Habitat Hosts in Europe 1950, DE AT, BE, CH, CZ, DE, G Fagus, Castanea, FR, HU, IT, NL, PL, Buxus, Ficus, RU, SI Carpinus, Quercus, Juglans, Picea, Pinus (pp: broadleaf trees, conifers) 1916, AT, AT, CZ, FR, GB, IT J100 greenhouse orchids CZ, FR, as Dendrobium (pp: GB broadleaf trees) I2 References Henin and Versteirt (2004), Kirkendall and Faccoli (2010) Kirkendall and Faccoli (2010), Reitter (1916) Barranco et al. (1996), Bitton and Nakache (2000), EPPO (2006), FREDON-Corse (2007), Kehat (1999), Kontodimas et al. (2006), MAPA (2006), Sacchetti et al. (2005) Agave (pp: Agavaceae, Germain et al. (2008b), Longo Dracaenaceae) (2007) Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Dryophthoridae Cosmopolites sordidus (Germar 1824) A Feeding Native habits range phytoAsia phagous (xmp) 254 Family / subfamily Species Xylosandrus germanus (Blandford 1894) Family / subfamily Species Sitophilus linearis (Herbst 1797) Status A Feeding Native habits range phytoTropical, phagous subtropical (spe) A phytophagous (spe) Sitophilus zeamais Motschulsky 1855 C phytophagous (spe) A phytophagous (rbw) phytophagous (lbw) Erirhinidae Lissorhoptrus oryzophilus Kuschel 1952 Stenopelmus rufinasus Gyllenhal 1835 A AsiaTropical Habitat J1 1896, SE AL, AT, BG, BY, CH, J1 CY, CZ, DE, DK, EE, ES, ES-CAN, FI, FR, FR-COR, GB, GL, HR, HU, IS, IT, IT-SAR, IT-SIC, LT, LV, MT, NL, NO, PL, PT, PT-AZO, RO, SE, UA Cryptogenic 1927, DE AD, AL, AT, BE, BG, J1 CH, CZ, DE, DK, EE, ES-CAN, FI, FR, GB, IT, IT-SAR, IT-SIC, PL, PT, PT-AZO, PT-MAD, RU, SE North America 2004, IT IT North America 1900, FR BE, DE, ES, FR, GB, IE, IT, NL I1 Hosts Tamarindus indica (mp: Tamarindus indica) grain (op: cereal grain) grain (op: cereal grain) Oryza, Carex (pp: Gramineae, Cyperaceae) C1, C2 Azolla (mp: Azolla) References Abbazzi et al. (1994), Essl and Rabitsch (2002), Hoffmann (1954), Machado and Oromí (2000), Tomov et al. (2009) Abbazzi et al. (1994), Balachowsky (1963), Essl and Rabitsch (2002), Hoffmann (1954), Joakimow (1904), Machado and Oromí (2000), Silfverberg (2004a), Silfverberg (2004b), Teodorescu et al. (2006), Tomov et al. (2009), Wittenberg (2005) Balachowsky (1963), Dal Monte (1972), Essl and Rabitsch (2002), Haghebaert (1991), Lundberg (1995), Machado and Oromí (2000), Obretenchev et al. (1990), Tomov et al. (2009), Wittenberg (2005) Caldara et al. (2004) Baars and Caffery (2008), Dana and Viva (2006), Fernandez Carrillo et al. (2005), Hill et al. (2005), Janson (1921) Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 Sitophilus oryzae (Linnaeus 1763) 1st record Invaded countries in Europe 1954, AL, AT, ES-CAN, FR, FR-COR FR-COR, IT, PL 255 Family / subfamily Species Anthribidae Bruchela rufipes (Olivier 1790) Habitat Hosts phytophagous (spe) Europe GB I2 N/A (mp: Reseda lutea) Hill et al. (2005), Morris (1990) phytophagous (her) Europe, Mediterranean, Asia phytophagous (her) Europe, Mediterranean phytophagous Europe, West Mediterranean phytophagous (spe) Europe, Mediterranean, Asia phytophagous (spe) Mediterranean ES-CAN, PT-AZO N/A N/A (op: Malvaceae) ES-CAN N/A N/A (mp: Trifolium) Base de dados da biodiversidade dos Açores, Machado and Oromí (2000) Machado and Oromí (2000) ES-CAN N/A N/A (op: Fabaceae) Machado and Oromí (2000) ES-CAN N/A N/A (mp: Ononis) Machado and Oromí (2000) ES-CAN N/A N/A (mp: Lotus) Machado and Oromí (2000) ES-CAN N/A N/A (mp: Trifolium) Machado and Oromí (2000) GB I2, H5 phytophagous (her) Europe, Mediterranean ES-CAN, PT-AZO N/A Viscum album (mp: Viscum album) N/A (mp: Mercurialis) Duff (2008), Foster et al. (2001) Base de dados da biodiversidade dos Açores, Machado and Oromí (2000) Brachyceridae Brachycerus plicatus Gyllenhal 1833* phytophagous (her?) Mediterranean ES-CAN N/A N/A (op: Liliaceae?) Machado and Oromí (2000) Catapion pubescens (W. Kirby 1811)* Eutrichapion (Cnemapion) vorax (Herbst 1797)* Holotrichapion (Holotrichapion) ononis (W. Kirby 1808)* Native range Ischnopterapion (Ischnopterapion) plumbeomicans (Rosenhauer 1856)* Ischnopterapion (Chlorapion) virens phytophagous (her) Europe, (Herbst 1797)* Mediterranean, Asia Ixapion variegatum (Wencker 1864) phytophagous (her) Europe Kalcapion semivittatum (Gyllenhal 1833)* References Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Invaded countries Apionidae Aspidapion (Aspidapion) radiolus (Marsham 1802)* Feeding habits 256 Table 8.2.2. Characteristics of the Curculionoidea species alien in Europe. See Table 8.2.1 legend. Native range: „Mediterranean“ refers to southern Europe, North Africa and western Asia; „West Mediterranean“ refers to southern Europe and North Africa. Family / subfamily Feeding habits Species Curculionidae Bagoinae Bagous exilis Jacquelin du Val 1854* phytophagous Habitat Hosts References West Mediterranean ES-CAN B N/A (coastal shrubs: Frankenia, Chenopodiaceae) Machado and Oromí (2000) West Mediterranean phytophagous (her) West Mediterranean ES-CAN N/A N/A (N/A) Machado and Oromí (2000) ES-CAN I, J N/A (op: Cruciferae) Machado and Oromí (2000) phytophagous (spe) Europe, West Mediterranean phytophagous (spe) West Mediterranean phytophagous (rbo) Europe, West Mediterranean phytophagous (rbo) Europe, West Mediterranean, Asia PT-AZO I, J Borges et al. (2005) ES-CAN E N/A (op: Brassica, Cruciferae) N/A (mp: Erica) PT-AZO I, G Echium (mp: Echium) Borges et al. (2005) FÖ E, I N/A (mp: Rumex) N/A phytophagous (xyl) Europe, West Mediterranean phytophagous (xyl) Europe PT-AZO I2 N/A (op: Pinaceae) Borges et al. (2005) PT-AZO B, E Stüben (2003) phytophagous (xyl) Europe PT-AZO N/A phytophagous (xyl) Europe PT-AZO G marine driftwood (pp: decaying wood) N/A (pp: decaying wood) N/A (pp: dead wood) phytophagous (her?) West Mediterranean ES-CAN, PT-AZO N/A Baridinae Melaleucus sellatus (Boheman 1844)* phytophagous Melanobaris quadraticollis (Boheman 1836)* Ceutorhynchinae Ceutorhynchus assimilis (Paykull 1800) Micrelus ferrugatus (Perris 1847)* Mogulones geographicus (Goeze 1777) Rhinoncus pericarpius (Linnaeus 1758) Cossoninae Brachytemnus porcatus (Germar 1824) Pselactus spadix (Herbst 1795) Pseudophloeophagus aeneopiceus (Boheman 1845)* Rhopalomesites tardyi (Curtis 1825) Cryptorhynchinae Dichromacalles (Dichromacalles) dromedarius (Boheman 1844)* Machado and Oromí (2000) Base de dados da biodiversidade dos Açores Borges et al. (2005) N/A (op: Compositae) Base de dados da biodiversidade dos Açores, Machado and Oromí (2000) 257 Invaded countries Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 Native range Habitat Hosts References ES-CAN I, J N/A (mp: Plantago) Machado and Oromí (2000) ES-CAN I, J ES-CAN, PT-AZO ES-CAN I, J N/A N/A (op: Machado and Oromí (2000) Scrophulariaceae) Plantago (mp: Plantago) Borges et al. (2005), Machado and Oromí (2000) N/A (N/A) Machado and Oromí (2000) ES-CAN ES-CAN N/A N/A N/A (N/A) Limonium (N/A) Machado and Oromí (2000) Machado and Oromí (2000) ES-CAN N/A N/A (N/A) Machado and Oromí (2000) ES-CAN N/A Machado and Oromí (2000) Smicronyx albosquamosus Wollaston phytophagous (her?) West 1854* Mediterranean Smicronyx brevicornis Solari 1952* phytophagous (her) West Mediterranean Tychius (Tychius) cuprifer (Panzer phytophagous (spe) Europe, 1799) Mediterranean Tychius (Tychius) picirostris phytophagous (spe) Europe, (Fabricius 1787) Mediterranean, Asia Tychius (Tychius) stephensi phytophagous (spe) Europe, Schonherr 1836* Mediterranean, Asia ES-CAN, PT-MAD ES-CAN N/A N/A (pp: Caryophyllaceae, Plumbaginaceae, Thymelaeaceae) N/A (N/A) N/A N/A (mp: Cuscuta) PT-AZO I1 N/A (mp: Trifolium) PT-AZO I1, E N/A (mp: Trifolium) Borges et al. (2005), Stüben (2003) Borges et al. (2005) ES-CAN N/A N/A (mp: Trifolium) Machado and Oromí (2000) Hoffmann (1958), Machado and Oromí (2000) Machado and Oromí (2000) Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Invaded countries 258 Family / subfamily Feeding habits Native range Species Curculioninae Mecinus circulatus (Marsham 1802)* phytophagous (her) Europe, Mediterranean Mecinus longiusculus Boheman phytophagous (her) West 1845* Mediterranean Mecinus pascuorum (Gyllenhal phytophagous (spe) Europe, 1813) Mediterranean Pachytychius aridicola (Wollaston phytophagous (spe) Mediterranean 1864)* Philernus farinosus Gyllenhal 1835* phytophagous Europe, Asia Sibinia (Dichotychius) albosquamosa phytophagous (spe?) Mediterranean Pic 1904* Sibinia (Dichotychius) planiuscula phytophagous (spe?) Mediterranean (Desbrochers 1873)* Sibinia (Sibinia) primita (Herbst phytophagous (spe) Europe, West 1795)* Mediterranean Feeding habits Native range Habitat Hosts References phytophagous (spe) Mediterranean Invaded countries ES-CAN N/A N/A (mp: Ononis) Machado and Oromí (2000) phytophagous Mediterranean ES-CAN N/A Opuntia (N/A) Machado and Oromí (2000) phytophagous phytophagous (rbw?) phytophagous Europe Europe FÖ IS G I N/A Ólafsson (1991) Europe PT-AZO F5 N/A (N/A) Medicago (mp: Medicago) Pittosporum? (N/A) phytophagous (rbw) Europe, GB Mediterranean phytophagous (rbw) Europe (southern) DE, DK, FR, GB Otiorhynchus (Arammichnus) cribricollis Gyllenhal 1834 phytophagous (rbw) West Mediterranean Otiorhynchus (Arammichnus) dieckmanni Magnano 1979 phytophagous (rbw) Europe (western) phytophagous (rbw) Europe (Alps) J100 phytophagous (rbw) Mediterranean N/A I2 N/A (Cyclamen) Hill et al. (2005) I Pyrus? (N/A) ES-CAN, PT-AZO I, J N/A (mp: Artemisia) DK, SE G, I2 N/A (N/A) Barclay (2001), Lucht (1985), Palm (1996), Valladares and Cocquempot (2008) Borges et al. (2005), Machado and Oromí (2000), Stüben (2003) Borisch (1997), Runge (2008), Silfverberg (2004a), Silfverberg (2004b) DK 259 phytophagous (rbw) Europe (southern) SE N/A (pp: Acer, Camelia, Heijerman et al. (2003), Hill Prunus, Rhododendron) et al. (2005), Runge (2008) N/A (Alnus) Borisch (1997), Hill et al. (2005) Fragaria, Vitis, Carduus, Borisch (1997), Silfverberg Rumex (N/A) (2004a), Silfverberg (2004b) N/A (N/A) Runge (2008) phytophagous (rbw) Europe (central) DK, GB, MT, I2 NL, SE GB, SE I2, J4 Stüben (2003) Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 Family / subfamily Species Tychius (Tychius) striatulus Gyllenhal 1836* Cyclominae Gronops fasciatus Kuster 1851* Entiminae Barynotus squamosus Germar 1824 Barypeithes (Exomias) pellucidus (Boheman 1834) Cathormiocerus (Cathormiocerus) curvipes (Wollaston 1854) Otiorhynchus (Otiorhynchus) apenninus Stierlin 1883 Otiorhynchus (Otiorhynchus) armadillo (Rossi 1792) Otiorhynchus (Nehrodistus) armatus Boheman 1843 Otiorhynchus (Otiorhynchus) aurifer Boheman 1843 Otiorhynchus (Pocodalemes) crataegi Germar 1824 Otiorhynchus (Nehrodistus) corruptor (Host 1789) Sitona (Sitona) lepidus Gyllenhal 1834 Sitona (Sitona) lineatus (Linnaeus 1758)* Sitona (Sitona) macularius (Marsham 1802)* Native range Invaded countries CH Habitat Hosts References J Vincetoxicum (N/A) Germann (2004) phytophagous (rbw) Europe, West Mediterranean phytophagous (rbw) Europe PT-AZO N/A FÖ, IS I2 N/A (pp: Rumex, Dactylis, Trifolium...) N/A (N/A) Base de dados da biodiversidade dos Açores Ólafsson (1991) phytophagous (rbw) Europe PT-AZO F5 phytophagous (rbw?) phytophagous Europe, West Mediterranean Europe (southcentral, southeastern) phytophagous (rbw) Mediterranean PT-AZO I,G Pittosporum? (pp: Vitis...) N/A (Ammophila) Borges et al. (2005), Stüben (2003) Borges et al. (2005) SE I1 N/A (N/A) Lundberg (2006) ES-CAN N/A Machado and Oromí (2000) phytophagous (rbw) West Mediterranean phytophagous (rbw) Mediterranean ES-CAN N/A N/A (pp: Resedaceae, Cruciferae) N/A (mp: Astragalus) PT-AZO I, G Borges et al. (2005) phytophagous (rbw) Mediterranean ES-CAN, PT-AZO ES-CAN, PT-AZO I, G N/A (op: Lotus, Trifolium, Fabaceae) N/A (mp: Medicago) N/A N/A (mp: Lupinus) PT-AZO I, J ES-CAN, PT-AZO I, J N/A (op: Lotus, Trifolium, Fabaceae) N/A (op: Fabaceae) ES-CAN I, J N/A (mp: Trifolium) phytophagous (rbw) Europe (eastern) phytophagous (rbw) Mediterranean, Asia phytophagous (rbw) Europe, Mediterranean phytophagous (rbw) Europe, Mediterranean, Asia phytophagous (rbw) Europe, Mediterranean, Asia Machado and Oromí (2000) Borges et al. (2005), Machado and Oromí (2000) Base de dados da biodiversidade dos Açores, Machado and Oromí (2000) Borges et al. (2005) Base de dados da biodiversidade dos Açores, Machado and Oromí (2000) Machado and Oromí (2000) Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Rhytideres (Rhytideres) plicatus (Olivier 1790)* Sitona (Charagmus) cachectus Gyllenhal 1834* Sitona (Sitona) cinnamomeus Allard 1863 Sitona (Sitona) discoideus Gyllenhal 1834 Sitona (Charagmus) gressorius (Fabricius 1792)* Feeding habits 260 Family / subfamily Species Otiorhynchus (Padilehus) pinastri (Herbst 1795) Otiorhynchus (Zustalestus) rugosostriatus (Goeze 1777)* Otiorhynchus (Metopiorrhynchus) singularis (Linnaeus 1767) Otiorhynchus (Dorymerus) sulcatus (Fabricius 1775) Philopedon plagiatum (Schaller 1783) Psallidium (Psallidium) maxillosum (Fabricius 1792) Feeding habits Native range phytophagous (rbw) Mediterranean phytophagous (rbw) Mediterranean Sitona (Sitona) puncticollis Stephens phytophagous (rbw) Europe, 1831 Mediterranean, Asia Sitona (Charagmus) variegatus phytophagous (rbw) West Fåhraeus 1840* Mediterranean Strophosoma (Strophosoma) phytophagous Europe melanogrammum melanogrammum (rbw?) (Forster 1771) Trachyphloeus (Trachyphloeus) phytophagous Europe angustisetulus Hansen 1915* (rbw?) Trachyphloeus (Trachyphloeus) phytophagous Mediterranean laticollis Boheman 1843* (rbw?) Trachyphloeus (Trachyphloeus) phytophagous (rbw) Europe, Asia spinimanus Germar 1824* Hyperinae Coniatus (Coniatus) tamarisci phytophagous Mediterranean (Fabricius 1787)* Donus (Antidonus) lunatus phytophagous (lbw) Europe, (Wollaston 1854)* Mediterranean, Asia Hypera (Hypera) melancholica phytophagous (lbw) Europe, (Fabricius 1792)* Mediterranean, Asia Invaded countries ES-CAN Habitat Hosts References N/A N/A (Fabaceae?) Machado and Oromí (2000) ES-CAN, PT-AZO, PT-MAD I, J N/A (mp: Lotus) FÖ, PT-AZO I N/A (op: Trifolium, Melilotus?) Borges et al. (2005), Hoffmann (1950), Machado and Oromí (2000), Stüben (2003) Borges et al. (2005) ES-CAN N/A N/A (mp: Astragalus) Machado and Oromí (2000) PT-AZO G, I2 N/A (pp: Rumex, Aira...) Borges et al. (2005) ES-CAN N/A N/A (N/A) Machado and Oromí (2000) ES-CAN N/A Machado and Oromí (2000) ES-CAN N/A Mercurialis, Bidens (N/A) N/A (mp: Cynodon) ES-CAN N/A N/A (mp: Tamarix) Machado and Oromí (2000) ES-CAN E N/A (op: Geraniaceae) Machado and Oromí (2000) ES-CAN I, J N/A (op: Medicago, Trifolium) Machado and Oromí (2000) Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 Family / subfamily Species Sitona (Sitona) ocellatus Kuster 1849* Sitona (Sitona) puberulus Reitter 1903 Machado and Oromí (2000) 261 Feeding habits Native range Habitat Hosts References E, J N/A (op: Ononis, Trifolium) Machado and Oromí (2000) ES-CAN E N/A (mp: Ononis) Machado and Oromí (2000) ES-CAN, PT-AZO I, J N/A (op: Fabaceae) Borges et al. (2005), Machado and Oromí (2000) ES-CAN, PT-AZO N/A N/A (N/A) West Mediterranean phytophagous (rbo) Mediterranean, Africa phytophagous Mediterranean ES-CAN N/A N/A (N/A) Base de dados da biodiversidade dos Açores, Machado and Oromí (2000) Machado and Oromí (2000) ES-CAN B Machado and Oromí (2000) ES-CAN N/A N/A (op: Chenopodiaceae) N/A (N/A) phytophagous (her) West Mediterranean phytophagous (her) West Mediterranean phytophagous (her) Europe, Mediterranean phytophagous (her) Mediterranean, Asia phytophagous (her) Europe, Mediterranean phytophagous (her) Europe, Asia, North Africa ES-CAN E? Machado and Oromí (2000) ES-CAN N/A N/A (op: Cheiranthus, Sinapis) N/A (mp: Atriplex) ES-CAN I, J N/A (mp: Carduus) Machado and Oromí (2000) ES-CAN N/A Machado and Oromí (2000) ES-CAN I, J N/A (op: Chenopodiaceae) N/A (mp: Rumex) ES-CAN N/A N/A (op: Malvaceae, Fabaceae) Machado and Oromí (2000) phytophagous (lbw) Europe, Mediterranean, Asia Hypera (Hypera) ononidis (Chevrolat phytophagous (lbw) Europe, West 1863)* Mediterranean Hypera (Hypera) postica (Gyllenhal phytophagous (lbw) Europe, 1813) Mediterranean, Asia Lixinae Coniocleonus excoriatus (Gyllenhal phytophagous Europe, 1834)* Mediterranean Coniocleonus variolosus (Wollaston 1864)* Conorhynchus (Pycnodactylus) brevirostris (Gyllenhal 1834)* Conorhynchus (Pycnodactylus) conicirostris (Olivier 1807)* Lixus (Compsolixus) anguinus (Linnaeus 1767)* Lixus (Eulixus) brevirostris Boheman 1835* Lixus (Epimeces) filiformis (Fabricius 1781)* Lixus (Compsolixus) juncii Boheman 1835* Lixus (Dilixellus) linearis Olivier 1807* Lixus (Dilixellus) pulverulentus (Scopoli 1763)* phytophagous Machado and Oromí (2000) Machado and Oromí (2000) Machado and Oromí (2000) Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Invaded countries ES-CAN 262 Family / subfamily Species Hypera (Hypera) nigrirostris (Fabricius 1775)* Family / subfamily Species Rhinocyllus conicus (Froelich 1792) Molytinae Anisorhynchus hespericus Desbrochers 1875* Ita crassirostris Tournier 1878* Liparus (Liparus) glabrirostris Küster 1849 Pissodes (Pissodes) castaneus (De Geer 1775)* Native range phytophagous (spe) Europe, Mediterranean phytophagous (her) Europe, Mediterranean, Asia phytophagous phytophagous phytophagous Invaded countries LT, LV, SE Habitat Hosts References E, I N/A (op: Carduus, Cirsium, Galactites, Cynara...) Gillerfors (1988), Lundberg (2006) GB G3 Pinus (mp: Pinus) Hill et al. (2005) N/A N/A (N/A) Machado and Oromí (2000) N/A G N/A (N/A) N/A (mp: Heracleum) Machado and Oromí (2000) Hansen (1996) ES-CAN, PT-AZO G N/A (mp: Pinus) Base de dados da biodiversidade dos Açores, Machado and Oromí (2000) ES-CAN G1, I2 GB G3 GB G3 ES-CAN, PT-MAD GB G1 G3 Laurus (pp: Pistacia, Schedl et al. (1959) Cotinus, Olea, Smilax) Picea (op: Pinus, Abies, Alexander (2002) Picea) Picea (mp: Picea) Alexander (2002), Hill et al. (2005) Laurus (pp: Laurus, Schedl (1963), Schedl et al. Alnus) (1959) Pinus (mp: Pinus) Alexander (2002) GB, PT-AZO G3, I2 Pinus (mp: Pinus) Europe ES-CAN (southwestern) Europe (southern) ES-CAN Europe (Alps) DK phytophagous (phl) Europe, Mediterranean, Asia Scolytinae Chaetoptelius vestitus (Mulsant & phytophagous (phl) Mediterranean, Rey 1860)* Asia Crypturgus subcribrosus Eggers 1933 phytophagous (phl) Europe (central, eastern) phytophagous (phl) Europe, Asia Dendroctonus micans (Kugelann 1794) Dryocoetes villosus (Fabricius 1792)* phytophagous (phl) Europe, West Mediterranean Hylastes angustatus (Herbst 1793) phytophagous (phl) Europe (southern, central), Asia Hylastes ater (Paykull 1800) phytophagous (phl) Europe, Asia Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 Mesoptiliinae Magdalis (Magdalis) memnonia (Gyllenhal 1837) Feeding habits Alexander (2002), Bright (1987) 263 Hylastes cunicularius Erichson 1836 Hylastes linearis Erichson 1836* Native range phytophagous (phl) Europe, Mediterranean, Asia phytophagous (phl) Europe, Asia phytophagous (phl) Europe, West Mediterranean phytophagous (phl, Europe, West rbo) Mediterranean Hylurgops palliatus (Gyllenhal 1813) phytophagous (phl) Europe, Mediterranean, Asia Hylurgus ligniperda (Fabricius phytophagous (phl) Europe, 1787)* Mediterranean, Asia Hypoborus ficus Erichson 1836* phytophagous (phl) Europe, West Mediterranean Ips cembrae (Heer 1836) phytophagous (phl) Europe (central) Ips duplicatus (Sahlberg 1836) phytophagous (phl) Europe (northeastern, Russia) phytophagous (phl) Europe, Mediterranean, Asia phytophagous (phl) Mediterranean (eastern) phytophagous (phl) Europe, West Mediterranean Orthotomicus erosus (Wollaston 1857)* Phloeosinus armatus Reitter 1887 Phloeosinus aubei (Perris 1855)9 Invaded Habitat Hosts countries GB, PT-AZO, G3 Pinus (mp: Pinus) PT-MAD References GB ES-CAN, PT-MAD ES-CAN, PT-MAD G3 G3 GB G3 Alexander (2002), Bright (1987), Mandelshtam et al. (2006) Picea (mp: Picea) Alexander (2002) Pinus (mp: Pinus) Schedl (1963), Schedl et al. (1959) Cytisus, Laurus, Schedl (1963), Schedl et al. Castanea (op: Trifolium, (1959) Fabaceae); N/A (op: Pinaceae) Alexander (2002) ES-CAN, PT-AZO, PT-MAD ES-CAN, PT-AZO, PT-MAD DK, GB, NL G3 Pinus (mp: Pinus) Bright (1987), Schedl (1963), Schedl et al. (1959) I2 Echium, Ficus (mp: Ficus) Bright (1987), Schedl (1963), Schedl et al. (1959) G3 AT, BE, SK G3 Larix (op: Larix, Pinus cembra) Picea abies (mp: Picea) PT-MAD G3 Pinus (mp: Pinus) EPPO (2005), Hill et al. (2005), Stauffer et al. (2001) Essl and Rabitsch (2002), OPIE (2002), Piel et al. (2006) Schedl (1963) IT FA, G5 ES-CAN, NL G3 Cupressus (op: Cupressaceae) Juniperus (op: Cupressaceae) F5, F7 Covassi (1991) Moraal (2006), Oromí and García (1995) Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Hylastinus obscurus (Marsham 1802)* Feeding habits 264 Family / subfamily Species Hylastes attenuatus Erichson 1836 Family / subfamily Species Phloeosinus thujae (Perris 1855) Habitat Hosts AT, CZ, FR FA, G5 Phloeotribus cristatus (Fauvel 1889)* phytophagous (phl) West Mediterranean Phloeotribus rhododactylus phytophagous (phl) Europe, West (Marsham 1802)* Mediterranean Phloeotribus scarabaeoides (Bernard phytophagous (phl) Europe, West 1788)* Mediterranean Pityophthorus traegardhi Spessivtseff phytophagous (phl) Europe 1921 (northern), Asia Polygraphus poligraphus (Linnaeus phytophagous (phl) Europe (central, 1758) northern, eastern) Pteleobius kraatzii (Eichhoff 1864)* phytophagous (phl) Europe, West Mediterranean Scolytus amygdali Guérin-Méneville phytophagous (phl) Europe, 1847* Mediterranean, Asia Scolytus laevis Chapuis 1869 phytophagous (phl) Europe ES-CAN F5, F7 PT-MAD F5, F7 ES-CAN I2 AT Scolytus pygmaeus (Fabricius 1787) phytophagous (phl) Europe GB Scolytus rugulosus (Muller 1818)* phytophagous (phl) Europe, Mediterranean, Asia PT-AZO phytophagous (phl) Europe, West Mediterranean phytophagous (phl) Europe (eastern), Asia FA, G5 References Juniperus (op: Alexander (2002), Machado Cupressaceae) and Oromí (2000) Fraxinus (mp: Fraxinus) Bouget and Noblecourt (2005), Essl and Rabitsch (2002), Schott and Callot (1994) N/A: Fabaceae? (op: Machado and Oromí (2000) Fabaceae) Cytisus (op: Fabaceae) Schedl (1963) Machado and Oromí (2000) G3 N/A: Oleaceae? (op: Oleaceae) Picea (mp: Picea) GB G3 N/A (op: Pinaceae) Alexander (2002) ES-CAN I2, G1, G5, FA I2 N/A: Ulmus? (mp: Ulmus) Prunus (op: Rosaceae trees) Pfeffer (1995) ES-CAN GB G1, G5, Ulmus (mp: Ulmus) I2 G1, I2, Ulmus (mp: Ulmus) FA, FB I2 N/A (op: Rosaceae trees) Holzschuh (1994) Israelson (1969) Hill et al. (2005) Hill et al. (2005) Bright (1987) This species was incorrectly reported from the Canary Islands (Oromí and García 1995) as P. gillerforsi Bright, an Azores endemic. Specimens so identified have been examined by Kirkendall, and they belong to the common Mediterranean species P. aubei. 265 9 Native range Weevils and Bark Beetles (Coleoptera, Curculionoidea). Chapter 8.2 Invaded countries ES-CAN, GB Phloeotribus caucasicus Reitter 1891 Feeding habits Feeding habits Habitat Hosts References G3, I2 Pinus (mp: Pinus) Schedl (1963) Europe, Mediterranean, Asia ES-CAN, PT-AZO, PT-MAD I2 Laurus, Pinus, Castanea Bright (1987), Schedl et al. (pp: broadleaves, (1959) conifers) West Mediterranean ES-CAN E6 N/A (N/A) Machado and Oromí (2000) Europe, Mediterranean ES-CAN N/A N/A (N/A) Machado and Oromí (2000) phytophagous (her) Europe, Mediterranean phytophagous (spe) West Mediterranean ES-CAN N/A N/A (mp: Lythrum) Machado and Oromí (2000) ES-CAN N/A N/A (op: Juniperus, Cupressus) Machado and Oromí (2000) phytophagous (spe) Europe, Mediterranean, Asia GB G3 Pinus sylvestris (mp: Pinus sylvestris) Duff (2008) phytophagous (phl) Europe, Asia phytophagous (xmp) Nemonychidae Cimberis attelaboides (Fabricius 1787) 10 11 Early records from Madeira refer to T. piniperda, but specimens collected by Kirkendall in 1999 are T. destruens; as the two species had been mixed up for a long time we think all records correspond to T. destruens This species has been improperly recorded in the Canary Islands as Xyleborus xylographus. Xyleborus xylographus (Say 1826), an oak specialist from the eastern United States, does not occur in any recent collections from the archipelago (or elsewhere in Europe), whereas X. saxesenii does (Kirkendall, unpublished data). The presence of X. xylographus on all Canary Islands species lists (Schedl et al. 1959, Oromi and Garcia 1995, Machado and Oromi 2000, Izquierdo et al. 2004), and the absence of X. saxesenii, seems to stem from an early mistaken treatment of X. saxesenii as a junior synonym of X. xylographus (Schedl 1970). To verify this, Kirkendall located one specimen recently determined as X. Xylographus (Oromi and Garcia 1995), and confirmed that it is X. saxesenii. Daniel Sauvard et al. / BioRisk 4(1): 219–266 (2010) Invaded countries PT-MAD Dryophthoridae Sphenophorus meridionalis Gyllenhal phytophagous 1838 (rbo?) Erirhinidae Procas armillatus (Fabricius 1801)* phytophagous Nanophyidae Dieckmanniellus nitidulus (Gyllenhal 1838)* Nanodiscus transversus (Aube 1850) Native range 266 Family / subfamily Species Tomicus destruens (Wollaston 1865)10 Xyleborinus saxesenii (Ratzeburg 1837)11 A peer reviewed open access journal BioRisk 4(1): 267–292 (2010) RESEARCH ARTICLE doi: 10.3897/biorisk.4.52 BioRisk www.pensoftonline.net/biorisk Leaf and Seed Beetles (Coleoptera, Chrysomelidae) Chapter 8.3 Ron Beenen1, Alain Roques2 1 Universiteit van Amsterdam, Zoölogisch Museum Amsterdam, Plantage Middenlaan 64, 1018 DH, Amsterdam, The Netherlands 2 INRA, UR633 Zoologie Forestière, 2163 Av. Pomme de pin, 45075 Orléans, France Corresponding author: Ron Beenen (r.beenen@wxs.nl) Academic editor: David Roy | Received 4 February 2010 | Accepted 22 May 2010 | Published 6 July 2010 Citation: Beenen R, Roques A (2010) Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 267–292. doi: 10.3897/biorisk.4.52 Abstract The inventory of the leaf and seed beetles alien to Europe revealed a total of 25 species of which 14 seed beetles (bruchids) and 11 leaf beetles mostly belonging to the subfamilies Alticinae and Chrysomelinae. At present, aliens account for 9.4% of the total fauna of seed beetles in Europe whereas this percentage is less than 1% for leaf beetles. Whilst seed beetles dominated the introductions in Europe until 1950, there has been an exponential increase in the rate of arrival of leaf beetles since then. New leaf beetles arrived at an average rate of 0.6 species per year during the period 2000–2009. Most alien species originated from Asia but this pattern is mainly due to seed beetles of which a half are of Asian origin whereas leaf beetles predominantly originated from North America (36.4%). Unlike other insect groups, a large number of alien species have colonized most of Europe. All but one species have been introduced accidentally with either the trade of beans or as contaminants of vegetal crops or stowaway. Most aliens presently concentrate in man-made habitats but little affect natural habitats (<6%). Highly negative economic impacts have been recorded on stored pulses of legumes and crops but very little is known about possible ecological impact. Keywords Coleoptera, Chrysomelidae, Bruchidae, seed beetle, leaf beetle, bioinvasion, alien, Europe, translocation, introduction Copyright R. Beenen, A. Roques. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 268 Ron Beenen & Alain Roques / BioRisk 4(1): 267–292 (2010) 8.3.1 Introduction The family Chrysomelidae is one of the largest Coleopteran families, including ca. 37 000 described species in the world and perhaps the same number as yet undescribed (Jolivet and Verma 2002). Bruchidae, or seed beetles, is a relatively small family. Kingsolver (2004), referring to the most recent world catalogue, mentions 1,346 valid bruchid species. Although there are good arguments to treat Bruchidae as a subfamily of Chrysomelidae and raise some leaf beetle subfamilies to family rank (Reid 1995), this is still not common practice among leaf beetle researchers. We treat Bruchidae and Chrysomelidae in this contribution as families, merely for practical reasons. According to Fauna Europaea, the fauna presently observed in Europe includes 1532 leaf beetles and 145 seed beetles. Except for important agricultural pests such as the Colorado potato beetle, Leptinotarsa decemlineata, and more recently, the western corn rootworm, Diabrotica virgifera virgifera, little was known about introductions of alien leaf beetles until Beenen (2006) revealed that 126 species have been translocated at least once from one continent to another. More information on alien seed beetles has been available in the literature mainly because of their potential impact on stored products (Southgate 1979). In the present work, we will show that 25 non-native species of leaf and seed beetles of which one is of unknown origin (cryptogenic) have already established in Europe (Table 8.3.1). Thus, aliens still represent only a very small proportion (1.5%) of the total fauna of leaf and seed beetles in Europe. By comparison, approximately 71 alien leaf beetle species have been recorded from North America (Beenen 2006, Beenen, unpubl.). Within Europe, changes in the distribution of native leaf beetles have also been noticed which can be partly associated either to human activity or to natural trends such as delayed post-glacial expansion and global warming. For example, the recent northwards expansion of a flea beetle, Longitarsus dorsalis, seems to result from both the introduction of a rapidly expanding invasive plant originating from South Africa, Senecio inaequidens DC., on which L. dorsalis thrives (Beenen 1992), and from increasing temperatures during the past years. However, the role of human activity is often difficult to ascertain in such observed range expansions of native species. We will essentially consider the species alien to Europe, a summary of the species alien in Europe (Table 8.3.2) and will present their characteristics at the end of the chapter. 8.3.2 Taxonomy A total of 25 alien species of which 14 seed beetles and 11 leaf beetles have been recorded as established in Europe (Table 8.3.1). Thus, bruchids represent more than a half (56.0%) of the alien species whereas they account for only 8.1% of the native fauna of seed and leaf beetles (Figure 8.3.1). This arrival of alien seed beetles has Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 269 Figure 8.3.1. Comparison of the relative importance of the subfamilies of Chrysomelidae and Bruchidae in the alien and native entomofauna in Europe. Subfamilies are presented in a decreasing order based on the number of alien species. The number right to the bar indicates the number of species per family. significantly modified the composition of the total fauna of seed beetles observed in Europe, where aliens account for 9.4% at present. The pattern is rather different for Chrysomelidae. Although this family includes 13 subfamilies in Europe the alien entomofauna is only distributed among five of these subfamilies. Large differences are observed in the contribution of each subfamily without any apparent correlation to its numerical importance in the native fauna. The recent arrival in France of an alien palm hispine beetle, Pistosia dactylifera (Drescher and Martinez 2005), largely modified the composition of the Hispinae subfamily which includes only three native species (Fauna Europaea 2009). However, aliens represent much less so for the two major subfamilies of leaf beetles, Alticinae flea beetles (four species- 0.7% of the total) and Chrysomelinae (four species- 1.3% of the total). Other alien species include one skeletonizing leaf beetle (Galerucinae) and one tortoise leaf beetle (Cassidinae). The same subfamily pattern is observed for translocations of leaf beetles at world level but Beenen (2006) also noticed other species belonging to Hispinae (e.g. Brontispa palm leaf beetles) and Criocerinae. It is noticeable that representatives from some important subfamilies such as Cryptocephalinae and Donaciinae have never been introduced, or never established at least. Leaf beetles and seed beetles largely differ in biological traits that may be involved in the relative success of seed beetle invaders compared to other groups. Seed beetles have several ways of egg-laying. Most species deposit their eggs on mature pods of legumes (Fabaceae), the eggs being cemented to the pod or dropped in a self- made 270 Ron Beenen & Alain Roques / BioRisk 4(1): 267–292 (2010) hole in the pod wall. Other species lay eggs on mature seeds that are still attached to the inside of a partly opened pod. A third group of species oviposit on mature seeds that have fallen to the ground from a fully dehisced pod. However, some species such as Acantoscelides obtectus use different life history strategies. Early in the season in this species, oviposition occurs on green pods of Phaseolus, while later in the season, the eggs are deposited on mature seeds that have fallen to the ground. These biological features make A. obtectus fully capable of completing cycle after cycle on naked seeds in storage (Kingsolver 2004). The larvae of seed beetles entirely develop within the seeds until pupation and their presence cannot be recognized before adult emergence, unless the seed is X-rayed. In contrast, leaf beetles show a large variety of reproductive traits. Many Galerucinae (e.g., Diabrotica species) and Alticinae larvae (e.g., Epitrix species) develop in or at the roots of plants and adults feed from leaves of a specific host plant or a wide variety of plant species. Other Chrysomelidae feed both as larva and adult externally on leaves of their host plants. Although practically no plant species is free of leaf beetles, most leaf beetles need fresh plant products in all or at least in the adult stage. Stored dry plant products are not suitable for leaf beetles to complete their life cycle. 8.3.3 Temporal trends Chrysomelids probably began to be introduced thousands of years ago. It is likely that leaf beetles associated with crops have taken the same route as herbs associated with cereals which are supposed to have entered Europe from the Near East (Pinhasi et al. 2005). Beenen (2006) argued that the combination of Buglossoides arvensis (L.) Johnston and Longitarsus fuscoaeneus Redtenbacher 1849 might have taken the route from southwest Asia where they spread with agriculture to large parts of the temperate parts of the Northern hemisphere. Thus, a number of species which are at present considered as native may indeed be originally alien. Bruchidae must have infested pulses grown by man since the dawn of agriculture. Southgate (1979) also mentioned infestations of lentils from the Egyptian Ptolemaic period (305 BC – 30 BC). Relatively little is known of these ancient introductions. More recent ones are much better documented as in the case of the potato Colorado beetle (Leptinotarsa decemlineata) (see factsheet 14.10). From a global point of view, new records of alien species in Europe were relatively important during the 2nd half of the 19th century, due to seed beetle species. The most important being Acanthoscelides obtectus, Callosobruchus chinensis and C. maculatus. However, these species may have been introduced well before their first record. Since ca. 1900, the rate of seed and leaf beetle introductions severely decreased until 1975 when it began to increase again with globalization, essentially through the arrival of leaf beetles. The last seven years since 2000 corresponded to an acceleration of introductions, with an average of 0.8 new species of Chrysomelidae per year, again mostly leaf beetles (Figure 8.3.2) Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 271 Figure 8.3.2. Temporal changes in the mean number of new records per year of seed and leaf beetle species alien to Europe from 1800 to 2009. The number right to the bar indicates the total number of seed and leaf beetle species recorded per time period. 8.3.4 Biogeographic patterns Asia supplied the major proportion of the alien seed and leaf beetles that have established in Europe (Figure 8.3.3). However, this pattern is mainly due to seed beetles of which a half are of Asian origin whereas leaf beetles predominantly originated from North America (36.4%). No seed and leaf beetle species of Australasian origin have yet established in Europe. Alien species are not evenly distributed in Europe, and leaf and seed beetles do not show the same pattern of expansion. Half of the alien seed beetles have colonized more than ten countries with four of them present in more than 50 countries and the main islands of Europe. In contrast, 63.6% of the alien leaf beetles have not yet spread out of the country where they have been initially introduced. Only two species, Leptinotarsa decemlineata and Diabrotica virgifera, are presently encountered in 38 and 20 countries respectively (EPPO 2009, Gödöllo University 2004, Grapputo et al. 2005, Purdue University 2008) (see maps in the spreadsheets 8 and 10). Owing to climate change, L. decemlineata may extend its range to Finland (Valosaari et al. 2008). Alien seed and leaf beetles appear to be concentrated in southern Europe with 18 species observed in mainland Italy and more than 10 species in continental France 272 Ron Beenen & Alain Roques / BioRisk 4(1): 267–292 (2010) Figure 8.3.3. Comparative origin of seed and leaf beetle species alien to Europe and mainland Greece. Central Europe usually hosts less than 10 species except Czech Republic (11 species), whereas aliens have been little recorded in Northern Europe (Figure 8.3.4). 8.3.5 Main pathways and vectors to Europe All alien species of seed and leaf beetle except one (i.e., 95.7%) have been introduced accidentally to Europe. Unlike North America and South Africa, where a number of alien species were released for biological control of weeds (Beenen 2006), only the ragweed leaf beetle, Zygogramma suturalis, has been intentionally introduced from North America for the biological control of common ragweed, Ambrosia artemisifolia L., since 1978 in Russia (Reznik et al. 2004) and several countries of southeastern Europe, and subsequently established in the wild especially in the Caucasus (Kovalev 2004). A flea beetle native of Continental Europe, Altica carduorum (Guérin- Méneville), has also been introduced in Britain and Wales in 1969–1970 to control creeping thistles, Cirsium arvense (L.) Scop. but none apparently established (Baker et al. 1972, Cox 2007). Although it is difficult to ascertain the exact pathway of introduction for most of the Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 273 Figure 8.3.4. Colonization of continental European countries and main European islands by seed and leaf beetle species alien to Europe. other species introduced accidentally, the general behaviour of chrysomelids suggests that most introductions are related to trade of plants and stored products, although some may have arrived as stowaways in all forms of packaging and transport, or even as wind-borne organisms. The world trade of beans for agricultural purposes is probably responsible for the nowadays wide distribution in Europe of most alien species of seed beetles, such as Acanthoscelides obtectus, Bruchus species Callosobruchus species and Zabrotes subfasciatus (Figure 8.3.8) which develop in legume seeds of the subfamily Papilionoideae (Phaseolus, Lathyrus, Pisum, Vicia) (Böhme 2001, Kingsolver 2004). However, the arrival of other seed beetles of the genera Bruchidius, Caryedon, Megabruchidius and Mimosestes seems to be more related to the trade in legume tree seeds of Mimosoideae (Albizzia, Acacia) and Caesalpinoideae (Cassia, Cercis, Tamarindus) used as ornamentals in parks and gardens. Megabruchidius tonkineus was at first suspected to have been introduced from Vietnam to Germany with white beans (Wendt 1980) but it was later found to 274 Ron Beenen & Alain Roques / BioRisk 4(1): 267–292 (2010) be associated with pods of honey locust trees, Gleditsia triacanthos L. (Papilionoideae), and not capable of complete development in beans (Guillemaud et al. 2010). Similarly, Acanthoscelides pallidipennis was probably introduced with seeds of false indigo bush (Amorpha fructicosa L., Papilionoideae) (Tuda et al. 2006) and Bruchidius siliquastri with these of redbuds (Cercis; Caesalpinoideae) from China (Kergoat et al. 2007). Seeds imported for ornamental purposes may also serve as the vector of seed beetles. Specularius impressithorax (Pic) sustained several generations indoors in the Netherlands after having been introduced from South Africa along with seeds of Erythrina (Papilionoideae) used for decoration, but did not eventually establish (Heetman and Beenen 2008) (Figure 8.3.7). Most alien leaf beetles are associated with vegetable crops (Solanaceae, Brassicaceae, Gramineae including maize). With both larvae and adults feeding on foliage, these species probably entered Europe as plant contaminants (eggs, larvae) or crop contaminants (adults). The Colorado potato beetle has frequently been intercepted with potato plants and tubers, but also in all forms of packaging and transport. For example, it usually arrived to Great Britain with commercial freight among vegetable crops such as lettuce, Lactuca sativa L., or on ships, aircraft or private cars traveling from the continent (Cox 2007). Indeed, fresh vegetables grown on land harbouring overwintering beetles are common means of beetle transport in international trade (Bartlett 1980). The African tortoise beetle Aspidimorpha fabricii (= A. cincta Fabricius) was believed to be imported in Italy as a contaminant of bananas in the late 1950s but it became a problem in cultures of Beta vulgaris L. (Zangheri 1960). A hispine palm leaf beetle, Pistosia dactyliferae was also probably introduced as a contaminant of palms imported for ornamental purposes (Drescher and Martinez 2005). The means of introduction appears different when larvae are root-feeding as in Diabrotica and Epitrix species. Unless soil infested with larvae has been imported with host plants, which is usually prohibited, these species probably travel as stowaways. The western corn rootworm, Diabrotica virgifera virgifera, proved to have been translocated from North America to Europe at least three times in aircraft laden with goods and materials, but probably not with maize plants (Ciosi et al. 2008, Miller et al. 2005). The outbreaks in Northwestern Italy and Central Europe probably resulted from introductions of individuals originating in northern USA (Delaware) (Guillemaud et al. 2010). However, another pest species related to tobacco, Epitrix hirtipennis, is assumed to have arrived in Europe as aerial plankton with easterly trade winds blowing from the New World to Europe (Döberl 1994b). Similarly, Jolivet (2001) reported the translocation of the Sweet potato flea beetle, Chaetocnema confinis Crotch, from the USA to several tropical destinations by hurricanes. Adults of Colorado potato beetle are also assumed to be capable of migrating across the Channel although this beetle does not fly strongly (Cox 2007) or from Russia (the St Petersburg region) to Finland (Grapputo et al. 2005). The collection and trade of orchids for greenhouses has also resulted in the arrival of several species which caused severe damage without persisting such as a flea beetle, Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 275 Acrocrypta purpurea Baly, a species from Southeast Asia which was accidentally introduced with plant collections into a greenhouse of Leiden University in the Netherlands (Döberl 1994a). Likewise, larvae of a criocerine species, the yellow orchid beetle Lema pectoralis Baly, were imported to the Netherlands with an orchid collected in 1988 in Thailand (Beenen, unpubl.). Originating of the Peninsula Malaysia and Singapore (Mohamedsaid 2004), L. pectoralis is a major pest (‘orchid lema’) of orchid cultures, particularly Vanda and Dendrobium, in the Philippines (de la Cruz 2003). Pathways within Europe are a source of particular concern because of the waiver of formerly routine phytosanitary inspections on goods transported within the European Union. Thus, alien species once introduced into one European country along with alien plants or seeds, can freely move to other European countries. Spread may combine long-distance, human-mediated dispersal and natural dispersal by adult flight, as it is the case for Leptinotarsa decemlineata (Grapputo et al. 2005). Another significant example is the present northwards expansion of a species alien in Europe, Chrysolina americana. This leaf beetle originates from the Mediterranean Basin where it is associated to Rosmarinus and Lavendula. Because both plants are popular garden plants throughout Europe, C. americana has been translocated outside its native range along with its host plants, e.g. to the Netherlands along with potted Lavendula plants imported from Italy (Beenen, unpubl.). Once introduced, this species, which has good flight capacities, disperses naturally by flight. 8.3.6 Most invaded ecosystems and habitats All alien Chrysomelidae are phytophagous. As expected from the numerical importance of Bruchidae within aliens, seeds constitute the most important larval feeding niche (56.0%), far more important than leaves (24.0%) and roots (20.0%). Almost all these species are only present in man-made habitats which represent 94.1% of the colonized habitats, essentially agricultural lands, parks and gardens, glasshouses, and warehouses for seed beetles (Figure 8.3.5). Natural and semi-natural habitats have been very little colonized yet. In addition to these strong habitat trends, about 40% of the alien chrysomelid species remain strictly related to their original, alien plants. This is especially true for leaf beetles, where only Epitrix hirtipennis out of the 11 alien species has been observed to shift onto native Solanaceae in Italy (Beenen 2006). In contrast, most alien seed beetles found outdoors have already switched to seeds of native plants, for example Bruchidius siliquastri on the native redbud, Cercis siliquastrum, in France (Kergoat et al. 2007), and Acanthoscelides obtectus and Callosobruchus chinensis on wild legumes (Tuda et al. 2001). Under outdoor conditions, a strict dependency to the original alien host was only observed for two Megabruchidius species, M. tonkineus and M. dorsalis, associated with seeds of honey locust tree, Gleditsia triacanthos, in parks and gardens. However, a number of seed beetle species still confined to greenhouses and warehouses only develop on alien hosts of tropical origin, such as Caryedon serratus associated with 276 Ron Beenen & Alain Roques / BioRisk 4(1): 267–292 (2010) Figure 8.3 5. Main European habitats colonized by the established alien species of Chrysomelidae and Bruchidae. The number over each bar indicates the absolute number of alien species recorded per habitat. Note that a species may have colonized several habitats. groundnuts (Arachis hypogaea L.), tamarind (Tamarindus indica L.) and other seeds of alien Caesalpinioideae (Kingsolver 2004). Such species still cannot establish outdoors because none of their alien hosts can survive in the wild at the present time. 8.3.7 Ecological and economic impact Threats due to alien chrysomelid species were first pointed out by Linnaeus in a lecture in 1752, referring to his observation of asparagus plants (Asparagus officinalis L.) that were heavily infested in the vicinity of Hamburg by Crioceris asparagi, a species introduced from Russia at this time (Aurivillius 1909). Alien chrysomelid species are better known for their economic impact than for their ecological impact. Indeed, possible ecological impacts on native flora and fauna are very little documented. Positive impact can be appreciated for only one alien species, Zygogramma suturalis, a strict monophagous species deliberately introduced to Europe for the control of the invasive ragweed (cf above). Negative economic impacts have been recorded in seven of the alien seed beetle species which may severely affect stored pulses of economically-important legumes (Acanthoscelides obtectus, A. pallidipennis, Bruchus pisorum, B. rufimanus, Callosobruchus chinensis, C. maculatus, C. phaseoli, and Zabrotes subfasciatus; see (Borowiec 1987, Hoffmann et al. 1962)). Most of them are capable of re-infesting stored legumes until the Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 277 food reserves are exhausted. In leaf beetles, large economic impacts have been shown for the Colorado potato beetle, L. decemlineata, affecting potato crops (see factsheet 14.10) and the western corn rootworm, D. virgifera virgifera affecting maize roots and foliage (see factsheet 14.8). However, It must be stressed that economic damage has only been seen on maize in Serbia, and in some bordering areas in Croatia, Hungary, Romania, and small areas in Bosnia-Herzegovina and Bulgaria (EPPO 2009). In the United Kingdom, yield losses to be expected from the arrival and spread of D. virgifera virgifera have been estimated to range from 0.9 to 4.1 million € over 20 years in absence of obligatory campaign to prevent spread of western corn rootworm but the costs of such a campaign could also range from 3.7 to 10.5 million € (Central Science Laboratory 2007). Epitrix hirtipennis may also impact tobacco crops (Sannino et al. 1984, Sannino et al. 1985) as well as E. cucumeris these of potato and tomato (Borges and Serrano 1989), and Phaedon brassicae the cabbage crops (Limonta and Colombo 2004). Alien foliage-feeding chrysomelids may also act as vectors for plant diseases, for example D. virgifera which transmits several cowpea virus strains in North America (Lammers 2006). However, little is yet known in this field (Jolivet and Verma 2002). Besides such economic damage, aesthetic impacts are recorded on ornamental plants, such as these of the leaf beetle Pistosia dactylifera on palm trees in southern France (Drescher and Martinez 2005). 8.3.8 Expected trends Introduction of alien chrysomelids is still an ongoing process, especially through the trade of ornamentals via garden centers. For example, an alien species of the genus Luperomorpha was recently imported to Europe. L. xanthodera, originating from China, was first found in Great Britain feeding in flowers of several plant species in garden centers (Johnson and Booth 2004). Later it was observed in Switzerland (F. Köhler, personal communication), Germany (Döberl and Sprick 2009) and the Netherlands (Beenen et al. 2009), and also in garden centers, especially on rose flowers (Figure 8.3.6). Other alien specimens of Luperomorpha observed in Italy (Conti and Raspi 2007) and France (Doguet 2008) were first identified as L. nigripennis, from India and Nepal, but finally identified as L. xanthodera (Döberl and Sprick 2009). Plants cultivated in the Mediterranean area, then transported without severe pest control and sold in Central, Western and Northern Europe also constitute a serious threat for the expansion of species alien in Europe. The risks associated to this pathway were estimated for Norway (Staverløkk and Saethre 2007). Species originating from subtropical and tropical regions have also been translocated such as Aspidimorpha nigropunctata (Klug) from tropical Africa to The Netherlands and Macrima pallida (Laboissière) from the Himalayan region to Cyprus. These introductions usually have not led to establishment (Beenen 2006). However, they do indicate a potential risk, especially in the context of global warming which may facilitate establishments of such species in the near future. The arrival in southern Europe of additional species associated with ornamental palms such as the hispine leaf beetle, Brontispa longis- 278 Ron Beenen & Alain Roques / BioRisk 4(1): 267–292 (2010) Figure 8.3.6. Adult of alien flea beetle, Luperomorpha xanthodera (Credit: Urs Rindlisbacher- Foto: www.insektenwelt.ch) Figure 8.3.7. Adult of alien seed beetle, Specularius impressithorax; a- dorsal view; b- lateral view (credit: C. van Achterberg; photo taken using Olympus stereomicroscope SZX12 with AnalySIS Extended Focal Imaging software). Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 279 Figure 8.3.8. Adult of Mexican bean weevil, Zabrotes subfasciatus. a- dorsal view; b- lateral view (credit: C. van Achterberg; photo taken using Olympus stereomicroscope SZX12 with AnalySIS Extended Focal Imaging software) sima (Gestro), already invasive in other parts of the world (Nakamura et al. 2006), is thus probable, considering the current increase in alien pests related to palms (see Chapter X). Finally, it is difficult to make serious predictions about the results of future translocations because the species may react differently to the new habitats and hosts when compared with the situation in their native environment. Furthermore, translocations may enhance evolutionary changes partly because of founder effects and genetic bottlenecks and partly because of the triggering of evolution by new environmental factors (Whitney and Gabler 2008). 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Simmonds MSJ, Blaney WM, Birch ANE (1989) Legume Seeds: the Defences of Wild and Cultivated Species of Phaseolus Against Attack by Bruchid Beetles. Annals of Botany 63: 177–184. Southgate BJ (1979) Biology of the Bruchidae. Annual Review of Entomology 24: 449–473. Staverløkk A, Saethre MG (2007) Stowaways in imported horticultural plants: alien and invasive species – assesing their bioclimatic potential in Norway. Bioforsk Report 2, N°66. 70 pp. Stephens JF (1839) A Manual of British Coleoptera or Beetles. London: Longman, Orme Brown, Green & Longmans. 443 pp. Strejček J (1990) Brouci celedí Bruchidae, Urodontidae a Anthribidae. Praha : Academia. 88 pp. Strejček J (1993) Faunistic records from the Czech Republic 9. Klapalekiana 29: 169–171 Szentesi, A. (1999) Predispersal seed predation of the introduced false indigo, Amorpha fruticosa L. in Hungary. Acta Zool. Acad. Sci. Hung. 45: 125–141. Tomov R, Trencheva K, Trenchev G, Kenis M (2007) A review of the non-indigenous insects of Bulgaria. Plant Science 44: 199–204. Tuda M, Shima K, Johnson CD, Morimoto K (2001) Establishment of Acanthoscelides pallidipennis (Coleoptera: Bruchidae) feeding in seeds of the introduced legume Amorpha fruticosa, with a new record of its Eupelmus parasitoid in Japan. Applied Entomology and Zoology 36: 269–276. Tuda M, Rönn J, Buranapanichpan S, Wasano N, Arnqvist G (2006) Evolutionary diversification of the bean beetle genus Callosobruchus (Coleoptera: Bruchidae): traits associated with stored-product pest status. Molecular Ecology 15: 3541–3551. Valosaari KR, Aikio S, Kaitala V (2008) Spatial simulation model to predict the Colorado potato beetle invasion under different management strategies. Annales Zoologici Fennici 45: 1–14. Wendt H (1980) Erstmaliges Auftreten des Vorratsschädlings, Bruchidius tonkineus (Pic, 1904) in der DDR. Deutsche Entomologische Zeitschrift 27: 317–318. Wendt, H. (1981) Eine für Sudost-Europa neue Samenkäfer-Art (Coleoptera: Bruchidae). Folia Entomologica Hungarica 42: 223–226. Whitney KD, Gabler CA (2008) Rapid evolution in introduced species, ‘invasive traits’ and recipient communities: challenges for predicting invasive potential. Diverstity and Distributions 14: 569–580. Zangheri S (1960) L’Aspidimorpha cincta F. (Coleoptera Chrysomelidae, Cassidinae) in Italia. Bollettino di Zoologia agraria e di Bachicoltura Ser. II 3: 219–220. Table 8.3.1. List and characteristics of the established Chrysomelidae species alien to Europe. Status: A Alien to Europe C cryptogenic species. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Last update 1 February 2010. Status Regime A Epitrix hirtipennis (Melsheimer, 1847) A Epitrix similaris Gentner, 1944 Luperomorpha xanthodera (Fairmaire, 1888) A Bruchidae – seed beetles Acanthoscelides obtectus Say, 1831 A A Native range Phyto- Central phagous and South America Phyto- Southern phagous USA, Central and South America. Phyto- USA phagous Phyto- China, phagous Korea Phyto- C & S phagous America 1st record in Europe Invaded countries Habitat* 1987, PT- PT-AZO AZO I1 1984, IT I1 BG, GR, IT, MK, PT-AZO 2008, PT PT I1 2003, GB CH, DE, FR, GB, IT, NL I2 1889, IT Nicotiana and other Solanaceae Nicotiana and other Solanaceae References Borges and Serrano (1989) Döberl (1994b), Döberl (2000), Sannino et al. (1984), Sannino et al. (1985) Solanum tuberosum Iris and Euonymus roots (larva); adult polyphagous Doguet (2009), Oliviera et al. (2008) Beenen, unpubl., Conti and Raspi (2007), Del Bene and Conti (2009), Delobel and Delobel (2003), Doguet (2008), Johnson and Booth (2004) Phaseolus seeds, wild and cultivated legumes outdoors Borges et al. (2005), Delobel and Delobel (2003), Hoffmann et al. (1962), Tomov et al. (2007) 285 AL, AD, AT, BA, BE, BG, BY, J1, I CH, CY, CZ, DE, DK, EE, ES, ES-BAL, ES-CAN, FR, FR-COR, GB, GR, GR-CRE, GR-NEG, GR-SEG, HR, HU, IE, IL, IS, IT, IT-SAR, IT-SIC, LI, LT, LU, LV, MD, MK, MT, NL, NO, NO-SVL, PT, PT-AZO, PT-MAD, RO, RS, RU, SE, SI, SK, UA Hosts Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 Family or subfamily Species Alticinae- flea beetles Epitrix cucumeris (Harris, 1851) Status Regime A C Bruchus rufimanus Bohemann, 1833 A A Phytophagous Phytophagous 1st record Invaded countries Habitat* Hosts in Europe 1980, BG AT, BA, BG, CH, CZ, DE, HR, I, J Amorpha HU, IT, LU, MK,PL, RO, RS fruticosa (indigobush) and other legumes Crypto2003, FR FR I2 genic Asia1850, CZ AD, AL, AT, BA, BE, BG, BY, I, J1 Temperate CH, CY, CZ, DE, DK, EE, ES, ES-BAL, ES-CAN, FI, FR, FRCOR, GB, GR, GR-CRE, GRNEG, GR-SEG, HR, HU, IE, IS, IT, IT-SAR, IT-SIC, LI, LT, LU, LV, MD, MK, MO, MT, NL, NO, NO-SVL, PL, PT, PTAZO, PT-MAD, PT, RO, RS, RU, SE, SI, SK, UA Phyto- Africa 1894, PT AD, AL, AT, BA, BE, BG, BY, I, J1 phagous CH, CY, CZ, DE, DK, EE, ES, ES-BAL, ES-CAN, FI, FR, FRCOR, GB, GR, GR-CRE, GRNEG, GR-SEG, HR, HU, IE, IS, IT, IT-SAR, IT-SIC, LI, LT, LU, LV, MD, MK, MT, NL, NO, NO-SVL, PL, PT, PT-AZO, PT-MAD, RO, RS, RU, SE, SI, SK, UA Cercis seeds References Borowiec (1983), Borowiec (1988), Migliaccio and Zampetti (1989), Szentesi (1999), Wendt (1981) Kergoat et al. (2007) Dried peas; Lathyrus, Pisum, Vicia Delobel and Delobel (2003), Fauna Europaea (2009), Gobierno de Canarias (2010), Hoffmann (1945), Sainte-Claire Deville (1938) Stored beans; Phaseolus, Vicia, Lathyrus, Lupinus, Pisum, Lens, Cicer (wild and cultivated) Delobel and Delobel (2003), Fauna Europaea (2009), Gobierno de Canarias (2010), Hoffmann (1945), Sainte-Claire Deville (1938) Ron Beenen & Alain Roques / BioRisk 4(1): 267–292 (2010) Bruchidius siliquastri Delobel 2007 Bruchus pisorum (Linnaeus, 1758) Native range Phyto- North phagous America 286 Family or subfamily Species Acanthoscelides pallidipennis (Motschulsky, 1874) Family or subfamily Species Callosobruchus chinensis (Linnaeus, 1758) Status Regime A Callosobruchus maculatus (Fabricius, 1775) A Callosobruchus phaseoli (Gyllenhal, 1833) A Phyto- Asia1945, FR phagous Temperate Caryedon serratus (Olivier, 1790) A Phyto- Africa phagous 1900, CZ CY, CZ, DE, GR, GR-CRE I1, I2, F, J1 Megabruchidius dorsalis (Fahreus, 1839) Megabruchidius tonkineus György 2007 A Phytophagous Phytophagous 1989, IT I2 A Asia (Japan) Asiatropical (Vietnam) AL, CZ, ES, FR, GR, GR-CRE, IL, IT, IT-SIC IT 2001, HU HU I, J1 I2 Phaseolus, Lupinus and other stored legumes (capable of re-infesting) Acacia, Cassia, Prosopis seeds Gleditsia seeds Gleditsia seeds References Biondi et al. (1994), Essl and Rabitsch (Eds) (2002), Fauna Europaea (2009), Gobierno de Canarias (2010), Hoffmann (1945), Sainte-Claire Deville (1938), Tomov et al. (2007) Binaghi (1947), Delobel and Delobel (2003), Fauna Europaea (2009), Gu et al. (2009), Hoffmann (1945), Tomov et al. (2007) Delobel and Delobel (2003), Hoffmann (1945), Tomov et al. (2007) Delobel and Delobel (2003) Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 Native 1st record Invaded countries Habitat* Hosts range in Europe I, J1 Stored Phyto- Asia1878, FR AD, AL, AT, BA, BE, BG, BY, legumes phagous Temperate CH, CY, CZ, DE, DK, EE, ES, (capable of ES-BAL, ES-CAN, FI, FR, FRre-infesting) COR, GB, GR, GR-CRE, GRNEG, GR-SEG, HR, HU, IE, IL, IS, IT, IT-SAR, IT-SIC, LI, LT, LU, LV, MD, MK, MT, NL, NO, NO-SVL, PT, PT-AZO, PT-MAD, RO, RS, RU, SE, SI, SK, UA Phyto- Africa 1878, FR AL, BG, CZ, ES, FR, GR, GRI, J1 Phaseolus and phagous CRE, IL, IT, IT-SIC, IT, PT, other stored PT-AZO legumes (capable of re-infesting) Migliaccio and Zampetti (1989) György (2007), Jermy et al. (2002) 287 Status Regime A Native 1st record Invaded countries range in Europe Phyto- Asia1945, FR DE, DK, FR, IT phagous Temperate Habitat* J1 Hosts References Hansen (1996), Hoffmann (1945) Pseudopachymerina spinipes (Erichson, 1833) A Phyto- C & S phagous America 1919, ES ES, FR, GR, GR-CRE, IT, ITSIC I2 Zabrotes subfasciatus (Bohemann, 1833) A Phyto- C & S phagous America 1858, FR AL, CZ, ES, ES-CAN, FR, GR, GR-CRE, IT, IT-SIC, NL, PT, PT-AZO J1 Bouchelos and Chalkia (2003), Fauna Europaea (2009), Ramos et al. (2007) Phaseolus and Delobel and Delobel other stored (2003), Hoffmann legumes (1945) (capable of re-infesting) Phyto- Africa phagous 1957, IT IT I1 Beta vulgaris Zangheri (1960) Solanum tuberosum and other Solanaceae CABI/EPPO (2003), EPPO (2006), Fauna Europaea, Grapputo et al. (2005), Tomov et al. (2007) Brassicaceae Limonta and Colombo (2004) Sida rhombifolia Jolivet (2001) Cassidinae – Tortoise leaf beetles Aspidomorpha fabricii A Sekerka, 2008 Chrysomelinae – leaf beetles Leptinotarsa decemlineata A (Say, 1824) Phaedon brassicae Baly, 1874 A Calligrapha polyspila (Germar, 1821) C Phyto- North and 1922, FR phagous Central America Phyto- China, phagous Japan, Taiwan, Vietnam. Phyto- North phagous America 2000, IT > 2001, PT-AZO I1 AD, AL, AT, BA, BE, BG, BY, CH, CZ, DE, EE, ES, ES-BAL, FR, FR-COR, GR, HR, HU?, IT, IT-SAR, IT-SIC, LI, LT, LU, LV, MD, MK, MO, NL, PL, PT, RO, RS, RU, SE, SI, SK, UA IT I1 PT-AZO Ron Beenen & Alain Roques / BioRisk 4(1): 267–292 (2010) Acacia, Phaseolus, Vicia, Ciser (chickpea) seeds Acacia farnesiana seeds 288 Family or subfamily Species Mimosestes mimose (Fabricius, 1781) Hispinae – Hispine leaf beetles Pistosia dactyliferae (Maulik, 1919) A Native range North America 1st record in Europe 1985, HR HR Central America 1992, RS 2004, FR Phyto- India phagous Invaded countries Habitat* Hosts References Ambrosia artemisiifolia Igrc et al. (1995) AT, BA, BE, BG, CH, CZ, DE, I1 FR, GB, HR, HU, IT, MO, NL, PL, RO, RS, SI, SK, UA. Zea mays. Baca (1994), Ciosi et al. (2007), EPPO (2009), Gödöllo University (2009), Guillemaud et al. (2010), Purdue University (2009) FR Palms Drescher and Martinez (2005) I2 Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 Family or subfamily Status Regime Species Zygogramma suturalis A Phyto(Fabricius, 1775) phagous Galerucinae – Skeletonizing leaf beetles Diabrotica virgifera virgifera A PhytoLeConte, 1868 phagous 289 Regime Phytophagous Phytophagous Phytophagous Phytophagous Phytophagous Phytophagous Phytophagous Phytophagous Phytophagous Phytophagous Phytophagous Phytophagous Native range Invaded countries Habitat* Hosts References Western, Southern and Central Europe Continental Europe PT-AZO I Vitis GB I2 Continental Europe PT-AZO I Lathyrus pratensis Cox (2007) (meadow vetchling) Graminae Borges and Serrano (1989) Continental Europe PT-AZO I Solanum Borges and Serrano (1989) Continental Europe PT-AZO I Plantago Borges and Serrano (1989) Mediterranean region PT-AZO I Borago officinalis and Borges and Serrano (1989) other Boraginaceae Continental Europe GB I2 Thymus, Rosmarinus Cox (2007) Alps DK G3, G4 Cirsium Hansen (1964) Continental Europe, PT-AZO Caucasus Continental Europe PT-AZO I Asteraceae and Poaceae Brassicaceae Borges and Serrano (1989) Continental Europe GB I2 Spergula arvensis (Corn spurrey) Cox (1995), Cox (2007) Continental Europe PT-AZO I1 Sarothamnus scoparius seeds Borges et al. (2005) I Borges and Serrano (1989) Borges and Serrano (1989) Ron Beenen & Alain Roques / BioRisk 4(1): 267–292 (2010) Family or subfamily Species Alticinae- flea beetles Altica ampelophaga GuérinMéneville, 1858 Altica carinthiaca Weise, 1888 Chaetocnema hortensis (Geoffroy, 1785) Epitrix pubescens (Koch, 1803) Longitarsus kutscherae (Rye, 1872) Longitarsus lateripunctatus lateripunctatus (Rosenhauer, 1856) Longitarsus obliteratoides Gruev, 1973 Neocrepidodera brevicollis (J. Daniel, 1904) Neocrepidodera ferruginea (Scopoli, 1763) Psylliodes chrysocephalus (Linnaeus, 1758) Psylliodes cucullata (Illiger, 1807) Bruchidae – seed beetles Bruchidius foveolatus (Gyllenhal, 1833) 290 Table 8.3.2. List and characteristics of the Chrysomelidae species alien in Europe. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Last update 1 February 2010. Family or subfamily Species Bruchidius lividimanus (Gyllenhal, 1833) Bruchidius varius (Olivier) Native range Invaded countries Habitat* Phytophagous Phytophagous Mediterranean region PT-AZO I1 Continental Europe E, G Phytophagous Mediterranean region BE, CH, CZ, DE, I, J1 DK, ES-CAN, FI, GB, HU, IE, LI, LU, LV, NL, NO, PT-AZO, PT-MAD, RO, SE, SK, UA Southern Europe ES-CAN I, J1 Bruchus lentis Fröhlich, 1799 Phytophagous Bruchus rufipes Herbst, 1783 Phytophagous Bruchus signaticornis PhytoGyllenhal, 1833 phagous Criocerinae- leaf beetles Crioceris asparagi (Linnaeus, Phyto1758) phagous West Palaearctic GB PT-AZO, ES- CAN I Mediterranean region BE, CH, CZ, DE, I, J1 DK, EE, FI, GB, HU, IE, LI, LT, LU, LV, MD, NL, NO, RU, SE, SK, UA Continental Europe, GB Central Asia I, J Hosts References Genistea, Ononis, Borges et al. (2005) Cytisus seeds Cox (2007), Hodge (1997) Trifolium pratens (red clover), T. medium (zig-zag clover), Ulex europaeus (gorse), Bolboschoenus maritimus (sea clubrush) seeds Lens seeds Fauna Europaea (2009), Gobierno de Canarias (2010), Strejček (1990) Lens, Vicia seeds Gobierno de Canarias (2010), Igrc et al. (1995) Lathyrus, Pisum, Borges et al. (2005), Gobierno de Vicia seeds Canarias (2010) Lathyrus, Lens, Vicia Strejček (1990) seeds Cox (2007), Hill et al. (2005) 291 Asparagus officinalis officinalis (garden asparagus), A. officinalis prostratus (wild asparagus) Leaf and Seed Beetles (Coleoptera, Chrysomelidae). Chapter 8.3 Bruchus ervi Frölich, 1799 Regime Native range Invaded countries Habitat* Hosts References GB, IE I2, I1 Lilium, Fritillaria and other Liliaceae; Arum maculatum Cox (2007), Stephens (1839) Western Mediterranean PT- AZO I2 Pulmonaria Borges and Serrano (1989) Mediterranean region BE, GB, NL I1, I2 Beenen and Winkleman (2001), Cox (2007), Johnson (1963), Lays (1988) Chrysolina bankii (Fabricius, Phyto1775) phagous Mediterranean region GB I2 Gonioctena fornicata Phyto(Bruggemann, 1873) phagous Galerucinae- Skeletonizing leaf beetles Xanthogaleruca luteola Phyto(Müller, 1766) phagous Eastern Europe IT I Rosmarinus, Lavandula, Salvia, Thymus Plantago lanceolata (ribwort plantain), Ballota nigra (black horehound), Mentha spp., and other Lamiaceae Medicago Michieli (1957) Europe GB I2 Ulmus Buckland and Skidmore (1999) Cryptocephalinae – casebearers Cryptocephalus sulphureus G. PhytoA. Olivier, 1808 phagous Chrysomelinae – leaf beetles Chrysolina americana PhytoLinnaeus, 1758 phagous Cox (2007) Ron Beenen & Alain Roques / BioRisk 4(1): 267–292 (2010) Continental Europe 292 Family or subfamily Regime Species Lilioceris lilii (Scopoli, 1763) Phytophagous A peer reviewed open access journal BioRisk 4(1): 293–313 (2010) RESEARCH ARTICLE doi: 10.3897/biorisk.4.49 BioRisk www.pensoftonline.net/biorisk Ladybeetles (Coccinellidae) Chapter 8.4 Helen Roy1, Alain Migeon2 1 NERC Centre for Ecology & Hydrology, Biological Records Centre, Crowmarsh Gifford, Oxfordshire, OX10 8BB, United Kindgom 2 INRA, UMR CBGP (INRA/IRD/Cirad/Montpellier SupAgro), Centre de Biologie et Génétique des populations, CS 30016, 34988 Montferrier- sur-Lez Cedex, France Corresponding authors: Helen Roy (hele@ceh.ac.uk), Alain Migeon (migeon@supagro.inra.fr) Academic editor: Alain Roques | Received 26 January 2010 | Accepted 22 May 2010 | Published 6 July 2010 Citation: Roy H, Migeon A (2010) Ladybeetles (Coccinellidae). Chapter 8.4. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 293–313. doi: 10.3897/biorisk.4.49 Abstract The majority of Coccinellidae are beneficial predators and they have received considerable research attention because of their potential as biological control agents. Indeed the role of coccinellids as predators of pest insects has been a major factor in the movement of coccinellids between countries. The commercial production of coccinellids by biological control companies and local producers led to a rapid increase in distribution thoughout the 1990’s. To date, 13 alien coccinellid species have been documented in Europe; 11 of these are alien to Europe (two are alien to Great Britain and Sweden but native within Europe). The distribution of alien coccinellids in Europe mirrors the biogeographical distribution and patterns of introduction. Some species have dispersed widely; Harmonia axyridis has spread rapidly from countries where it was deliberately introduced to many others across Europe. The ecological and economic impacts of alien coccinellids are not well documented. In this chapter we provide an overview of the temporal and spatial patterns of alien coccinellids in Europe. Keywords Coccinellid, ladybird, alien, Europe, biological control agent, Harmonia axyridis, distribution patterns 8.4.1 Introduction The Coccinellidae are commonly referred to as ladybirds (Britain, Australia, South Africa), ladybugs (North America) or ladybeetles (various countries). Coccinellids have received considerable research attention because of their role as predators of pest Copyright H. Roy, A. Migeon. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 294 Helen Roy & Alain Migeon / BioRisk 4(1): 293–313 (2010) insects. The Coccinellidae comprises over 4200 species worldwide (Iperti 1999, Majerus et al. (2006a)). Audisio and Canepari 2009 report that there are approximately 253 species and subspecies of Coccinellid in Europe. However, a review in 1999 documented only 110 species including species acclimatized through the 1900s: Rodolia cardinalis, Cryptolaemus montrouzieri, Rhyzobius (Lindorus) lophanthae, Rhyzobius forestieri and Serangium parcestosum (Iperti 1999). The discrepancy in species number from these different sources can not solely be accounted for by the addition of new species arriving in Europe but is an indication of the dynamic state of coccinellid taxonomy and the difficulty of establishing a checklist for Europe. Not only is the taxonomy of coccinellids under review but also the arrival of new species is ongoing; recently the UK Ladybird Survey (www.ladybird-survey.org) reported the first British record of Cynegetis impunctata (Thomas et al. 2009). There is also considerable variation in reported coccinellid diversity between countries. Great Britain is relatively species poor with only 46 species (Majerus et al. 2006b) whereas in contrast the Netherlands have 86 native coccinellid species. The proportion of alien species for this group is quite high in Europe, with 13 species observed in the wild to date. Two of these are native to Europe but alien within Great Britain (Henosepilachna argus, Scymnus impexus) and Sweden (Scymnus impexus). For the remainder of this section only the 11 species alien to Europe (and not the three alien species in Europe) will be considered. The majority of coccinellid species (about 90 %) are beneficial predators (others are phytophagous or mycophagous); consequently coccinellids have played a significant role in the development of biological control strategies (Berthiaume et al. 2007, Brown and Miller 1998, Galecka 1991, Gurney and Hussey 1970, Iperti 1999, Obrycki and Kring 1998). This has been a major factor in the movement of coccinellids between countries worldwide. 8.4.2 Taxonomy of the coccinellid species alien to Europe The family Coccinellidae belongs to the coleopteran superfamily Cucujoidea and is a member of the phylogenetic branch of Coleoptera termed the Cerylonid complex of families (Cerylonidae, Discolomidae, Alexiidae, Corylophidae, Endomychidae and Lathridiidae). Worldwide there are six subfamilies of Coccinellidae: Sticholotidinae, Chilocorinae, Scymninae, Coccidulinae, Coccinellinae and Epilachninae although a recent phylogeny suggests a seventh subfamily, Ortaliinae (Fürsch 1990, Kovář 1996). European species are mainly represented by three subfamilies: Scyminae, Chilocorinae and Coccinellinae. There are very few European Sticholotinae, very few Coccidulinae and only three species of Epilachninae (Iperti 1999). Although the species list for Coccinellidae in Fauna Europaea (Audisio and Canepari 2009) includes representatives from all six subfamilies. Species alien to Europe are quite evenly represented between five of the six subfamilies. Three species are observed in the subfamily Coccidulinae (two Coccidulini Ladybeetles (Coccinellidae). Chapter 8.4 295 and one Noviini) and in the Scymninae (two Scymnini and one Hyperaspidini). Two species are in the Chilocorinae (two Chilocorini) and Coccinellinae (two Coccinellini). One species is in the Sticholotidinae (Sticholotidini). There are no Epilachninae that are alien to Europe (although Henosepilachna argus is alien in Europe). Most species in the Epilachninae are phytophagous, while the majority of species in the other subfamilies are predatory. The preferred diets of the two feeding stages in the life-cycle, the larval and adult stages, are generally the same. Most predatory ladybirds feed on either aphids or coccids (a few feed on both), however some predatory species feed on mites, adelgids, aleyrodids, ants, chrysomelid larvae, cicadellids, pentatomids, phylloxera, mycophagous coccinellids and psyllids (Dixon 2000). Indeed, a small number of species within the Coccinellinae and Epilachninae are mycophagous, feeding on the hyphae and spores of fungi. There is also considerable variability in the degree of dietary specialisation between species (Hodek 1996). Some species have a very narrow preferred prey range, such as a single species of mite, aphids of a single genus, or plants of a single family, other species have a wide prey range. Harmonia axyridis, for example, will feed on aphids, coccids, adelgids, psyllids, and the eggs and larvae of many other insects, including other coccinellids and lepidopterans (Legaspi et al. 2008, Ware and Majerus 2008). Ladybirds exhibit complex adaptations to specific or more general diets such as mandibular dentition, gut length and structure, and morphological features that affect mobility (Hodek 1996). Many predatory coccinellids will feed on alternative foods, such as pollen, nectar, honey-dew and fungi (many also resort to cannibalism) when preferred prey are scarce (De Clercq et al. 2005, Hodek 1996). Coccinellids are distinguished from the remainder of the Cerylonid complex of families by a number of adult characteristics: five pairs of abdominal spiracles, tentorial bridge is absent, anterior tentorial branches are separated, frontoclypeal suture absent, apical segment of maxillary palpus never aciculate, galea and lacinia separated, mandible with reduced mola, front coxal cavities open posteriorly, middle coxal cavities open outwardly, metaepimeron parallel-sided, femoral lines present on abdominal sternite 2, tarsal formula 4-4-4 or 3-3-3, tarsal segment 2 usually strongly dilated below (Kovář 1996). In Europe, the diagnostic features of the family Coccinellidae can be considered in more simple terms (Majerus 2004). They are small to medium sized beetles (1.3–10 mm in length). There body shape is oval, oblong oval or hemispherical (upper surface convex and lower surface flat). They have large, compound eyes. The antennae are often 11-segmented but this figure varies and can be as low as seven. The mouthparts consist of large, strong mandibles; four-segmented maxillary palps (terminal segment axe shaped) behind the mandibles; labium divided into the pre-labium and post-labium; three-segmented labial palps; and the labrum. The head can be partly withdrawn under the pronotum. The pronotum is broader than long and has anterior extensions at the margin. The legs are short and can be retracted into depressions under the body. The tarsi are usually four segmented but the third segment is small and hidden in the end of the second segment. Each tarsus bears two claws. The abdomen has ten segments (Kovář 1996, Majerus et al. 2006a). 296 Helen Roy & Alain Migeon / BioRisk 4(1): 293–313 (2010) 8.4.3 Temporal trends of introduction in Europe of alien coccinellids The first species of coccinellid to be introduced into Europe was the vedalia beetle, R. cardinalis, for the control of the cottony cushion scale (coccid), Icerya purchasi (Figure 8.4.1). Two further species were introduced during the early twentieth century (mainly to the Mediterranean regions including France, Portugal and Italy) but there then followed a period of stagnation and respect to biological control in general. This correlates with the trend towards chemical insect pest control with the development of synthetic pesticides. From the 1980’s onwards there were a considerable number of introductions on an extensive scale across Europe through the use of tropical coccinellids to control glasshouse pest insects. 8.4.4 Biogeography of the coccinellid species alien to Europe Each continent has a specific fauna of coccinellidae. Belicek (1976) stated that “many species develop their cycles in life zones delineated by the general physiography of the continents (mountainous barriers) and climatic patterns combined with the types of vegetation in a given zone”. Glaciation had profound effects on the distribution of coccinellids and the level of endemism is further controlled by ecological factors including temperature, food and natural enemies. The temperate zones of Europe and North America are heavily infested by Aphidae and grasslands in these regions contain coccinellids from the tribus Coccinellini (Coccinella spp., Adalia spp., Harmonia spp.) and Hippodamiini, Cheilomenini and Scymnini. Open deciduous and coniferous forests in this temperate zone contain other genera of Coccinellini (Anatis spp., Myrrha spp., Myzia spp.). Tropical zones in central and South Africa, South America, India and China where Coccidae are abundant are characterised by coccinellids from the tribus Chilocorini (Chilocorus spp., Exochomus spp., Brumus spp.), Scymnini, Hyperaspini, Coccidulini and Noviini. In the Mediterranean regions of Europe, aphids and coccids are found together and are attacked by coccinellids from the temperate and tropical zones (Iperti 1999). It is interesting to note that coccinellids native to temperate zones enter either simple quiescence or intense diapause as adults. In contrast, exotic species such as Rhyzobius lophanthae and Cryptolaemus montrouzieri do not enter quiescence or diapause but instead resist drastic changes in climate by reducing the speed of development during winter but not entirely stopping it (Iperti 1999). The early introductions of alien coccinellids were characteristically as classical biological control agents; the predatory coccinellid originated from the same country as the target pest insect. So, for example, both R. cardinalis and I. purchasi originated from Australia; R. lophanate and various Diaspididae (Pseudolacaspis pentagona, Quadraspidiotus perniciosus, Chrysomphalus dictyospermi, Parlatoria blanchardi) from Australia and New Zealand; C. montrouzieri and Planococcus citri from Australia. Notably all these species are from tropical regions and were introduced into Mediterranean regions for Ladybeetles (Coccinellidae). Chapter 8.4 297 Figure 8.4.1. Temporal trends in the mean number of new records per year of coccinellid species alien to Europe from 1875 to 2008. The number above the bar indicates the total number of alien species newly recorded during the considered time period. control purposes (Figures 8.4.1 and 8.4.2). In contrast, the coccinellid species selected to reinforce the activity of native natural enemies in temperate regions of Europe are from temperate regions of the globe for example, temperate Asia (H. axyridis) or North America (Hippodamia convergens). 8.4.5 Distribution of alien Coccinellids in Europe The distribution of alien coccinellids in Europe mirrors the biogeographical distribution and patterns of introduction (Figure 8.4.3). Some species have dispersed widely; H. axyridis has spread rapidly from countries where it was deliberately introduced to many others across Europe. Furthermore, the commercial production of coccinellids by biological control companies and local producers led to a rapid increase in distribution thoughout the 1990’s. 8.4.6 Use of alien coccinellids for biological control in Europe The ecosystem service that predatory coccinellids provide in consuming pest insects has been recognised for over a century. The vedalia ladybird, R. cardinalis, is considered to have initiated modern biological pest control. It was released as a classical bio- 298 Helen Roy & Alain Migeon / BioRisk 4(1): 293–313 (2010) Figure 8.4.2. Origin of the 11 alien coccinellid species established in Europe. logical control agent (native to Australia) in 1887 to control an alien cottony cushion scale (coccid), I. purchasi, which was threatening the citrus industry of California. The vedalia ladybird and the cottony cushion scale are still present in Californian citrus groves but the ecological balance between predator and prey ensures that the pest is no longer a problem (Caltagirone 1989, Majerus et al. 2006a). The successful introduction of R. cardinalis for the control of I. purchasi resulted in considerable focus on Coccinellidae for importation programmes worldwide (Obrycki and Kring 1998). Over 40 species of coccinellid were introduced to North America following R. cardinalis during a period colloquially referred to as the “ladybird fantasy” (Caltagirone 1989, Dixon 2000). This worldwide phenomenon was mainly ineffectual; only four of over 40 species introduced to North America during this time established (Caltagirone 1989). In recent times there have been 155 attempts to control aphids and 613 to control coccids worldwide through the introduction of ladybirds (Dixon 2000). On a scale of success (complete, substantial, partial or no control) only one attempt to control aphids using coccinellids has been ranked as substantially successful and none have been completely successful (Dixon 2000). In contrast, 23 complete and 30 substantial successes have been achieved against coccids (Dixon 2000). In a few cases the introduced coccinellid species has had farreaching, unacceptable impacts on biodiversity and so has been deemed an invasive species. Harmonia axyridis, harlequin ladybird, is the only such example in Europe (Brown et al. 2008a). All of the 11 alien coccinellids in Europe have been intentionally released as biological control agents of pest insects. The first coccinellid to be introduced to Europe was R. cardinalis as a predator of I. purchasi in 1888 (Portugal), 1901 (Italy) and 1912 (Italy and France). This species was subsequently released through the mid and late 1900s to Italy, Portugal, Israel, France, Spain, Malta, Great Britain, Albania, Cyprus, Switzerland and the Ukraine. Cryptolaemus montrouzieri, native to Australia, was intentionally released to control mealybugs (Pseudococcidae), Planococcus citri, from 1908 in Italy. Subsequent releases were made in Spain (1926), Corsica (1970), France Ladybeetles (Coccinellidae). Chapter 8.4 299 Figure 8.4.3. Colonisation of European countries and islands by coccinellids alien to Europe where known. Scale = total number of recorded alien coccinellids. (1974), Portugal (1984) and Sweden (2001). This species is considered established in all the countries where it has been released other than Sweden (for which the status of this species is unknown). Cryptolaemus montrouzieri has been used extensively through augmentation (release of reared adults) and was the first coccinellid used to demonstrate an inoculative approach (whereby the aim is introduce a small number of individuals into a crop system with the expectation that they will reproduce and their offspring will continue to provide control of the target pest for an extended period of time). Cryptolaemus montrouzieri is easy and cheap to culture on mealybugs (Majerus 2004). Rhyzobius lophanthae is a species native to New Zealand which was introduced to Italy in 1908 for the control of Diaspididae (armoured scale insects). It has been released widely in European countries including: Portugal (1930 and 1984), Spain (1958), Sardinia (1973), France (1975), Greece (1977) and Germany (2000). This species has recently been reported as established in London, Great Britain (Natural History Museum, 2008). 300 Helen Roy & Alain Migeon / BioRisk 4(1): 293–313 (2010) Figure 8.4.4. Harlequin ladybeetle (Harmonia axyridis). Credit: Mark Bond 8.4.6.1 Control of Scale Insects A number of coccinellid species have been used in historically significant and successful projects for the biological control of scale (Borges et al. 2006, Erler 2001, Katsoyannos 1997) including R. cardinalis and R. lophanthae. Other species introduced to Europe for control of scales include Rhyzobius forestieri, Nephus reunioni, Chilocorus nigritus and Chilocorus kuwanae. Rhyzobius forestieri (native to Australia) has established in Italy, France, Greece and Albania. In the Cambos coastal plain of Greece this species is now considered the most abundant species of coccinellid within the coccidophagous guild (Katsoyannos 1997). Nephus reunioni (native to Africa) was intentionally released in a number of countries (Italy, Portugal, France, Greece, Albania and Spain) and is now considered to be established in Italy and Portugal. Chilocorus nigritus is native to the Indian sub-continent and South East Asia and is a candidate biological control agent for the control of species within the Coccoidea including three economically important families (Diaspididae, Pseudococcidae and Coccidae). It has a recent history, 1985 onwards, of introduction to a number of countries: Italy, Denmark, France, Germany, Netherlands, Great Britain and Albania. Chilocorus kuwanae is a biological control agent of scale insects and was introduced to Europe (Albania and Italy) from Asia in 1989. 8.4.6.2 Control of Aleyrodidae The family Aleyrodidae comprises the commonly referred to whiteflies. Over fifty species of coccinellidae attack eggs and immature stages of whitefly pests (Obrycki and Kring 1998, Yigit et al. 2003). There is interesting variation in the preda- Ladybeetles (Coccinellidae). Chapter 8.4 301 Figure 8.4.5. Adults of Cynegetis impunctata. Credit: Gilles San Martin tory behaviour of these polyphagous coccinellids; some are mobile, seeking out prey, and others are sedentary, and complete preimaginal development on one leaf (Obrycki and Kring 1998). In Europe one species, Serangium parcesetosum, has been introduced for the control of whitefly (Bemisia tabaci). Serangium parcesetosum was introduced from its native range of Asia and the Indian subcontinent to France including Corsica (Majka and McCorquodale 2006). A further species Delphastus catalinae, native to North America, has been introduced in glasshouses within Albania and Russia for the control of Bemisia tabaci and Trialeurodes vaporariorum (Kutuk and Yigit 2007, Legaspi et al. 2008). However, this species has not established in the wild. Studies on the thermal biology of D. catalinae, assessing the effects of temperature on development, voltinism and survival in the laboratory and field (non-indigenous range), indicate a strong correlation between survival in the laboratory at 5 ºC and in the field in winter (Simmons and Legaspi 2004, Simmons and Legaspi 2007). Delphastus catalinae died out quickly in winter temperatures and this suggests that the probability of establishment is low in regions that experience low temperatures and scarcity of suitable food for part of the year (van Lenterenet et al. 2003). In the absence of studies on cold tolerance it is insufficient to assume that, on the basis of climate matching, winter would be an effective barrier to establishment of species originating from warmer climatic zones (van Lenteren et al. 2006). Risk assessments should also be sufficiently detailed to encompass strain specific parameters; the release of a non-diapausing strain versus a diapausing strain could result in very different impacts (van Lenteren et al. 2006). Furthermore, impacts through consumption of non-target hosts and dispersal require considerable attention (van Lenterenet et al. 2003). So, for example, although D. catalinae is not anticipated to survive winter temperatures in northern Europe, it is oligophagous 302 Helen Roy & Alain Migeon / BioRisk 4(1): 293–313 (2010) Figure 8.4.6. Adult of the phytophagous bryony ladybeetle, Henosepilachna argus. Credit: Mike Majerus. and reported as an intra-guild predator of the aphelinid parasitoid Encarsia sophia (Zang and Liu 2007). 8.4.6.3 Control of Aphids Hippodamia convergens and H. axyridis have both been released extensively throughout Europe for the control of aphids. Hippodamia convergens is native to America and several billion are collected annually from overwintering sites in California and sold throughout America. This practice has been shown to be highly ineffective because of adult dispersal (Dixon 2000, Roy and Majerus, unpubl.). Furthermore, removal of H. convergens is considered to have adverse effects on local populations and, in America, is responsible for the distribution of two ladybird parasites (the braconid wasp, Dinocampus coccinellae and the microsporidian, Nosema hippodamiae) (Saito and Bjornson 2006) and vectoring of plant pathogens (dogwood anthracnose fungus) (Bjornson 2008). This coccinellid has been released in Belgium, Sweden, Denmark, Albania and the Czech Republic in the 1990s and early 2000. It is unknown whether or not it is established. The use of H. axyridis as an augmentative biological control agent (mass reared and released) has been widespread (Berkvens et al. 2008, Brown et al. 2008a). In 1982 it was introduced into France and has since been reared continuously over 100 generations on industrially produced eggs of the moth, Ephestia kuehniella (Brown et al. 2008a). It has since been introduced to a number of countries across Europe and also spread to others which had not intentionally released it (Table 8.4.3). Ladybeetles (Coccinellidae). Chapter 8.4 303 Figure 8.4.7. Larva of Henosepilachna argus. Credit: Gilles San Martin 8.4.7 Ecosystems and habitats invaded in Europe by alien Coccinellids Coccinellid species can be classified as stenotopic or eurytopic (Hodek 1993, Iperti 1991). Microclimate is considered to be a particularly important feature of a coccinellid habitat. Many species of ladybird exhibit a preference for specific vegetation types or certain strata of the habitat. Coupled with this is the requirement for suitable food in sufficient abundance. Habitat preference varies seasonally as the microclimatic characteristics of a habitat change, which in turn influences the distribution of prey populations and the behaviour of coccinellids. Iperti (1999) documents the succession of aphid outbreaks in south eastern France; during a normal year aphids first appear on low plants and shrubs, they then progress to cultivated low plants and early deciduous trees and develop on cultivated trees and shrubs. However, climatic conditions vary annually and so it is difficult to predict the behaviour of coccinellids, particularly in a period of climate change. There is a strong trend for alien coccinellids to occur in urban or cultivated habitats in Europe. Almost all species are most prevalent in recently cultivated agricultural, horticultural and domestic habitats, gardens and parks and greenhouses (EUNIS categories I I1, I2, J100; see appendix II). Harmonia axyridis, the most invasive of the alien coccinellids in Europe, follows this pattern although there have been a considerable number of records in Great Britain from natural habitats (Brown et al. 2008b). Indeed, H. axyridis is documented from both woodlands and forest habitats, small anthropogenic woodlands, parks and gardens, agricultural and horticultural habitats as well as from buildings in cities, towns and villages. The abundance of native and alien coccinellid species in urban habitats and their tendency to aggregate in large numbers during autumn and winter enhances their 304 Helen Roy & Alain Migeon / BioRisk 4(1): 293–313 (2010) visibility to people. This aggregation behaviour can be exploited by biological control practitioners through the collection and release of large numbers of beetles but species that exhibit this behaviour, such as H. axyridis, are increasingly seen as nuisance insects in domestic dwellings (Roy and Majerus 2006, Roy et al. 2008). 8.4.8 Ecological and economic impacts of alien coccinellids The ecological and economic impacts of alien coccinellids are not well documented. Many authors have noted the low success rate of coccinellids as biological control agents of aphids (Dixon 2000, Iperti 1999, Majerus et al. 2006a). The success of coccinellids as biological control agents of coccids is higher than that of aphids but still relatively low at only 40 % of cases studied being designated as exerting complete or substantial control (Iperti 1999). Rodolia cardinalis has been heralded as a success story for biological control (Caltagirone 1989). This species has been introduced into 33 countries to control I. purchasi and has yielded complete control in 26 countries (North America, Argentina, Peru, Chile, Portugal, Uruguay, Venezuela, France, Italy, Spain, Greece, Morocco, Tunisia, Turkey, Egypt, India, Japan and New Zealand); substantial control in four countries (Russia, Libya, the Bahamas, Ecuador) and partial control in two countries (Seychelles and Mauritius). A similar rate of success was achieved through the acclimatization of C. montrouzieri to control Pseudococcus spp. (Iperti 1999). Therefore, R. cardinalis and C. montrouzieri have contributed economic benefits through the ecosystem service they provide. Indeed, the initial cost of the R. cardinalis introduction programme in California 1888 was $1 500 with a return in just over a year of millions of dollars (Majerus 2004). The lack of success of aphidophagous coccinellids has been attributed to asynchrony between the reproductive and development rates of the predatory coccinellids and their aphid prey (Dixon 2000). Furthermore, many aphidophagous coccinellids, in temperate climates, are univoltine whereas aphids are multivoltine. Coccidophagous coccinellids tend to stay in a localised area throughout their life cycle and, in contrast, aphidophagous coccinellids disperse widely (Iperti 1999). Most intentional insect introductions do not cause ecological or economic problems, indeed of all the intentionally introduced insects to North America only 1.4 % have caused problems (van Lenteren et al. 2003). Indeed insect introductions are considered to be relatively safe: less than 1 % cause a population level effect in nontargets and only 3–5 % may have caused smaller scale effects (van Lenterenet et al. 2003). However, a number of coccinellids are documented as having non-target effects (van Lenterenet et al. 2003). Cryptolaemus montrouzieri is reported to lower the effectiveness of an introduced natural enemy (Dactylopius opuntiae) for weed control (Goeden and Louda 1976). The most infamous coccinellid introduction is undoubtedly H. axyridis (Majerus et al. 2006b, Roy and Majerus 2006, Roy et al. 2005, Roy and Wajnberg 2008). Ladybeetles (Coccinellidae). Chapter 8.4 305 Harmonia axyridis has been released as a classical biological control agent in North America since 1916. It has been commercially available in Europe since the 1980s and has many attributes that contribute to its economic viability, including its polyphagous nature. Harmonia axyridis preys on a wide variety of tree-dwelling homopteran insects, such as aphids, psyllids, coccids, adelgids and other insects (Koch et al. 2006). In North America, as well as offering effective control of target pests, such as aphids in pecans (Tedders and Schaefer 1994), H. axyridis is also providing control of pests in other systems such as Aphis spiraecola in apple orchards (Brown and Miller 1998) and several citrus pests (Michaud 2002). In both Asia and North America, H. axyridis has been reported to contribute to control of aphids on sweet corn, alfalfa, cotton, tobacco, winter wheat and soybean (Longo et al. 1994). The spread and increase of H. axyridis throughout Europe could, therefore, prove to be beneficial to ecosystem services through the reduction in aphid numbers below economically damaging levels and subsequent reduction in the use of chemical pesticides. The polyphagous nature of H. axyridis means that negative impacts on non-target prey species would appear to be inevitable (Majerus 2006, Pell et al. 2008). However, there is very limited empirical evidence on this subject and studies considering the effects of H. axyridis on the population demography of non-target aphids, coccids and other prey species away from crop systems have not been conducted. Harmonia axyridis has been implicated as a potential predator of immature monarch butterflies, Danaus plexippus, an aposematic species that contains defensive chemicals (Koch et al. 2003). Laboratory studies have also indicated the potential for H. axyridis to engage in intra-guild predation (Pell et al. 2008, Roy et al. 2008, Ware and Majerus 2008). It is likely that many other species will be directly or indirectly affected by the arrival of H. axyridis. Indeed, intraguild predation is thought to be an important force in structuring aphidophagous ladybird guilds (Yasuda et al. 2004) and so H. axyridis has the potential to dramatically disrupt native guilds in Europe. Harmonia axyridis is a large, aggressive, polyphagous coccinellid (with a tendency for intraguild predation) that could impact on the abundance of native coccinellids and reduce their available niches (Legaspi et al. 2008). The wide dietary range of H. axyridis coupled with its ability to disperse rapidly, forage widely and continuously breed gives this species the potential to significantly reduce European populations of coccids and aphids. This is, of course, considered beneficial in crop and horticultural systems, but not in other habitats where such direct competition for prey may result in a reduction in biodiversity and declines in native beneficial predators and parasitoids of aphids and coccids (Majerus 2006). Majerus et al. (2008) noted that the negative effects of H. axyridis on other aphidophages are likely to be the result of a complex range of interactions, with H. axyridis in general having a competitive edge through resource competition, intraguild predation and a more plastic phenotype. A more rapid development rate, continual breeding ability and lack of diapause requirement, efficient chemical defence and relatively large size would provide H. axyridis with a significant reproductive advantage over many native British species. The pattern is anticipated to be widespread throughout Europe (Brown et al. 2008a). 306 Helen Roy & Alain Migeon / BioRisk 4(1): 293–313 (2010) 8.4.10 Conclusions Coccinellids have been introduced widely throughout Europe for the biological control of pest insects. Some of these species have established and for others the status is unknown. It is difficult to estimate the proportion of alien coccinellids in Europe for two reasons: there is not a definitive European check list for coccinellids and the status of some of the alien species is unknown. However, the proportion of alien coccinellids appears to be higher (approximately 5–10 %) than the proportion of aliens for other taxonomic groups (3.1 % alien Diptera). Only one species (H. axyridis) is considered to be invasive. Acknowledgements HER is based in the Biological Records Centre (within the NERC Centre for Ecology & Hydrology) and receives co-funding from the Joint Nature Conservation Committee and the Natural Environment Research Council. AM is funded by the Institut National de la Recherche Agronomique (INRA). The authors gratefully acknowledge Stephanie Ames for production of the European distribution map. Michael E.N. Majerus and Peter M.J. Brown are thanked for insightful discussions on the European species check list. 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Biocontrol 53: 1–4. 310 Helen Roy & Alain Migeon / BioRisk 4(1): 293–313 (2010) Saito T, Bjornson S (2006) Horizontal transmission of a microsporidium from the convergent lady beetle, Hippodamia convergens Guerin-Meneville (Coleoptera: Coccinellidae), to three coccinellid species of Nova Scotia. Biological Control 39: 427–433. Simmons AM, Legaspi JC (2004) Survival and predation of Delphastus catalinae (Coleoptera: Coceinellidae), a predator of whiteflies (Homoptera: Aleyrodidae), after exposure to a range of constant temperatures. Environmental Entomology 33: 839–843. Simmons AM, Legaspi JC (2007). Ability of Delphastus catalinae (Coleoptera: Coccinellidae), a predator of whiteflies (Homoptera: Aleyrodidae), to survive mild winters. Journal of Entomological Science 42: 163–173. Smith SF, Krischik VA (2000) Effects of biorational pesticides on four coccinellid species (Coleoptera: Coccinellidae) having potential as biological control agents in interior landscapes. 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Ware RL, Majerus MEN (2008) Intraguild predation of immature stages of British and Japanese coccinellids by the invasive ladybird Harmonia axyridis. Biocontrol 53: 169–188. Yasuda H, Evans EW, Kajita Y, Urakawa K, Takizawa T (2004) Asymmetric larval interactions between introduced and indigenous ladybirds in North America. Oecologia 141: 722–731. Yigit A, Canhilal R (2005) Establishment and dispersal of Serangium parcesetosum Sicard (Coleoptera, Coccinellidae), a predatory beetle of citrus whitefly, Dialeurodes citri Ashm. (Homoptera, Aleyrodidae) in the East Mediterranean region of Turkey. Zeitschrift Fur Pflanzenkrankheiten Und Pflanzenschutz-Journal of Plant Diseases and Protection 112: 268–275. Yigit A, Canhilal R, Ekmekci U (2003). Seasonal population fluctuations of Serangium parcesetosum (Coleoptera: Coccinellidae), a predator of citrus whitefly, Dialeurodes citri (Homoptera: Aleyrodidae) in Turkey’s eastern Mediterranean citrus groves. Environmental Entomology 32: 1105–1114. Zang LS, Liu TX (2007) Intraguild interactions between an oligophagous predator, Delphastus catalinae (Coleoptera: Coccinellidae), and a parasitoid, Encarsia sophia (Hymenoptera: Aphelinidae), of Bemisia tabaci (Homoptera: Aleyrodidae). Biological Control 41: 142–150. Table 8.4.1. List and main characteristics of the Coccinellidae species alien to Europe. Status: A Alien to Europe C cryptogenic species. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Phylogeny after (2 0, 35). Last update 01/03/2010. Rodolia cardinalis (Mulsant, 1850) Scymninae Hyperaspis pantherina Fürsch, 1975 Cryptolaemus montrouzieri Mulsant, 1853 Nephus reunioni Fürsch, 1974 Chilocorinae Chilocorus kuwanae Silvestri, 1909 Chilocorus nigritus (Fabricius, 1798) Status Regime A A Native range 1st record in Europe Parasitic/ Australasia 1982, IT Predator Parasitic/ Australasia 1908, IT Predator A Parasitic/ Australasia 1888, PT Predator A Parasitic/ Africa Predator A Parasitic/ Australasia 1908, IT Predator A Parasitic/ Africa Predator A A Invaded countries Habitat AL, FR, GR, IT I AL, DE, ES, ES-BAL, FR, FRCOR, GB, GR,GR-CRE, IT, IT-SAR, IT-SIC, IL, MT, PT, PT-AZO, PT-MAD, I, J100 AL, CH, CY, DE, ES, ES-BAL, I, J100 ES-CAN, FR, FR-COR, GB, GR, GR-CRE, IL, IT, IT-SAR, IT-SIC, MT, PT, PT-AZO, PT-MAD, UA 2002, PT- PT-MAD MAD U Hosts References Coccids (Scale Katsoyannos (1997) insects) Coccids Erler (2001) (Scale insects specifically Diaspididae) Coccids (Scale Caltagirone (1989), Frank and insects) McCoy (2007) Booth et al. (1995), Fowler Orthezia insignis (Scale (2004) insect) Mealybugs Hamid and Michelakis (1994), Smith and Krischik (2000) I Coccids (Scale Izhevsky and Orlinsky (1988) insects) Parasitic/ Asia Predator 1989, IT AL, IT I Parasitic/ Asia Predator 1994, IT AL, ,IT I, J100 Coccids (Scale Ponsonby and Copland insects) (2007b), Ricci et al. (2006) Coccids (Scale Booth (1998), Ponsonby and insects) Copland (2007a), Ponsonby and Copland (2007b) 311 I, J100 1983, FR AL, ES, ES-CAN, FR, FR-COR, GR,GR-CRE, IL, IT, IT-SAR, IT-SIC, PT, RU, SE, AL, ES, FR, GR, IT-SAR, PT Ladybeetles (Coccinellidae). Chapter 8.4 Subfamily Species Coccidulinae Rhyzobius forestieri (Mulsant, 1853) Rhyzobius lophanthae (Blaisdell, 1892) Native range 1st record in Europe Invaded countries Habitat Hosts References A Parasitic/ Asia Predator 1986, FR- FR, FR-COR COR I Aleyrodidae Yigit and Canhilal (2005), Yigit et al. (2003) A Parasitic/ Asia Predator 1991, BE I Polyphagous insect predator particularly aphids and coccids A Parasitic/ North Predator America 1992, CZ AL, BE, CZ, DK, SE FA, J100 Aphids Adriaens et al. (2003), Adriaens et al. (2008), Brown et al. (2008a), Brown et al. (2008b), Koch et al. (2003), Majerus (1994), Roy et al. (2005), Roy and Wajnberg (2008) Bjornson (2008), Phoofolo et al. (2008), Saito and Bjornson (2006) AL, AT, BE, BG, BY, CH, CZ, DE, DK, ES, ES-CAN, FR, FR-COR, GB, GR, GR-CRE, GR-ION, GR-SEG, HU, IL, IT, IT-SIC, , LI, LU, NL, NO, PT, RO, RU, SE, SK, UA Helen Roy & Alain Migeon / BioRisk 4(1): 293–313 (2010) Hippodamia convergens GuerinMeneville, 1842 Status Regime 312 Subfamily Species Sticholotidinae Serangium parcesetosum Sicard, 1929 Coccinellinae Harmonia axyridis (Pallas, 1773) Ladybeetles (Coccinellidae). Chapter 8.4 313 Table 8.4.2. List and main characteristics of the Coccinellidae species alien within Europe. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Phylogeny after Fürsch (1990), Koch et al. (2006). Last update 01/03/2010. SubFamily Regime Native Species range Scymninae Scymnus Parasitic/ West impexus Predator Palearctic Mulsant, 1850* Epilachninae Henosepilachna Phytoargus (Geoffroy, phagous 1762)* West Palearctic Invaded Habitat* countries Hosts References GB, SE G, I2 Dreyfusia Humble (1994), piceae on Majka and spruce and fir McCorquodale (2006) GB E5, I2, FA White bryony Hill et al. (2005) (Bryonia dioica) Table 8.4.3. Summary of release dates and records from wild populations of Harmonia axyridis across Europe. Adapted from Brown et al. (2008a). Updated: 01/03/2010 Country Ukraine Belarus Portugal France Greece Germany Belgium Netherlands Spain Switzerland Luxembourg England and Channel Isl. Italy Czech Republic Austria Denmark Wales Norway Poland Liechtenstein Sweden Northern Ireland Scotland Serbia Slovakia Hungary Bulgaria Romania Year of release (blank if not released) 1964 1968 1984 1982 1994 1997 1997 1996 1995 1996 1990s 2003 2000s Year of first record in the wild Unknown Unknown 1991 1998 1999 2001 2002 2003 2004 2004 2004 2006 2006 2006 2006 2006 2006 2007 2007 2007 2007 2007 2008 2008 2008 2009 2009 A peer reviewed open access journal BioRisk 4(1): 315–406 (2010) RESEARCH ARTICLE doi: 10.3897/biorisk.4.61 BioRisk www.pensoftonline.net/biorisk Coleoptera families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae Chapter 8.5 Olivier Denux1, Pierre Zagatti2 1 INRA, UR633 Zoologie Forestière, 2163 Av. Pomme de pin, 45075 Orléans, France, 45075 Orléans Cedex 2 INRA – Centre de recherche de Versailles, Unité PISC, Route de Saint-Cyr, 78026 Versailles Cedex, France Corresponding authors: Olivier Denux (olivier.denux@orleans.inra.fr), Pierre Zagatti (pierre.zagatti@ver- sailles.inra.fr) Academic editor: David Roy | Received 4 February 2010 | Accepted 23 May 2010 | Published 6 July 2010 Citation: Denux O, Zagatti P (2010) Coleoptera families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae. Chapter 8.5. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 315–406. doi: 10.3897/biorisk.4.61 Abstract Here we consider 274 alien Coleoptera species belonging to 41 of the 137 beetle families in Europe (Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae are treated separately elsewhere). Among the families we consider as having invaded the European fauna, Acanthocnemidae and Ptilodactylidae represent new arrivals. Many species-rich families have surprisingly few aliens, whereas some relatively minor families such as Dermestidae, Nitidulidae and Anobiidae have a relatively high representation of alien species. Since the start of the 19th century, the number of coleopteran aliens introduced into Europe has continued to increase. Alien species colonizing Europe derive from a wide range of geographic regions as well as ecozones, but the most important source area is Asia. The countries with the largest number of alien species established are France, Germany and Italy. The majority have been introduced accidentally via international transport mechanisms. The most important route for importation is stored products and crops, followed by transport of wood, then horticultural and ornamental plants. Most alien species in these families are found within anthropogenic habitats in Europe. The introduction of invasive alien beetles in these families has had significant economic impacts, particularly as pests of stored foodstuffs, as well as serious ecological impacts. For example, the buprestid species Agrilus planipennis, recently recorded in Russia, is an important potential economic threat which may also impact the biodiversity associated with ash trees. Keywords Europe, beetles, Dermestidae, Nitidulidae, Anobiidae, alien species, invasive species, stored products, pests Copyright O. Denux, P. Zagatti. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 316 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) 8.5.1. Introduction Introductions of alien species in Europe started in ancient times (Genovesi and Shine 2003), but this phenomenon has grown rapidly during the two last centuries. This is considered largely to be a consequence of the globalization of trade (Smith et al. 2007). Among these introductions, Coleoptera dominate the alien terrestrial invertebrates in Europe, where the fauna consists of over 27,000 species in 137 families (Fauna Europaea Web Service). In addition to the alien species observed in the families Cerambycidae, Curculionidae (sensu lato), Chrysomelidae (sensu lato) and Coccinelidae, which were treated in the preceding chapters, 274 other beetles of exotic or cryptogenic origin have been established to date in Europe (Table 8.5.1). These alien species belong to 41 different families. Additionally, 237 species are considered to have been introduced through human activity from one region of Europe to another (Table 8.5.2). However, the cause of such movements are often difficult to ascertain, particularly where the original range is poorly known. Thus, the analyses detailed below will mostly consider the species alien to Europe. 8.5.2 Diversity of alien coleopteran species The Coleoptera families treated here with the greatest number of species in Europe are Staphylinidae (rove beetles), Carabidae (ground beetles) and Tenebrionidae (darkling beetles) but these have proportionally few alien species (figure 8.5.1). These three families constitute an important component of the European ground fauna (Dajoz 2002). Conversely, the families with the most aliens in Europe and significant economic impact tend to be families with relatively few native species such as Dermestidae (carpet beetles), Nitidulidae (sap-feeding beetles) and Anobiidae (death-watch beetles). Two of the 41 families do not have any native species in Europe and they are new arrivals for the European fauna: Acanthocnemidae (little ash beetles) and Ptilodactylidae (toewinged beetles). The following presentation of families follows the taxonomic classification of Fauna Europaea (Fauna Europaea Web Service) and of the Tree of Life Web Project (Maddison et al. 2007) (for Ptilodactylidae, not included in Fauna Europaea). ADEPHAGA The Carabidae, are widespread and known to colonize a great diversity of ecological niches (Denux et al. 2007, Holland 2002). They are typically predators (as larvae and adults), although some groups (e.g. Harpalinae) have evolved toward granivory (feeding on seeds). These life traits do not favour passive transportation by humans, and thus, only eight alien species have been established in Europe, accounting for approximately 0.2% of the European carabid fauna. Among these, Somotrichus unifasciatus, Trechicus nigriceps and Plochionus pallens have benefited from the global trade in food products to become cosmopolitan, being introduced with cargos of groundnuts, rice, broad beans, Coleoptera families other than Cerambycidae, Curculionidae... 317 Figure 8.5.1. Relative importance of the Coleoptera families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae families in the alien and native fauna in Europe. Right - Relative importance of the families in the alien entomofauna. Families are presented in a decreasing order based on the number of alien species. Species alien to Europe include cryptogenic species. The number over each bar indicates the number of alien species observed per family. Left - Species richness of the same families in the native European entomofauna. The number over each bar indicates the total number of species observed per family in Europe. cocoa, etc. (Jeannel 1942, Weidner et al. 1984). Only one species is established throughout Europe: Trechicus nigriceps (recorded in 30 countries). This species seems to have been imported from the Eastern coast of Africa several centuries ago (Jeannel 1942). The Dytiscidae (predaceous diving beetles) are all aquatic carnivores. Only one dytiscid beetle has been reported in our database (DAISIE). This large South American species, Megadytes costalis, has been recorded once in Great Britain, but there is no data on its establishment in the wild. POLYPHAGA STAPHYLINIFORMIA The Hydrophilidae (water scavenger beetles) are another family of aquatic beetles, easily distinguished from the Dytiscidae by the length of their maxillary palpi. One tribe, the Sphaeridiini, is exceptional due to its terrestrial, saprophagous and coprophagous habits. Many species share mammal dung with scarab beetles. Significantly, among eight hydrophilids reported as aliens in Europe, seven belong to the Sphaeridiini. 318 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) The Histeridae (clown beetles) are mainly predators, specializing on saprophagous, coprophagous or necrophagous prey. Eight species have been reported in the database, but little is known about their life traits, except for the widespread, cryptogenic Carcinops pumilio, which is common everywhere in natural and anthropized habitats. The Ptiliidae (featherwinged beetles) are a very small family (120 species in Europe and 180 in the world) of which 12 alien species have been recorded in Europe. These are very tiny beetles, including the smallest of all, with a length of just 0.5 mm, whilst even the largest members of the family do not exceed 2 mm. Adults and larvae are usually found in rotting organic material in a wide range of habitats. Their small size and lifestyle means that they are easily dispersed via the movements of soil. Staphylinidae is the most important group of Coleoptera in Europe and the second richest in the world (with over 46,000 species), but the number of alien species (31) in Europe is proportionally low, representing 0.7% of the whole of the Europeans staphylinid fauna. Many genera were not included in Fauna Europaea (Fauna Europaea Web Service), due to the lack of taxonomic expertise. Staphylinidae alien species found in Europe are essentially predatory (Coiffait 1972, Paulian 1988) and mainly species associated with compost, humus and decomposing matter (Cho 2008, Ødegaard and Tømmerås 2000, Tronquet 2006), such as Bisnius parcus, Lithocharis nigriceps and Oxytelus migrator. One predatory species, Philonthus rectangulus, has been reported from 36 countries/ islands. Originating from temperate East Asia, it may have expanded westward naturally. POLYPHAGA SCARABAEIFORMIA The Trogidae (hide beetles) are a small family of beetles related to the scarabs. They feed on mammal skins and furs, or on bird feathers, either as late arriving necrophages on carrion, or as commensals of vertebrates in their nests. Two species from Australasia have been recorded in Spain in our database. The Aphodiidae (dung beetle) are mainly small dung beetles, frequently included in the Scarabaeidae. Four species have been recorded as aliens, in one country only. Both Saprosites species introduced in Great Britain seem to be saproxylic beetles (Angus et al. 2003). The Rutelidae (leaf chafers) are a family of brightly-coloured beetles, especially diverse in the tropics. Only one species of this family has been found in the Azores, the well-known Japanese beetle, Popilia japonica, which is considered as a severe pest in the United States, where it was introduced from Japan in 1912. POLYPHAGA ELATERIFORMIA The Clambidae (minute beetles) are very small beetles that have the capability to roll into a ball. One species is listed here, the Australian Clambus simsoni, a saprophagous species which seems to be rapidly expanding in western Europe. Coleoptera families other than Cerambycidae, Curculionidae... 319 The Buprestidae (metallic wood-boring or jewel beetles) are a well-known family of xylophagous beetles. In most cases, the larvae develop in living wood, and a few species became major pests in orchards or forests. Only three buprestid species have been reported as aliens in the database, each observed in only one country. The Ptilodactylidae, the “toed-winged beetles”, are a group of elateriform Coleoptera, which was formely treated as part of the Dascilloidea and included in Byrrhoidea (Maddison et al. 2007). Little is known of the biology of adults (Aberlenc and Allemand 1997). The habit of soil-leaf litter dwelling of both the adults and larvae facilitates their distribution with potted plants (Mann 2006). The Elateridae (click beetles) are a large family of beetles with quite diverse life history traits. Some species have soil-living larvae, either predators or rhizophages, with reported agricultural pests in the latter category. Other species are saproxylic (predators or saprophages), some of which are very specialized, and have high conservation value. Three species are reported as aliens here, occuring in one country each. The life history traits of these species remain unknown. POLYPHAGA BOSTRICHIFORMIA The European Dermestidae comprise only 139 species (less than 1% of the European Coleoptera fauna) yet they are the largest contributor to the database, with 40 species reported as aliens. Many species are synanthropic and associated with animal remains, leathers and skins, dried meats, woollens and furs (Delobel and Tran 1993), such as Dermestes vorax, D. frischi, D. maculatus, D. lardarius and Anthrenus flavidus. Some species eat stored seeds such as Trogoderma granarium. The protraction of the number of larval stages and longevity in suboptimal nutritive media (Delobel and Tran 1993), as well as the relevance of the food product trade, explain partly how the damaging pests of this family have easily conquered new territories. The Lyctidae (true powder-post beetles) are a very small family (13 species in Europe) closely related to the Bostrichidae. All species are wood-borers, specializing on hardwoods. They usually attack dry wood that is less than five years old, and may become important pests of structural wood or furniture. As inhabitants of raw or manufactured wood products, they are easily transported. Six species have been reported as aliens in Europe, but only one, Lyctus brunneus, has been established throughout the continent for more than 150 years. The Bostrichidae (horned powder-post beetles) are a small family (37 native species in Europe). The native species are saproxylophages, whereas the aliens are either wood-borers or grain-feeders (apparently, some species show both feeding habits) (Lesne 1901). Seven species have been reported as aliens, and have been found in many countries. The wood-borers may cause important damage to manufactured objects, but the stored-product feeders (Dinoderus spp., Rhyzopertha dominica) are the most economically harmful. Among these, the lesser grain borer, Rhyzopertha dominica, has been observed in 34 countries/islands. 320 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) The Anobiidae have 19 alien species compared to 402 native species in Europe. About 11 species are associated with stored food products and include devastating pests such as Lasioderma sericorne which attacks a wide variety of dried products of animal or vegetable origin (Espanol 1992, Weidner et al. 1984). Several species attack soft woody matter: wood in the case of Ernobius mollis, but also books in the case of Nicobium castaneum, which can cause irrepairable damage. Many cryptogenic anobiid species are established in Europe for centuries, and may be found in many countries. POLYPHAGA CUCUJIFORMIA The Nitidulidae have 26 aliens compared with 219 native species in Europe. A third of these have occurred as far west as Macaronesia, but the other species have expanded their range in many countries of mainland Europe. As the majority of species are polleneaters, phytophagous, mycetophagous or predatory, they have a particular agronomic importance, damaging crops and stored food products. Among these, the 13 aliens species of the genus Carpophilus cause damage to dried fruits (Weidner et al. 1984). The Cybocephalidae are a very small family, frequently subsumed within Nitidulidae. Cybocephaline beetles are well known predators of armoured scale insects (Coccoidea: Diaspididae) throughout tropical, sub-tropical and temperate regions of the world (Kirejtshuk et al. 1997).They are minute beetles, very convex and able to roll into a ball, as for Clambidae. The Silvanidae (silvanid flat bark beetles) are a small family (34 native species in Europe) of flat beetles, formerly included in the Cucujidae. These insects were originally mycetophages, living under the bark of trees, but the feeding habits of many species have adapted to grain and fruit feeding, so that they have become synanthropic pests of stored products (Ratti 2007). Nine species are listed in the database, among which three are cryptogenic, long-established species occuring in several countries, such as the sawtoothed grain beetle, Oryzaephilus surinamensis. The Laemophloeidae (lined flat bark beetles) are a small family of flat beetles with 23 native species in Europe, which was formerly included in the Cucujidae. They are closely related to the Silvanidae, and show the same life history traits. Six species, belonging to the genus Cryptolestes, are reported as aliens in Europe. They have established successfully in many countries. The Phalacridae (shining flower beetles) are a small family of minute, rounded beetles. One North American species of Phalacrus has been recorded in the Azores, whose biological traits remain unknown (many species are micro-mycetophages). The Cryptophagidae (silken fungus beetles) are an important family of mycetophagous insects with 228 native species in Europe, living in various habitats. Ten species have been reported as aliens in Europe, which are now established in many countries (the Cryptophagidae have the widest species range). The majority of these species (Cryptophagus spp.) are cryptogenic, feeding on fungal spores or decaying vegetal material, sometimes on stored products. Coleoptera families other than Cerambycidae, Curculionidae... 321 The Languriidae (lizard beetles) are a small family (12 native species in Europe) of phytophagous or saprophagous beetles. Three alien species are considered here, with a rather low dispersal rate. Nevertheless, Cryptophilus integer and Pharaxonotha kirschii are reported as pests of stored products. The Erotylidae (pleasing fungus beetles) are a small family of mycetophagous beetles, with many species in saproxylic habitats. One Japanese species, Dacne picta, has possibly been introduced in Central Europe. The Cerylonidae (minute bark beetles) are a small family of saproxylic beetles. They just appear here because a well-known pest of stored grain, Murmidius ovalis, is now included in this family (formerly Murmidiidae). This is a cosmopolitan species probably originating from tropical Asia. The Endomychidae (handsome fungus beetles) are a small family of mycetophagous beetles (Shockley 2009, Shockley et al. 2009b), closely related to the Coccinellidae. Two very small species (Holoparamecus spp.) are cryptogenic and may be found in many countries worldwide. The Corylophidae (minute hooded beetles) are another small family of micromycetophagous beetles, which occur in a variety of habitats. One species, Orthoperus aequalis, from Australia, has now established in 10 countries within Europe. The Latridiidae (minute hooded beetles) are also a small family with 171 native species in Europe and 17 aliens which are essentially mycetophagous and associated with stored food products, such as Dianerella filum or Cartodere nodifer. These species are also plaster beetles which occupy wet places in the plastered walls of houses (Bouget and Vincent 2008). However, these latridiids do not appear to have an economic impact (Delobel and Tran 1993) and merely indicate bad food storage conditions. The Trogositidae (bark-gnawing beetles) are a small family of saproxylic insects, living as saprophages or predators of other insects under the bark of trees. The three species reported here are predators of cosmopolitan pests of stored products. The Cleridae (checkered beetles) are a conspicuous family of brightly coloured insects. Nearby all species are predators of other insects. Seven species are reported as aliens in the database, some of them (Necrobia spp.) established in Europe for a long time. These are either predators of xylophagous beetles or predators of stored product insects, and thus likely to be transported everywhere with their prey. We include here in the Cleridae the small family Thanerocleridae, which shows life traits similar to the typical Cleridae, with one introduced species, Thaneroclerus buqueti. The Acanthocnemidae, have only one alien species: Acanthocnemus nigricans which is attracted by forest fires (Schmitz et al. 2002). The recent worldwide expansion of this species is due to the commercial export of Australian wood (Kreiss et al. 2005). The Mycetophagidae (hairy fungus beetles) are a family of saproxylic insects, feeding on tree fungi. Two species, specialized on fungi growing on rotten vegetal material, are reported in the database. Typhaea stercorea is a well-known cryptogenic species, whereas Litargus balteatus is an American species found only recently in Europe The Ciidae (minute tree-fungus beetles) are another family of saproxylic insects feeding on tree fungi. Only one species (out of 76 occurring in Europe) is reported 322 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) here as alien, Xylographus bostrichoides. This small insect probably originates from Asia and has to date been found in 19 European countries. The Mordellidae (tumbling flower beetles) are a large family (256 native species in Europe) of flower-dwelling insects, with endophytic larvae. Only one species, Mordellistena cattleyana, is considered as an alien in Europe. This is a neotropical insect whose larvae develop inside tissues of ornamental orchids (Costa Lima 1955). This behaviour may have enabled its importation through the horticultural trade, since it has been found in Germany and the Netherlands. The Ripiphoridae, formerly spelled Rhipiphoridae (wedge-shaped beetles), are a small family of strange parasitic insects. Their larvae develop in other insect orders, namely Hymenoptera, Orthoptera or Dictyoptera. One species, Ripidius pectinicornis, has sometimes been found in harbours, along with its host cockroaches (mainly Blatta orientalis). The Zopheridae (ironclad beetles) were previously included in the Colydiidae. This is a family of saproxylic, bark-living insects with 125 native species in Europe. The three species reported as aliens in Europe are probably predators of other saproxylic insects. They are established in one country only, or a small number of countries in the case of Pycnomerus inexpectus, a species found in tropical greenhouses. The Tenebrionidae is mainly composed of saprophagous species. Many species are xerophiles or thermophiles, which explains their predominance in areas with hot climate and their low representation in more temperate zones (Dajoz 2002). About 15 tenebrionid alien species are present in Europe (1.1% of European tenebrionid fauna). The majority of these species are associated with spoiled or wet cereals (Weidner et al. 1984). They include very damaging pests, such as species of Tribolium, which enter cracks in wet or already damaged seeds, and Alphitobius spp., which feed on mildewed food products. The Salpingidae (narrow-waisted bark beetles) are a small family of saproxylic beetles with 18 native species in Europe. One species only is mentioned here, Aglenus brunneus, formerly included in the Colydiidae (Zopheridae). It is a very small, blind insect, often found in stables or poultry houses, where it feeds on animal waste (Dajoz 1977). The Anthicidae (antlike flower beetles) are small beetles resembling ground beetles. Four species are considered as aliens, among 310 native species living in Europe. These insects typically feed on rotten vegetal material, which has been heated through fermentation. These life history traits probably enable a wide tolerance to cold temperatures, and some species are cosmopolitan, found everywhere in the world, from tropical to boreal climates, e.g. Omonadus floralis, recorded in 40 countries. 8.5.3 Temporal trends Some Coleoptera species were introduced to Europe a very long time ago. Fossils of alien species have even been found in archeological sites, such as the blind flightless beetle Aglenus brunneus in Iceland (Buckland et al. 2009) and Amara aulica (alien but native in Europe), which arrived in the Faroe islands with the Viking settlers Coleoptera families other than Cerambycidae, Curculionidae... 323 (Brandt 2006). But the first date of introduction of a new species into a country is often difficult to establish. A species could have been present for years without its presence being noticed immediately. Particularly relevant here are small or inconspicuous species lacking agronomic or economic impact (e.g. Ptiliidae), and members of neglected or hard to identify taxonomic groups (e.g. Cryptophagidae and Staphylinidae). The precise date of the first record is available for 201 species (i.e. 73.1% of aliens). The first statistical data derives from the beginning of the 19th century with the introduction of the nitidulid Carpophilus hemipterus in 1800 by the historical opening of trade routes (Audisio 1993). Then comes the trogossitid Tenebroides mauritanicus in 1803, and the anobiid Nicobium castaneum in 1807. The endomychid Holoparamecus depressus arrived in 1843 and the anobiid Lasioderma sericorne in 1848. These detritivores are all associated with stored food products or wood. We observed an accelerating increase in the number of new records per year (figure 8.5.2), from 0.1 p.a. between 1800–1849 to 3.5 p.a. during 2000–2007, with an intermediate level of 1.3 p.a. during the period 1900–1924. During this last period, 33 new alien species were recorded, including 14 alone for the year 1900. This unexpected increase coincides with the industrial revolution of the first developing European countries (Cosseron and Faverjon 1991) (Great Britain, Belgium, France, and Germany) and with the increase in imports ensuing from it. 8.5.4 Biogeographic patterns 8.5.4.1 Origin of alien species Alien species come from all continents except Antarctica (figure 8.5.3) (arthropods most represented on this continent are Collembola and mites rather than beetles) (Schulte et al. 2008). The considerable periods of environmental stress in Antarctic (Benoit et al. 2009) limit the diversity of insects, even though a very few beetles do occur there (Vernon et al. 1999), such as the ground-beetles Amblysogenium pacificum and A. minimum. These factors explain easily the absence of invasives coming from Antarctic. About 82 aliens have origins currently considered cryptogenic. These are cosmopolitan species or distributed mainly in on one or more ecozones, with a tendency to become cosmopolitan. This is particularly the case with the cryptophagid Cryptophagus cellaris, a holarctic species which has become practically cosmopolitan following international commercial exchanges (Delobel and Tran 1993). Asia is the most important source of aliens, with 58 species established in Europe (21%), comprising Dermestidae (13 spp.), Staphylinidae (8 spp.), Nitidulidae (6 spp.), Anthicidae (4 spp.) and Carabidae (3 spp.). These families are generally associated with stored products, crops, decomposing matter such as compost, and to a lesser extent with wood. The 16 other families number one or two species of aliens each. 324 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Figure 8.5.2. Temporal changes in the mean number of new records per year of alien Coleoptera species of families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae, from 1800 to 2007. The number over each bar indicates the absolute number of species newly recorded per time period. About 35 aliens come from Africa and these comprise Nitidulidae (5 spp.), Carabidae (3 spp.), Histeridae (3 spp.), Hydrophilidae (3 spp.) and Tenebrionidae (3 spp.). Nitidulidae and Tenebrionidae have been transported through stored food products. The mode of introduction is unknown for Carabidae and Hydrophilidae. There are also 14 other families having one or two alien species, which are partly associated with stored food products and wood. The 55 aliens coming from the American continent (20% of the all alien species to Europe), include 24 species from North America and 31 species from Central and South America. From North America, the principal families are Dermestidae (7 spp.), Nitidulidae (6 spp.) and Tenebrionidae (4 spp.). Four species of Staphylinidae and four species of Ptiliidae derive from Central and South America. As for Asia and Africa, the neoarctic and neotropic aliens are mainly associated with foodstuffs and cultures. About 16 other families coming from America with one or two alien species have also been recorded in Europe. Relatively few aliens originate from Australia. The 25 species of Australian origin include Latridiidae (4 spp.), Ptiliidae (4 spp.) and Staphylinidae (3 spp.). These species have no economic impact. The 12 other families include one or two alien species each, among which are species of the stored food products (Ptinus ocellus, Anthrenus oceanicus, Brachypeplus mauli) or living under the tree bark (Ptinella cavelli and P. errabunda). The aliens with a specifically tropical origin (Pantropical) are the least presented in Europe with 20 species, that is to say 7% of all exotic species to Europe. The families Coleoptera families other than Cerambycidae, Curculionidae... 325 Figure 8.5.3. Origin of the Coleoptera species alien to Europe of families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae with the most species are Anobiidae (3 spp.), Bostrichidae (3 spp.) and Tenebrionidae (3 spp.). The eight other families have only one or two species each. These tropical aliens are associated with stored food products and fruits. During different time slices, the origin of alien species has increasingly diversified (figure 8.5.4). The number of ecozones represented has increased from three (Africa, Asia, Pantropical) during 1800–1849 to six since 1950–1974 (Africa, Asia, Australasia, Central and South America, North America, Pantropical). The geographic source has also varied temporally although Asia has always been both an important and early region of origin. This situation can be explained by the opening of the trade route between Europe and India by the Cape of Good Hope at the end of the 15th century (which was also the sole sea route before the opening of the Suez Canal in 1869) and the strong Western influence which followed, the opium wars and the East India Companies, which revolutionized methods and the extent of the trade with Asia. We highlight especially two ambiguous periods for biological invasions: 1850–1899 and 1925–1949. During the first period, no new record of an alien from Africa was recorded in Europe. The same goes for the second period with a fall of the number of new arrivals detected from South America (nine in 1900–1924 and only two in 1925– 1949). These phenomena may coincide with the Great Depression, the result of the economic crisis of 1929 (Cosseron and Faverjon 1991, Gravereau and Trauman 2001), which affected both the level of protectionism on trade routes and the overall volume of international economic exchange between Europe and its colonies. The consequence for South America, Asia and Africa was “the crisis of dessert products”, coinciding with the fall of the purchasing power in Europe and North America. Thus in Brazil for example, in an attempt to control the market, coffee was burned in engines (Launay 1999). 326 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Figure 8.5.4. Temporal changes in the origin of the Coleoptera species alien to Europe of families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae The late arrival of aliens to Europe from North America is remarkable (first record in 1935) and probably corresponds to weak exports of foodstuffs towards Europe (except cereals). For forest biotopes especially, the North American component of species is small and of limited economic impact in Europe (Dajoz 2007). 8.5.4.2 Distribution of alien species within Europe and their range expansion The majority of European countries have been directly affected by alien species (figure 8.5.5), but there is a very great mismatch in the number of species present in one country versus another. The archipelago of Svalbard, with an insect fauna of a meagre 230 species (Coulson 2007), seems free from aliens. As in the case of Antarctica, the strong environmental contraints (harsh temperatures, shortened seasons and strong winds) have evidently limited the colonization of insects (Hulle et al. 2008) and geographical isolation has posed a barrier. For Macedonia there is a lack of readily accessible data (Tomov 2009), which has prevented us updating the situation there. The countries/islands most affected by aliens are France (126), Germany (107), Italy (101), Austria (98), Great Britain (97), Switzerland (91), the archipelago of Azores (92), Denmark (89) and the Czech Republic (84). Coleoptera families other than Cerambycidae, Curculionidae... 327 Figure 8.5.5. Comparative colonization of continental European countries and islands by by the Coleoptera species alien to Europe of families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae. Archipelago: 1 Azores 2 Madeira 3 Canary islands. The number of aliens per country is not significantly correlated with Global Domestic Product per capita (International Monetary Fund), latitude, nor longitude of the centroid of the country. In contrast, the number of aliens per country is significantly correlated with import (Spearman-Rho 0.650, P-value < 0.001) from 2003 to 2008 (The World Factbook) and also more weakly with area (Spearman-Rho 0.432, P-value < 0.01). In spite of its geographical isolation (1500km from Europe, 1450km from Africa and 3900km from North America) and its small area, the archipelago of Azores has a large number of aliens. Since their historical discovery, the geographic position of the Azores has made the islands a strategic harbour for transatlantic ships, resulting in the introduction overall of several hundreds of taxa (Haggar 1988, Heleno 2008). Twenty-four alien species have been recorded exclusively in the Azores archipelago. 328 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Indeed, alien native species in Europe have colonized islands more than other continental countries. The archipelago of Azores is the most affected with 126 alien species to Europe, followed by Great Britain (with 58 aliens), Faroe Islands (32 aliens) and Canary Islands (32 aliens). Perhaps surprisingly, Austria is the most important continental country affect by alien native to Europe, with 13 species. 8.5.5 Main pathways to Europe The most important pathways for accidental invasions of exotic species to Europe are those closely bound to international transport, whereas the most important processes relating to deliberate introductions are the biological control of agricultural pests and the pollination of crops (Ruiz and Carlton 2003). Rapidly developing international trade and the reduction of travel times by air to less than two days, have meant that a living insect can be transported almost any part of the world (Mouchet et al. 1995). Only three species have been introduced intentionally in Europe, two for biological control. The first is the cybocephalid beetle Cybocephalus nipponicus, originating in South Korea (Evans et al. 2005) and introduced into Italy for the control of cochineals bugs (Diaspididae) (Lupi 2002). The second species is Ripidius pectinicornis (Ripiphoridae), a parasitoid of the german cockroach Blattella germanica (Falin 2001) which was released from culture and is now present in several European countries (Bétis 1912). The third species is the tenebrionid Zophobas morio which has been used for bird and especially lizard food (Thomas 1995). About 98.9% of aliens have been introduced accidently in Europe. The exact pathway of introduction is difficult to establish. The introduction vector is unknown for 123 aliens out of the total of 275. Theses aliens are essentially detritiphagous, saproxylophagous or predatory species. The first clearly identified means of importation is via stored products and crops (approximately 120 aliens, or 40%). This can be explained by the importance of the international stored products trade (cereals, fruits and vegetables) and the primary position of Coleoptera as pests of stored products (Delobel and Tran 1993). About 20 Coleoptera have been implicated directly in the transport of woods. Some species have been found in wood derivatives such as Dinoderus minutus, a bostrichid introduced with furniture and bamboo-work (Lesne 1901). Few species have been identified as transported with horticultural or ornamental products, despite the increase of economic importance of ornamental pot plants (Lawson 1996), in sharp contrast for example to the situation in Lepidoptera (see Chapter 11). However, the level may be underestimated for this route, as some Coleoptera tend to occur in compost and may pass unnoticed via the pot plant trade. The extruded starch products used as impact protection for fragile packing can even be a food source for stored grains pests (Fraga et al. 2009) as for Cryptolestes ferrugineus, Lasioderma serricorne and Tribolium castaneum. Thus starch-packings could become a new vector of introductions in the future. Coleoptera families other than Cerambycidae, Curculionidae... 329 8.5.6 Most invaded ecosystems and habitats The anthropogenic habitats most strongly colonized by coleopteran alien species (figure 8.5.6), are buildings (50%), cultivated lands (20%) and forest habitats (10%). The large proportion of species associated with foodstuffs explains this relation. Conversely, the weak colonization of pseudo-natural habitats can be explained by the near-absence of phytophagous, and more particularly phyllophagous species among the coleopteran families treated here. This result contrasts with the situation for other groups of predominantly phytophagous insects (Cerambycidae, Chrysomelidae, Lepidoptera: Chapter 8.1, 8.3, 11). In spite of the popularity of exotic species for the aquatic animal and plant trade (Leppäkoski et al. 2002) and the fact that migrating waterfowl can transport aquatic invertebrates or their eggs (Figuerola et al. 2003), surprisingly no water beetle has been introduced into Europe, except for the dytiscid Megadytes costalis (again contrasting with the situation for Lepidoptera, the aquatic Pyraloidea: Chapter 11). This low importance of the aquatic route in Coleoptera is also observed in the United States, where only 2.2% of the invasive arthropods are aquatics (Pimentel et al. 2005). 8.5.7 Ecological and economics impacts Most alien species do not become invasive in their new locations (Genovesi and Shine 2003). It is often difficult to predict whether a new introduction will actually become established (Streito and Martinez 2008). However, the subset of alien species that are invasive may have significant environmental, economic and public health impacts and threaten the wholesale homogenisation of ecosystems (Sefrova 2005). Invasive alien species are now considered to be the second greatest cause of global biodiversity loss after direct habitat destruction (Simberloff 2001) and have adverse environmental, economic and social impacts from the local level upwards. The invasion of most Coleoptera treated here bears a direct relation to human presence (synanthropic species). Their impact is essentially with stored foodstuffs which they can extensively damage (Sefrova 2005). Coleoptera damaging stored food products on a global economic scale are very few (Delobel and Tran 1993), but include several species of aliens in Europe, among which are Cryptolestes ferrugineus, C. pusillus, Lasioderma serricorne, Oryzaephilus surinamensis, Rhyzopertha dominica, Tribolium castaneum, T. confusum and Trogoderma granarium. The impact of insect pests in a given situation can widely fluctuate depending on various parameters, in particular on production levels and the commercial value of those products infested both in time and in a geo-economic context. However, these synanthropic species are not known to have a direct effect on biodiversity. The situation for agronomic and forest species can be different. The buprestid Agrilus planipennis, recently recorded in European Russia, is a very good example. This xylophagous East Asian species is presently causing significant damage to ash trees (Fraxinus spp.) in North America (Baranchikov et al. 2008). A. planipennis has killed 330 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Figure 8.5.6. Main European habitats colonized by the Coleoptera species alien to Europe of families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae. The number over each bar indicates the absolute number of alien coleopterans recorded per habitat. Note that a species may have colonized several habitats. over 15 million forest and ornamental trees in several US States in less than 10 years (Poland and McCullough 2006). It is alarming that European ash trees are not more resistant than those of North America (Baranchikov et al. 2008). Agrilus planipennis could become a serious pest in Europe with a dramatic economy impact as well as potentially for biodiversity associated with Fraxinus. Many species are associated with compost and even while their economical impact may be negligible (as mainly predators or detritivores), ecological disruption may still occur. This appears to be the case with the Staphylinid Lithocharis ochracea. This native beetle has declined, supplanted by the alien species L. nigriceps (Ødegaard and Tømmerås 2000, Tronquet 2006). Even if the eradication of invasive species seems possible in Europe and in particular for mammals (Genovesi 2005), the possibility of eradication of invasive Coleoptera appears much more remote. 8.5.8 Conclusion On of the most striking consequences of globalization is the increase in the problem of invasive species (Perrings et al. 2005). The volume of international trade and travel is now so great, and the modes of entry so varied, that not all consignments or routes of entry can be screened (Levine and D’Antonio 2003). Three categories are particularly important to highlight for the coleopteran alien species treated here: synantropic Coleoptera families other than Cerambycidae, Curculionidae... b 331 c a d e h g f l k i j 1 mm Figure 8.5.7. Habitus of some Coleoptera species alien to Europe. a Ernobius mollis b Tribolium castaneum c Oryzaephilus surinamensis d Alphitobius diaperinus e Cryptolestes duplicatus f Dermestes lardarius g Gnathocerus cornutus h Rhizopertha dominica i Necrobia ruficollis j Trechicus nigriceps k Lyctus brunneus l Gibbium psylloides (Credit: Pierre Zagatti). 332 Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) habitats with essentially stored products, compost (probably that associated with ornamental plants), and forest or wood-derived products. Acknowledgements We thank Alain Roques and David Roy for their useful comments on the manuscript and David Lees for linguistic improvements. 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Status: A Alien to Europe C Cryptogenic. Country codes abbreviations refer to ISO 3166 (see Appendix I). Habitat abbreviations refer to EUNIS (see Appendix II). Family species Regime Native range 1st record in Europe Invaded countries Habitat Host References A phytophagous Australasia 1922, CY, FR, FR-COR, DE, IT, FR-COR IT-SAR, IT-SIC, PT, ES I2 timber, wood Alonso-Zarazaga et al. (2003), (Kreiss et al. (2005) A phytophagous Tropical, Unknown PT-AZO subtropical J1 stored products C detritivorous Cryptogenic Bercedo et al. (2008), Borges et al. (2005), Espanol (1979), Mendonça and Borges (2009) Tomov (2009), Wittenberg et al. (2006) Ernobius mollis (Linnaeus, 1758) C phytophagous Cryptogenic Gibbium aequinoctiale Boieldieu, 1854 Gibbium psylloides (Czempinski, 1778) A detritivorous Tropical, Unknown MT subtropical C detritivorous Cryptogenic 1861, DE AT, BE, BA, BG, HR, CZ, J1 DK, EE, FI, FR, FR-COR, DE, HU, IS, IE, IT, LV, LT, LU, MD, NL, NO, PL, PT, PT-AZO, RO, RU, RS, SK, SI, ES, SE, CH, UA, GB Unknown PT-AZO J, G barns, cowsheds, animal burrows soft wood, sawmills, books stored products 1900, CZ AL, AT, BE, BA, BG, HR, J1 CY, CZ, DK, EE, FI, FR, FR-COR, DE, GR, HU, IE, IT, IT-SAR, IT-SIC, MT, MD, NL, PL, PT, PT-MAD, RO, RU, RS, SK, ES, ESBAL, SE, CH, UA, GB houses, hotels, stored products Bellés (1985), Bellés and Halstead (1985), Duff (2008), Freude et al. (1969), Šefrova and Lastuvka (2005), Wittenberg et al. (2006) 343 J1 Borges et al. (2005), Espanol (1992), Mendonça and Borges (2009) Bellés and Halstead (1985) Coleoptera families other than Cerambycidae, Curculionidae... Acanthocnemidae Acanthocnemus nigricans (Hope 1845) Anobiidae Calymmaderus oblongus (Gorham, 1883) Epauloecus unicolor (Piller and Mitterpacher) Status Status Regime Native range 1st record in Europe Invaded countries Habitat Host References A phytophagous Tropical, 1848, PT AL, AT, BG, CZ, DK, EE, subtropical HU, IT, IT-SAR, IT-SIC, LV, MT, PT, RS, CH Mezium affine Boieldieu 1856 C detritivorous Cryptogenic Unknown AT, DK, DE, PT-AZO, PT- J MAD, ES, ES-CAN, SE Mezium americanum Laporte de Castelnau, 1840 Nicobium castaneum (Olivier, 1790) A detritivorous North America Unknown IT, IT-SAR, MT, PT-AZO J C phytophagous Cryptogenic J soft wood furniture, old books Espanol (1992), Freude et al. (1969), Mendonça and Borges (2009), Šefrova and Lastuvka (2005) Ozognathus cornutus (LeConte, 1859) Pseudeurostus hilleri (Reitter 1877) A detritivorous North America 1807, PT AT, BA, HR, CY, CZ, FR, FR-COR, DE, GR, IT, ITSAR, IT-SIC, MT, PL, PT, PT-AZO, PT-MAD, RO, SI, ES, ES-BAL, ES-CAN, CH, UA 2005, ES MT, RO, ES J dead wood Allemand (2008), Bercedo et al. (2005), Zahradnik and Mifsud (2005) A detritivorous Asia1993, DE DK, DE Temperate J Ptilineurus marmoratus (Reitter, 1877) A phytophagous Asia G likely scavenger and inhabitant of residues, potential minor pest of feed mills and warehouses trees Imperial Institute of Entomology (1930) 1999, FR FR, SE J1 tobacco, stored products mills, stored products, bird nests stored products Borges et al. (2005), Espanol (1992), Freude et al. (1969), Glavendekic et al. (2005), Šefrova and Lastuvka (2005), Wittenberg et al. (2006) Bellés (1985), Freude et al. (1969) Bellés (1985), Borges et al. (2005) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Lasioderma sericorne Fabricius, 1792 344 Family species Family species Status Regime Native range C detritivorous Cryptogenic Ptinus clavipes Panzer, 1792 C detritivorous Cryptogenic Ptinus fur (Linnaeus 1758) C detritivorous Cryptogenic Ptinus latro Fabricius, 1775 C detritivorous Cryptogenic Invaded countries Habitat Host References J1 1856, FR AT, BY, BE, BA, BG, HR, CZ, DK, EE, FI, FR, FRCOR, DE, HU, IT, IT-SAR, LV, NL, NO, PL, RO, RU, RS, SK, SI, ES, SE, CH, UA Unknown EE, LV, MT, ES-CAN, GB J1 stored products Freude et al. (1969) stored products, fur 1940, BG AL, AD, AT, BY, BE, BA, J1, J6 BG, HR, CY, CZ, DK, EE, FÖ, FI, FR, FR-COR, DE, GR, HU, IS, IE, IT, IT-SAR, IT-SIC, LV, LI, LT, LU, MT, MD, NL, NO, PL, PT, PT-AZO, PT-MAD, RO, RU, RS, SK, SI, ES, ES-BAL, ES-CAN, SE, CH, UA, GB 1850, CZ AL, AT, BY, BE, BA, BG, J HR, CY, CZ, DK, EE, FI, FR, FR-COR, DE, GR, GR-CRE, HU, IE, IT, ITSAR, IT-SIC, LV, LI, LT, LU, MT, MD, NL, NO, PL, PT, PT-AZO, PT-MAD, RO, RS, SK, SI, ES, ESCAN, SE, CH, UA, GB waste, dried vegetals Duff (2008), Freude et al. (1969), Machado and Oromi (2000) Bengtson (1981), Borges et al. (2005), Duff (2008), Mendonça and Borges (2009), Tomov (2009) old wood, synanthropic Borges et al. (2005), Freude et al. (1969), Šefrova and Lastuvka (2005), Tomov (2009) Coleoptera families other than Cerambycidae, Curculionidae... Ptinus bicinctus Sturm 1837 1st record in Europe 345 Status Regime 1st record in Europe Invaded countries Habitat Host References stored products Allemand (2008), Bengtson (1981), Duff (2008), Wittenberg et al. (2006) J Freude et al. (1969), Šefrova and Lastuvka (2005) A detritivorous C&S America 1939, CZ CZ, DK, GB J seeds, stored products; crataegus in native fields dried animal products, insects, herbarium, stored products A unknown Asia Unknown CY, GR, GR-SEG, MT, PT-MAD J A unknown Asia 1982, IT IT U A detritivorous AsiaTropical 1951, HR, BG AL, AT, BA, BG, HR, CY, J6 CZ, DK, EE, FI, FR, FRCOR, DE, GR, GR-CRE, GR-ION, GR-SEG, HU, IE, IT, IT-SAR, IT-SIC, LV, LI, LT, MT, NL, NO, PL, PT, PT-AZO, PT-MAD, RO, RU, SK, ES, ES-BAL, ES-CAN, SE, CH A detritivorous Tricorynus tabaci (Guérin-Méneville, 1850) A Trigonogenius globulus Solier, 1849 Duff (2008), Ratti. Coleotteri alieni in Italia., Šefrova and Lastuvka (2005) anthropophilous, Pollock and Ivie (1996) larva scavenger Degiovanni and Pezzi (2007) vegetal decay, detritiphage, mycophage, adult predator Freude et al. (1969), Hemp and Dettner (2003), Machado and Oromi (2000), Mendonça and Borges (2009), Tomov (2009) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) J1 phytophagous Australasia 1916, DE AT, BY, BE, BA, BG, HR, CY, CZ, DK, EE, FÖ, FI, FR, DE, GR, HU, IE, IT, LV, LT, LU, MD, NL, NO, PL, PT, PT-AZO, RU, RS, SK, SI, ES, SE, CH, UA, GB C&S 1965, CZ HR, CZ, DK, FR, DE, IT America Ptinus tectus Boieldieu 1856 Anthicidae Anthicus crinitus La Ferte-Senectere, 1849 Anthicus czernohorskyi Pic, 1912 Omonadus floralis (Linnaeus 1758) Native range 346 Family species Family species Stricticomus tobias (De Marseul 1879) Status Regime Native range 1st record in Europe Invaded countries Habitat Duff (2008), Freude et al. (1969), Machado and Oromi (2000), Telnov (1996), Wittenberg et al. (2006) E dung Baraud (1985) Australasia 1921, GB GB I2 Africa 1982, GB GB I2 rotting wood; in Baraud (1992), Duff (2008), borings of Dorcus Paulian and Baraud (1982) and Sinodendron beetles rotting wood Duff (2008) North America 1976, GB GB U Baraud (1992), Duff (2008) Asia 1944, IT A detritivorous detritivorous Africa Unknown PT-AZO detritivorous detritivorous A phytophagous Tropical, Unknown FR, FR-COR, IT-SAR, ITsubtropical SIC, ES G3, I2 Bostrychoplites cornutus (Olivier 1790) A phytophagous Africa J Dinoderus bifoveolatus (Wollaston, 1858) A phytophagous Tropical, Unknown AT, BE, HR, DK, DE, NL, J subtropical PT-MAD, SK, ES, SE, CH, GB Saprosites natalensis (Peringuey, 1901) Tesarius caelatus (Laconte, 1857) Bostrichidae Apate monachus Fabricius, 1775 A A AT, BY, BE, CZ, DK, EE, FR, FR-COR, DE, HU, IT, IT-SAR, IT-SIC, LV, LT, MT, MD, NL, PT, PTMAD, SK, ES, ES-CAN, SE, CH, GB Unknown DK, DE, IT, ES, SE Freude et al. (1969), Lesne (1901) Freude et al. (1969), Ratti. Coleotteri alieni in Italia.) Duff (2008), Freude et al. (1969), Lesne (1901) 347 polyphagous stem borer, fruit trees, Acacia timber (ethnic carved wooden bowls and ornaments ) bamboo borer (N); dried cassava chips and stored products Coleoptera families other than Cerambycidae, Curculionidae... rotten vegetal tissues detritivorous A References I, J1 A Aphodiidae Aphodius gracilis Boheman, 1857 Saprosites mendax Blackburn, 1892 Host Status Regime Native range 1st record in Europe Invaded countries Habitat A phytophagous Tropical, 1965, CZ AL, BE, CZ, DK, FR, DE, subtropical GR, IT, IT-SAR, IT-SIC, NL, PL, SK, SE, GB Rhyzopertha dominica (Fabricius, 1792) A phytophagous AsiaTropical J1 1900, CZ AL, AT, BY, BE, BG, HR, CY, CZ, DK, EE, FI, FR, FR-COR, DE, GR, GRSEG, IE, IT, IT-SAR, ITSIC, LV, MT, NL, PL, PT, PT-AZO, RO, SK, ES, ESBAL, ES-CAN, SE, CH, GB Sinoxylon senegalense Karsch, 1831 A phytophagous Africa Unknown DE J Heterobostrychus hamatipennis (Lesne, 1895) Buprestidae Agrilus planipennis Fairmaire, 1888 Buprestis decora Fabricius, 1775 A phytophagous Asia 2005, BE BE J A phytophagous phytophagous Asia 2003 I2 North America Unknown ES-CAN A RU J,I2 I2 References bamboo, manioc (Cassava), stored products (intro) Duff (2008), Freude et al. (1969), Lesne (1901),Lesne (1904), Šefrova and Lastuvka (2005) stored products, Borges et al. (2005), Cobos mainly cereals (1986), Duff (2008), Freude et al. (1969), Lesne (1901), Lesne (1904), Machado and Oromi (2000), Mendonça and Borges (2009), Šefrova and Lastuvka (2005), Tomov (2009), Wittenberg et al. (2006) Acacia wood borer Lesne (1901) (N); imported construction wood xylophagous, Lesne (1901) Salix, osier goods Fraxinus Baranchikov et al. (2008) Cobos (1986), Machado and Oromi (2000) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Dinoderus minutus (Fabricius, 1775) Host 348 Family species Family species Status Regime Native range 1st record in Europe Invaded countries Habitat Host A phytophagous Africa 1986, ES ES A parasitic/ predator Africa Unknown FR, FR-COR, IE, PT-AZO, B, J, ES-CAN, GB H1 Leistus nubivagus Wollaston, 1864 Notiobia cupripennis (Germar, 1824) Plochionus pallens (Fabricius, 1775) A parasitic/ predator phytophagous Africa Unknown ES-CAN U C&S America Unknown ES-CAN I2 seeds of Amaranthus A parasitic/ predator C&S America 2000, NL DK, FR, DE, HU, IT, NL J Pterostichus caspius (Ménétriés, 1832) A parasitic/ predator Asia1980, CZ BG, CZ Temperate U Somotrichus unifasciatus (Dejean, 1831) A parasitic/ predator Africa J in ports, transported with peanuts, raisin storages Predator in various environments, pyrophilous predator of beetles in stored products, avian droppings A Unknown FR, IT F5, I Ratti. Coleotteri alieni in Italia.) littoral in ports, cellars caves Anderson et al. (2000), Arndt (2006), Borges et al. (2005), Duff (2008), Jeannel (1942), Luff (1998), Luff (2007), Machado (1976), Machado and Oromi (2000), Mendonça and Borges (2009), Perrault (1981), Perrault (1984) Machado (1976), Machado and Oromi (2000), Perrault (1981) Machado and Oromi (2000), Perrault (1984) Trautner and Geigenmuller (1987), Valemberg (1997) Hurka (1996), (Šefrova and Lastuvka (2005), Valemberg (1997) Coleoptera families other than Cerambycidae, Curculionidae... Chrysobothris dorsata (Fabricius, 1787) Carabidae Laemostenus complanatus (Dejean, 1828) References Jeannel (1942), (Valemberg (1997) 349 Trechicus nigriceps (Dejean, 1831) Philothermus montandoni Aube, 1843 Ciidae Xylographus bostrychoides (Dufour 1843) Clambidae Clambus simsoni Blackburn 1902 Regime Native range 1st record in Europe Invaded countries Habitat Host References A parasitic/ predator AsiaTropical 1902, DE AT, BE, BA, BG, HR, CZ, I1, I2, J1, J6 DK, FI, FR, DE, HU, IT, LV, LI, LU, MD, NL, NO, PL, PT-AZO, PT-MAD, RS, SK, SI, ES, ES-CAN, SE, CH, UA, GB Borges et al. (2005), Darlington compost, predator, gardens; (1964), Duff (2008), Hurka also in peanuts (1996), Luff (2007), Machado and Oromi (2000), Mendonça and Borges (2009), Neculiseanu and Matalin (2000), Serrano et al. (2003), Tomov (2009), Trautner and Geigenmuller (1987), Valemberg (1997), Wittenberg et al. (2006) A detritivorous Asia Unknown AL, AT, DK, FR, DE, HU, IT, PL, CH, GB A detritivorous Tropical, Unknown FR, IT subtropical X11 stored products (few damageports) botanical garden A detritivorous Asia? I feeds on fungi Tomov (2009) A detritivorous Australasia 1987, SE AT, FR, DE, NL, SE, GB G forest, firewood, compost; mycophagous Duff (2008), Tamisier (2004) Unknown AT, BY, BA, BG, HR, CZ, DK, FR, FR-COR, GR, HU, IT, IT-SAR, IT-SIC, PL, RO, SK, ES, UA J1 Duff (2008), Wittenberg et al. (2006), Moncoutier (2002) Stoch: Checklist of the species of the italian fauna) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Cerylonidae Murmidius ovalis (Beck 1817) Status 350 Family species Family species Cleridae Necrobia ruficollis (Fabricius 1775) Status Regime Native range 1st record in Europe Invaded countries Cryptogenic Necrobia rufipes (De Geer 1775) A parasitic/ predator Tropical, 1935, LT AT, BG, DK, EE, FI, DE, subtropical LT, NO, PT, PT-AZO, SE, CH J1, J6 Necrobia violacea (Linnaeus 1758) Opetiopalpus scutellaris (Panzer 1797) Paratillus carus (Newman, 1840) Tarsostenus univittatus (Rossi, 1792) Thaneroclerus buqueti (Lefebvre, 1835) C parasitic/ predator parasitic/ predator Cryptogenic Africa J1, J6 parasitic/ predator parasitic/ predator Australasia 1933, GB FR, GB G,I2 Cryptogenic 1990, CZ AT, CZ, CH J A parasitic/ predator Asia 1963, CZ CZ, DE, IT, PL J A detritivorous Australasia Unknown HR, FR, FR-COR, IT, PT-AZO, PT-MAD, ES, ES-CAN, CH, GB Corylophidae Orthoperus aequalis Sharp 1885 A C 1976, LT AT, DK, FI, HU, LT, NO, SE, CH Unknown AT, EE, FR, DE, ES J1, J6 J G, I2 References Borges et al. (2005), Du Chatenet (2000), Freude et al. (1979), Mendonça and Borges (2009), Wittenberg et al. (2006) predator, Borges et al. (2005), Du necrophage, seeds Chatenet (2000), Freude et al. with oil content (1979), Haines and Rees (1989), (copra, soya), Tomov (2009), Wittenberg et al. dried fish (2006) old bones, prey Freude et al. (1979), Wittenberg dry carrion et al. (2006) old timber houses Du Chatenet (2000), Freude et al. (1979) predator on old bones, decaying animals predator on Lyctiidae predator on Bostrychidae, Anobiidae predator on insects on tobacco, rice (Lasioderma, Areaocerus) Du Chatenet (2000), Duff (2008) Du Chatenet (2000), Freude et al. (1979), Šefrova and Lastuvka (2005), Wittenberg et al. (2006) Du Chatenet (2000), Freude et al. (1979), Šefrova and Lastuvka (2005) Borges et al. (2005), Bowestead (1999), Duff (2008), Machado and Oromi (2000), Ratti. Coleotteri alieni in Italia.) 351 parasitic/ predator Host Coleoptera families other than Cerambycidae, Curculionidae... C A 1976, LT AT, DK, EE, FI, HU, LT, NO, PT-AZO, SE, CH Habitat Cryptophagidae Atomaria lewisi Reitter, 1877 Status Regime Native range detritivorous Asia Caenoscelis subdeplanata C.Brisout de Barneville, 1882 A detritivorous North America Cryptophagus acutangulus Gyllenhall, 1828 C detritivorous Cryptogenic Cryptophagus affinis Sturm 1845 C detritivorous Cryptogenic Cryptophagus cellaris (Scopoli, 1763) C detritivorous Cryptogenic Invaded countries Habitat Host mycophage; compost, In decaying plant material G, X11, mycophage; I2, FB forests In decaying wood and plant material Duff (2008), Freude et al. (1967), Ødegaard and Tømmerås (2000), Šefrova and Lastuvka (2005), Wittenberg et al. (2006) Duff (2008), Falcoz (1929), Freude et al. (1967), Ratti. Coleotteri alieni in Italia., Tomov (2009), Wittenberg et al. (2006) J attic, mills Falcoz (1929), Freude et al. (1967), Tomov (2009) J fungi, dry fruits 1939, PT AL, AT, BY, BE, BA, BG, J HR, CZ, DK, FI, FR, DE, GR, HU, IT, IT-SIC, LV, MT, MD, NL, NO, PL, PT, PT-AZO, PT-MAD, RO, SK, SI, ES-CAN, SE, CH, UA, GB mycophagous, stored products, herbariums, insects Borges et al. (2005), Duff (2008), Falcoz (1929), Freude et al. (1967), Machado and Oromi (2000), Mendonça and Borges (2009), Tomov (2009) Borges et al. (2005), Duff (2008), Falcoz (1929), Freude et al. (1967), Machado and Oromi (2000), Moncoutier (2002), Tomov (2009) 1937, GB AL, AT, BY, BE, HR, CZ, DK, EE, FI, DE, IT, LV, LT, MD, NO, PL, PT-AZO, SK, SE, CH, UA, GB 1950, GB BY, HR, CZ, EE, FI, FR, FR-COR, DE, IT, LV, LT, LU, MT, MD, NL, NO, PL, PT-MAD, RU, SI, ES, ESCAN, SE, CH, UA, GB 1956, BG AL, AT, BY, BE, BA, BG, CZ, DK, EE, FI, FR, DE, IT, LV, LT, PL, RO, RS, SK, SI, SE, CH, UA, GB 1956, BG AL, BG, CZ, FR, GR, IT, IT-SIC, LV, MT, PT-AZO, PT-MAD, RO, ES-CAN, GB I2, J1, G References Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) A 1st record in Europe 352 Family species Family species Status Regime Native range 1st record in Europe Invaded countries Habitat Host References stored products Duff (2008), Freude et al. (1967), Šefrova and Lastuvka (2005), Tomov (2009) attic 1956, BG AD, AT, BY, BE, BA, BG, J1 CZ, DK, EE, FI, FR, FRCOR, DE, IT, IT-SAR, LV, LT, MD, NL, NO, PL, PT-AZO, PT-MAD, SK, SI, ES-CAN, SE, CH, UA, GB 1997, IT DE, IT, MT, ES, ES-CAN U dry fruits, nuts Bengtson (1981), Borges et al. (2005), Enckell et al. (1987), Falcoz (1929), Freude et al. (1967), Mendonça and Borges (2009), Tomov (2009) Duff (2008), Falcoz (1929), Freude et al. (1967), Tomov (2009) North America Unknown BY, BE, DK, FR, DE, NL, SE, GB J1 stored products Duff (2008), Falcoz (1929), Freude et al. (1967), Ratti. Coleotteri alieni in Italia.) Africa 1912, GB ES-CAN, GB G, F,I2, J potters bar Duff (2008), Machado and Oromi (2000) C detritivorous Cryptogenic Cryptophagus pilosus Gyllenhal 1828 C detritivorous Cryptogenic Cryptophagus subfumatus Kraatz, 1856 C detritivorous Cryptogenic Curelius japonicus (Reitter, 1877) C detritivorous Cryptogenic Henoticus californicus (Mannhereim 1843) Cybocephalidae Aglyptinus agathidioides Blair 1930 A detritivorous A parasitic/ predator probably a fungus Peck (2009) feeder Coleoptera families other than Cerambycidae, Curculionidae... J1 1900, CZ AL, AT, BY, BE, BA, BG, CZ, DK, EE, FI, FR, DE, IE, IT, IT-SIC, LV, LT, MT, NL, PL, RO, RS, SK, SI, SE, CH, UA, GB 1956, BG BY, BG, FÖ, FR, LV, PTJ1 AZO, PT-MAD Cryptophagus fallax Balfour-Browne, 1953 353 Regime Native range 1st record in Europe A parasitic/ predator Asia2002, IT Temperate A detritivorous Anthrenus caucasicus Reitter, 1881 A Anthrenus flavidus Solsky, 1876 Invaded countries IT Habitat Host References J100 predator of scales Evans et al. (2005), Lupi (2002), Ratti. Coleotteri alieni in Italia.) Australasia 1933, GB FR, NL, GB J1 clothes detritivorous Asia 1941, LV AT, LV, PL J1, I2, E larva scavenger; adult on flowers A detritivorous Asia 1935, PL DE, PL J1, E Anthrenus flavipes LeConte, 1854 C detritivorous Cryptogenic 1955, PL BG, CZ, DK, IT-SAR, ITSIC, PL, CH, GB J1, G Anthrenus oceanicus Fauvel, 1903 A detritivorous Australasia 2004, CZ CZ, MT wood, paper, leather and woven fabrics in collections in museums domestic, feeds on furnitures, fabrics, etc., adult pollinophage; larva necrophagous (faeces, cadavers, pine processionnary nests) stored products J1, E Duff (2008), Freude et al. (1979), Hava (2003), Hava. A Catalogue of World Dermestidae., Reemer (2003) Freude et al. (1979), Hava. A Catalogue of World Dermestidae., Ruta et al. (2004) Freude et al. (1979), Hava. A Catalogue of World Dermestidae.) Duff (2008), Freude et al. (1979), Hava (2003), Hava. A Catalogue of World Dermestidae., Ratti. Coleotteri alieni in Italia., Šefrova and Lastuvka (2005), Tomov (2009), Wittenberg et al. (2006) Hava (2003), Hava. A Catalogue of World Dermestidae., Šefrova and Lastuvka (2005) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Cybocephalus nipponicus Endrody-Younga, 1971 Dermestidae Anthrenocerus australis (Hope, 1843) Status 354 Family species Family species Regime Native range 1st record in Europe Invaded countries Habitat Host References A detritivorous C&S America Unknown DE J stored products C detritivorous Cryptogenic 1927, DE BG, DE, IT, MT, GB J1, J6 Attagenus gobicola Frivaldszky, 1892 A detritivorous AsiaUnknown SE Temperate J necrophagous, in Duff (2008), Freude et al. vegetal (1979), Hava (2003), Ratti. Coleotteri alieni in Italia., Tomov (2009) stored products Hava (2003) Attagenus lynx (Mulsant & Rey, 1868) Attagenus smirnovi Zhantiev, 1973 A detritivorous AsiaUnknown PL Temperate J stored products Hava (2003) C detritivorous Cryptogenic 1973, RU BY, CZ, DK, LV, NO, PL, RU, CH, GB J1 pest of animalorigin material (skin, furs, wool) but also buildings, entomological collections Barsevskis et al. (2004), Duff (2008), Hava (2003), Ruta et al. (2004), Šefrova and Lastuvka (2005) C detritivorous Cryptogenic 1978, GB BG, CZ, DK, LV, PL, CH, GB J1, J6, E domestic, feeds mainly on fabrics, adult pollinophage; larva necrophagous and cereals Borges et al. (2005), Duff (2008), Freude et al. (1979), Hava (2003), Hermann and Baena (2004), Kadej (2005), Tomov (2009), Wittenberg et al. (2006) Attegenus unicolor Brahm 1791 Hava (2003) Coleoptera families other than Cerambycidae, Curculionidae... Attagenus diversepubescens Pic, 1936 Attagenus fasciatus (Thunberg, 1795) Status 355 Status Regime Native range 1st record in Europe Invaded countries Habitat Host References C detritivorous Cryptogenic 1868, GB AT, BG, EE, FR, DE, LT, MT, PL, PT-AZO, ESCAN, CH, GB J1, J6 Dermestes bicolor Fabricius, 1781 A detritivorous Asiatemperate Unknown ES-CAN J Dermestes carnivorus Fabricius, 1775 Dermestes coronatus Steven 1808 Dermestes frischi Kugelann, 1792 A detritivorous C&S America 1919, PL BE, FR, IE, PL, GB J1, J6, G A detritivorous detritivorous Asia Unknown PL E Duff (2008), Freude et al. (1979), Haines and Rees (1989), Hava (2003), Machado and Oromi (2000), Šefrova and Lastuvka (2005), Wittenberg et al. (2006) stored products Freude et al. (1979), Hava (2003), Machado and Oromi (2000) necrophagous in Freude et al. (1979), Haines and houses, bird nests, Rees (1989), Hava. A Catalogue dead fish of World Dermestidae.) grasslands Hava (2003) Cryptogenic 1862, GB BG, DK, EE, FR, IE, LV, LT, PT-AZO, GB J1, J6 domestic C Dermestes lardarius (Linnaeus, 1758) C detritivorous Cryptogenic 1880, BG BG, DK, EE, FR, HU, LT J1, J6 Dermestes leechi Kalík, 1952 A detritivorous Asia Unknown ES, GB J necrophagous Borges et al. (2005), Duff (2008), Freude et al. (1979), Haines and Rees (1989), Hava (2003), Hava. A Catalogue of World Dermestidae., Mendonça and Borges (2009), Tomov (2009) necrophagous but Camerini (2009), Freude et al. in vegetal matters (1979), Haines and Rees (1989), (peanuts, corn), Hava (2003), Hava. A Catalogue eggs predation of World Dermestidae., Tomov (2009) crushed bones Duff (2008), Hava (2003), Hava. A Catalogue of World Dermestidae.) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Dermestes ater De Geer 1774 356 Family species Family species Status Regime Native range 1st record in Europe Invaded countries Habitat C detritivorous Cryptogenic 1871, PL AL, AT, BG, FR, IE, LT, J1, J6 MT, PL, PT, PT-AZO, CH, GB Dermestes peruvianus Laporte de Castelnau, 1840 A detritivorous C&S America 1919, PL AT, CZ, FR, DE, IT, PL, ES-CAN, CH, GB Dermestes vorax Motschulsky, 1860 Novelsis horni (Jayne, 1882) Orphinus fulvipes Guerin-Meneville 1838 Phradonoma tricolor (Arrow, 1915b:431) Reesa vespulae (Milliron, 1939) A detritivorous detritivorous detritivorous AsiaUnknown IT Temperate C&S Unknown NL America Tropical, Unknown FR, GB subtropical J A detritivorous AsiaTropical Unknown DK, NL J A detritivorous North America 1977, GB CZ, DK, EE, FR, DE, IT, LV, NO, SE, CH, GB J1 A A J1, J6, G References domestic, on animal products, fabrics, necrophagous but in vegetal matter(corn kernels) domestic, on animal products, fabrics;, necrophagous but in vegetal matter (corn kernels) detrivorous Borges et al. (2005), Duff (2008), Freude et al. (1979), Haines and Rees (1989), 88180, Wittenberg et al. (2006) J J stored products Freude et al. (1979), Haines and Rees (1989), Hava (2003), Machado and Oromi (2000), Šefrova and Lastuvka (2005) Freude et al. (1979), Hava (2003) Hava (2003), Hava. A Catalogue of World Dermestidae.) Duff (2008), Freude et al. (1979), Hava (2003) Hava (2003), Hava. A Catalogue of World Dermestidae.) domestic places and in museum collections Duff (2008), Freude et al. (1979), Hava (2003), Martinez and Cocquempot (1985), Ratti. Coleotteri alieni in Italia., Šefrova and Lastuvka (2005), Wittenberg et al. (2006) Coleoptera families other than Cerambycidae, Curculionidae... Dermestes maculatus De Geer, 1774 Host 357 Status Regime Native range 1st record in Europe Invaded countries Habitat Host A detritivorous Africa 1998, PL FR, PL J1, J6 Telopes heydeni Reitter 1875 Thaumaglossa rufocapillata Redtenbacher, 1867 Thorictodes heydeni Reitter, 1875 A detritivorous parasitic/ predator Africa Unknown FR J1 Asia, Africa Unknown DE, NL U egg cases of mantids C detritivorous Cryptogenic IT J1 stored seeds, peanuts Thylodrias contractus Motschulsky, 1839 A detritivorous Asia1935, IT Temperate FR, IT, GB J1 animal materials Trogoderma angustum (Solier, 1849) A detritivorous C&S America 1921, PL AT, CZ, DK, DE, LV, LT, PL, SE, CH J1 Trogoderma glabrum (Herbst, 1783) C detritivorous Cryptogenic 1904, BG AT, BG, DK, FR, LV, LT, CH, GB J1 Trogoderma granarium Everts, 1898 A detritivorous Asia 1895, GB AL, AT, BG, CZ, DK, DE, HU, IE, IT, IT-SAR, ITSIC, PL, CH, GB J1 domestic situations and in museum collections domestic situations and in nests of solitary wasps stored products, especially cereals A 1958, IT fish bones, window sills, entomological collections Beal and Kadej (2008), Hava (2003), Hava. A Catalogue of World Dermestidae., Ruta et al. (2004) Freude et al. (1979), Hava (2003) Freude et al. (1979), Hava (2003) Ratti. Coleotteri alieni in Italia., Freude et al. (1979), Hava (2003) Duff (2008), Šefrova and Lastuvka (2005), Freude et al. (1979), Hava (2003) Barsevskis et al. (2004), Freude et al. (1979), Hava (2003), Ruta et al. (2006), Šefrova and Lastuvka (2005), Wittenberg et al. (2006) Duff (2008), Freude et al. (1979), Hava (2003), Tomov (2009), Wittenberg et al. (2006) Duff (2008), Freude et al. (1979), Hava (2003), Šefrova and Lastuvka (2005), Tomov (2009), Wittenberg et al. (2006) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Sefrania bleusei Pic 1899 References 358 Family species Family species Regime Native range 1st record in Europe Invaded countries Habitat Host References detritivorous North America 1956, GB AL, IT, PL, GB J1 psychophage, stored products A detritivorous detritivorous C&S America Asia Unknown FR J stored products 2005, CZ AL, CZ J1 stored products A detritivorous C&S America 1900, CZ AL, AT, CZ, FR, IT, NL, SE J1 Trogoderma variabile Ballion, 1878 A detritivorous Asia 1978, GB CZ, FI, IT, LV, SE, GB J1 Trogoderma versicolor (Creutzer, 1799) Dytiscidae Megadytes costalis Fabricius, 1775 Elateridae Cardiophorus taylori Cobos, 1970 Conoderus posticus (Eschscholtz) C detritivorous Cryptogenic Unknown AT J A parasitic/ predator C&S America Unknown GB U predator A phytophagous phytophagous Africa 1952, DE DE U unknown C&S America Unknown PT-AZO U Chrysanthemoides Borges (1990), Borges et al. monilifera (2005), Mendonça and Borges (2009) A A Duff (2008), Hava (2003), Hermann and Baena (2004), Ratti. Coleotteri alieni in Italia.) Hava (2003) Hava (2003), Hava. A Catalogue of World Dermestidae., Šefrova and Lastuvka (2005) insects in Freude et al. (1979), Hava collection (2003), Ratti. Coleotteri alieni in Italia., Šefrova and Lastuvka (2005) wheat, any dry Duff (2008), Hava (2003), vegetal and Hava. A Catalogue of World animal stored Dermestidae., Šefrova and products in Lastuvka (2005), Ratti. warehouse; major Coleotteri alieni in Italia.) pest eggs predation Camerini (2009), Freude et al. (1979) Duff (2008) 359 A Coleoptera families other than Cerambycidae, Curculionidae... Trogoderma inclusum LeConte, 1854 Trogoderma insulare Chevrolat, 1863 Trogoderma longisetosum Chao & Lee, 1966 Trogoderma megatomoides Reitter, 1881 Status Regime Native range 1st record in Europe Invaded countries Habitat A phytophagous Australasia 1981, GB GB U C detritivorous Cryptogenic 1937, FR, AT, BG, FR, FR-COR, DE, I, J, J6 FR-COR CH C detritivorous Cryptogenic 1843, FR DK, FR J, J6 A detritivorous Asia 1954, AL, CZ, FR, FR-COR, IT, FR-COR PL, ES C parasitic/ predator Cryptogenic Carcinops troglodytes (Paykull, 1811) A parasitic/ predator Chalcionellus decemstriatus Reichardt, 1932 Diplostix mayeti (Marseul, 1870) A A Host References unknown Duff (2008), Freude et al. (1979) on fungus, on decaying plant material, attic flour, dry fruits, medicinal plants, decayed wood Borges et al. (2005), Moncoutier (2002), Shockley et al. (2009a), Tomov (2009) Curtis (1836), Shockley et al. (2009a) J shitake mushrooms Iablokoff-Khnzorian (1975), Šefrova and Lastuvka (2005) 1995, LT AT, BG, DE, LV, LT, PTAZO, CH E cadavers, faeces, Dracunculus C&S America Unknown PT-AZO J parasitic/ predator Africa Unknown FR E predator on Tribolium, Sitophilus in manioc, poultry fly predator feces, cadavers Borges (1990), Borges et al. (2005), Freude et al. (1971), Mendonça and Borges (2009), Tomov (2009), Wittenberg et al. (2006) Borges et al. (2005) parasitic/ predator Africa Unknown FR I2 predator under bark and pods, peanuts, manioc Freude et al. (1971), Gomy (2006), Gomy (2008), Gomy (2009) Delobel and Tran (1993), Yélamos (1992) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Panspaeus guttatus Sharp, 1877 Endomychidae Holoparamecus caularum Aube, 1843 Holoparamecus depressus Curtis, 1833 Erotylidae Dacne picta Crotch, 1873 Histeridae Carcinops pumilio (Erichson, 1834) Status 360 Family species Family species Native range 1st record in Europe Invaded countries Habitat Host References parasitic/ predator parasitic/ predator Africa 1974, FR CY, FR, ES E dung Cryptogenic Unknown IT, PT-AZO B1 cadavers, faeces, sandy soil A detritivorous C&S America Unknown FR, ES, ES-CAN I A detritivorous North America, C&S America 1984, IT H decaying Opuntia Gomy (2008), Machado and in native range; Oromi (2000) straw and manure in invaded area cadavers, faeces Ratti. Coleotteri alieni in Italia.) A unknown Africa Unknown AT, HR, CZ, IT, PT-AZO, ES-CAN Cercyon laminatus Sharp, 1873 A parasitic/ predator Asia1950, Temperate CZ, IT Cercyon nigriceps (Marsham, 1802) A parasitic/ predator Asia? Saprinus lugens Erichson, 1834 Hydrophilidae Cercyon inquinatus Wollaston, 1854 A Regime C HR, FR, IT, IT-SAR, ITSIC, PT, ES U AL, AT, BE, CZ, DK, EE, E3, FI, FR, DE, IT, LT, NL, ES, F9, I SE, CH, GB Unknown CZ, PT-AZO U Mendonça and Borges (2009) decomposing Borges et al. (2005), Boukal et seaweed, rotting al. (2007), Machado and Oromi fruits, cave guano (2000), Ryndevich (2004) compost, predator, In various humid environments; wet grasslands Duff (2008), Freude et al. (1971), Ødegaard and Tømmerås (2000), Ratti. Coleotteri alieni in Italia., Wittenberg et al. (2006) Borges et al. (2005), Boukal et al. (2007), Freude et al. (1971), Mendonça and Borges (2009), Ryndevich (2004) Coleoptera families other than Cerambycidae, Curculionidae... Hister bipunctatus Paykull, 1811 Hypocaccus brasiliensis (Paykull, 1811) Paromalus luderti Marseul, 1862 Status 361 Status Regime Native range 1st record in Europe Invaded countries A parasitic/ predator Asia1950, IT Temperate Dactylosternum abdominale (Fabricius, 1792) A parasitic/ predator Africa Unknown HR, CY, FR, DE, GR, IT, PT-AZO, PT-MAD, ES, ES-CAN Oosternum sharpi Hansen, 1999 Pachysternum capense (Mulsant, 1894) A unknown Unknown PT-AZO A unknown North America Africa A unknown C&S America 1929, IT C detritivorous Cryptogenic 1990, FR AT, BY, CZ, DK, FR, DE, HU, PL C detritivorous, parasitic/ predator Cryptogenic 1875, CZ AT, BY, BE, BG, HR, CZ, J1, G DK, FI, FR, DE, GR, HU, IT-SIC, LV, LT, MT, PL, PT, PT-AZO, PT-MAD, RS, ES, SE, CH, UA, GB Unknown GR, IT, ES-CAN FR, IT Host E3, F9, I References compost, predator, In various humid environments C1+C2 thermophilic, standing water with plants; egg predator on banana weevil in Kenya C1, D in standing water Duff (2008), Freude et al. (1971), Ødegaard and Tømmerås (2000), Šefrova and Lastuvka (2005), Wittenberg et al. (2006) Borges et al. (2005), Machado and Oromi (2000), Mendonça and Borges (2009) J1, G1 under oak bark, stored products Santamaria et al. (1996) stored products, under bark Borges et al. (2005), Duff (2008), Mendonça and Borges (2009), Santamaria et al. (1996), Šefrova and Lastuvka (2005), Tomov (2009), Wittenberg et al. (2006) Borges et al. (2005), Mendonça and Borges (2009), Peck (2009) C1, D in standing water Boukal et al. (2007), Fikacek and Boukal (2004), Machado and Oromi (2000), Ratti. Coleotteri alieni in Italia.) D1-D4 plant held waters, Fikacek and Boukal (2004), ? J? or phytotelmata Sharp (1882–1887) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Cryptopleurum subtile Sharp, 1884 Pelosoma lafertei Mulsant, 1844 Laemophloeidae Cryptolestes duplicatus (Waltl 1834) Cryptolestes ferrugineus (Stephens, 1831) AL, AT, BE, CZ, DK, FI, FR, DE, HU, IT, NL, NO, SE, CH, GB Habitat 362 Family species Family species Status Regime C detritivorous Cryptolestes pusillus (Schönherr, 1817) A detritivorous Cryptolestes spartii (Curtis, 1834) C Cryptolestes turcicus (Grouvelle, 1876) Languriidae Cryptophilus integer (Heer, 1841) Invaded countries Habitat 1978, IT Host References AT, BE, HR, CZ, DK, FI, J FR, DE, GR, HU, IT, ITSIC, MT, PL, PT, PT-MAD, RS, ES, SE, UA, GB Tropical, 1875, CZ AL, AT, BY, BG, CZ, DK, J subtropical FR, DE, IT, IT-SAR, MT, PT-AZO stored products, psychophage/ mills Duff (2008), Ratti. Coleotteri alieni in Italia., Santamaria et al. (1996) synanthropic, grain, damage detritivorous Cryptogenic 1991, FR AL, EE, FR, FR-COR, DE, J1, F PT-AZO, ES, ES-CAN, CH C detritivorous Cryptogenic 1904, FR AL, AT, BE, HR, CZ, DK, J1 FI, FR, DE, GR, HU, IT, IT-SAR, IT-SIC, PL, PT, PT-AZO, PT-MAD, RS, ES, SE, CH, UA, GB corn flour; dry wood (Sarothamnus) dry fruits, grain, wheat, synanthropic Borges et al. (2005), Moncoutier (2002), Santamaria et al. (1996), Šefrova and Lastuvka (2005), Tomov (2009) Santamaria et al. (1996), Wittenberg et al. (2006) C detritivorous Cryptogenic Unknown AT, MT, PT-AZO, CH J1 A detritivorous Asia 1982, DE AT, DK, FR, DE I C detritivorous Cryptogenic 1900, CZ CZ C detritivorous Cryptogenic 1959, CZ CZ, FR, DE, CH, GB Borges et al. (2005), Duff (2008), Santamaria et al. (1996), Šefrova and Lastuvka (2005), Wittenberg et al. (2006) stored products; mycophagous, Vigna hay Borges et al. (2005), Mendonça and Borges (2009), Wittenberg et al. (2006) Callot (2003) J1 psychophage, grain, floour Šefrova and Lastuvka (2005) J1, I Tamarindus seeds, dry fruits, Feeds on fungus, found in herbariium Bouget and Vincent (2008), Duff (2008), Freude et al. (1967), Šefrova and Lastuvka (2005), Wittenberg et al. (2006) 363 Cryptophilus obliteratus Reitter,1874 Pharaxonotha kirschii Reitter, 1875 Latridiidae Adistemia watsoni (Wollaston, 1871) Cryptogenic 1st record in Europe Coleoptera families other than Cerambycidae, Curculionidae... Cryptolestes pusilloides (Steel & Howe, 1952) Native range Status Regime 1st record in Europe Invaded countries A Cartodere constricta (Gyllenhal, 1827) C detritivorous Australasia 2000, DE AT, BE, CZ, DK, FR, DE, NL, PT-MAD, SE, CH, GB C&S 1976, FR FR America Australasia 1850, DE AL, AT, BY, BE, BA, BG, HR, CY, CZ, DK, EE, FI, FR, FR-COR, DE, GR, GR-CRE, HU, IS, IE, IT, IT-SAR, IT-SIC, LV, LI, LT, LU, MT, MD, NL, NO, PL, PT, PT-AZO, PT-MAD, RO, RU, RS, SK, SI, ES, ES-BAL, ES-CAN, SE, CH, UA, GB Crypto1889, GB BY, FR, LV, NO, SE, GB genic Corticaria elongata(Gyllenhal 1827) C detritivorous Cryptogenic Corticaria fenestralis Linneaus, 1758) C detritivorous Cryptogenic A Habitat G, I2 I, J6 I, J6 J1, J6 1889, GB AT, BY, BE, BA, BG, HR, G, I, J CZ, DK, EE, FI, FR, FRCOR, DE, GR, HU, IT, IT-SAR, IT-SIC, LV, LT, LU, MD, ME, NL, NO, PL, PT, PT-AZO, RO, RS, SK, ES, SE, CH, UA, GB 1908, FR AT, BY, BG, FR, DE, CH G, I, J Host References mycophagous, under bark mycophagous, vegetal decay mycophagous, compost, attic, hay Bouget and Vincent (2008), Duff (2008), Reemer (2003) Bouget and Vincent (2008), Vincent (1999) Borges et al. (2005), Bouget and Vincent (2008), Duff (2008), Machado and Oromi (2000), Mendonça and Borges (2009), Tomov (2009) mycophagous, compost, dry fruits, remains, dust forest humus, rotten fruits, hay, firewood Bouget and Vincent (2008), Duff (2008), Telnov (1996) vegetal refuses, hotels, houses, pine bark Bouget and Vincent (2008), Duff (2008) Borges et al. (2005), Bouget and Vincent (2008), Duff (2008), Freude et al. (1967), Mendonça and Borges (2009), Moncoutier (2002), Telnov (1996), Tomov (2009) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Cartodere bifasciata (Reitter, 1877) Cartodere delamarei (Dajoz, 1960) Cartodere nodifer (Westwood, 1839) A detritivorous detritivorous detritivorous Native range 364 Family species Family species Status Regime Native range 1st record in Europe Invaded countries Habitat Host C detritivorous Cryptogenic 1874, FR AT, BY, BG, FR, DE, PTAZO, CH I, J6 Fungi on cacao, spices, cereals, decaying plant material Corticaria pubescens (Gyllenhal, 1827) C detritivorous Cryptogenic 1897, GB AT, BY, FR, DE, HU, LT, CH I, J6 Corticaria serrata (Paykull 1798) C detritivorous Cryptogenic 1997, LT AT, BY, BG, DE, LT, PTAZO, CH Dienerella argus (Reitter, 1884) C detritivorous Cryptogenic 1907, GB FR, LV, GB G mycophagous, mosses, old trees Dienerella costulata (Reitter, 1877) C detritivorous Cryptogenic 1900, CZ CZ, DK, FR J Dienerella filum (Aubé, 1850) C detritivorous Cryptogenic 1850, FR AT, BE, BG, CZ, FR, DE, IE, LV, MT, SE, CH, GB I, J Lathridius australicus Belon, 1887 A detritivorous Australasia Unknown PT-AZO foodstuffs, roots, cellars, appartments cereals, herbaria, yeast, on fungus, on decaying plant material unknown tobacco, medicinal plants, on fungus, on decaying plant material I, J1, J6 on fungus, on decaying plant material, corn, barley U Borges et al. (2005), Bouget and Vincent (2008), Duff (2008), Freude et al. (1967), Mendonça and Borges (2009), Tomov (2009), Wittenberg et al. (2006) Bouget and Vincent (2008), Freude et al. (1967), Wittenberg et al. (2006) Borges et al. (2005), Bouget and Vincent (2008), Freude et al. (1967), Mendonça and Borges (2009), Tomov (2009), Wittenberg et al. (2006) Bouget and Vincent (2008), Duff (2008), Moncoutier (2002), Telnov (1996) Bouget and Vincent (2008), Šefrova and Lastuvka (2005) Bouget and Vincent (2008), Duff (2008), Freude et al. (1967), Moncoutier (2002), Šefrova and Lastuvka (2005), Tomov (2009) Duff (2008), Freude et al. (1967), Mendonça and Borges (2009) Coleoptera families other than Cerambycidae, Curculionidae... Corticaria fulva (Comolli, 1837) References 365 Status Regime Native range 1st record in Europe Invaded countries Habitat C detritivorous Cryptogenic Metophthalmus serripennis Broun 1914 Migneauxia orientalis Reitter, 1877 Lyctidae Lyctus africanus Lesne, 1907 A detritivorous Australasia 1928, DE DE, GB J C detritivorous Cryptogenic 1993, DE AT, DK, FR, DE, PL, CH I, J A phytophagous Africa Unknown AT, FR, CH J1 Lyctus brunneus (Stephens, 1830) A phytophagous Asia 1850, FR AL, AT, BY, BG, CZ, DK, FR, DE, GR, IT, IT-SAR, LV, MT, PT, RS, CH J1 Lyctus cavicollis J. L. LeConte, 1805 Lyctus planicollis J. L. LeConte, 1858 A phytophagous phytophagous North America North America 1996, DE AT, FR, DE, CH J1 1935, FI J1 Lyctus sinensis Lesne, 1911 A phytophagous Asia Unknown GB A 1852, FR AT, BY, BG, FÖ, FR, FRI, J COR, DE, LV, LT, PT-AZO, CH, GB AT, FI, FR J1 cereals/ mills, cellars, attic, on fungus, on decaying plant material References Bengtson (1981), Borges et al. (2005), Bouget and Vincent (2008), Duff (2008), Enckell et al. (1987), Freude et al. (1967), Moncoutier (2002), Tomov (2009), Wittenberg et al. (2006) Duff (2008) fungi on straw, warehouses; dead leaves rice, on fungus, Bouget and Vincent (2008), on decaying plant Wittenberg et al. (2006) material ginger roots; sapwood in field Freude et al. (1969), Ratti. Coleotteri alieni in Italia., Wittenberg et al. (2006) manioc; sapwood Borges et al. (2005), Freude et al. (1969), Glavendekic et al. (2005), Mendonça and Borges (2009), Šefrova and Lastuvka (2005), Wittenberg et al. (2006) wood in houses Ratti. Coleotteri alieni in Italia., Wittenberg et al. (2006) Quercus, Fraxinus Freude et al. (1969), Ratti. (N), wood post in Coleotteri alieni in Italia.) houses timber yards, Duff (2008) rarely in the wild Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Latridius minutus (Linnaeus, 1767) Host 366 Family species Family species Minthea rugicollis (Walker, 1858) Status Regime Native range 1st record in Europe Invaded countries Habitat Host References A phytophagous Tropical, Unknown IT subtropical J1 timber-feeding beetle; attack wide-pored hardwood, broadleaf or coniferous trees and timber with starch levels of greater than 3% (Afzelia, Artocarpus, Avicennia, Bombax, Helicia, Koompassia, Shorea) Mordellidae Mordellistena cattleyana Champion, 1913 A phytophagous C&S America 1921, NL DE, LV, NL J100 Cattleya, Batten (1976), Lima (1955), Vandia, warm Telnov (1996) greenhouses. On flowers of Angelica silvestris in pine forest. Mycetophagidae Litargus balteatus Leconte, 1856 A detritivorous North America 1983, CZ AT, CZ, FR, IT, PT-AZO, CH I, J6 on fungus, on decaying plant material, Maize, dried grapes, stored products Abood and Murphy (2006), Halperin and Geis (1999) Coleoptera families other than Cerambycidae, Curculionidae... Borges et al. (2005), Ratti. Coleotteri alieni in Italia., Šefrova and Lastuvka (2005), Wittenberg et al. (2006) 367 Typhaea stercorea (Linnaeus, 1758) Status Regime Native range 1st record in Europe Invaded countries Habitat Host References detritivorous Cryptogenic 1955, BG AT, BG, FR, DE, IT, ITSAR, IT-SIC, LT, MT, PTAZO, CH I, J, J6 on fungus, on decaying plant material, waste, decay; mills, attic Borges et al. (2005), Freude et al. (1967), Mendonça and Borges (2009), Tomov (2009), Wittenberg et al. (2006) A detritivorous Africa 1999, FR FR I decaying fruits A detritivorous Australasia 2005, PT-AZO J1 stored products; under bark A phytophagous, detritivorous Africa Ratti. Coleotteri alieni in Italia., Mifsud and Audisio (2008), Moncoutier (2001) Audisio (1993), Borges (1990), Borges et al. (2005), Mendonça and Borges (2009) Mifsud and Audisio (2008) Carpophilus dimidiatus (Fabricius, 1792) A phytophagous, detritivorous C&S America Carpophilus freemani Dobson, 1956 A Tropical, 1976, IT subtropical AL, DK, FR-COR, GR, IT, I, J1 IT-SAR, IT-SIC, PT-AZO, ES dry fruits, maize in field Carpophilus fumatus Boheman, 1851 A phytophagous, detritivorous phytophagous, detritivorous Africa AL, IT, IT-SIC, PT, PTAZO Tamarindus seeds, dry fruits, granaries Nitidulidae Brachypeplus deyrollei Murray, 1864 Brachypeplus mauli Gardner & Classey, 1962 Carpophilus bifenestratus Murray, 1864 PT-AZO, PT-MAD 1993, AL, BA, BG, HR, CY, FR, I, J6 FR, FR-COR, GR, IT, IT-SAR, FR-COR IT-SIC, MT, ME, PT-MAD, RS, SI, ES, ES-BAL, ESCAN 1900, CZ AL, AT, BG, CZ, DK, EE, I, J1 FR, FR-COR, IT, IT-SAR, IT-SIC, MT, PL, PT-AZO, ES, CH 1977, IT J1 rotten fruits stored products, corn in fields Audisio (1993), Borges et al. (2005), Mendonça and Borges (2009), Mifsud and Audisio (2008), Moncoutier (2001), Šefrova and Lastuvka (2005), Tomov (2009) Audisio (1993), Borges (1990) Audisio (1993), Mendonça and Borges (2009), Ratti. Coleotteri alieni in Italia., Vieira et al. (2003) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) C 368 Family species Family species Status Regime Native range 1st record in Europe Invaded countries Habitat A phytophagous, detritivorous AsiaTropical 1800, IT AL, AT, BY, BG, CZ, FR, FR-COR, DE, IT, IT-SAR, IT-SIC, LT, MT, PL, PTAZO, ES, CH I, J1 Carpophilus ligneus Murray, 1864 A C&S America 1981, ES-CAN HR, FR, DE, GR, ES-CAN J1 Carpophilus marginellus Motschulsky, 1858 A phytophagous, detritivorous phytophagous, detritivorous AsiaTropical 1938, GB AT, BY, BE, BG, CZ, DK, J1 FI, FR, FR-COR, DE, GR, IT, IT-SAR, IT-SIC, MT, NL, NO, PL, PT-AZO, PT-MAD, ES, ES-CAN, SE, CH, GB Carpophilus mutilatus Erichson, 1843 A phytophagous, detritivorous C&S America 1900, CZ AT, BG, CZ, DK, FR, FRCOR, IT, IT-SAR, IT-SIC, LT, MT, PT-AZO Carpophilus nepos Murray, 1864 A phytophagous, detritivorous Tropical, Unknown AL, BA, BG, HR, CY, FR, J1, I subtropical FR-COR, GR, GR-CRE, GR-ION, GR-NEG, GRSEG, IT-SAR, IT-SIC, MT, PT, PT-AZO, RO, RU, SI, ES, ES-BAL, ES-CAN, UA J1, I decaying grapes, dry fruits, cereals in granaries, fruits on ground, mushrooms maize, dry fruits, granaries References Audisio (1993), Borges et al. (2005), Mendonça and Borges (2009), Mifsud and Audisio (2008), Šefrova and Lastuvka (2005), Tomov (2009), Wittenberg et al. (2006) Audisio (1993), Machado and Oromi (2000) mainly domestic; Audisio (1993), Borges et al. cereals, compost, (2005), Duff (2008), Machado saprophagous and Oromi (2000), Mendonça and Borges (2009), Ødegaard and Tømmerås (2000), Reemer (2003), Šefrova and Lastuvka (2005), Tomov (2009), Wittenberg et al. (2006) dry fruits Audisio (1993), Borges et al. (2005), Mendonça and Borges (2009), Mifsud and Audisio (2008), Šefrova and Lastuvka (2005), Tomov (2009) dry fruits, Borges et al. (2005), Machado outdoors in medi- and Oromi (2000), Mendonça terranean; houses and Borges (2009), Mifsud and in central europe Audisio (2008), Tomov (2009) Coleoptera families other than Cerambycidae, Curculionidae... Carpophilus hemipterus (Linnaeus, 1758) Host 369 Status A Carpophilus pilosellus Motschulsky, 1858 A Carpophilus succisus Erichson, 1843 A Carpophilus zeaphilus Dobson, 1969 A Epuraea luteola Erichson, 1843 A Epuraea ocularis Fairmaire, 1849 Glischrochilus fasciatus (Olivier, 1790) Native range 1st record in Europe Invaded countries Habitat phytophagous, detritivorous phytophagous, detritivorous phytophagous, detritivorous phytophagous, detritivorous detritivorous AsiaTropical 1895, CY, CZ, DK, FR, FR-COR, J1, I GR-CRE GR, GR-CRE, IT, IT-SAR, IT-SIC, MT, PT, ES, AsiaTropical 1983, CZ AT, HR, CZ, FR, IT, ITSAR, IT-SIC, PT-AZO, RS, SI J1, I C&S America 2005, PT-AZO PT-AZO Africa 1985, PT, ES AL, FR, IT, IT-SIC, PT, ES C&S America 1970, AL, FR, IT, IT-SAR, ITES-CAN, SIC, MT, MD, PT-MAD, PT-MAD ES-CAN A detritivorous AsiaTropical 1900, IT A phytophagous, parasitic/ predator North America 1977, DE DE, CH Host References rotten fruits outdoors, granaries (maize, corn) dry fruits, fruits on ground, poultry dung Audisio (1993), Mifsud and Audisio (2008), Šefrova and Lastuvka (2005) J1 maize Borges et al. (2005) J1, I maize Audisio (1993), Ratti. Coleotteri alieni in Italia.) G, I fruits (Prunus), mushrooms AL, AT, FR, DE, IT, IT-SIC, J MD, ES, ES-CAN, CH I Audisio (1993) Audisio (1993), Machado and Oromi (2000), Mifsud and Audisio (2008), Ratti. Coleotteri alieni in Italia., Tomov (2009) mycophagous; Machado and Oromi (2000), manioc, dry fruits Mifsud and Audisio (2008), Ratti. Coleotteri alieni in Italia.) bark beetle Audisio (1993) predator, vegetables, fruits Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Carpophilus obsoletus Erichson, 1843 Regime 370 Family species Family species Status Regime Native range 1st record in Europe Invaded countries Habitat Host phytophagous, parasitic/ predator North America 1950, DE AL, AT, BY, BA, BG, HR, CZ, FR, DE, GR, HU, IT, LI, LT, MD, ME, PL, RO, RU, RS, SK, SI, SE, CH, UA, GB I bark beetle predator, vegetables, fruits Meligethes ruficornis (Marsham, 1802) Nitidula carnaria (Schaller, 1783) C phytophagous detritivorous Cryptogenic Cryptogenic Unknown MT, GB G,I2 Ballota nigra pollen 2005, PT-AZO MT, PT-AZO J1 Omosita colon (Linnaeus, 1758) C detritivorous Cryptogenic 2005, PT-AZO PT-AZO E, G, I, J old bones left on the soil surface Omosita discoidea (Fabricius, 1775) C detritivorous Cryptogenic 2005, PT-AZO PT-AZO E, G, I, J cadavers, carrion Phenolia tibialis (Boheman, 1851) Stelidota geminata (Say, 1825) A Africa 2005, PT-AZO 1900, IT PT-AZO I2 FR, IT, PT-AZO, SI, ESCAN, CH I Urophorus humeralis (Fabricius, 1798) A detritivorous phytophagous, parasitic/ predator detritivorous AL, AT, BA, BG, HR, CY, FR, FR-COR, GR, GRCRE, GR-ION, GR-NEG, GR-SEG, IT, IT-SAR, IT-SIC, MT, ME, PT, PTMAD, RU, RS, SI, ES, ES-BAL, ES-CAN, UA J1 decaying and rotting fruits in insect galleries under oak bark, strawberries and other fruits dry fruits and vegetables C A C&S America AsiaTropical 1976, IT Audisio (1993), Glavendekic et al. (2005), Mendonça and Borges (2009), Ratti. Coleotteri alieni in Italia., Reemer (2003), Šefrova and Lastuvka (2005), Tomov (2009), Wittenberg et al. (2006) Audisio (1993), Duff (2008), Mifsud and Audisio (2008) Audisio (1993), Borges et al. (2005), Mendonça and Borges (2009), Mifsud and Audisio (2008) Audisio (1993), Borges et al. (2005), Mendonça and Borges (2009) Audisio (1993), Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005), Mendonça and Borges (2009) Audisio (1993), Borges et al. (2005), Mendonça and Borges (2009), Ratti. Coleotteri alieni in Italia.) Audisio (1993), Machado and Oromi (2000), Tomov (2009) 371 A Coleoptera families other than Cerambycidae, Curculionidae... Glischrochilus quadrisignatus (Say, 1835) References Passandridae Catogenus rufus (Fabricius, 1798) Status Regime Native range 1st record in Europe Invaded countries Habitat Host References parasitic/ predator North America 2007, AT AT F9 predator of wood- Mitter and Schuh (2008) boring Coleoptera in riverine forest A phytophagous North America Unknown PT-AZO I sweetcorn Borges et al. (2005), Mendonça and Borges (2009) A detritivorous North America 1966, GB DK, DE, NL, NO, SE, GB G, J6 compost Acrotrichis insularis (Maklin, 1852) A detritivorous North America 1965, NO, BG G, J6 compost, saprophagous, fungivore Acrotrichis josephi (Matthews, 1872) A detritivorous North America 1987, GB GB I Acrotrichis sanctaehelenae Johnson, 1972 A detritivorous Africa 1964, ES-CAN I, J6 grass moving; litter, roting organic material anthropogenic habitats, dung, compost, rotting organic substances Duff (2008), Freude et al. (1971), Reemer (2003), Sörensson and Johnson (2004) Borges et al. (2005), Duff (2008), Freude et al. (1971), Freude et al. (1989), Mendonça and Borges (2009), Ødegaard and Tømmerås (2000), Sörensson and Johnson (2004), Wittenberg et al. (2006) Duff (2008), Sörensson and Johnson (2004) Phalacridae Phalacrus politus Melsheimer, 1844 Ptiliidae Acrotrichis henrici (Matthews, 1872) AT, CZ, DK, FI, FR, DE, IE, NL, NO, PT-AZO, PTMAD, SE, CH, GB FR, IT, PT, ES-CAN, CH, GB Duff (2008), Machado and Oromi (2000), Ratti. Coleotteri alieni in Italia., Sörensson and Johnson (2004), Wittenberg et al. (2006) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) A 372 Family species Family species Status Regime Native range 1st record in Europe detritivorous Asia Ptinella cavelli (Broun, 1893) A detritivorous Australasia 1936, GB IE, GB Ptinella errabunda Johnson, 1975 A detritivorous Australasia 1925, GB DE, IE, NL, GB Ptinella simsoni (Matthews, 1878) A detritivorous Australasia 1929, GB GB Ptinella taylorae Johnson, 1977 Bambara contorta (Dybas, 1066) Bambara fusca (Dybas, 1966) Ptinella johnsoni Rutanen, 1985 Ptilodactylidae Ptilodactyla exotica Chapin, 1927 A detritivorous detritivorous detritivorous detritivorous Australasia 1967, GB IE, GB A A A detritivorous AT, BY, CZ, DK, FI, DE, HU, LV, NO, SK, SE Tropical, 1997, DE DE subtropical North 1997, DE DE America Asia 1978, FI, NO, SE FI, SE Africa 1971, IT FR, IT, SI, CH I, J Host compost, saprophagous, fungivore References Freude et al. (1989), Ødegaard and Tømmerås (2000), Ratti. Coleotteri alieni in Italia., Sörensson and Johnson (2004) Sörensson and Johnson (2004) G3, G4 under tight bark of dead broad-leaves and conifers G3 under tight bark of most species of dead trees G,I2 ? heap in crass cuttings in wooded areas around large coastal cities (e.g. London, Liverpool) G3, G4 under tight bark of dead trees E5 forest litter Duff (2008), Sörensson and Johnson (2004) Ryndevich (2004) E5 forest litter Sörensson and Johnson (2004) E5 taiga, litter Sörensson and Johnson (2004) J1, J100 Dracaena in Aberlenc and Allemand (1997), greenhouse; plants Mann (2006), Wittenberg et al. in appartments (2006) Freude et al. (1989), Reemer (2003), Sörensson and Johnson (2004) Sörensson and Johnson (2004) 373 A Habitat Coleoptera families other than Cerambycidae, Curculionidae... Baeocrara japonica (Matthews, 1884) A 1974, FI Invaded countries Regime Native range 1st record in Europe Invaded countries Habitat Host References C detritivorous Cryptogenic 1952, DE DE J100 greenhouse A parasitic/ predator Asia Unknown DK, FI, IT, NL J blatta parasitoid, synanthropic Bétis (1912), Falin (2001), Freude et al. (1969) A phytophagous Asia 2005, PT-AZO PT-AZO I2 polyphagous deciduous Borges et al. (2005), Mendonça and Borges (2009), Paulian and Baraud (1982) Salpingidae Aglenus brunneus (Gyllenhall) C detritivorous Cryptogenic 2005, PT- PT-AZO AZO J1 anthropophilic: attic, stables, poultry, damage cultivated mushrooms; rodent nests in forests Borges et al. (2005) Silvanidae Ahasverus advena (Waltl, 1832) A detritivorous C&S America 1875, CZ AT, BY, BG, CZ, DK, EE, FI, DE, LT, MT, PL, PTAZO, SE, CH I, J1 A detritivorous, parasitic/ predator Tropical, 1911, DE BE, DK, DE, NL, PT-AZO, G, I, J subtropical ES-CAN saprophagousstored products; compost, clethrophage in field banana, ananas; dead plants, bark, cadavers; larva predator Borges et al. (2005), Mendonça and Borges (2009), Ødegaard and Tømmerås (2000), Šefrova and Lastuvka (2005), Tomov (2009), Wittenberg et al. (2006) Borges et al. (2005), Machado and Oromi (2000), Mendonça and Borges (2009), Ratti (2007) Cryptamorpha desjardinsi (GuérinMéneville, 1844) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Ptilodactyla luteipes Pic, 1924 Ripiphoridae Ripidius pectinicornis Thunberg, 1806 Rutelidae Popilia japonica Newman, 1841 Status 374 Family species Family species Status Regime Native range 1st record in Europe Invaded countries Habitat Host C detritivorous Cryptogenic 1906, FR DK, FR, PT-AZO J1 stored products A detritivorous Asia 1980, GB GB J1 coconut, azadirachta seeds A detritivorous Tropical, 1962, CZ AT, BY, BG, CZ, DK, EE, subtropical HU, LV, MT, NO, PT, PTAZO, ES-CAN, CH J1 psychophage, stored products Oryzaephilus surinamensis (Linnaeus, 1758) C detritivorous Cryptogenic J1 psychophage, stored products Silvanus lateritius (Broun, 1880) A detritivorous Australasia Unknown PT-AZO J1 Silvanus lewisi Reitter, 1876 A detritivorous Asia Unknown MT J1, G Silvanus recticollis Reitter, 1876 Staphylinidae Acrotona pseudotenera (Cameron, 1933) A detritivorous Africa Unknown IT-SAR, IT-SIC J1 A parasitic/ predator Asia 1988, FI I 1894, PT AT, BY, BG, CZ, DK, EE, FR, DE, HU, LV, LT, MT, NO, PT, PT-AZO, RS, ESCAN, CH AT, DK, FI, DE, NO, SE, CH rice, manioc, stored products; under bark of dead trees in field Borges et al. (2005), Mendonça and Borges (2009), Ratti (2007), Moncoutier (2002) Duff (2008) Borges et al. (2005), Machado and Oromi (2000), Mendonça and Borges (2009), Šefrova and Lastuvka (2005), Tomov (2009), Wittenberg et al. (2006) Borges et al. (2005), Glavendekic et al. (2005), Machado and Oromi (2000), Mendonça and Borges (2009), Šefrova and Lastuvka (2005), Tomov (2009), Wittenberg et al. (2006) Borges et al. (2005), Mendonça and Borges (2009), Ratti. Coleotteri alieni in Italia.) Ratti (2007), Ratti. Coleotteri alieni in Italia.) Coleoptera families other than Cerambycidae, Curculionidae... Nausibius clavicornis (Kugelann, 1794) Oryzaephilus acuminatus Halstead, 1980 Oryzaephilus mercator (Fauvel, 1889) References Ratti. Coleotteri alieni in Italia.) Luka et al. (2009), Ødegaard and Tømmerås (2000), Wittenberg et al. (2006) 375 compost, predator, fungivorous Status Regime Native range 1st record in Europe Invaded countries Habitat Host A parasitic/ predator North America Unknown GB B Aleochara puberula Klug, 1833 C parasitic/ predator Cryptogenic Unknown AT, PT-AZO I1, J Anotylus nitidifrons (Wollaston, 1871) C parasitic/ predator Cryptogenic Unknown PT-AZO, ES-CAN I Atheta dilutipennis (Motschulsky, 1858) A parasitic/ predator Atheta mucronata (Kraatz, 1859) Bisnius palmi (Smetana, 1955) Bisnius parcus (Sharp, 1874) A parasitic/ predator parasitic/ predator parasitic/ predator Bohemiellina flavipennis (Cameron, 1921) Carpelimus bilineatus Stephens, 1834 C parasitic/ predator Borges et al. (2005), Machado and Oromi (2000), Mendonça and Borges (2009) Africa, Asia 1995, IT AL, IT, PT-AZO, ES-CAN U Borges (1990), Borges et al. (2005), Machado and Oromi (2000), Mendonça and Borges (2009) Tropical, 2002, ES IT, ES I2 decaying vegetals, Gamarra and Outerelo (2005), subtropical citrus groves Monzo et al. (2005) North Unknown AL, CZ, IT, IT-SIC I, J6 Newton. Staphylinini Species America Catalog Draft) Asia1950, FI, AL, AT, DK, FI, FR, DE, I, J6 compost, predator Cho (2008), Duff (2008), Korge Temperate DE IT, NO, ES-CAN, SE, CH, (2005), Luka et al. (2009), GB Ødegaard and Tømmerås (2000), Ratti. Coleotteri alieni in Italia., Tronquet (2006) Crypto1941, AT, BE, DK, FI, FR, DE, B1, E3 compost Ødegaard and Tømmerås (2000), genic FI, DE NO, SE, GB Tronquet (2006) C phytophagous Cryptogenic 2005, PT-AZO B1, E3 grassy coastal patches, sand dunes A PT-AZO Duff (2008) Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) Borges et al. (2005), Mendonça and Borges (2009), Tronquet (2006) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Adota maritima Mannerheim, 1843 A decomposing seaweed, predator flies predator of cyclorrhaphous Diptera (Musca) in stables predator on Delia (carrots) References 376 Family species Family species Status Regime Native range 1st record in Europe Invaded countries Habitat Host References C phytophagous Cryptogenic 2005, PT-AZO PT-AZO B1, E3 floodplains, river banks, sand beaches Borges (1990), Borges et al. (2005), Tronquet (2006) C parasitic/ predator Cryptogenic 2005, PT-AZO PT-AZO Borges et al. (2005), Tronquet (2006) C parasitic/ predator Cryptogenic 2005, PT-AZO PT-AZO C unknown Cryptogenic 2005, PT- PT-AZO AZO Carpelimus zealandicus (Sharp, 1900) Cilea silphoides (Linnaeus, 1767) A unknown Australasia 2000, DE AT, BE, DE, SE, CH, GB B1, E3 floodplains, river banks, sand beaches B1, E3 floodplains, river banks, sand beaches B floodplains, river banks, sand beaches E Sandy banks C parasitic/ predator Cryptogenic 2005 PT-AZO, ES-CAN U A unknown North America Unknown PT-AZO, PT-MAD, ESCAN U A unknown C&S America 1982, IT U Borges et al. (2005), Machado and Oromi (2000), Mendonça and Borges (2009), Tronquet (2006) Borges et al. (2005), Machado and Oromi (2000), Mendonça and Borges (2009) Ratti. Coleotteri alieni in Italia.) A parasitic/ predator Asia Unknown CH U Wittenberg et al. (2006) Coproporus pulchellus (Erichson, 1839) Diestota guadalupensis Pace, 1987 Leptoplectus remyi (Jeannel, 1961) IT cattle dung Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005), Duff (2008), Vorst et al. (2007) Cuppen (2003), Korge (2005), Luka et al. (2009) Coleoptera families other than Cerambycidae, Curculionidae... Carpelimus corticinus (Gravenhorst, 1806) Carpelimus gracilis (Mannerheim, 1830) Carpelimus pusillus (Gravenhorst, 1802) Carpelimus subtilis (Erichson, 1839) 377 Status Regime Native range 1st record in Europe Invaded countries Habitat Host A parasitic/ predator AsiaTropical 1912, CZ AL, AT, BE, CZ, DK, EE, I, J6 FI, FR, DE, HU, IT, LV, NL, NO, PL, PT-AZO, SK, ES, SE, CH, UA, GB Myrmecocephalus concinna (Erichson,1840) Myrmecopora brevipes Butler, 1909 Nacaeus impressicollis (Motschulsky, 1857) C detritivorous Cryptogenic 1970, DE DE, PT-AZO, PT-MAD, RU, ES-CAN, SE, GB G C parasitic/ predator Cryptogenic Unknown FR, IE, GB U A unknown Africa (or Asia?) 2005, PT-AZO I2,G? Oligota parva Kraatz, 1862 A detritivorous C&S America 1858, FR AT, BE, BA, HR, DK, EE, FI, FR, FR-COR, DE, GR, GR-CRE, IT, IT-SIC, NL, NO, PL, PT-AZO, PTMAD, ES-CAN, SE, CH, GB I, J6 compost, predator, fungivorous. Synanthropic Oxytelus migrator Fauvel, 1904 A detritivorous Asia 1975, DK AT, BE, CZ, DK, FR, DE, IT, LT, LU, NO, SE, CH I, J6 compost, saprophagous CZ, PT-AZO compost, predator. Borges et al. (2005), Duff (2008), Freude et al. (1964), Korge (2005), Luka et al. (2009), Ødegaard and Tømmerås (2000), Šefrova and Lastuvka (2005), Tronquet (2006) deadwood Duff (2008), Korge (2005), Machado and Oromi (2000), Tronquet (2006) in wet sand under Anderson (1997), Scheerpeltz plants (1972) Borges et al. (2005), Mendonça and Borges (2009), Rogé (2003), Tronquet (2006) Borges et al. (2005), Freude et al. (1974), Korge (2005), Luka et al. (2009), Machado and Oromi (2000), Mendonça and Borges (2009), Ødegaard and Tømmerås (2000), Reemer (2003), Wittenberg et al. (2006) Korge (2005), Luka et al. (2009), Ratti. Coleotteri alieni in Italia., Šefrova and Lastuvka (2005), Wittenberg et al. (2006) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Lithocharis nigriceps (Kraatz, 1859) References 378 Family species Family species Status Regime Native range 1st record in Europe Invaded countries Habitat Host References A detritivorous Australasia 1988, IT FR, DE, IT, IT-SIC, PT, ES, I, J6 ES-CAN, CH rotting fallen fruits of various trees, decaying vegetals Philonthus rectangulus Sharp, 1874 A parasitic/ predator Asiatemperate Philonthus spinipes Sharp, 1874 A parasitic/ Asia predator, detrivorous 1980, IT AL, AT, BE, BA, BG, HR, I, J6 CZ, DK, EE, FI, FR, DE, GR, HU, IT, IT-SAR, LV, LT, LU, MD, ME, NL, NO, PT, PT-AZO, PT-MAD, RO, RS, SK, SI, ES, ESCAN, SE, CH, UA, GB AL, AT, BG, CZ, DK, FR, J1, J6 IT, LT, RU, CH compost, predator Borges et al. (2005), Coiffait (1972), Korge (2005), Luka et al. (2009), Machado and Oromi (2000), Ødegaard and Tømmerås (2000), Šefrova and Lastuvka (2005), Tomov (2009), Tronquet (2006), Wittenberg et al. (2006) in stable litter, Callot (1993), Luka et al. (2009), cadavers Ratti. Coleotteri alieni in Italia., Šefrova and Lastuvka (2005), Tomov (2009), Tronquet (2006) Tachinus sibiricus Sharp, 1888 Trichiusa immigrata Lohse, 1984 A unknown Asia Unknown AT U A unknown North America 1975, DE AL, AT, BE, CZ, DK, FR, DE, IT, NO, ES-CAN, SE, CH I, I2 compost, predator, fungivorous Teropalpus unicolor (Sharp, 1900) A parasitic/ Australasia Unknown GB predator, detrivorous I2 halophilous 1920, IT Duff (2008), Korge (2005), Luka et al. (2009), Machado and Oromi (2000), Ratti. Coleotteri alieni in Italia., Tronquet (2006), Wittenberg et al. (2006) Korge (2005), Luka et al. (2009), Ødegaard and Tømmerås (2000), Ratti. Coleotteri alieni in Italia., Tronquet (2006), Wittenberg et al. (2006) Duff (2008), Kuschel (1990) Coleoptera families other than Cerambycidae, Curculionidae... Paraphloeostiba gayndahensis (Mac Leay, 1871) 379 Regime Native range 1st record in Europe Invaded countries Habitat A parasitic/ Tropical, 1921, ME AT, BG, DK, EE, FR, FRpredator, subtropical COR, DE, HU, IT, LV, LT, detrivorous MT, ME, NO, ES-CAN, CH, GB J1, J6, G Alphitobius laevigatus (Fabricius, 1781) A detritivorous Tropical, Unknown DK, EE, FR, MT, ES-CAN, J1, J6, subtropical GB G Alphitophagus bifasciatus (Say, 1823) C detritivorous Cryptogenic 1940, BG AL, AT, BG, HR, DK, FI, J1, J6, FR, DE, GR, HU, LT, NO, G RO, SE, CH Cynaeus angustus (Leconte, 1851) A detritivorous C&S America 1988, SE FI, FR, DE, SE J6 Cynaeus depressus Horn, 1870 A detritivorous C&S America 1988, SE SE U Host References minor pest of residues, common inhabitant of chicken houses; feeds on faeces and wastes; outdoors in rotten trunks and bird/ bat nests minor pest of residues; stored products; outdoors on fungi in trunks minor pest of residues; compost, Mainly domestic in rotten fruits; under bark old stumps saprophagous, waste heaps Borges et al. (2005), Duff (2008), Freude et al. (1969), Tomov (2009), Wittenberg et al. (2006) waste heaps Borges et al. (2005), Duff (2008), Freude et al. (1969), Machado and Oromi (2000) Freude et al. (1969), Ødegaard and Tømmerås (2000), Tomov (2009), Wittenberg et al. (2006) Ferrer (2004), Ferrer and Andersson (2002), Reibnitz and Schawaller (2006), Soldati (2007) Ferrer (2004), Mannerkoski and Ferrer (1992) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Tenebrionidae Alphitobius diaperinus (Panzer, 1797) Status 380 Family species Family species Status Regime Native range 1st record in Europe Invaded countries Habitat Host References detritivorous C&S America 1900, CZ AT, CZ, EE, FR, DE, IT, IT- J1 SAR, IT-SIC, LV, MT, PTAZO, ES-CAN, CH, GB cereal grains in warehouses Gnathocerus maxillosus (Fabricius, 1801) Latheticus oryzae Waterhouse, 1880 C detritivorous Cryptogenic 1977, IT AL, FR, FR-COR, IT, ESCAN J1 cereal grains in warehouses A detritivorous Asia 1973, BG, CZ AL, AT, BG, CZ, DK, EE, FR, IT, IT-SIC, RS, ESCAN, CH, GB J1 stored products, cereals in warehouses Lyphia tetraphylla (Fairmaire, 1856) Palorus ratzeburgi (Wissmann, 1848) A detritivorous detritivorous Asia 1934, CZ HR, CZ, FR, GR, ME U Duff (2008), Freude et al. (1969), Glavendekic et al. (2005), Machado and Oromi (2000), Šefrova and Lastuvka (2005), Tomov (2009), Wittenberg et al. (2006) Šefrova and Lastuvka (2005) Africa 1976, LT HR, DK, FR, GR, LT, ESCAN, GB J1 Palorus subdepressus (Wollaston, 1864) A detritivorous Africa 1975, BG BG, HR, CZ, DK, FR, GR, J1 MT, PT-AZO, ES-CAN, GB stored products, mainly cereals; mycophagous stored products, mainly cereals; mycophagous Tribolium castaneum (Herbst, 1797) C detritivorous Cryptogenic 1900, CZ AL, AT, BG, CZ, DK, EE, J1, J2 FR, FR-COR, DE, GR, HU, LV, LT, MT, ME, NO, PT, PT-AZO, RO, ES-CAN, CH, GB Borges et al. (2005), Duff (2008), Freude et al. (1969), Machado and Oromi (2000) Borges et al. (2005), Duff (2008), Freude et al. (1969), Machado and Oromi (2000), Šefrova and Lastuvka (2005), Tomov (2009) Borges et al. (2005), Duff (2008), Freude et al. (1969), Machado and Oromi (2000), Mendonça and Borges (2009), Šefrova and Lastuvka (2005), Tomov (2009), Wittenberg et al. (2006) A stored products Borges et al. (2005), Duff (2008), Freude et al. (1969), Machado and Oromi (2000), Mendonça and Borges (2009), Šefrova and Lastuvka (2005), Wittenberg et al. (2006) Machado and Oromi (2000), Tomov (2009) 381 A Coleoptera families other than Cerambycidae, Curculionidae... Gnathocerus cornutus (Fabricius, 1798) Status Regime Native range 1st record in Europe Invaded countries Habitat Host References A detritivorous Africa 1900, CZ AL, AT, BG, HR, CZ, DK, EE, FR, DE, GR, HU, IT, LV, LT, NO, PT-AZO, ESCAN, CH, GB J1, J2 stored products Tribolium destructor Uyttenboogaart, 1933 A detritivorous tropical 1927, DE AL, AT, BG, CZ, DK, EE, DE, HU, IT, LV, LT, NO, ES-CAN, CH, GB J1, J2 stored products Zophobas morio (Fabricius, 1776) Trogidae Omorgus subcarinatus (MacLeay, 1864) Omorgus suberosus (Fabricius, 1775) Trogossitidae Lophocateres pusillus (Klug, 1832) A detritivorous C&S America Unknown LV J used as food for reptile pets A detritivorous Australasia 1997, ES ES J1, J6 Bercedo (1997) A detritivorous Australasia 1997, ES ES J1, J6 Bercedo (1997) A detritivorous Asia 1962, CZ AL, CZ, DK, IT J1 Tenebroides maroccanus Reitter 1884 A parasitic/ predator Africa 2005, PT-AZO G PT-AZO psychophage, necrophagous; rice, stored products predator egg Lymantria dispar Borges et al. (2005), Duff (2008), Freude et al. (1969), Machado and Oromi (2000), Šefrova and Lastuvka (2005), Tomov (2009), Wittenberg et al. (2006) Duff (2008), Freude et al. (1969), Machado and Oromi (2000), Ratti. Coleotteri alieni in Italia., Šefrova and Lastuvka (2005), Tomov (2009), Wittenberg et al. (2006) Thomas (1995) Šefrova and Lastuvka (2005) Borges et al. (2005) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Tribolium confusum Jacquelin du Val, 1868 382 Family species Family species Tenebroides mauritanicus (Linnaeus, 1758) Regime Native range 1st record in Europe Invaded countries Habitat Host References A detritivorous Africa 1803, PT AT, BG, CZ, DK, EE, DE, IT, LV, LT, PT, PT-AZO, RS, CH J1, G psychophage, carnivorous; stored products, bark in field Borges et al. (2005), Glavendekic et al. (2005), Mendonça and Borges (2009), Šefrova and Lastuvka (2005), Tomov (2009) A unknown Africa Unknown MT U timber Schuh and Mifsud (2000) A unknown Australasia 1962, GB GB B2, I2 C unknown Cryptogenic J100 1901, IT AL, AT, BE, CZ, FR, IT, ES, GB Duff (2008) orchid greenhouses Ratti. Coleotteri alieni in Italia.) Coleoptera families other than Cerambycidae, Curculionidae... Zopheridae Microprius rufulus (Motschulsky, 1863) Pycnomerus fuliginosus Erichson, 1842 Pycnomerus inexpectus (Jaquelin Du Val, 1859) Status 383 Native range 1st record in Europe Invaded countries Habitat wooden furnitures; twigs References Europe Unknown PT-AZO, ES-CAN Mediterranean region Europe Unknown AT, DE, HU, PL, PT- G, J1 MAD, SK, ES-CAN, CH Unknown ES-CAN J1 wood broadleaved trees and furnitures Mediteranean Unknown DK, GB J psychophage; dry roots Duff (2008) unknown Palaearctic Unknown PT-AZO B1 sandy grounds Borges et al. (2005) unknown Europe Unknown PT-AZO U clayey ground detritivorous Europe, Unknown PT-AZO cosmopolitan almost J6 vegetal decay Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005), Mendonça and Borges (2009) phytophagous J Host Borges et al. (2005), Espanol (1992), Machado and Oromi (2000), Mendonça and Borges (2009) De Laclos and Büche (2009), Espanol (1992), Machado and Oromi (2000), Wittenberg et al. (2006) Machado and Oromi (2000) Oligomerus ptilinoides (Wollaston, 1854) Ptinus dubius Sturm, 1837 Sphaericus gibboides (Boieldieu, 1854) Anthicidae Cordicomus instabilis (Schmidt, 1842) Cyclodinus humilis (Germar, 1824) Omonadus formicarius (Goeze, 1777) Aphodiidae Calamosternus granarius (Linnaeus, 1767) Pleurophorus caesus (Creutzer, 1796) detritivorous North Africa, Unknown PT-AZO Europe E dung Borges et al. (2005), Mendonça and Borges (2009) detritivorous Eurasia, north America Unknown PT-AZO E dung Borges et al. (2005), Mendonça and Borges (2009) Buprestidae Agrilus angustulus (Illiger, 1803) phytophagous Europe 2005, PT-AZO G Quercus Borges et al. (2005), Cobos (1986), Freude et al. (1979), Schaefer (1949), Théry (1942) detritivorous detritivorous PT-AZO stored products Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Family Regime Species Anobiidae Anobium punctatum phytoDe Geer, 1774 phagous 384 Table 9.5.2. List and characteristics of the Coleoptera species alien in Europe of families other than Cerambycidae, Curculionidae sensu lato, Chrysomelidae sensu lato and Coccinelidae. Country codes abbreviations refer to ISO 3166 (see Appendix I). Habitat abbreviations refer to EUNIS (see Appendix II). Family Species Buprestis novemmaculata Linnaeus, 1758 Regime phytophagous Native range 1st record Invaded countries in Europe All over 2005, PT-AZO Europe PT-AZO Habitat Host I2 conifers References phytophagous holarctic Unknown GB F4 conifers phytophagous Central & southeast Europe Unknown FÖ E synathropic steppe; feeds on moss (Mnium) parasitic/ predator phytophagous Central Europe Palaearctic 1800, GB G Unknown PT-AZO, ES-CAN E, I Poaceae seeds Amara anthobia A. Villa & G.B. Villa, 1833 phytophagous Unknown GB F4, B1 Poaceae seeds; sandy soils Amara aulicus (Panzer, 1797) phytophagous Mediterranean region, Central Europe Palaearctic Unknown FÖ E, I Bengtson (1981), Enckell et al. (1987) Amara montivaga Sturm, 1825 phytophagous Central Europe, mountains Mediterranean region, Central Europe 1972, IE F4, B1, I compositea & carduaceae seeds, waste lands Poaceae seeds E3, I Apiaceae seeds Anderson et al. (2000), Duff (2008), Borges et al. (2005), Luff (2007), Mendonça and Borges (2009), Valemberg (1997) Melanophila acuminata (De Geer, 1774) Byrrhidae Simplocaria semistriata (Fabricius, 1794) Carabidae Abax parallelus Duftschmid, 1812 Amara aenea (De Geer, 1774) IE Unknown IS, IE, LI, PT-AZO, PT-MAD, GB Duff (2008), Jeannel (1942), Luff (2007), Valemberg (1997) Borges et al. (2005), Machado and Oromi (2000), Mendonça and Borges (2009), Valemberg (1997) Duff (2008), Luff (1998), Luff (2007) Anderson et al. (2000) 385 Anisodactylus parasitic/ binotatus (Fabricius, predator 1787) GB Bengtson (1981), Enckell et al. (1987), Freude et al. (1979) Coleoptera families other than Cerambycidae, Curculionidae... Borges et al. (2005), Cobos (1986), Freude et al. (1979), Mendonça and Borges (2009), Schaefer (1949), Théry (1942) Cobos (1986), Duff (2008), Freude et al. (1979), Schaefer (1949), Théry (1942) Regime Carabus auratus Linnaeus, 1758 Carabus cancellatus Linnaeus, 1758 parasitic/ predator parasitic/ predator Carabus convexus Fabricius, 1775 Carabus nemoralis O.F. Müller, 1764 Demetrias atricapillus (Linnaeus, 1758) Epaphius secalis (Paykull, 1790) parasitic/ predator parasitic/ predator parasitic/ predator Habitat References sandy soil, under felled trunks, bark, tree bases plains, waste lands, predator molluscs dry soil, field, forest edge Duff (2008), Luff (1998), Luff (2007) G forests West Unknown IS Palaearctic Eurosiberian Unknown ES-CAN I2, I1, G Duff (2008), Luff (2007), Turin et al. (2003) Libungan et al. (2008), Turin et al. (2003) Machado and Oromi (2000) parasitic/ predator Eurosiberian Unknown IS F9 Graniger femoralis (Coquerel, 1858) Harpalus distinguendus (Duftschmid, 1812) Leistus rufomarginatus (Duftschmid, 1812) phytophagous phytophagous Spain, Italy, Crimea Mediterranean Unknown ES-CAN H woodlands, fields, gardens in vegetal decays along rivers and bogs, Carex, Oenanthe along rivers, mountains (orophilous) seeds, under stones Machado and Oromi (2000) Unknown ES-CAN I seeds; dry soils, paths, Machado and Oromi (2000), fields, dunes Mendonça and Borges (2009) parasitic/ predator G, I mountains, forests, waste lands Leistus terminatus (Panzer, 1793) Licinus punctatulus (Fabricius, 1792) parasitic/ predator parasitic/ predator Eastern, 1942, GB GB central, western Europe Eurosiberian Unknown IS F9, G osieries Spain, North Unknown PT-AZO, ES-CAN Africa H5 under stones, arid, sandy environments Western Unknown GB Europe Western Unknown GB and Central Europe Eurosiberian 1836, GB GB B1, F9, G Host I1,E, G5 E5 F9, D Duff (2008), Luff (2007), Turin et al. (2003) Duff (2008), Luff (2007), Turin et al. (2003) Duff (2008), Luff (1998), Luff (2007) Borges et al. (2005), Machado and Oromi (2000), Mendonça and Borges (2009), Valemberg (1997) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) parasitic/ predator Native range 1st record Invaded countries in Europe Europe, Asia Unknown GB minor 386 Family Species Callistus lunatus (Fabricius, 1775) parasitic/ predator Native range 1st record Invaded countries in Europe Southern Unknown AT, HU, UA Europe Habitat Host References B, D waste, near littoral, bogs Valemberg (1997) G1 dry soil, under deciduous Duff (2008), Luff (1998), Luff (2007) salty marshes, along Machado and Oromi (2000), Ortuno rivers, lakes and Toribio (2005) humid environments, herbs, along rivers along rivers, coast Borges et al. (2005), Mendonça and Borges (2009) near bogs in forests Anderson et al. (2000) parasitic/ predator Palaearctic 1976, GB parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator Europe, Minor Asia Palaearctic Unknown ES-CAN D6, F9 Unknown IS F9 Europe, North Africa Europe, Asia minor Northern and Central Europe Europe Unknown PT-AZO F9, B 1900, IE IE G, D 1900, GB GB H, G associated with burnt Duff (2008), Luff (1998), Luff (2007) sites 1800, GB GB G, F9 under stones in fresh, humid woods waste in wet grasslands, near bogs in colonies in noncultivated fields parasitic/ predator parasitic/ predator parasitic/ predator GB Europe Unknown PT-AZO E3 Europe 1879, GB E2, I parasitic/ predator West Palaearctic Unknown IE, ES-CAN, GB J2 parasitic/ predator Holarctic Unknown ES-CAN G3 GB cellars, stables Duff (2008), Luff (1998), Luff (2007) Borges et al. (2005), Duff (2008), Mendonça and Borges (2009) Duff (2008), Luff (1998), Luff (2007) Anderson et al. (2000), Duff (2008), Machado and Oromi (2000), Luff (1998), Luff (2007), Valemberg (1997) under humid bark, in Machado and Oromi (2000) bark beetle galleries in Abies and Cedrus 387 Tachyta nana (Gyllenhal, 1810) Regime Coleoptera families other than Cerambycidae, Curculionidae... Family Species Lymnastis galilaeus Piochard de la Brûlerie, 1876 Microlestes minutulus (Goeze, 1777) Notaphus varius (Olivier, 1795) Ocydromus tetracolus (Say, 1823) Paranchus albipes (Fabricius, 1796) Philochthus guttula (Fabricius, 1792) Pterostichus angustatus (Duftschmid, 1812) Pterostichus cristatus (Dufour, 1820) Pterostichus vernalis (Panzer, 1796) Scybalicus oblongiusculus (Dejean, 1829) Sphodrus leucophthalmus (Linnaeus, 1758) parasitic/ predator phytophagous Native range 1st record Invaded countries in Europe 1940, IE IE, GB Spain, North Unknown ES-CAN Africa, Crimea detritivorous southern Europe, Minor Asia parasitic/ predator Medi1990, CZ CZ terranean Region Europe, Unknown PT-AZO North Africa Europe, Unknown PT-AZO North Africa Habitat J6, J2, I2 Host References near littoral; in compost in Ireland mountains under stones, arid, sandy environments; granivore Duff (2008), Anderson et al. (2000), Luff (1998), Luff (2007) Machado and Oromi (2000) in hollow Malus, debris in rotten stump, in moss among rotten logs Duff (2008) J6 predatory Freude et al. (1979), Šefrova and Lastuvka (2005) J buildings, prey anobiids timber, prey larvae anobiids, buildings Borges et al. (2005), Freude et al. (1979) Borges et al. (2005), Freude et al. (1979) H5 Unknown AL, DK, DE, HU, G IE, NL, SE, CH, GB Cleridae Enoplium serraticorne (Olivier, 1790) Opilo domesticus (Sturm, 1837) Opilo mollis (Linnaeus, 1758) Corylophidae Sericoderus lateralis (Gyllenhal, 1827) detritivorous palaearctic Unknown PT-AZO I, J1 moldy plant remains in warm places, especially garden compost and grass cuttings Borges et al. (2005), Bowestead (1999), Mendonça and Borges (2009) Cryptophagidae Atomaria apicalis Erichson, 1846 detritivorous Europe Unknown FÖ, PT-AZO J6 mycophage Atomaria bella Reitter, 1875 detritivorous Europe, north Africa 1967, GB G3 mycophage Bengtson (1981), Borges et al. (2005), Enckell et al. (1987), Falcoz (1929), Freude et al. (1967) Duff (2008) parasitic/ predator parasitic/ predator GB J Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Clambidae Clambus pallidulus Reitter, 1911 Regime 388 Family Species Trechus subnotatus Dejean, 1831 Tschitscherinellus cordatus (Dejean, 1825) Regime Atomaria testacea Stephens, 1830 Atomaria turgida Erichson, 1846 detritivorous detritivorous Cryptophagus dentatus (Herbst, 1793) Cryptophagus distinguendus Sturm 1845 detritivorous detritivorous detritivorous detritivorous detritivorous U Europe Unknown GB U Northern Europe Central Europe, Spain Eurasia Unknown GB U 1976, GB G3 Europe, north Africa Central, Northern Europe Europe, north Africa Central, southern Europe Europe Northern, Central Europe Palaearctic GB Habitat Host References mycophage; also adults damaging beet mycophage; also adults damaging beet mycophage Duff (2008), Falcoz (1929) Duff (2008) Duff (2008), Falcoz (1929), Freude et al. (1967) Duff (2008), Falcoz (1929) Unknown PT-AZO, GB J1 rotten wood debris abroad; mainly conifer forest attic Unknown GB J1 mycophage Borges et al. (2005), Falcoz (1929), Freude et al. (1967) Duff (2008), Falcoz (1929) Unknown GB J1 mycophage Duff (2008) Unknown IE, GB J2, I2 mycophage Unknown IE, GB J1 mycophage Duff (2008), Falcoz (1929), Freude et al. (1967) Duff (2008) Unknown GB J1 mycophage Duff (2008), Falcoz (1929) Duff (2008), Falcoz (1929), Freude et al. (1967) 1996, IE, GB IE, GB G3 mycophage 1937, PT-MAD PT-AZO, PT-MAD J1 flour, dry fruits Europe, Asia, Unknown FÖ Africa J1 Borges et al. (2005), Duff (2008), Falcoz (1929), Freude et al. (1967), Mendonça and Borges (2009) mills, stored products Bengtson (1981), Enckell et al. (1987), Falcoz (1929), Freude et al. (1967) 389 detritivorous detritivorous detritivorous detritivorous detritivorous detritivorous Native range 1st record Invaded countries in Europe Europe Unknown GB Coleoptera families other than Cerambycidae, Curculionidae... Family Species Atomaria fuscata (Schönherr, 1808) Atomaria fuscipes (Gyllenhal, 1808) Atomaria hislopi Wollaston, 1857 Atomaria lohsei Johnson & Strand, 1968 Atomaria munda Erichson, 1846 Atomaria nitidula Marsham, 1802 Atomaria punctithorax Reitter, 1887 Atomaria pusilla (Paykull, 1798) Atomaria strandi Johnson, 1967 Regime detritivorous Native range 1st record Invaded countries in Europe Europe, Unknown PT-AZO North Africa Habitat Host J1 attic References detritivorous Europe Unknown FÖ J1 grain, dry fruits detritivorous Eurasia Unknown PT-AZO J1 mammals and Vespa nests detritivorous Europe Unknown IE, PT-AZO, GB G1 ground, salix basis southern Europe, Minor Asia Mediterranean region Europe Unknown DK J1, E stored products Unknown CH, GB J1 domestic Duff (2008), Freude et al. (1979) Unknown IE, GB J1, E5, I2 animal materials Freude et al. (1979) Unknown CH J1 domestic Freude et al. (1979), Wittenberg et al. (2006) Unknown CH J1 domestic Wittenberg et al. (2006) Dermestidae Attagenus bifasciatus detriti(Olivier, 1790) vorous Attegenus brunneus Faldermann, 1835 detritivorous Attegenus pellio Linnaeus, 1758 Attagenus quadrimaculatus Kraatz, 1858 Attagenus rossi Ganglbauer, 1904 detritivorous detritivorous detritivorous southern Europe, Minor Asia Cosmopolitan (native? Europe, Africa, USSR) Borges et al. (2005), Duff (2008), Falcoz (1929), Mendonça and Borges (2009) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Borges (1990), Borges et al. (2005), Falcoz (1929), Freude et al. (1967), Mendonça and Borges (2009) Bengtson (1981), Enckell et al. (1987), Falcoz (1929), Freude et al. (1967) Borges et al. (2005), Falcoz (1929), Freude et al. (1967) 390 Family Species Cryptophagus saginatus Sturm, 1845 Cryptophagus scanicus (Linnaeus, 1758) Cryptophagus schmidti Sturm, 1845 Ephistemus globulus Paykull, 1798 Regime detritivorous detritivorous Anthrenus coloratus Reitter, 1881 detritivorous Anthrenus festivus Erichson, 1846 detritivorous Anthrenus museorum (Linnaeus, 1761) Anthrenus olgae Kalik, 1946 Dermestes murinus Linnaeus, 1758 Dermestes undulatus Brahm, 1790 detritivorous detritivorous detritivorous detritivorous Derodontidae Laricobius erichsonii parasitic/ Rosenhauer, 1846 predator phytophagous Melanotus dichrous (Erichson, 1841) phytophagous Invaded countries Habitat Host References SE J stored products DE, GB J stored products AT, GB J1, E skins, stuffed animals Duff (2008), Freude et al. (1979) AT, CH J1, E insects in collection; adults on flowers Freude et al. (1979), Wittenberg et al. (2006) PT-AZO J1, E insects in collection Borges et al. (2005), Freude et al. (1979) Duff (2008), Freude et al. (1979) Freude et al. (1979), Hermann and Baena (2004) Central Europe Europe Unknown AT, GB J1, E stored products Unknown PT-AZO, ES-CAN J Holarctic Unknown LV, PT-AZO, ESCAN J domestic on animal products domestic on animal products Borges et al. (2005), Freude et al. (1979), Machado and Oromi (2000) Borges et al. (2005), Freude et al. (1979), Machado and Oromi (2000), Mendonça and Borges (2009) G3 aphid predator Franz (1958), Freude et al. (1979) Unknown PT-AZO E5 roots cereals, potato Borges et al. (2005), Laibner (2000), Leseigneur (1972) Unknown PT-AZO F5 shrubs Borges et al. (2005), Leseigneur (1972), Mendonça and Borges (2009) europe 1971, GB (imported to USA) Western, central, Northern Europe southern Europe GB 391 Elateridae Athous haemorrhoidalis (Fabricius, 1801) Native range 1st record in Europe North Africa, Unknown Italy Unknown Mediterranean region East Medi1983, GB terranean region MediUnknown terranean region Holarctic Unknown Coleoptera families other than Cerambycidae, Curculionidae... Family Species Attagenus simplex Reitter, 1881 Attagenus trifasciatus (Fabricius, 1787) Saprinus acuminatus (Fabricius, 1798) Saprinus caerulescens (Hoffmann, 1803) Saprinus planiusculus Motschulsky, 1849 Saprinus semistriatus (Scriba, 1790) Saprinus subnitescens Bickhardt, 1909 Hydrophilidae Cercyon depressus Stephens, 1829 Cercyon haemorhoidalis (Fabricius, 1775) Native range 1st record in Europe Invaded countries References Palaearctic parasitic/ predator parasitic/ predator europe south Unknown PT-AZO U Mediterranean Region Mediterranean Region eurocentrosasiatic Europe Unknown PT-AZO B1 Unknown DK B1 cow dung, nr litoral Mazur (1989) Unknown PT-AZO U Unknown PT-AZO U Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005) palaearctic Unknown PT-AZO B fish decaying, cadavers, feces, Arum fish decaying, cadavers, feces, Arum fish decaying, cadavers, feces, Arum Borges et al. (2005), Mendonça and Borges (2009) palaearctic Unknown PT-AZO B Europe Unknown PT-AZO B fish decaying, cadavers, feces, Arum fish decaying, cadavers, feces, Arum Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005), Mendonça and Borges (2009) Northern, Central Europe Europe Unknown PT-AZO B rotting seaweed on seashores Borges et al. (2005), Mendonça and Borges (2009) Unknown PT-AZO J6 decaying organic matter, flood debris Borges et al. (2005), Mendonça and Borges (2009) parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator detritivorous parasitic/ predator parasitic/ predator E Host parasitic/ predator parasitic/ predator Unknown PT-AZO, ES-CAN Habitat cow, horse dung Borges et al. (2005), Machado and Oromi (2000), Mendonça and Borges (2009) Borges et al. (2005), Mendonça and Borges (2009) cadavers, feces, vegetal Borges et al. (2005), Mendonça and decays, sandy soil Borges (2009) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Halacritus punctum (Aube, 1843) Hypocaccus dimidiatus (Illiger, 1807) Macrolister major (Linnaeus, 1767) Regime 392 Family Species Histeridae Acritus nigricornis (Hoffmann, 1803) Family Species Cercyon obsoletus (Gyllenhal, 1808) Sphaeridium bipustulatum Fabricius, 1781 U unknown Europe Unknown PT-AZO U unknown All over Europe central, western, southern Europe Western, Central Europe Unknown PT-AZO B halophil Unknown PT-AZO C1, D in standing waters Unknown PT-AZO E parasitic/ predator parasitic/ predator, phytophagous parasitic/ predator Habitat Host mainly in dung of larger herbivores, but also recorded from arrion and manure References Vorst (2009) Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005) parasitic/ predator Eurasia Unknown PT-AZO E mammal dung, Borges et al. (2005), Mendonça and decaying organic Borges (2009) matter, fungi, and on plant sap dung Borges et al. (2005) phytophagous Mediterranean Region West Mediterranean Region 1926, GB BE, LI, LU, NL, GB E, I2 Antirrhinum, Linaria Audisio (1993), Borges et al. (2005), Duff (2008) 1929, GB AT, BE, CZ, DE, LI, CH, GB E, I2 Antirrhinum, Linaria Audisio (1993), Duff (2008), Šefrova and Lastuvka (2005) 1962, CZ AL, AT, BE, CZ, DK, J1 FI, DE, HU, PL, SE, UA, GB grain and grain products, nuts, oilseeds, dried root crops Borges et al. (2005), Duff (2008), Šefrova and Lastuvka (2005) phytophagous detritivorous Mediterranean Region 393 Sphaeridium scarabaeoides (Linnaeus, 1758) Kateretidae Brachypterolus antirrhini (Murray, 1864) Brachypterolus vestitus (Kiesenwetter, 1850) Laemophloeidae Cryptolestes capensis (Waltl, 1834) Native range 1st record Invaded countries in Europe Northern, Unknown PT-AZO Central Europe Coleoptera families other than Cerambycidae, Curculionidae... Cercyon quisquilius (Linnaeus ,1761) Enochrus bicolor (Fabricius, 1792) Helochares lividus (Forster, 1771) Regime Dienerella ruficollis (Marsham, 1802) detritivorous Thes bergrothi (Reitter, 1880) detritivorous Leiodidae Catops fuliginosus Erichson 1837 detritivorous Western, Central, Southern Europe parasitic/ predator, phytophagous Malachiidae Axinotarsus marginalis (Laporte de Castelnau, 1840) Monotomidae Monotoma bicolor A. Villa & G. B. Villa, 1835 Native range 1st record in Europe Europe Invaded countries Unknown PT-AZO, PT-MAD Habitat Host References FB under populus bark Borges et al. (2005), Rücker (1995) G3 conifer specialist (douglas-fir, abies) Freude et al. (1967) J1 dry plants, flour Borges et al. (2005), Bouget and Vincent (2008), Duff (2008) I, J on fungus, on decaying plant material, attic; flour, dattes Duff (2008) Unknown FÖ F fungi Bengtson (1981), Duff (2008) Eurasia Unknown IT-SAR E adult floricolous, parasite Acrididae detritivorous Europe Unknown GB G saproxilic/ woodland Duff (2008) detritivorous Europe 2005, PT-AZO E, J mole nest, vegetal waste Borges et al. (2005) Unknown PL, GB Central northern Europe Medi1889, GB DE, IT-SIC, PTterranean AZO, GB region northeastern Unknown GB Europe PT-AZO Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) detritivorous detritivorous Meloidae Mylabris variabilis (Pallas, 1781) Regime 394 Family Species Latridiidae Cartodere norvegica (Strand, 1940) Corticaria abietorum Motschulsky, 1867 Family Species Monotoma longicollis (Gyllenhal, 1827) Monotoma picipes Herbst, 1793 Regime detritivorous detritivorous Native range 1st record Invaded countries in Europe Europe 2005, PT-AZO PT-AZO Europe 2005, PT-AZO PT-AZO Habitat Host vegetal waste J, J6 saprophage/ mycophage; vegetal waste decaying grains detritivorous Eurasia 2005, PT-AZO PT-AZO J, J6 detritivorous parasitic/ predator Europe 2005, PT-AZO 1983, GB PT-AZO J GB Mycetophagidae Berginus tamarisci Wollaston, 1854 detritivorous Eulagius filicornis (Reitter, 1887) detritivorous Nitidulidae Carpophilus quadrisignatus Erichson, 1843 Epuraea aestiva (Linnaeus, 1758) Epuraea biguttata (Thunberg, 1784) phytophagous, detrivorous detritivorous detritivorous Borges et al. (2005) G3 paddy residues, paddy storage predator Dendroctonus- Picea stands Borges et al. (2005), Mendonça and Borges (2009) Bouget and Moncoutier (2003), Duff (2008) southern Unknown AT, CH Europe, Canary Isls southern 1993, GB GB France, North Africa G3 Tamarix, on pine Borges et al. (2005), Freude et al. (1967) G3 with the fungus Stereum hirsutum growing on dead branches of broadleaved trees. Duff (2008) Medi2000, DE terranean region Europe, Asia 2005, PT-AZO Northern 2005, Europe PT-AZO AT, DE, PT-AZO J1 dry fruits PT-AZO G, I Audisio (1993), Borges et al. (2005), Freude et al. (1967), Mendonça and Borges (2009) Audisio (1993), Borges et al. (2005) PT-AZO J1, I mushrooms Audisio (1993), Borges et al. (2005), Freude et al. (1967), Mendonça and Borges (2009) Europe 395 Monotoma quadrifoveolata Aube, 1837 Monotoma spinicollis Aubé, 1837 Rhizophagus grandis Gyllenhal, 1827 Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005), Mendonça and Borges (2009) Coleoptera families other than Cerambycidae, Curculionidae... J, J6 References Native range 1st record Invaded countries in Europe Eurasia 2005, PT-AZO PT-AZO Meligethes aeneus (Fabricius, 1775) phytophagous Europe Meligethes incanus Sturm, 1845 Nitidula flavomaculata Rossi, 1790 Pocadius adustus Reitter, 1888 Oedemeridae Nacerdes melanura (Linnaeus, 1758) Phalacridae Phalacrus corruscus (Panzer, 1797) phytophagous detritivorous Southeastern 1867, Europe PT-AZO southern 1900, CZ Europe detritivorous Eurasia detritivorous Ptiliidae Acrotrichis cognata (Matthews, 1877) Habitat References PT-AZO, ES-CAN I1 PT-AZO, GB FA, E5 Audisio (1993), Borges (1990), Borges et al. (2005), Mendonça and Borges (2009) rape, rosaceae, pollen- Audisio (1993), Borges et al. (2005), feeding Duff (2008), Freude et al. (1967), Machado and Oromi (2000), Mendonça and Borges (2009) Nepeta cataria Audisio (1993), Borges et al. (2005) CZ J1, J6 bones vertebrates 2004, GB GB E2 epigeous gastermyctes Audisio (1993), Duff (2008) specialist Europe 2005, PT-AZO PT-AZO B driftwood on beaches, Borges et al. (2005), Mendonça and moist wood Borges (2009) phytophagous Europe Unknown PT-AZO I seeds of yellow sowthistle Sonchus arvensis Borges et al. (2005) detritivorous Europe 1932, SE E5, J6 Duff (2008), Freude et al. (1971) detritivorous detritivorous Europe Unknown PT-AZO U dung, rotting fungi, carcasses, compost near forests unknown Europe Unknown PT-AZO U detritivorous Europe 2005, PT-AZO E 2005, PT-AZO AT, DK, FI, DE, IE, NL, NO, SE, GB PT-AZO J Host dung Audisio (1993), Freude et al. (1967), Šefrova and Lastuvka (2005) Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005), Mendonça and Borges (2009) Baraud (1992), Borges et al. (2005), Bunalski (1999), Mendonça and Borges (2009) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) detritivorous Actinopteryx fucicola (Allibert, 1844) Ptenidium pusillum (Gyllenhal, 1808) Scarabaeidae Onthophagus illyricus (Scopoli, 1763) Regime 396 Family Species Epuraea longula Erichson, 1845 Native range 1st record Invaded countries in Europe Europe 2005, PT-AZO PT-AZO Onthophagus vacca (Linnaeus, 1767) detritivorous Europe 2005, PT-AZO Oryctes nasicornis (Linnaeus, 1758) Scydmaenidae Stenichnus collaris (Muller & Kunze, 1822) Silphidae Ablattaria laevigata (Fabricius, 1775) detritivorous southern Europe detritivorous parasitic/ predator Aclypea opaca (Linnaeus, 1758) phytophagous Host E dung E dung 1880, DK DK, FI, HU, LT, NO, SE J saprophagous, compost Europe Unknown FÖ I2 mosses, leaves Western & southcentral Europe Central, Northern, Eastern Europe Unknown EE E, I1 snail predator, fields 2005, PT-AZO E, I1 chenopodiacées PT-AZO PT-AZO References Baraud (1992), Borges et al. (2005), Bunalski (1999), Mendonça and Borges (2009) Baraud (1992), Borges et al. (2005), Bunalski (1999), Mendonça and Borges (2009) Baraud (1992), Bunalski (1999) Bengtson (1981) Borges et al. (2005) detritivorous Europe Unknown PT-AZO J1 detritivorous europe 2005, PT-AZO U mycophage parasitic/ predator Palaearctic Unknown PT-AZO I solitary ectoparasitoids Borges et al. (2005), Freude et al. of cyclorrhaphous (1974), Mendonça and Borges (2009) Diptera (Delia) PT-AZO Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005), Freude et al. (1967), Mendonça and Borges (2009) 397 Silvanidae Silvanus unidentatus (Olivier, 1790) Sphindidae Sphindus dubius (Gyllenhal, 1808) Staphylinidae Aleochara bipustulata (Linnaeus, 1761) Habitat Coleoptera families other than Cerambycidae, Curculionidae... Family Regime Species Onthophagus taurus detriti(Schreber, 1759) vorous Regime parasitic/ predator Habitat I1, J6 Europe Unknown FÖ I1, J Amischa analis parasitic/ (Gravenhorst, 1802) predator Italy Unknown PT-AZO U Anotylus nitidulus (Gravenhorst 1802) Anotylus speculifrons (Kraatz 1857) parasitic/ predator parasitic/ predator 2005, PT-AZO 2005, PT-AZO PT-AZO U PT-AZO U Atheta acuticollis Fauvel, 1907 Atheta amicula (Stephens,1832) parasitic/ predator parasitic/ predator Europe, cosmopolitan Europe, Asia Minor, North Africa palaearctic PT-AZO U PT-AZO, PT-MAD, ES-CAN U Europe 2005, PT-AZO 2005, PT-AZO Atheta atramentaria parasitic/ (Gyllenhal,1810) predator Europe Unknown PT-AZO, PT-MAD, ES-CAN U Atheta castanoptera (Mannerheim, 1830) Atheta coriaria (Kraatz, 1858) parasitic/ predator Europe 2005, PT-AZO PT-AZO U parasitic/ predator Europe 2005, PT-AZO PT-AZO, ES-CAN U Host References feed on decaying meat, fly maggots and also on fly puparia predator of cyclorrhaphous Diptera (Musca) in stables Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) Bengtson (1981), Enckell et al. (1987), Freude et al. (1974) Borges (1990), Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005), Mendonça and Borges (2009) Borges (1990), Borges et al. (2005), Freude et al. (1974), Machado and Oromi (2000) Borges (1990), Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) predator, biological control soil-dwelling larvae of small Diptera Borges et al. (2005), Freude et al. (1974), Machado and Oromi (2000), Mendonça and Borges (2009) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) parasitic/ predator Native range 1st record Invaded countries in Europe Palaearctic 2005, PT-AZO PT-AZO 398 Family Species Aleochara clavicornis Redtenbacher, L., 1849 Aleochara sparsa Heer, 1839 Family Species Atheta divisa (Maerkel, 1844) Regime parasitic/ predator Native range 1st record Invaded countries in Europe Europe 2005, PT-AZO PT-AZO Habitat U Europe Unknown FÖ, PT-AZO, PTMAD, ES-CAN I1 Atheta gregaria (Casey, 1910) Atheta harwoodi Williams, 1930 parasitic/ predator parasitic/ predator europe Unknown FÖ U europe Unknown FÖ, GB J6 Atheta luridipennis (Mannerheim, 1830) Atheta nigra (Kraatz,1856) parasitic/ predator 2003, ES FÖ, PT-AZO, ES C3 parasitic/ predator Central, Northern Europe Europe 2005, PT-AZO PT-AZO, ES-CAN U Atheta nigricornis (Thomson,1852) Atheta oblita (Erichson,1839) Atheta palustris (Kiesenwetter,1844) Atheta sordida Marsham,1802 parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator Atheta triangulum (Kraatz,1856) parasitic/ predator Northern Europe Northern Europe Morocco, France Italy southern Europe, Minor Asia Europe Unknown FÖ U 2005, PT-AZO 2005, PT-AZO 2005, PT-AZO PT-AZO U PT-AZO, PT-MAD U PT-AZO, PT-MAD, ES-CAN U PT-AZO U 2005, PT-AZO References bird and animal nest Borges (1990), Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) predator, carrot fields Bengtson (1981), Borges et al. (2005), Enckell et al. (1987), Freude et al. (1974), Machado and Oromi (2000), Mendonça and Borges (2009) Bengtson (1981), Enckell et al. (1987), Freude et al. (1974) bird nest, compost Bengtson (1981), Duff (2008), Enckell et al. (1987), Freude et al. (1974) streambanks Bengtson (1981), Borges et al. (2005), Enckell et al. (1987), Freude et al. (1974), Mendonça and Borges (2009) Borges et al. (2005), Freude et al. (1974), Machado and Oromi (2000), Mendonça and Borges (2009) fungi Meripilus Bengtson (1981), Enckell et al. giganteus (1987), Freude et al. (1974) Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) Borges (1990), Freude et al. (1974), Mendonça and Borges (2009) Coleoptera families other than Cerambycidae, Curculionidae... Atheta fungi parasitic/ (Gravenhorst,1806) predator Host Borges et al. (2005), Freude et al. (1974) 399 Creophilus maxillosus (Linnaeus, 1758) Cypha pulicaria (Erichson,1839) Edaphus beszedesi Reitter, 1914 Euplectus infirmus Raffray, 1910 Gabrius nigritulus (Gravenhorst, 1802) Gabronthus thermarum (Aubé, 1850) Gyrophaena bihamata Thomson,1867 Gyrophinus fracticornis (O. Müller, 1776) parasitic/ predator parasitic/ predator unknown unknown unknown unknown unknown detritivorous unknown unknown parasitic/ predator unknown unknown Native range 1st record Invaded countries in Europe europe Unknown FÖ, PT-MAD, ESCAN Europe, Asia, North Africa Minor Asia, Bulgaria Europe, Asia, Africa Northern Europe 2005, PT-AZO PT-AZO Unknown DK 2005, PT-AZO PT-AZO Unknown PT-AZO, PT-MAD, ES-CAN Europe (intro 2005, NAm) PT-AZO Europe 2005, PT-AZO southern Unknown Europe Southern 2005, Europe PT-AZO Eurasia 2005, PT-AZO Europe 2005, PT-AZO Central, Northern Europe euroMediterranean Habitat Host U I, J6 compost, predator References Bengtson (1981), Enckell et al. (1987), Freude et al. (1974), Machado and Oromi (2000) Borges et al. (2005), Mendonça and Borges (2009) U U U PT-AZO U PT-AZO U AT, EE, CH J6 PT-AZO U PT-AZO U PT-AZO I, J6 2005, PT-AZO PT-AZO U 2005, PT-AZO PT-AZO J6 Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005), Freude et al. (1974), Machado and Oromi (2000), Mendonça and Borges (2009) Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) compost, rotting plant Luka et al. (2009), Wittenberg et al. material (2006) Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) compost, predator Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) waste, decay Borges et al. (2005), Mendonça and Borges (2009) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Bisnius sordidus (Gravenhorst, 1802) Brachygluta paludosa (Peyron, 1858) Cafius xantholoma (Gravenhorst, 1806) Cordalia obscura (Gravenhorst,1802) Regime 400 Family Species Atheta trinotata (Kraatz,1856) Habitat G, J6 Host humus References Duff (2008) 2005, PT-AZO PT-AZO U Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) 2005, PT-AZO PT-AZO, PT-MAD, ES-CAN U Northern and Central Europe, siberia Europe (introAF, AUS) Eurasia Unknown FÖ D Borges et al. (2005), Freude et al. (1974), Machado and Oromi (2000), Mendonça and Borges (2009) bogs, mires, wet fields Bengtson (1981), Enckell et al. (1987), Freude et al. (1974) 2005, PT-AZO PT-AZO U Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) Unknown PT-AZO U Unknown AT, CH J6 rotten vegetals unknown Southern Europe, Caucasus Europe Borges et al. (2005), Mendonça and Borges (2009) Luka et al. (2009), Wittenberg et al. (2006) Unknown ES-CAN J6 rotten vegetals unknown Europe Unknown FÖ U unknown Europe 2005, PT-AZO PT-AZO, ES-CAN U unknown Europe 2005, PT-AZO PT-AZO U Lathrobium unknown fulvipenne (Gravenhorst, 1806) Leptacinus pusillus (Stephens, 1833) unknown Lithocharis ochracea (Gravenhorst, 1802) Micropeplus marietti Jacquelin du Val, 1857 Mycetoporus nigricollis (Stephens, 1832) Myllaena brevicornis (Matthews,1838) Myrmecopora sulcata (Kiesenwetter,1850) unknown unknown Machado and Oromi (2000) Bengtson (1981), Enckell et al. (1987), Freude et al. (1974) Borges et al. (2005), Freude et al. (1974), Machado and Oromi (2000), Mendonça and Borges (2009) Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) 401 Northern, Central Europe Europe Heterota plumbea unknown (Waterhouse,1858) Myrmecopora uvida (Erichson, 1840) Native range 1st record Invaded countries in Europe Western 1989, GB GB Europe Coleoptera families other than Cerambycidae, Curculionidae... Family Regime Species Hadrognathus unknown longipalpis (Mulsant & Rey, 1851) Halobrecta flavipes unknown Thomson,1861 Omalium excavatum Stephens, 1834 Omalium rivulare (Paykull, 1789) Oxypoda haemorrhoa (Mannerheim, 1830) Oxytelus sculptus Gravenhorst, 1806 Phacophallus parumpunctatus (Gyllenhal, 1827) Philonthus cephalotes (Gravenhorst, 1802) Philonthus concinnus (Gravenhorst, 1802) unknown unknown Habitat Host U Borges et al. (2005), Mendonça and Borges (2009) U Borges et al. (2005) Bengtson (1981), Enckell et al. (1987), Freude et al. (1974) Europe 2005, PT-AZO Northern, Central Europe Northern Europe Northern & Central Europe Europe, caucasus Europe Unknown FÖ U 2005, PT-AZO PT-AZO Unknown FÖ U mite predator D bogs Unknown FÖ E, J nests micromammals Unknown FÖ J6 vegetal decay PT-AZO unknown Northern, Central Europe Unknown FÖ U unknown Europe PT-AZO U unknown Europe 2005, PT-AZO 1854, IE IE, PT-AZO, GB U parasitic/ predator parasitic/ predator Holarctic Unknown FÖ Eurasia (intro 2005, Nam) PT-AZO PT-AZO References U U Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) Bengtson (1981), Enckell et al. (1987) Bengtson (1981), Enckell et al. (1987) Bengtson (1981), Enckell et al. (1987) Bengtson (1981), Enckell et al. (1987), Freude et al. (1974) Borges et al. (2005), Mendonça and Borges (2009) Anderson (1997), Borges et al. (2005), Duff (2008), Mendonça and Borges (2009) Bengtson (1981), Enckell et al. (1987) Borges et al. (2005) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Oligota pusillima parasitic/ (Gravenhorst,1806) predator Olophrum fuscum unknown (Gravenhorst, 1806) Native range 1st record Invaded countries in Europe Europe (intro 2005, PT-AZO NAm) PT-AZO 402 Family Regime Species Neobisnius unknown lathrobioides (Baudi, 1848) Neobisnius unknown procerulus (Gravenhorst, 1806) Ocalea picata unknown (Stephens,1832) Philonthus fenestratus Fauvel, 1872 Philonthus fimetarius (Gravenhorst, 1802) Philonthus longicornis Stephens, 1832 Philonthus marginatus (O. Muller, 1764) Philonthus politus (Linnaeus, 1758) Philonthus quisquiliarius (Gyllenhal, 1810) Philonthus umbratilis (Gravenhorst, 1802) Phloeopora angustiformis Baudi, 1870 Phloeopora teres (Gravenhorst, 1802) Phloeopora testacea (Mannerheim, 1830) parasitic/ predator Europe, caucasus 2005, PT-AZO parasitic/ predator parasitic/ predator Palaearctic Unknown FÖ G Eurasia 2005, PT-AZO U parasitic/ predator Europe, Siberia Unknown FÖ parasitic/ predator parasitic/ predator Europe 2005, PT-AZO Eurasia, 2005, North Africa PT-AZO parasitic/ predator PT-AZO PT-AZO Habitat U U Host References Borges et al. (2005), Freude et al. (1974), Machado and Oromi (2000), Mendonça and Borges (2009) Borges et al. (2005) Bengtson (1981), Enckell et al. (1987) Borges et al. (2005), Mendonça and Borges (2009) U Bengtson (1981), Enckell et al. (1987) PT-AZO E PT-AZO U Borges et al. (2005), Mendonça and Borges (2009) Borges et al. (2005), Mendonça and Borges (2009) Europe (intro 2005, NAm) PT-AZO PT-AZO, ES-CAN U unknown Europe 2005, PT-AZO PT-AZO U unknown Europe PT-AZO U unknown Northern Europe 2005, PT-AZO 2005, PT-AZO PT-AZO U Borges et al. (2005), Machado and Oromi (2000), Mendonça and Borges (2009) Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) Borges (1990), Borges et al. (2005), Freude et al. (1974), Mendonça and Borges (2009) 403 Native range 1st record Invaded countries in Europe Eurasia, 2005, PT-AZO, ES-CAN North Africa PT-AZO Coleoptera families other than Cerambycidae, Curculionidae... Family Regime Species Philonthus discoideus parasitic/ (Gravenhorst, 1802) predator detritivorous Native range 1st record Invaded countries in Europe Palaearctic Unknown FÖ Habitat J6 Host rotten vegetals References Bengtson (1981), Enckell et al. (1987), Gamarra and Outerelo (2009) parasitic/ predator parasitic/ predator unknown Alps, Central Europe southern Europe Europe Unknown FÖ U 2005, PT-AZO PT-AZO Unknown FÖ U unknown Europe Unknown PT-AZO U Borges et al. (2005), Freude et al. (1974) detritivorous unknown Europe Unknown FÖ U Bengtson (1981), Enckell et al. (1987) Bengtson (1981), Enckell et al. (1987) Borges et al. (2005), Mendonça and Borges (2009) unknown unknown detritivorous parasitic/ predator parasitic/ predator Europe (intro Unknown FÖ NAm) Eurasia 2005, PT-AZO PT-AZO Europe (Int AUS) Southern Europe All over Europe Europe Bengtson (1981), Enckell et al. (1987) Borges et al. (2005), Mendonça and Borges (2009) Freude et al. (1974) E,G1 U U 2005, PT-AZO U PT-AZO Unknown AT, DK, EE, CH, GB I,J6 Mendonça and Borges (2009) waste land, compost Unknown FÖ, PT-AZO E, G, I2 stones, mosses, fungi Unknown PT-AZO U Luka et al. (2009), Wittenberg et al. (2006) Bengtson (1981), Borges et al. (2005), Enckell et al. (1987), Freude et al. (1974), Mendonça and Borges (2009) Borges et al. (2005) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) Xantholinus longiventris Heer, 1839 Regime 404 Family Species Proteinus brachypterus (Fabricius, 1792). Quedius mesomelinus (Marsham, 1802) Remus pruinosus (Erichson, 1840) Geostiba circellaris (Gravenhorst, 1806) Sunius propinquus (Brisout de Barneville, 1867) Tachinus laticollis Gravenhorst, 1802 Tachinus signatus Gravenhorst, 1802 Tachyporus chrysomelinus (Linnaeus, 1758) Tachyporus nitidulus (Fabricius, 1781) Thecturota marchii (Dodero,1922) Xantholinus linearis (Olivier, 1795) Family Species Xylodromus concinnus (Marsham, 1802) Xylodromus depressus (Gravenhorst, 1802) Tenebrionidae Blaps gigas (Linnaeus, 1758) Regime parasitic/ predator Europe Unknown FÖ G, I2 detritivorous 1888, CZ J6 Borges et al. (2005), Šefrova and Lastuvka (2005) Blaps lethifera Marsham, 1802 Blaps mortisaga (Linnaeus, 1758) detritivorous detritivorous Mediterranean region Europe Unknown PT-AZO, GB J1, J2 Borges et al. (2005), Duff (2008) Unknown GB J1, J2 Blaps mucronata Latreille, 1804 detritivorous Unknown IE, GB J1, J2 Corticeus linearis (Fabricus, 1790) Corticeus pini (Panzer, 1799) Scaurus punctatus Fabricius, 1798 detritivorous detritivorous detritivorous Eastern and Central Europe Europe, Mediterranean Europe Unknown GB G3 Europe Unknown GB G3 Unknown ES-CAN U Tenebrio obscurus Fabricius, 1792 Trachyscelis aphodioides Latreille, 1809 detritivorous detritivorous Mediterranean region Europe Unknown IE, PT-AZO, ESCAN, GB Unknown ES-CAN J1, J2 stored products J stored products parasitic/ predator Native range 1st record Invaded countries in Europe Europe Unknown FÖ Host References G, F, I2, J1 forests, gardens, cellars Bengtson (1981), Enckell et al. (1987) bark, wet wood detrivorous Bengtson (1981), Enckell et al. (1987) Duff (2008), Ferrer and Martinez Fernandez (2008) Duff (2008) old broadleaved forests Duff (2008) Machado and Oromi (2000) Borges et al. (2005), Duff (2008), Machado and Oromi (2000) Borges et al. (2005), Machado and Oromi (2000) Coleoptera families other than Cerambycidae, Curculionidae... Mediterranean region CZ, DK, PT-AZO Habitat 405 Regime Native range 1st record in Europe Invaded countries Habitat Host References Europe 2005, PT-AZO PT-AZO G bark, in forest Borges et al. (2005), Mendonça and Borges (2009), Freude et al. (1979) detritivorous Eurasia 2005, PT-AZO PT-AZO U nests Borges (1990), Borges et al. (2005), Mendonça and Borges (2009) unknown Mediteranean Unknown GB U Duff (2008) Olivier Denux & Pierre Zagatti / BioRisk 4(1): 315–406 (2010) detritivorous 406 Family Species Throscidae Throscus dermestoides (Linnaeus, 1766) Trogidae Trox scaber (Linnaeus, 1767) Zopheridae Aulonium ruficorne (Olivier, 1790) A peer reviewed open access journal BioRisk 4(1): 407–433 (2010) doi: 10.3897/biorisk.4.44 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk True Bugs (Hemiptera, Heteroptera) Chapter 9.1 Wolfgang Rabitsch Environment Agency Austria, Dept. Biodiversity & Nature Conservation, Spittelauer Lände 5, 1090 Vienna, Austria. Corresponding author: Wolfgang Rabitsch (wolfgang.rabitsch@umweltbundesamt.at) Academic editor: David Roy | Received 24 January 2010 | Accepted 23 May 2010 | Published 6 July 2010 Citation: Rabitsch W (2010) True Bugs (Hemiptera, Heteroptera). Chapter 9.1. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 407–403. doi: 10.3897/biorisk.4.44 Abstract The inventory of the alien Heteroptera of Europe includes 16 species alien to Europe, 25 species alien in Europe and 7 cryptogenic species. This is approximately 1.7% of the Heteroptera species occurring in Europe. Most species belong to Miridae (20 spp.), Tingidae (8 spp.), and Anthocoridae (7 spp.). The rate of introductions has exponentially increased within the 20th century and since 1990 an approximate arrival rate of seven species per decade has been observed. Most of the species alien to Europe are from North America, almost all of the species alien in Europe originate in the Mediterranean region and were translocated to central and northern Europe. Most alien Heteroptera species are known from Central and Western Europe (Czech Republic, Germany, Netherlands, Great Britain). Ornamental trade and movement as stowaways with transport vehicles are the major pathways for alien Heteroptera. Most alien Heteroptera colonize habitats under strong human influence, like agricultural, horticultural, and domestic habitats, parks and gardens. A few species prefer woodland including plantations of non-native forest trees. Impacts of alien Heteroptera in Europe are poorly investigated. A few species are considered pests in agriculture, forestry, or on ornamentals. More research is needed for a better understanding of the ecological and economic effects of introduced Heteroptera. Keywords alien, non-native, Hemiptera, Heteroptera, Europe Copyright W. Rabitsch. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 408 Wolfgang Rabitsch / BioRisk 4(1): 407–433 (2010) 9.1.1 Introduction The Heteroptera, or true bugs, is a highly diverse insect taxon with approximately 42,300 described species worldwide, separated into seven infraorders and 75–89 families (Henry 2009, Schuh and Slater 1995). Their body size ranges from less than 1 mm to 10 cm. True bugs feed on many different resources (e.g., haemolymph of insects, blood of endotherms, fungi cytoplasma, phloem-, xylem- or parenchym-sap of mosses, ferns, monocotyledons, mostly dicotyledons, algae, the endosperm of seeds, plant pollen). Heteropterans live in virtually all terrestrial and aquatic ecosystems from Antarctic birds’ nests to rainforest canopies, from the open surface of the ocean (almost uniquely for insects), to torrential and stagnant rivers, from ephemeral rain pools and phytotelmata to large lakes, and in aphotic caves and man-made buildings (Schuh and Slater 1995). Among the characteristic features are the mouthparts, which evolved as sucking stylets for the uptake of liquid food and the injection of secretions from the salivary gland; restricted diets are commonly observed. Most species are phytophagous, some feed exclusively on particular plant species, genera or families, whereas others are polyphagous species feeding on dozens to hundreds of different host plants. Some species are of considerable economic concern in agriculture or (more rarely) forestry, many species are predatory and some are used as biocontrol agents against agricultural pests (Schaefer and Panizzi 2000). Although some heteropteran species have reduced wings or wing musculature, and some are sexually dimorphic in this respect, many species are good flyers and capable of negotiating long distances. Subsequent spread after introduction by humans into a new area is commonly observed. Eggs and nymphs are translocated with host plants over long distances. Unlike the situation in many other Hemiptera, sexual reproduction prevails, with only one parthenogenetic species known in the European fauna, and depending on the species, one to several generations develop under temperate conditions. Many species deposit their eggs inside the host plant, which effectively fosters passive translocation and facilitates spread. 9.1.2 Methods Previously published information on alien Heteroptera species is available for some countries, e.g., Germany (Geiter et al. 2002) but see Hoffmann (2003) for a critical review, Austria (Essl and Rabitsch 2002), Switzerland (Kenis 2005), Czech Republic (Kment 2006b, Šefrová and Laštùvka 2005), and the Azores (Borges et al. 2005). Comparison of these lists is hindered by the use of different terminology and criteria for selecting species. The first attempt at a comprehensive treatment of the alien Heteroptera of Europe was published recently Rabitsch (2008) and serves as basis for this work, but is supplemented by new data (up to May 2009 including a few works in press). The reader is referred to Rabitsch (Rabitsch 2008) for a more detailed account on the history of introductions for each species. True Bugs (Hemiptera, Heteroptera). Chapter 9.1 409 This present chapter deals with species alien to Europe and species alien in Europe, but excludes continental European species alien to European islands. For example, Borges et al. (2005) stated that Tingis cardui (Linnaeus, 1758) and Gastrodes grossipes (De Geer, 1773), which both feeding on non-native host plants, are alien to the Azores. On the contrary, Heiss & Péricart (2007) argued that Aradus canariensis Kormilev, 1954 may have been introduced to Mallorca. The anthropogenic contribution of some recent range changes of continental “European” species to Great Britain and to Scandinavia, and hence their alien status, is particularly difficult to identify. For example, Ødegaard & Endrestøl (2007) present three hypotheses, not mutually exclusive, for the recent occurrence of Chilacis typhae (Perris, 1857) in Norway. For the time being, only Deraeocoris lutescens is here considered alien in Sweden and Norway, but the status of additional species needs careful re-examination, e.g. Pinalitus atomarius (Meyer-Dür, 1843) in Sweden (Lindskog and Viklund 2000), Chilacis typhae and Heterogaster urticae (Fabricius, 1775) in Norway (Ødegaard and Endrestøl 2007). Kirby et al. (2001) review several similar cases for Great Britain. 9.1.3 Taxonomy of the alien Heteroptera of Europe Alien Heteroptera are non-uniformly distributed across the seven infraorders. There are no alien species in Enicocephalomorpha and Dipsocoromorpha, the basal infraorders with 420 and 340 species worldwide, respectively. These predatory, usually tiny and fragile species live their secret lives in seclusion of riparian habitats and ground litter. No alien Gerromorpha are known in Europe; members of this predatory infraorder with more than 2100 species worldwide are commonly known as “Jesus-bugs” due to their ability to move on the surface of running and standing waters. Among Nepomorpha, the aquatic true bugs, with 2300 species worldwide, and Leptopodomorpha, the “shore bugs”, with 380 species worldwide, there is a single alien species in each infraorder, Trichocorixa verticalis and Pentacora sphacelata, both originally from North America, being introduced to the western Mediterranean region. Most alien Heteroptera belong to the most species-rich infraorders Cimicomorpha (20,500 species worldwide, 37 alien species in/to Europe) and Pentatomomorpha (16,200 species worldwide, 9 alien species in/to Europe). Within Hemiptera, Heteroptera constitute only a small fraction of alien species compared to aphids and scales (see chapters 9.2 and 9.3). At the end of the chapter, Table 9.1.1 and 9.1.2 list 48 Heteroptera species considered alien in this study of which 16 species are alien to Europe (i.e., species introduced from outside Europe), 25 species are alien in Europe (i.e., species introduced from one part of Europe to another), and seven cryptogenic species are of unknown origin. According to Aukema & Rieger (1995–2006), there are approximately 2860 Heteroptera species (including subspecies) in Europe, which means that 1.7% of the European fauna is alien. 410 Wolfgang Rabitsch / BioRisk 4(1): 407–433 (2010) At the family level, Miridae (20 spp.) and Tingidae (8 spp.) prevail, followed by Anthocoridae (7 spp.) and Lygaeidae sensu lato (5 spp.) (Figure 9.1.1). The systematic classification of Lygaeidae is still under discussion. While most heteropterists agree that Lygaeidae are paraphyletic (Henry 1997), there is no consensus on how to arrange them. The most species-rich family is Miridae, both in the native and the non-native faunas. Species of nine families are represented in the alien fauna, which is only 10% of the known families worldwide. Genera with more than one alien species are Amphiareus (2), Anthocoris (2), Corythucha (2), Deraeocoris (2), Orthotylus (4), Stephanitis (4), and Tuponia (5). Whereas all alien species belong to families present in Europe, 10 genera (13 genera including the cryptogenics, asterisked here) are alien at the genus level (Amphiareus, Belonochilus, *Buchananiella, Corythucha, Halyomorpha, *Nesidiocoris, Nezara, Pentacora, Perillus, *Taylorilygus, Trichocorixa, Tropidosteptes, Tupiocoris). Anthocoridae All Anthocoridae (flower bugs or minute pirate bugs) are small insects (< 5 mm body size) and most species are predatory, actively searching and hunting for their prey, which regularly consist of soft-bodied Sternorrhyncha. About 450 species are known at the world level (Henry 2009) of which 75 are considered native in Europe (Aukema and Rieger 1995–2006). The alien Heteroptera of Europe only include 4 species alien to Europe and 3 alien in Europe (Figure 9.1.1). Hence several species, especially in the genera Anthocoris and Orius, are successfully used commercially in biological control programs in greenhouses and sometimes in the wild, e.g., (Lattin 1999, Schaefer and Panizzi 2000). Apparently, only one species, the western and southern European Orius laevigatus is established outside its natural range in the Netherlands (Aukema and Loomans 2005) although these authors do not rule out the possibility that this species has shifted northwards due to climate change. Similarly, the true cause of the recent westward spread of the East-Palaearctic Amphiareus obscuriceps cannot definitely be identified. Although predatory, several anthocorid species are specialized to host plants, where they search for prey, e.g., Anthocoris butleri on Buxus and A. sarothamni on Cytisus. Both host plants are widely used as ornamentals and introduction of the Heteroptera with the host plants, as well as a range shift from western to eastern Europe, is possible. The origin of the pan-tropical Buchananiella continua is unknown. It is known from western Europe and appears to have spread both in Great Britain and in continental Europe (Aukema 2007, Aukema and Hermes 2009, Kirby 1999). Likewise, the origin of the cosmopolitan Amphiareus constrictus is unclear. It was introduced to the Netherlands (Aukema and Hermes 2009) and may further spread in Europe. The alien status of Lyctocoris campestris in Europe is debatable. True Bugs (Hemiptera, Heteroptera). Chapter 9.1 411 Figure 9.1.1. Taxonomic overview of the alien Heteroptera of Europe at the family level. Species alien to Europe include cryptogenics. Coreidae The leaf-footed or squash bugs is a species-rich family with species of medium to large body size. A total of 1900 species have been described throughout the world (Henry 2009), including 52 in Europe (Aukema and Rieger 1995–2006) but only one alien species has so far established on the continent. For several reasons, this single alien species, Leptoglossus occidentalis, is of particular interest. The native range is presumed to be west of the Rocky Mountains and following its spread in North America since the 1950s, it was introduced to Europe only in the late 1990s. The first date recorded in European record was 1999 in northern Italy (Bernardinelli and Zandigiacomo 2001) and the species rapidly spread over most of Europe (Dusoulier et al. 2007, Rabitsch 2008) with no foreseeable stop (Lis et al. 2008). This spread is likely to be the result of multiple introductions into Europe, and secondary translocations within it. When feeding on conifer seeds, fertility of the seeds is reduced, causing an economic impact for forestry. Recently, infrared receptive organs were found in L. occidentalis, orienting specimens towards conifer cones (Takács et al. 2009). Because individuals aggregate in autumn seeking hibernation sites in buildings, this species may also become a nuisance to people. Recently, it was found in Japan (Tokyo) (Ishikawa and Kikuhara 2009). Corixidae The family has about 600 described species in the world (Henry 2009), and 72 in Europe (Aukema and Rieger 1995–2006). The single aquatic species yet recognized as alien to Europe, Trichocorixa verticalis, is of nearctic origin and was introduced to 412 Wolfgang Rabitsch / BioRisk 4(1): 407–433 (2010) Europe (Portugal) between 1997 and 2003 (Sala and Boix 2005). Its pathway and potential impact is not known, but it may well have been introduced as a stowaway with mosquitofish (Gambusia sp.) and may outcompete native corixids and lead to a simplification of the aquatic community (Kment 2006a, Millán et al. 2005, Rodríguez-Pérez 2009). Lygaeidae sensu lato Lygaeidae or seed-bugs are a species-rich group of about 4000 species (Henry 2009) of medium body size that include both seed-feeding and predatory species with economic impact that is sometimes significant (Schaefer and Panizzi 2000). A total of 363 species are native to Europe (Aukema and Rieger 1995–2006) but only two species are alien to Europe, Nysius huttoni from New Zealand, and Belonochilus numenius from North America. Both species currently are locally distributed, but have the potential to spread over large parts of Europe. The former is known from the Netherlands, Belgium, northern France and Great Britain, where it occurs in ruderal sites, waste grounds and abandoned fields (Smit et al. 2007). N. huttoni feeds on several weeds and crops and attains pest status in its native area (Sweet 2000). The latter has been found in Corsica and mainland southern France in the vicinity of a railway station and at a university campus (Montpellier) (Matocq 2008) as well as in Catalonia, Spain (Castelldefels, Barcelona) (Gessé et al. 2009) on or near ornamental sycamore (Platanus sp.). These almost simultaneous findings and the fact that its host plant is regularly planted in urban parks and gardens, indicates that the species is already much more widely distributed and that further spread in Europe is very likely. Three further lygaeid species are here considered alien in Europe. The first is Arocatus longiceps, an eastern Mediterranean species living on sycamore, whose occurrence is restricted to urban settings where it sometimes reaches high abundance causing a nuisance to people. Due to its variability, heteropterists debate its separation from native Arocatus species, considering possible hybridization and post-invasion colour changes (Hoffmann 2008). The second, Orsillus depressus, is a Mediterranean species living on Cupressaceae. Its adaptation to ornamental Thuja, Chamaecyparis, and Juniperus promoted its northward spread. Intraguild competition on native Juniperus-stands is likely, but so far not investigated. Lastly, Oxycarenus lavaterae is a western Mediterranean species living on Malvaceae s.l. with a preference for lime trees (Tilia sp.). The species builds spectacular large aggregations of millions of individuals, also sometimes causing nuisance to people, e.g., at market places in cities or when entering buildings. Miridae With more than 10,000 described species (Henry 2009) of which 1036 in Europe (Aukema and Rieger 1995–2006), Miridae or plant bugs is the most species-rich family True Bugs (Hemiptera, Heteroptera). Chapter 9.1 413 within Heteroptera. Plant bugs include tiny to large, soft-bodied, dull to brightly coloured, phytophagous, zoophytophagous and predatory species (Wheeler 2001). Only 5 species alien to Europe have established whereas 15 species are considered alien in Europe (Figure 9.1.1.). Whereas some species are considered serious agricultural pests, others are used in biological control programmes. Closterotomus trivialis and Dicyphus escalerae are examples of Mediterranean species occurring locally in central Europe, the latter recently also found in Great Britain (Kirby et al. 2009), being introduced with their host plants. The same is most likely true for Deraeocoris lutescens, a western Palaearctic species introduced to Scandinavia. Another predatory, remarkably fast spreading species, is the Mediterranean Deraeocoris flavilinea, that presumably has been introduced unintentionally along transportation routes. Tupiocoris rhododendri was described from specimens collected in 1971 in Kew Gardens, London, but it originally comes from North America. Recently, this predatory species was found in continental Europe, and its further spread is to be expected (Aukema 2007, Aukema et al. 2005a). One of the most recent members of the European alien Heteroptera fauna is Tropidosteptes pacificus from North America, collected on European ash (Fraxinus excelsior) in a natural environment in the Netherlands (Aukema et al. 2009a). Three Orthotylus species live zoophytophagously on Cytisus and probably were introduced with their host plant to central and eastern Europe. The mediterranean Orthotylus caprai was only recently observed in central and western Europe on Cupressaceae, and is considered an alien species in Europe north of the Alps. Five Tuponia species, living phytophagously on Tamarix, were most likely introduced with their ornamental host plants. Pentatomidae Pentatomidae or stink bugs are a species-rich and medium to large body-sized heteropteran family with often stout and colourful bodies. About 4700 species have been recognized (Henry 2009), including 187 species in Europe (Aukema and Rieger 1995– 2006). Members of one subfamily (Asopinae) are predatory and some are used in biocontrol programmes. This is true for Perillus bioculatus, native to North America and used against the Colorado potato beetles Leptinotarsa decemlineata in several European countries (De Clercq 2000). However, successful establishment in the wild apparently so far only occurred in Turkey and Greece. Recently, the Brown Marmorated Stink Bug Halyomorpha halys, native to Asia, was introduced to Switzerland (see factsheet 14.49) (Wermelinger e al. 2008). This species lives on ornamentals, vegetables and fruit trees where it may become a pest and it is regarded as a nuisance when seeking hibernation sites. The Southern Green Stink Bug Nezara viridula, a polyphytophagous pest species on several crops, is presumably of African and/or Mediterranean origin. Nezara viridula is a clear case of establishment of populations outside its original distribution in Germany, Hungary, Great Britain, and northern Switzerland. In addition, this species is found regularly in other parts of Europe, and is regularly intercepted by plant quarantine (Malumphy and Reid 2007). 414 Wolfgang Rabitsch / BioRisk 4(1): 407–433 (2010) Reduviidae Reduviidae, the assassin bugs, are a species-rich and morphologically highly diverse predatory heteropteran family including 6900 species in the world (Henry 2009) of which 110 occur in Europe (Aukema and Rieger 1995–2006). However, only two cryptogenic, pantropical species are included here. Empicoris rubromaculatus is found in southwestern Europe with isolated records in Belgium, Croatia and Greece; the latter records may reflect a recent eastward range shift, but maybe this species was previously overlooked in the eastern Mediterranean region. Ploiaria chilensis is known from Macaronesia and Spain, with doubtful records from the central and eastern Mediterranean. Saldidae Shore bugs or Saldidae are a species-poor (340 species in the world (Henry 2009)), medium-sized, predatory family, living in littoral habitats along the edges of running and standing waters, marine shoreline and bogs. Whereas the native fauna includes 47 species (Aukema and Rieger 1995–2006), there is only one species alien to Europe. This single species, Pentacora sphacelata, is known since the 1950s from the Iberian Peninsula and Sardinia. This is a halophilous species living in the tidal-zone and close to saline waters. Tingidae Lace bugs or Tingidae are a species-rich, small-sized (< 8 mm body size), phytophagous family, with characteristic ornate and lacelike hemelytra and pronotum. Most species live on or near their host plants with a usually tight preference to particular plant species or families. About 2100 species are recognized in the world (Henry 2009) but only 171 are native to Europe (Aukema and Rieger 1995–2006). Thus, the alien fauna which includes 5 species alien to Europe is proportionally a little more important than in Miridae (2.9% of the total fauna vs. 0.5%; Figure 9.1.1). Both Corythucha-species were introduced from North America to Italy and live arboreally on their host plants, including the oak lace bug C. arcuata on Quercus (see factsheet 14.51) and the sycamore lace bug C. ciliata on Platanus (see factsheet 14.52). The former species was introduced a decade ago and only started to spread (Dioli et al. 2007), whereas the latter was introduced in the 1960s and nowadays is very widespread across Europe. Stephanitis pyrioides and S. takeyai were introduced from Japan and S. rhododendri from North America with ornamental Ericaceae (Rhododendron, Azalea, Pieris). Dictyonota fuliginosa and Elasmotropis testacea are both considered alien in parts of Europe where the host plants are also alien, although unambiguous evidence on their introduction status often is lacking. The alien status of Stephanitis oberti in Central Europe is debatable. True Bugs (Hemiptera, Heteroptera). Chapter 9.1 415 9.1.4 Temporal trends of introduction of alien Heteroptera in Europe The (published) year when first recorded is known for all species (Table 9.1.1 and 9.1.2; see also Rabitsch (2008) for all country records), although it is evident that this need not be identical with the year of introduction. Usually it takes a few years for introduced insects to increase in abundance above a certain threshold to establish reproducing populations and to get recognized. This time-lag is known as a common characteristic of biological invasions and it can extend over long time periods in some organisms, e.g. decades or even centuries in some plants (Kowarik 1995). For insects, however, this time-lag usually extends over much shorter periods, but several years may still elapse since an alien species is discovered and information is communicated. Some Heteroptera were already introduced in ancient times, such as the notorious bed bug Cimex lectularius Linnaeus, 1758 and maybe some others following human expansion associated with agricultural land reclamation. Those ancient introductions were rarely if ever documented and are therefore excluded in this study. However, there is no doubt that the rate of introductions has exponentially increased within the 20th century and reached unprecedented magnitudes in the 21st century (Figure 9.1.2). Since 1990, an approximate arrival rate of seven species per decade has been observed (Rabitsch 2008). Currently, Heteroptera alien to and alien in Europe both establish at a rate of 0.33 species per year; this means that on average every third year an Heteroptera species from outside Europe arrives in Europe. Even within the last eight years, five species have been detected: Corythucha arcuata, Tropidosteptes pacificus and Belonochilus numenius from North America (2000, 2007, 2008, respectively), Nysius huttoni from New Zealand (2002) and Halyomorpha halys from East Asia (2007). Some species are suspected of having been introduced in the 19th century together with ornamental plants, e.g. Anthocoris butleri on Buxus sempervirens, Anthocoris sarothamni, Orthotylus adenocarpi, O. concolor, O. virescens, Dictyonota fuliginosa on Cytisus scoparius, and Macrolophus glaucescens, Elasmotropis testacea on Echinops sphaerocephalus. More recently, several Tuponia species were introduced with the increasing use of ornamental Tamarix species in public and private gardens. The time of introduction for cryptogenic species into Europe is unclear and may well be several centuries before present. Most are pan-tropically distributed, zoophagous species. 9.1.5 Biogeographic patterns of the alien Heteroptera of Europe 9.1.5.1 Origin of alien species A total of 16 species are alien to Europe, 10 of these from North America, 4 from the eastern Palaearctic and East Asia and one each from South America and Oceania. Almost all of the 25 species alien in Europe originate in the Mediterranean region and 416 Wolfgang Rabitsch / BioRisk 4(1): 407–433 (2010) Figure 9.1.2. Temporal trends in the mean number of new records per year for Heteroptera species alien to Europe and alien in Europe from 1492 to 2008. Cryptogenic species are excluded. The number above the bar indicates the absolute number of species in this time period. were translocated to central and northern Europe. Seven species are considered cryptogenic with unknown origin and cosmopolitan distribution (Figure 9.1.3). Rabitsch (2008) mentioned the increasing trend of North American species arriving in Europe (Figure 9.1.4). This is corroborated by the most recent introductions of Tropidosteptes pacificus in the Netherlands (Aukema et al. 2009a) and Belonochilus numenius in Corsica, continental France and Spain (Gessé et al. 2009, Matocq 2008). Few species have been introduced from Oceania (New Zealand, Nysius huttoni, see factsheet 14.47) and South America (Fulvius borgesi). The latter species was only recently described as new to science, based on specimens collected in banana plantations at low altitudes on the Azores (Chérot et al. 2006). The authors argued, based on morphological characters, that the species was introduced from South America. Nezara viridula is considered the only alien species of African origin, although some were previously intercepted during plant health inspections, e.g. the Grain Chinch Bug, Macchiademus diplopterus (Distant, 1903) (Lygaeidae) and Natalicola pallidus (Westwood, 1837) (Tessaratomidae) at Heathrow Airport, London, on fruits and plants imported from South Africa (Malumphy and Reid 2007, 2008). Suitable climate seems to be a significant factor for the establishment of Heteroptera alien to Europe since 87% (14 species) come from temperate climates and only two species were introduced from the southern hemisphere. True Bugs (Hemiptera, Heteroptera). Chapter 9.1 417 Figure 9.1.3. Geographic origin of the alien Heteroptera species of Europe. 9.1.5.2 Distribution of alien species in the European countries Most alien Heteroptera species are known from Central Europe (Czech Republic: 22 species, that is 47% of all species, Germany: 20 species) and Western Europe (Netherlands: 20 species, Great Britain: 17 species) (Figure 9.1.5). One reason for the subordinate relevance of South Europe as a recipient for alien Heteroptera lies in the fact that almost all species alien in Europe originate in the Mediterranean region and were translocated north. This is likely a consequence of the increasing north-south exchange of people and merchandise (e.g., summer holiday tourism, fruits, vegetables) (Rédei and Torma 2003). A west-east pattern, however, can be found in suspected previous introductions of species living on western European ornamental plants, which were later widely planted across Europe. This concerns species living on Buxus sempervirens, Cytisus scoparius, and Echinops spp. Those plants are nowadays widely planted in cemeteries and private gardens and host monophagous Heteroptera species (e.g. Anthocoris butleri, A. sarothamni, Dictyonota fuliginosa, Elasmotropis testacea, Macrolophus glaucescens and Orthotylus spp.). This northwest-southeast gradient is also demonstrated by a significant negative rank correlation of alien species numbers and longitude when the diversity of alien heteropterans is tentatively correlated to environmental and economic variables using a Spearman rank correlation (ρ= -0.548; P < 0.001; Rabitsch, unpublished data). Whereas the number of native Heteropteran species per country appears to be significantly correlated with both the number of native plant species (ρ= 0.887; P < 0.001) and the country size (ρ= 0.576; P < 0.001), the number of alien Heteroptera species does not (ρ= -0.548 and ρ= 0.093, respectively, n.s.). On the contrary, whereas the number of alien Heteroptera is positively correlated with some economic variables (GDP per capita, ρ = 0.417; P < 0.01; average trade import 1990–1997, ρ= 0.748; 418 Wolfgang Rabitsch / BioRisk 4(1): 407–433 (2010) Figure 9.1.4. Numbers of established alien Heteroptera species of Europe by period of introduction and geographic origin. Cryptogenic species are excluded. P < 0.001), the number of native species is not (ρ= -0.049, n.s.). The distribution patterns of alien Heteropterans also seem to match these of alien plants (ρ= 0.394; P<0.05) and alien terrestrial invertebrates (ρ= 0.703; P<0.001); this likely is a fact of the overwhelming importance of urbanisation and trade import for the establishment of alien terrestrial invertebrate species in Europe (Roques et al. 2008). The Netherlands must be regarded as an invasion focus for the alien Heteroptera of Europe, with seven species having been first recorded in this country (Tables 9.1.1 and 9.1.2). A more sophisticated statistical analysis with several explanatory variables and taking into account area and sample effects, autocorrelation, multicollinearity, etc. will be presented elsewhere (Rabitsch and Moser, in prep.). 9.1.6 Pathways of introduction of the alien species of Heteroptera Heteroptera are rarely intercepted (Roques and Auger-Rozenberg 2006) or at least rarely reported, in part due to their ancillary role as pest organisms. Recently, however, a number of such cases were published from regular plant health inspections in Great Britain. For example, Natalicola pallidus (Tessaratomidae) was found on Crassula multicava from South Africa (Malumphy and Reid 2008) and one specimen of Leptoglossus occidentalis was found in a timber shipment from the USA (Malumphy et al. 2008) indicating multiple introductions of this species into Europe. Ornamental trade and movement as stowaways with transport vehicles are the major pathways for alien Heteroptera (Rabitsch True Bugs (Hemiptera, Heteroptera). Chapter 9.1 419 Figure 9.1.5. Numbers of established alien Heteroptera species per European country. Data rely on Tables 9.1.1 and 9.1.2. Aliens with doubtful status are included. Archipelago: 1 Azores 2 Madeira 3 Canary islands. 2008), also confirmed by the interruption of introductions between 1925 and 1949 (Figure 9.1.4). 9.1.7 Ecosystems and habitats invaded by alien Heteroptera in Europe Most alien Heteroptera colonize habitats under strong human influence, like agricultural, horticultural, and domestic habitats (51%) and parks and gardens (27%) (Figure 9.1.6). Some species prefer woodland including plantations of non-native forest trees. It is worth mentioning that Leptoglossus occidentalis has not only spread across Europe, but has also 420 Wolfgang Rabitsch / BioRisk 4(1): 407–433 (2010) Figure 9.1.6. Main habitats colonized by alien Heteroptera species in Europe. The number above each bar indicates the absolute number of alien species recorded per habitat. Note that a species may have colonized several habitats. expanded its occupied habitat: first records in most countries are indoors, from cities and harbours, but increasingly records in the field are observed at higher elevations. In France, L. occidentalis has twice been captured above 1000 m (Dusoulier et al. 2007) and in Austria (Styria) there is a documented record at 1500 m (Gepp, in litt.) (see factsheet 14.42). 9.1.8 Ecological and economic impact of alien Heteroptera in Europe Impacts of alien Heteroptera in Europe are poorly investigated (Rabitsch 2008). A few species are considered pests in agriculture or forestry, e.g. Nysius huttoni, and Leptoglossus occidentalis, or on ornamental plants, e.g. Corythucha ciliata and Stephanitis takeyai, but damage is only locally reported in Europe to date. No data are available on any negative ecological impact on native species either due to predation, hybridization, competition or pathogen-transfer. However, as mentioned by Rabitsch (2008), no one has yet looked at such effects. It may be worth investigating intraguild competition within the juniper-feeding guild or the effects of Trichocorixa verticalis in aquatic communities. 9.1.9 Conclusion It is essential to observe and document range changes of species. Clearly, the number of introduced Heteroptera will increase. Climate change and habitat modification will further promote establishment of additional species. Some introduced species, currently considered as not established, were excluded in this study, but may establish populations in the near True Bugs (Hemiptera, Heteroptera). Chapter 9.1 421 a e b f d c Figure 9.1.7. Adults of some alien Heteroptera species: a Arocatus longiceps (Credit: Wolfgang Rabitsch) b Leptoglossus occidentalis feeding on Scots pine (Credit: Alain Roques) c Oxycarenus lavaterae aggregating on trunk (Credit: Wolfgang Rabitsch) d Oxycarenus lavaterae detail (Credit: Wolfgang Rabitsch) e Stephanitis takeyai (Credit: Wolfgang Rabitsch) f Tupiocoris rhododendri (Credit: Ab Baas). 422 Wolfgang Rabitsch / BioRisk 4(1): 407–433 (2010) future; e.g., Orius flagellum Linnavuori, 1968 in the Netherlands (Aukema and Hermes 2009), Xylocoris flavipes (Reuter, 1875) in several European countries (Péricart 1972, 1996). Also, recent range changes of some continental European species need to be carefully reconsidered when new data become available as some of these may deserve alien status; e.g. Ødegaard & Endrestøl (2007), see Rabitsch (2008) for additional examples. 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Streito JC (2006) Note sur quelques espèces envahissantes de Tingidae: Corythucha ciliata (Say, 1832), Stephanitis pyrioides (Scott, 1874) et Stephanitis takeyai Drake & Maa, 1955 (Hemiptera Tingidae). L’Entomologiste 62: 31–36. Sweet MH (2000) Seed and Chinch Bugs (Lygaeoidea). In: Schaefer CW, Panizzi AR (Eds) Heteroptera of Economic Importance. Boca Raton, Florida: CRC Press, pp. 143–264. Takács S, Bottomley H, Andreller I, Zaradnik T, Schwarz J, Bennett R, Strong W, Gries G (2009) Infrared radiation from hot cones on cool conifers attracts seed-feeding insects. Proceedings Royal Society London B 276: 649–655. Velimirovic V, Durovic Z, Raicevic M (1992) Bug Oxycarenus lavaterae Fabricius (Lygaeidae, Heteroptera) new pest on lindens in southern part of Montenegro. Zastita Bilja 43: 69–72. Voigt K (1977) Bemerkenswerte Wanzenfunde aus Baden-Württemberg, mit einem Erstnachweis für Deutschland. Beiträge zur naturkundlichen Forschung in Südwestdeutschland 36: 153–158. Wermelinger B, Wyniger D, Forster B (2005) Massenauftreten und erster Nachweis von Oxycarenus lavaterae (F.) (Heteroptera, Lygaeidae) auf der Schweizer Alpennordseite. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 78: 311–316. Wermelinger B, Wyniger D, Forster B (2008) First records of an invasive bug in Europe: Halyomorpha halys Stål (Heteroptera: Pentatomidae), a new pest on woody ornamentals and fruit trees? Mitteilungen der Schweizer Entomologischen Gesellschaft 81: 1–8. Werner DJ (2005) Nezara viridula (Linnaeus, 1758) in Köln und in Deutschland (Heteroptera, Pentatomidae). Heteropteron 21: 29–31. Wheeler AG (2001) Biology of the Plant Bugs (Hemiptera: Miridae). Pests, Predators, Opportunists. Ithaca: Cornell University Press. 506 pp. Coreidae Leptoglossus occidentalis Heidemann, 1910 A Phytophagous Native range 1st record in Europe East Palaearctic 1987, BG Cosmopolitan 2007, NL Invaded countries AT, BE, BG, BY, CZ, DE, EE, FI, HU, IT, NL, SK NL Habitat E, I – E – 1880, PT- BE, ES, ES-CAN, FR, I, X MAD GB, IT, NL, PT, PTAZO, PT-MAD West ? AL, AT, BA, BE, BG, I Palaearctic? BY, CH, CZ, DE, DK, Cosmopolitan EE, ES, ES-CAN, FI, FR, GB, GR, HR, HU, IE, IT, IT-SAR, IT-SIC, LT, LU, LV, MD, ME, MK, MT, NL, NO, PL, PT, PT-AZO, PTMAD, RS, SE, SI, SK, UA Pantropical North America 1999, IT AT, BE, BG, CH, CZ, DE, ES, FR, GB, GR , HR, HU, IT, ME, NL, PL, RO, RS, SI, SK Hosts G, I, X Refs Aukema (2007), Aukema et al. (2005a), Hradil et al. (2008), Péricart and Stehlík (1998) Aukema and Hermes (2009) – Aukema and Hermes (2009), Aukema et al. (2009b), Kirby (1999) – Péricart (1972) Pinaceae (Pinus, Pseudotsuga, Picea, Abies), Cupressaceae (Libocedrus) Aukema (2008), Bernardinelli and Zandigiacomo (2001), Dusoulier et al. (2007), Hradil (2008), Kment et al. (2005), Malumphy et al. (2008), Protić (2008), Ruicănescu (2009) Wolfgang Rabitsch / BioRisk 4(1): 407–433 (2010) Family Status Feeding Species Regime Anthocoridae A ZooAmphiareus obscuriceps (Poppius, phagous 1909) Amphiareus C Zooconstrictus (Stål, phagous 1860) Buchananiella C Zoocontinua (White, phagous 1880) Lyctocoris campestris C Zoo(Fabricius, 1794) phagous 428 Table 9.1.1. List and main characteristics of Heteroptera species alien to Europe. Status: A Alien to Europe C cryptogenic species. For details see Rabitsch (2008). ? = occurrence doubtful, * = probably not established. New data since Rabitsch (2008) are given in bold. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Only selected references are given. Last update May 2009. Family Status Feeding Species Regime Corixidae Trichocorixa verticalis A Omni(Fieber, 1851) vorous Lygaeidae Nysius huttoni A PhytoF.B.White, 1878 phagous Native range 1st record in Europe North America 1997, PT Invaded countries Habitat Refs C – New Zealand 2002, NL BE, FR, GB, NL I North America ES, FR, FR-COR I Poaceae, Aukema et al. (2005b), Cuming Brassicaceae (2008) and others (polyphagous) Platanaceae Gessé et al. (2009), Matocq (2008) (Platanus) A Zoophyto- South phagous? America 2003, PT- PT-AZO AZO C Zoophyto- Pantropical phagous ? Taylorilygus apicalis (Fieber, 1861) C Phytophagous Pantropical ? Tropidosteptes pacificus Van Duzee, 1921 Tupiocoris rhododendri (Dolling, 1972) A Phytophagous North America 2007, NL CY, ES, ES-CAN, FR, I GR, GR-CRE, IT, MT, PT-MAD AL, BA, BG, CY, ES, I ES-CAN, FR, FRCOR, GR, HR, IT, ITSAR, IT-SIC, MT, PT, PT-AZO, PT-MAD, SI, UA NL G A Zoophagous North America 1971, GB BE, DE, GB, NL 2008, FR I I, X – Chérot et al. (2006) – Kerzhner and Josifov (1999) Asteraceae Kerzhner and Josifov (1999) and others (polyphagous) Oleaceae (Fraxinus excelsior) Ericaceae (Rhododendron) Aukema et al. (2009a) Aukema et al. (2005a), Aukema et al. (2007), Dolling (1972) 429 Phytophagous Kment (2006a), Sala and Boix (2005) True Bugs (Hemiptera, Heteroptera). Chapter 9.1 ES, PT A Belonochilus numenius (Say, 1831) Miridae Fulvius borgesi Chérot, J. Ribes & Gorczyca, 2006 Nesidiocoris tenuis (Reuter, 1895) Hosts Status Feeding Regime Native range 1st record in Europe Invaded countries Habitat A Phytophagous East Asia 2007, CH CH I, X Perillus bioculatus (Fabricius, 1775) Reduviidae Empicoris rubromaculatus (Blackburn, 1889) Ploiaria chilensis (Philippi, 1862) Saldidae Pentacora sphacelata (Uhler, 1877) Tingidae Corythucha arcuata (Say, 1832) A Zoophagous North America 1992, TU GR, TU G, I C Zoophagous Pantropical ? – Aukema et al. (2009b) C Zoophagous Pantropical ? BE, ES, ES-CAN, FR, I FR-COR, GR, HR, IT, PT, PT-AZO, PT-MAD ?CY, ES, ES-CAN, ?IT, I PT-AZO, PT-MAD – Putshkov and Putshkov (1996) A Zoophagous North America 1953, ES ES, IT, PT B – Carapezza (1980) A Phytophagous North America 2000, IT CH, IT G Dioli et al. (2007), Forster et al. (2005) Corythucha ciliata (Say, 1832) A Phytophagous North America 1964, IT I, X Stephanitis pyrioides (Scott, 1874) A Phytophagous Japan 1904, NL AT, BE, BG, CH, CZ, DE, ES, FR, GB, GR, HR, HU, IT, ME, NL, PT, RS, SK, SI CH, *FR, GR, IT, NL Fagaceae (Quercus, Castanea) Platanaceae (Platanus) Stephanitis rhododendri Horváth, 1905 Stephanitis takeyai Drake & Maa, 1955 A Phytophagous North America A Phytophagous Japan <1900, NL *BE, BG, CH, CZ, DE, I, X DK, *FI (J100), *FR, GB, NL, *PL, SE 1994, NL BE, CZ, DE, FR, GB, I, X IT, NL, PL Refs fruit trees and Wermelinger et al. (2008) ornamentals (polyphagous) – Kivan (2004) Ericaceae (Rhododendron) Ericaceae (Rhododendron) Ericaceae (Pieris, Rhododendron) Aukema and Hermes (2009), Kment (2007), Servadei (1966), Stehlík (1997), Streito (2006) Kment (2007), Streito (2006) Halstead and Malumphy (2003), Jindra and Kment (2006), Simov and Pencheva (2007) Aukema (1996), Halstead and Malumphy (2003), Ishikawa and Kikuhara (2009), Streito (2006) Wolfgang Rabitsch / BioRisk 4(1): 407–433 (2010) I, X Hosts 430 Family Species Pentatomidae Halyomorpha halys (Stål, 1855) Table 9.1.2. List and characteristics of the Heteroptera species alien in Europe. For details see Rabitsch (2008). ?N = Alien status doubtful (species could be native), ? = occurrence doubtful, * = probably not established. New data since Rabitsch (2008) are given in bold. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Only selected references are given. Last update May 2009. Family Species Native range Phytophagous East Mediterranean Orsillus depressus (Mulsant & Rey, 1852) Phytophagous Mediterranean Oxycarenus lavaterae (Fabricius, 1787) Phytophagous West Mediterranean Miridae Closterotomus trivialis (A. PhytoCosta, 1853) phagous Mediterranean Zoophagous Hosts References Buxaceae (Buxus) Kment et al. (2006) 1953, CZ I, X Kment (2006b) 2005, NL NL I Fabaceae (Cytisus) – 1962, CZ West Mediterranean Southwest Europe Habitat I, X Southwest Europe Zoophagous Zoophagous Invaded countries AT, BE, CH (?N), CZ, DE (?N), IE, LU, NL, SE (Gotland), SK *AT, CZ Zoophagous Anthocoris sarothamni Douglas & Scott, 1865 Orius laevigatus (Fieber, 1860) Lygaeidae Arocatus longiceps Stål, 1872 Deraeocoris lutescens (Schilling, 1837) 1st record in invaded areas 1990, HU AT, BE, CH, CZ, DE, I, X Platanaceae ES, FR (N?), GB, HU, (Platanus) NL, PT, SI (?N), SK 1971, DE AT (?N), BE, CZ, DE, E, I, X Cupressaceae *FI, GB, HU (?N), LU, NL, SK G, I, X Malvaceae 1985, ME AT, BG, CH (north), CZ, DE, *FI, (Tilia) FR(north), HU, ME, *NL, RO, RS, SI, SK 1998, NL NL West Palaearctic 1990, NO NO, SE I I, X Weeds, olive trees, Citrus (polyphagous) Malvaceae (Tilia) Aukema and Loomans (2005) Göricke (2008), Kondorosy (1997), Nau and Straw (2007), Ribes and Pagola-Carte (2008) Hradil et al. (2002), Voigt (1977) Hradil et al. (2008), Kment (2009), Kondorosy (1997), Rabitsch and Adlbauer (2001), Velimirovic et al. (1992), Wermelinger et al. (2005) True Bugs (Hemiptera, Heteroptera). Chapter 9.1 Anthocoridae Anthocoris butleri Le Quesne, 1954 Feeding Regime Aukema (1999), Aukema and Hermes (2009) Lindskog and Viklund (2000), Ødegaard and Endrestøl (2007) 431 Feeding Regime Native range Habitat Deraeocoris flavilinea (A. Costa, 1862) Zoophagous Mediterranean Dichrooscytus gustavi Josifov, 1981 Phytophagous European – Cryptogenic Dicyphus escalerae Lindberg, 1934 Phytophagous West Mediterranean Macrolophus glaucescens Fieber, 1858 Orthotylus adenocarpi (Perris, 1857) Orthotylus caprai Wagner, 1955 Orthotylus concolor (Kirschbaum, 1856) Orthotylus virescens (Douglas & Scott, 1865) Tuponia brevirostris Reuter, 1883 Tuponia elegans (Jakovlev, 1867) Zoophagous Zoophytophagous Zoophytophagous Zoophytophagous Zoophytophagous Phytophagous Phytophagous Mediterranean <1858, CZ CZ E West Mediterranean Mediterranean <1892?, CZ 2006, GB CZ (?N) E, G, I DE, GB I West Mediterranean West Mediterranean West Mediterranean Central Asia <1892?, *AT, CZ (?N) CZ 2003, HU CZ (?N), HU 1964, HU AT, CZ, HU, SK I, X Tuponia hippophaes (Fieber, 1861) Tuponia macedonica Wagner, 1957 Phytophagous Phytophagous Mediterranean <1982, SK CZ, BE, SK I, X East Mediterranean 2003, SK I, X 2001, GB References I, X Many trees and shrubs Kment et al. (2006), Péricart (1965) I Cupressaceae Bryja and Kment (2002), Hradil et al. (2008) I Veronicaceae (Antirrhinum majus) Asteraceae (Echinops) Fabaceae (Cytisus) Cupressaceae Hollier and Matocq (2004), Kirby et al. (2009), Servadei (1966) E, G, I E, G, I DE, GB, GR (?N), HR I, X SK Hosts Fabaceae (Cytisus) Fabaceae (Cytisus) Tamaricaceae (Tamarix) Tamaricaceae (Tamarix) Tamaricaceae (Tamarix) Tamaricaceae (Tamarix) Kment (2006b) Kment (2006b) Nau (2007), Simon (2007) Frieß and Rabitsch (2009), Kment (2006b) Kment (2006b), Kondorosy (2005) Barclay and Nau (2003), Simon (2007) Benedek and Jászai (1968), Bryja and Kment (2002), Hradil et al. (2008), Rabitsch (2002) Bryja and Kment (2002), Hradil et al. (2008) Hradil et al. (2008) Wolfgang Rabitsch / BioRisk 4(1): 407–433 (2010) 1st record Invaded countries in invaded areas 1961, FR- AL, AT, BE, CH, CZ, COR DE, FR (Alsace), FRCOR, GB, LU, MT, NL, SE, SI 1934, DE AT, BE, CZ, DE, FI, FR, GB, HU, ?IT, LU, NL, SK 1994, DE CH, DE, GB 432 Family Species Family Species Feeding Regime Native range 1st record Invaded countries in invaded areas 1979, GB DE, GB, SI (?N) Habitat MediterraneanCentral Asia Phytophagous Mediterranean and/or Africa 1979, DE *AT, *BE, BG (?N), CH I, X (north), DE, *FI, GB, HU, *UA Barclay (2004), Rédei and Torma Fabaceae, cultivated and (2003), Wheeler (2001) uncultivated plants (polyphagous) Tingidae Dictyonota fuliginosa A. Costa, 1853 Elasmotropis testacea (Herrich-Schäffer, 1830) Stephanitis oberti (Kolenati, 1857) Phytophagous Phytophagous Phytophagous West Mediterranean Palaearctic 1954, CZ CZ Fabaceae (Cytisus) Asteraceae (Echinops) Ericaceae (Rhododendron, Vaccinium) North Palaearctic <1906?, DE E, I *AT, BE (?N), CZ (?N), I, X DE (?N), NL (?N) Nau (1980), Simon (2007) Kment (2006b) Kment (2006b) Bruers and Viskens (1999) True Bugs (Hemiptera, Heteroptera). Chapter 9.1 Phytophagous E, G, I Tamaricaceae (Tamarix) References Tuponia mixticolor (A. Costa, 1862) Pentatomidae Nezara viridula (Linnaeus, 1758) <1844, CZ CZ, DE (?N), ?PL I, X Hosts 433 A peer reviewed open access journal BioRisk 4(1): 435–474 (2010) RESEARCH ARTICLE doi: 10.3897/biorisk.4.57 BioRisk www.pensoftonline.net/biorisk Aphids (Hemiptera, Aphididae) Chapter 9.2 Armelle Cœur d’acier1, Nicolas Pérez Hidalgo2, Olivera Petrović-Obradović3 1 INRA, UMR CBGP (INRA / IRD / Cirad / Montpellier SupAgro), Campus International de Baillarguet, CS 30016, F-34988 Montferrier-sur-Lez, France 2 Universidad de León, Facultad de Ciencias Biológicas y Ambientales, Universidad de León, 24071 – León, Spain 3 University of Belgrade, Faculty of Agriculture, Nemanjina 6, SER-11000, Belgrade, Serbia Corresponding authors: Armelle Cœur d’acier (coeur@supagro.inra.fr), Nicolas Pérez Hidalgo (nperh@unileon.es), Olivera Petrović-Obradović (petrovic@agrif.bg.ac.rs) Academic editor: David Roy | Received 1 March 2010 | Accepted 24 May 2010 | Published 6 July 2010 Citation: Cœur d’acier A (2010) Aphids (Hemiptera, Aphididae). Chapter 9.2. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 435–474. doi: 10.3897/biorisk.4.57 Abstract Our study aimed at providing a comprehensive list of Aphididae alien to Europe. A total of 98 species originating from other continents have established so far in Europe, to which we add 4 cosmopolitan species of uncertain origin (cryptogenic). The 102 alien species of Aphididae established in Europe belong to 12 different subfamilies, five of them contributing by more than 5 species to the alien fauna. Most alien aphids originate from temperate regions of the world. There was no significant variation in the geographic origin of the alien aphids over time. The average introduction rate was 0.5 species per year since 1800. The mean number of newly recorded species per year decreased since 2000 but this pattern may change in the following years. Keywords alien, Hemiptera, Aphid, Aphididae, Europe 9.2.1. Introduction About 4700 species of Aphididae have been described worldwide (Remaudière and Remaudière 1997). About one third of these species are present in Europe. As for many other taxonomic groups, very few checklists of alien Aphididae have been available for Europe until recently. In 2002, Geiter et al. (2002) published a list of 131 species Copyright A. Cœur d’acier. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 436 Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) considered non-indigenous in Germany and Nobanis (2005) listed 34 species of nonnative Aphididae in its geographic area in 2005. Lampel and Gonseth (2005) listed 37 species alien to Switzerland in 2005 whilst Rabitsch and Essl (2006) listed 40 alien aphid species from Austria in 2006. The differences in the number of species considered non-indigenous clearly reflect differences in the composition of the fauna of each country, but also reflect differences in the definition of ‘alien’. Lampel and Gonseth (2005) considered only species of non-European origin whereas Geiter et al. (2002) included all species considered non-native to Germany. The goal of this work is to provide a first comprehensive list of Aphididae alien to Europe. Aphid species originating from one European country and introduced into another, i.e. species alien in Europe such as Diuraphis noxia (Kurdjumov, 1913) and Brachycorynella asparagi (Mordvilko, 1929), will not be considered in this work. These species may have an invasive status in the area where they were introduced but it appeared difficult to disentangle human- mediated introductions from natural expansion. To define the species present in Europe, we used the list of European Aphididae elaborated by Nieto Nafria for Fauna Europaea (Nieto Nafria et al. 2007). We compiled information about each species from published sources and experts to define their origin, i.e. European vs non-European. Among the references consulted, the lists cited above and the three comprehensive books by Blackman & Eastop (Blackman and Eastop 1994, 2000, 2006) proved to be particularly useful. Once a first list of alien aphids had been defined, we sought additional information, such as the date of first occurrence in Europe. June 2008 was the cut-off date for our literature survey. All the information collected for the 102 species considered is provided in Table 9.2.1. 9.2.2.Taxonomy of alien species The delineation of the taxa included under the family name Aphididae has varied over the last 50 years. Here, we use Aphididae sensu Eastop and Hille Ris Lambers (1976) and Remaudière and Remaudière (1997). Therefore, we did not consider Adelgidae and Phylloxeridae in this chapter. Taxonomy and nomenclature are as described by Remaudière and Remaudière (1997), Nieto Nafria et al. (1998), Quednau (1999, 2003), and Eastop and Blackman (2005). Some of the names cited in published studies could not be clearly attributed to a currently valid taxon and were therefore excluded. A total of 98 species present in Europe but originating from another continent have been listed to date, to which we can add four cosmopolitan species of uncertain origin (cryptogenic) (Table 9.2.1). For comparison, the European aphid fauna currently includes 1,373 species (Nieto Nafria et al. 2007), meaning that 7.4 % of the European aphid fauna is of alien origin. The 102 alien species of Aphididae established in Europe belong to 12 different subfamilies, most of which are already represented among the native entomofauna (Figure 9.2.1). However, three subfamilies (Greenideinae, Lizerinae and Neophyllaphidinae) were not known in Europe before introductions. Each of these three subfamilies Aphids (Hemiptera, Aphididae). Chapter 9.2 437 is represented by a single species. Greenidea ficicola is a member of the Greenideinae subfamily widespread in eastern regions and living on several species of Ficus. It was introduced into Italy in 2004 and seems to be widespread in Southern Europe (Italy, Spain and Malta) (Barbagallo et al. 2005a, Mifsud 1998). Paoliella eastopi, a species belonging to the Lizerinae described from Kenya, has only been found in one European country, England. All Paoliella species are of African origin. Neophyllaphis podocarpi, the only Neophyllaphidinae species known in Europe, originates from Asia and was recorded on Podocarpus in Italy in 1990 (Limonta 1990) but appears to have spread. Five subfamilies contribute more than five species to the alien fauna (Figure 9.2.1). The subfamily Aphidinae predominates, accounting for 59% of the alien Aphididae, followed by Calaphidinae (16%), Lachninae (5.8%), Eriosomatinae (4.8%) and Chaitophorinae (4.8%). These five subfamilies are also the most species-rich in native species. Each of the other seven subfamilies accounts for less than 1% of the alien Aphididae (Figure 9.2.1). The Hormaphidinae is the only subfamily represented by more alien than native species (4 species vs 1). The taxonomic composition of the alien entomofauna is highly diverse at genus level. The 102 alien species belong to 58 different genera (Table 9.2.1). Thirty-two (55%) of these genera are represented in Europe by only non-native species and 40 (69%) contribute only one species to the alien fauna. The genus Aphis is the most represented, with eight species. This is not surprising, given that this genus contains more than 10% of the world’s Aphididae and is abundant in all biogeographical regions of the world. This is not the case for another two species-rich genera, the North American Illinoia (seven alien species in Europe and 54 species worldwide) and the Asian Tinocallis (six alien species in Europe and 25 species worldwide). Although the genus Cinara is the second most species-rich genus in the world, with 222 species worldwide, three quarters of which being of non-European origin, surprisingly only three alien species from this genus are present in Europe 9.2.3.Temporal trends The date of the first record in Europe is known, with various degrees of precision, for 94 of the 102 alien aphid species (Table 9.2.1). The precise date of arrival is unknown for most species because their introduction was unintentional (see below 9.2.5) and large delays may occur between the date of introduction and the date of reporting. However, in certain cases, introduction is relatively well documented, available data suggesting that the date of the first report was close to the date of introduction. This is the case for recent introductions, such as the species detected and monitored by the permanent aerial suction-trap network “Euraphid”. This system of aphid flight surveys, based on a 12.2 m.-high suction trap, was developed by the Rothamsted Experimental Station in the 1960s (Taylor and Palmer 1972). This device is now used in several European countries, as part of integrated control networks, and has also proved useful for studies of the long-range dispersal of alates and for the regular detection of 438 Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) Figure 9.2.1. Taxonomic overview of the aphid species alien to Europe compared to the native European fauna and the world fauna. Subfamilies are presented in a decreasing order based on the number of alien species. Species alien to Europe include cryptogenic species. Data about native European aphids from Fauna europaea (Nieto Nafria et al. 2007); world data from Remaudière and Remaudière (1997). The number over each bar indicates the number of species observed per subfamily. aphid species new to the national or European fauna (Hullé et al. 1998). In France, a network of five such traps spread over the territory has been monitoring the aphid species trapped since 1978. This system detected four species new to Europe between 1984 and 1988 (Hullé et al. 1998): Essigella californica (Turpeau and Remaudière 1990), Klimaszewskia salviae (Leclant and Remaudière 1986), Myzocallis walshii (Remaudière 1989), and Tinocallis takachihoensis (Leclant and Remaudière 1986), and has monitored the extension of their geographical range in France. In a very small number of cases, more ancient introductions have also been documented, generally for important pest species. For example, the occurrence of Eriosoma lanigerum, a pest of apple trees originating from North America, was noted for the first time in a nursery in the outskirts of London in 1787 (Balachowsky and Mesnil 1935). The species was described by Hausmann in 1802, based on material from Germany, where aphids had been found in nurseries, causing extensive damage. In 1812, the species was found in France, by 1841, it was found in Italy and in 1870 it was reported in Switzerland. E. lanigerum has subsequently spread gradually to all temperate countries of the world (Balachowsky and Mesnil 1935, Marchal 1928). For most alien species, the date of first report sighting may not correspond to the date of introduction and secondary expansion. For example, the pest species Myzus persicae, Panaphis juglandis, and Chromaphis juglandicola were all reported for the Aphids (Hemiptera, Aphididae). Chapter 9.2 439 first time in Europe between 1800 and 1849, but they were probably introduced long before along with their host plants. The primary host of Myzus persicae, the peach tree, grown since classical times in the Mediterranean basin, was imported to Europe from Persia, but probably originated from western China, where it has been cultivated since 5,000 yr BP (Faust and Timon 1995). The host plant of Chromaphis juglandidola and Panaphis juglandis, the walnut, may have been introduced to Europe from Persia during the classical era, but this remains a matter of debate (Huntley and Birks 1983). Even for more recent introductions, the time lag between introduction and the first reported sighting may be considerable, particularly if the species concerned is not a pest. The date on which a taxonomic group was first recorded is therefore more likely to refer to the period during which it was studied for the first time. Börner between 1930 and 1952 made the largest single advance to studies of the aphid fauna of Europe, with the publication of “Europae Centralis Aphid” (Börner 1952). This catalysed intensive studies of the aphid fauna in various European countries over the following 20 years. The increase in the number of introduced species observed between 1950 and 1974 is partly attributable to this increase in taxonomic and faunistic activity. Bearing these biases in mind, and taking the first recorded sighting as a proxy for the date of introduction, the mean rate of introduction since 1800 was 0.5 species per year. A similar rate has also been reported for a more recent period (0.42 between 2000 and 2007). The number of introductions increased in the second half of the 20th century (Figure 9.2.2). The mean number of new records increased from 0.3–0.4 per year before 1950 to more than 1.3 per year between 1950 and 1974. The mean number of introductions per year has decreased since 2000, but this pattern may change again in the future. The three most recent alien aphid species introduced to Europe are Aphis illinoisensis, a Nearctic species and a pest of vineyards introduced into Crete in 2005 (Tsitsipis et al. 2005), Prociphilus fraxiniifolii, also of Nearctic origin, introduced into Europe in 2003, (Remaudière and Ripka 2003), and Greenidea ficicola, a tropical species, probably originating from Asia, introduced into Sicily in 2004 (Barbagallo et al. 2005a). 9.2.4. Biogeographic patterns 9.2.4.1 Origin of alien species A precise continent of origin was ascertained for 90.2% (92 species) of the alien Aphididae species, whereas 5.9% (six species) of the alien species were known only to be native to tropical or subtropical regions and 3.9% (four species) were of unknown origin (cryptogenic, Table 9.2.1, Figure 9.2.3). The cryptogenic species include the polyphagous pest species Myzus persicae and M. cymbalariae, which have a cosmopolitan distribution. Data concerning their host plant relationships and the distribution of other species of the genus Myzus, strongly 440 Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) Figure 9.2.2. Changes over time in the mean number of first sightings per year of aphid species alien to Europe from 1492 to 2007. The number to the right of the bar indicates the absolute number of species reported for the first time during the corresponding time period. suggest that these species originate from a continent other than Europe. Many other cosmopolitan species are not included in this list because they are thought to be of European origin, e.g. Acyrthosiphon pisum, Brevicoryne brassicae, although their origin is unclear and it remains possible that they were introduced into Europe by humans a long time ago. Most of the alien aphid species in Europe originate from temperate regions of the world. Asia and North America have contributed the largest numbers (each 43.1%, Figure 9.2.3). Most of the Asian species originated from temperate zones (32 species), and only four species (Cerataphis brasiliensis, Cerataphis orchidearum, Greenidea ficicola, and Stomaphis mordvilkoi) are known to have originated from tropical Asia. Only four alien species in Europe are of African origin. Two of these species come from North Africa (Cinara laportei and C. cedri) and two from subSaharan regions (Aloephagus myersi and Paoliella eastopi). No alien aphid species has yet been introduced into Europe from Australasia or South America. The proportions of aphids of different geographical origins in the alien aphid fauna of Europe have remained fairly constant over time (Figure 9.2.4) and seem to reflect the species diversity of the donor continents. Most of the described aphid species are of temperate origin, with Aleyrodidae and Coccoidea appearing to replace aphids in the tropics and subtropics (Dixon 1998). With only 219 (Remaudière et al. 1985) and 180 (Hales 2005) species, respectively, sub-Saharan Africa and Australia have a very poor aphid fauna. By contrast, 1,416 species are found in North America (Foottit et al. 2006) and 1,007 species are found in China (Qiao and Zhang 2004). Thus, the origins of the alien species in Europe might reflect regional species di- Aphids (Hemiptera, Aphididae). Chapter 9.2 441 Figure 9.2.3. Geographic origin of the alien species of Aphididae established in Europe. versity rather than preferential routes of introduction from North America and temperate Asia. 9.2.4.2. Distribution of alien species in Europe Alien Aphididae species are not evenly distributed within Europe (Figure 9.2.5). The number of alien species present in a country is significantly and positively correlated with the number of native species recorded in that country (r=0.6226, p<0.001). This may reflect differences in sampling intensity and in the number of local taxonomists. The number of alien species also seems to be weakly positively correlated with the total area covered by each country (r=0.3361, p=0.0182). Similarly, the number of native species is strongly correlated with the area of the country (r=0.6803, p<0.001). The top ten countries/regions within Europe with the largest numbers of recorded alien aphid species are: Great Britain (64), mainland France (63), mainland Italy (58), mainland Spain (56), Sicily (Italy) (45), Germany (44), Switzerland (37), Madeira (Portugal) (36), mainland Portugal (31), Czech Republic (29). Alien aphid species are well distributed across Europe, with 58% present in at least five European countries and 38% occurring in more than 10 countries or regions. The polyphagous pest species, Myzus persicae, Macrosiphum euphorbiae and Aphis gossypii are the most widely distributed alien species: they have been recorded in 43, 41 and 40 countries or regions, respectively. Only one of the 15 species occurring in more than half of the countries of Europe, Acyrthosiphon caraganae, is not considered to be a pest of crop plants. This species, probably originating from the Altai region, is now found in temperate regions throughout the Northern hemisphere, where it lives on woody Leguminosae, particularly Caragana and Colutea species. In 442 Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) Figure 9.2.4. Cumulative numbers of alien aphid species established in Europe, by year and by geographic origin most cases, it is not known whether the species expanded naturally after its establishment in a country, or whether the extension of its distribution was driven by repeated introductions from abroad. Thirteen of the 19 species present in only two European countries have discontinuous distributions, probably resulting from independent introductions. Thus, for example Ericaphis wakibae has been found in Great Britain and the Czech Republic, Chaitophotus populifolii in Germany and Serbia and Macrosiphum ptericolens in Poland and Great Britain. A continuous but restricted area may be accounted for by recent introductions, as for Aphis illinoisensis Shimer, 1866, a pest of grapevines introduced into Greece in 2005 (Tsitsipis et al. 2005). This species has extended its range from Crete to continental Greece and recently (2007) to the Mediterranean part of Montenegro (Petrovic, personal communication). Eight alien aphid species have each been found in only one European country. Four of these species are confined to England, two to Italy, one to Swirtzerland and one to the Ukraine. These species were all introduced before 2000 and have not spread elsewhere since. They may be unable to colonise a wider geographical area in Europe, they may have disappeared or they may simply have been overlooked. 9.2.5. Main routes and vectors for introduction into Europe No cases of intentional introduction of aphids into Europe are known. All alien species were therefore introduced accidentally. In a very small number of cases, the pathway Aphids (Hemiptera, Aphididae). Chapter 9.2 443 Figure 9.2.5. Comparative colonization of continental European countries and islands by Aphididae species alien to Europe. Archipelago: 1 Azores 2 Madeira 3 Canary islands. and vector are precisely known. For example, two Japanese aphids, Tinocallis ulmiparvifoliae and T. zelkovae were introduced into Europe in 1973 with their hosts, bonsai trees that were imported into Great Britain directly from Japan. The infested bonsai trees had been in Great Britain for about six months before the aphids were detected, and were growing in slatted wood buildings providing no effective physical barrier to insect dispersal (Prior 1971). In most cases, it is difficult to identify the vector of accidental introductions; most have been inferred from the known biological requirements of the aphid species. Most Aphididae have a high level of host-plant specificity and most alien species are therefore thought to have been introduced into Europe with their host plants. For example, the Takecallis species included in our list feed on bamboos of Asian origin. The Ne- 444 Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) a b c Figure 9.2.6. Some alien aphids. a Spiraea aphid, Aphis spiraephaga. (Credit: Olivera Petrović-Obradović) b Walnut aphid, Chromaphis juglandicola. (Credit: Olivera Petrović-Obradović) c Woolly apple aphid, Eriosoma lanigerum. (Credit: Olivera Petrović-Obradović). arctic aphid Prociphilus fraxinifolii has recently been detected in Budapest (Hungary) (Remaudière and Ripka 2003), but only on the North American red ash tree, Fraxinus pennsylvanica Marsh. This aphid has not been found on European ash planted in the same area. Two oriental species, Reticulaphis distylii and Greenidea ficicola, live on several species of Ficus, all originating from tropical regions. These Ficus species have been planted as ornamental trees in the warmest areas of the Mediterranean basin (Barbagallo et al. 2005a). These two species of aphids are found on tropical fig trees, but never on Ficus carica, the only European species of this genus. All these alien species are thought to have been introduced into Europe through trade, but the aphid species may have been introduced several years after their hosts. Impatientinum asiaticum is a species originating from Central Asia. It was introduced into Europe in 1967, whereas its host, Impatiens parviflora DC. was introduced into Europe much earlier, in the 19th Century, subsequently escaping from botanic gardens to establish itself as a common weed. The aphid was not introduced at the same time as its host plant in this case because the host plant is an annual, which was imported in the form of seeds. The aphid arrived more than 100 years later, probably on an aeroplane (Holman 1971, TambsLyche and Heie 1973). Another example is provided by Rhopalosiphoninus latysiphon, Aphids (Hemiptera, Aphididae). Chapter 9.2 445 a pest species particularly damaging to potato. This species was not introduced into Europe until the end of the 1st World War, long after the introduction of its host plant, and was transported with potatoes from the USA. It was subsequently found in Italy (1921), the Netherlands (1930), Germany (1943), England (1945), Switzerland and Austria (1949) (Remaudière 1952). Finally, we cannot exclude the possibility that some species originating in areas close to Europe may have been transferred into Europe by wind, air streams or windstorms. For example, it is difficult to determine whether Cinara laportei and C. cedri were transferred with their host, the Atlas cedar, which was planted in Europe, or whether these species colonised Europe following their introduction via wind or air streams. 9.2.6.The ecosystems and habitats most frequently invaded All aphids are phytophagous and their distribution is limited by the presence of their host plants. Aphid species with a limited spectrum of host plants of exotic origin, not present at natural sites, are restricted to artificial habitats, such as agricultural land, greenhouses and parks and gardens. For example, Illinoia liriodendri and Neophyllaphis podocarpi feed on exotic trees (Liriodendron tulipifera L. and Podocarpus spp., respectively). As a result, these aphids are restricted to parks, gardens and city areas in which these trees have been planted in Europe. Similarly, Cinara cedri and C. laportei which feed specifically on Cedrus are restricted to forest areas in which their hosts have been planted. Other species restricted to artificial habitats include tropical and subtropical aphids present only in indoor conditions in Europe. These species were included in the list because it is clear that they have become established in Europe. For example, Cerataphis spp., particularly C. lataniae and C. orchidearum have repeatedly been found in European greenhouses (Chapin and Germain 2005). Similarly, Sitobion luteum and Pentalonia nigronervosa are considered to have been introduced into hothouses in Europe (Blackman and Eastop 2000). Another subtropical Cerataphis, C. brasiliensis, has recently been found established outdoors in the south of the France (Chapin and Germain 2005, 2004). Some aphid species have a less limited host range spectrum. They can adapt to new hosts when introduced and may disperse in natural habitats. Cinara curvipes, a species recently introduced into Europe, is known to feed on various species of Abies in its native area (North America). In Europe, it is found on North American Abies species, but also on native Abies species and has recently been reported on many other conifers, including Picea, Tsuga, and Pinus (Scheurer and Binazzi 2004). C. curvipes is found in parks, gardens and forests. It could potentially colonise all European coniferous forests. Finally, polyphagous aphids, notably Myzus persicae, M. ascalonicus, M. ornatus, Macrosiphum euphorbiae and Aphis gossypi, have established themselves on many native plants in natural habitats. Most of the alien aphids seem to have become established in the European environment and habitats. However, some species, such as Paoliella eastopi and Macrosi- 446 Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) phum ptericolens have been recorded only once or twice, and it remains unclear whether these species are truly established. Other species, such as Rhopalosiphum parvae Hottes & Frison (1931), a North American aphid found in Sicily in 1982 (Barbagallo and Stroyan 1982), or Tuberocephalus higansakurae hainnevilleae Remaudière & Sorin, 1993, detected in France in 1990 on trees of Prunus subhirtella Miq. var. pendula Y.Tanaka imported from Japan (Remaudière and Sorin 1993), have been observed in Europe but have since been eradicated. Such species are not included in our list. 9.2.7. Ecological and economic impact Most of the alien Aphididae are recognised pests, feeding on crops, ornamental plants and forest trees in Europe. Other alien Aphididae species may have remained undetected because they feed on plants that are not commercially exploited. As for most insects, much more is known about the economic impact of aphids than about their ecological impact. Aphids cause direct (sap-feeding, deformation of their hosts) and indirect (transmission of plant diseases, deposition of honeydew on the leaves) damage. The economic impact of each species depends on (i) the type and extent of the damage caused and (ii) the economic importance of the host. Of the 102 alien aphid species in Europe, 52 are recognised pests of agricultural and horticultural crops (Blackman and Eastop 2000). The polyphagous species Myzus persicae, Macrosiphum euphorbiae and Aphis gossypii attack a wide range of vegetable crops, both indoors and outdoors. They are vectors of many viral diseases and are probably the aphids with the greatest economic impact in vegetable crops (Lampel and Gonseth 2005). European orchards are attacked by several alien aphid species. Apple trees can be severely damaged by the North American wolly aphid Eriosoma lanigerum and the Asian species Aphis spiraecola. The recent introduction of Toxoptera citricidus into the Iberian Peninsula (Portugal and Spain) (Ilharco et al. 2005) poses a serious threat to Mediterranean citrus fruit production because this aphid is the principal vector of the Triteza closterovirus of Citrus. Citrus trees in Europe are also the hosts of Aphis spiraecola and Toxoptera aurantii, two polyphagous species also capable of transmitting this closterovirus, albeit with a lower efficiency. The recent introduction and rapid dispersion of Aphis illinoiensis, a grapevine aphid, poses a particular threat to viticulture in the Mediterranean area (Remaudière et al. 2003, Tsitsipis et al. 2005). Some alien aphids attack agricultural crops, often as potential virus vectors. Rhopalosiphum maidis is known as a pest of maize and other grain crops in Europe and transmits the persistent luteovirus “yellow dwarf ” virus of barley. The grass aphid, Hysteroneura setariae Thomas, 1878, has recently been recorded in Spain (Meliá Masiá 1995). Its impact it difficult to predict because it usually lives on wild grass species, but it may occasionally infect cereals and can transmit several viral diseases to these crops. Macrosiphum albifrons is a widespread species in North America that has been introduced into Europe (Stroyan 1981) where the damage it causes to Aphids (Hemiptera, Aphididae). Chapter 9.2 447 lupins (Ferguson 1994) has stimulated recent research (Blackman and Eastop 2000). Finally, Acyrthosiphon kondoi, which currently has a restricted distribution in Europe, is known to be a serious pest of lucerne (Blackman and Eastop 2000). Exotic Aphididae are not considered to be serious pests of forest species in Europe (EUROFOR 1994) by contrast to the major damage caused to agricultural and horticultural crops. However, some species may cause economic losses. For example, the North African species Cinara cedri and C. laportei have been reported to damage plantations of Cedrus in southern France (Emonnot et al. 1967, Fabre 1976). Finally, in addition to their measurable economic impact, some alien aphids may have an aesthetic impact. The production of abundant honeydew and the distortions induced by feeding may significantly modify the appearance of the foliage of ornamental plants in parks and private gardens. Appendiseta robiniae has such an aesthetic impact on Robinia pseudacacia L., as does Prociphilus fraxinifolii on the red ash tree Fraxinus pennsylvanica and Illinoia liriodendri on Liriodendron tulipifera. 9.2.8. Conclusion There are several possible reasons for the overrepresentation of Aphididae in the alien insect fauna of Europe. First, aphids are phytophagous insects and many are pests of economically important host plants (Blackman and Eastop 2000). For this reason, many studies are carried out on the distribution, taxonomy and biology of this family. New alien species of Aphididae are therefore more likely to be detected than new members of other taxonomic groups, and this effect is enhanced by standard phytosanitary procedures. Second, aphids have the ability to reproduce both parthenogenetically and sexually. Several species can reproduce exclusively by parthenogenesis, and all species can potentially maintain parthenogenetic populations throughout the year in areas of mild climate. Consequently, very few introduction events, and theoretically even the introduction of a single parthenogenetic female, may lead to the development of a population and the establishment of an alien species. Third, although aphids, as a group, are cosmopolitan, they are most strongly represented in temperate regions. Consequently, most of the World’s aphids live in climatic conditions similar to those of Europe and are therefore preadapted to establishment where suitable hostplants are present. Moreover, global warming is also likely to promote the survival of alien tropical and subtropical species, at least locally (e.g. along the Mediterranean coast). Finally, aphids are small insects easily transported around the globe with plant materials. 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II 11: 177–193. 458 Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) Szelegiewicz H (1978) Przeglad systematyczny mszyc polski. Zeszyty problemowe postepow nauk rolniczych. Polska Akademia Nauk 208: 40pp. Tambs-Lyche H, Heie O (1973) An Asian aphid suddenly appearing in large numbers on Impatiens parviflora in Europe. Entomologiske Meddelelser 41: 167–173. Tashev D (1961) Novi listni vushki (Homoptera, Aphididae) za faunata na Bulgaria, godishnik na SU, biologo-geologo-geografski fakultet. 1-biologia 1958/1959, 53: 157–162. Tashev D (1964) Novi za faunata na Bulgaria listni bushki (Homoptera Aphididae), godishnik na SU biologo-geologo-geografski fakultet. 1-biologia 1961/1962, 56: 179–190. Tashev D (1982) A list of the aphids from Bulgaria. Annuaire de Universite de Sofia “Kliment Ohridski” Faculte de Biologie; Livre 1 – Zoologie: 20–35. Tavares JS (1900) As zoocecidias Portuguesas. Anais das Ciencias Naturaes 7: 15–108. Tavares JS (1905) Synopse das zoocecidias portuguesa. 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Status Feeding Regime Native range A phytophagous Acyrthosiphon Acyrthosiphon kondoi Shinji, 1938 A phytophagous Acyrthosiphon Acyrthosiphon primulae Theoblad 1913 A phytophagous Asia1913, GB Temperate BG, CH, CZ, DK, ES, FR, DE, GB, GR, IE, IT, IT-SIC, NL, PT, SE, SK I2, J100 Primula Aloephagus myersi Essig, 1950 A phytophagous Africa 1937, GB ES, FR, GB, GR, IT, IT-SIC I2, J100 Aloe, Haworthia, Gasteria Aphis Aphis forbesi Weed, 1889 A phytophagous North America 1928, FR I1, J100 Fragaria Aphis Aphis gossypii Glover 1877 A phytophagous Tropical, subtropical <1758 Unknown AL, AT, BE, BG, CH, CZ, DE, DK, EE, ES, FR, HR, HU, IT, LV, MD, PL, RO, RS, SK AL, AT, BE, BG, CH, CY, CZ, DE, DK, EE, ES, ESBAL, ES-CAN, FI, FR, FRCOR, GB, GR, GR-CRE, HR, HU, IL, IT, IT-SAR, ITSIC, LT, LV, MD, MK, NO, PL, PT, PT-AZO, PT-MAD, RO, RS, RU, SE, SK, UA I2, I1, J100, E, F Polyphagous (mainly Cucurbitaceae, Rutaceae and Malvaceae) References Cholodkovsky (1907), Hržič (1996), Mordvilko (1914), Petrović (1998), Remaudière (1951), Tashev (1982) Eastop (1971), Nieto Nafria e al. (2007), Tsitsipis et al. (2007) Heie (1994), Remaudière (1952), Theobald (1913), Tsitsipis et al. (2007) Eastop (1956), Leclant (1978), Micieli De Biase (1988), Tsitsipis et al. (2007) Balachowsky (1933), Heie (1986), Paillot (1928) Blackman and Eastop (2006), Buckton (1879), Theobald (1927), Tschorbadjiev (1924), Vasilev (1910) Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) Acyrthosiphon Acyrthosiphon caraganae Cholodkovsky 1908 1st record Invaded countries Habitat Hosts in invaded areas F, I2 Caragana. Asia1907, RU AL, AT, BG, CH, CZ, DE, Temperate DK, EE, ES, FI, FR, GB, GR, other Fabaceae HU, IT, IT-SIC, LT, LV, MK, NO, NL, NO, PL, RO, RS, RU, SE, SI, SK, UA Asia< 2004, FR-COR, GR E, I1 Medicago Temperate FR-COR Species 460 Table 9.1.1. List and main characteristics of Aphididae species alien to Europe. Status: A: Alien to Europe; C: cryptogenic species. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Only selected references are given. Last update February 2010. Species Status Feeding Regime Native range 1st record Invaded countries in invaded areas 2005, GR- GR-CRE, ME CRE A phytophagous North America Aphis Aphis spiraecola Patch, 1914 A phytophagous Asia1961, PT Temperate Aphis Aphis spiraephaga F.P. Müller, 1961 A phytophagous Asia1955, CZ Temperate Aphis Aphis spiraephila Patch, 1914 Aphis Bursaphis oenotherae oenotherae Oestlund 1887 Aphis catalpae Mamontova, 1953 A phytophagous phytophagous phytophagous North America North America Asia A A FB E, I2, FA, FB, G Hosts References Vitis Petrović-Obradović et al. (in press), Tsitsipis et al. (2005) Polyphagous Blackman and Eastop (Citrus, apple, (2000), Blackman and Spiraea) Eastop (2007), Ilharco (1968b) I2 Spiraea 1955 UA AT, BG, CH, DE, ES, ESBAL, ES-CAN, FR, FR-COR, GB, GR, HR, IL, IT, IT-SAR, IT-SIC, MT, PT, PT-AZO, PT-MAD, RS, UA AL, AT, CH, CZ, DE, DK, ES, FI, FR, HR, HU, IT-SIC, LT, LV, MD, MK, PL, PT, RO, RU, SE, SI, SK, UA UA I2 Spiraea 1972, DE FR, DE, GB, IT-SIC, PL, RS G3, I2 Oenothera 0 HU, UA I2 Catalpa A phytophagous North America 1978, IT BE, BG, CH, CZ, DE, ES, ES-BAL, FR, FR-COR, GB, GR, HR, HU, IT, IT-SIC, NL, RS, SK I2, G5 Robinia Brachycaudus Mordvilkomemor rumexicolens (Patch, 1917) A phytophagous North America 1953, GB BE, CZ, DE, DK, ES, ESCAN, FI, FR, GB, IT, ITSAR, IT-SIC, MK, NL, NO, PL, PT, PT-MAD, RO, RU, SE, SK, UA H5, I1 Rumex; other Polygonaceae 461 Appendiseta robiniae (Gillette, 1907) Heie (1986), Holman (1971), Ilharco (1968b), Ilharco (1973), Tashev (1964) Holman (1971), Nieto Nafria et al. (2007) Barbagallo (1994), Műller (1974) Mamontova (1955), Petrović-Obradović et al. (in press), Ripka (2001) Arzone and Vidano (1990), Lampel (1983), Leclant and Remaudière (1986), Micieli De Biase and Calambuca (1979), Pati and Tomatore (1988), Petrović (1998) Barbagallo (1994), Barbagallo and Stroyan (1982), Heie (1973), Holman (1965), Ilharco (1974), Stroyan (1956) Aphids (Hemiptera, Aphididae). Chapter 9.2 Aphis Aphis illinoisensis Shimer 1866 Habitat Status Feeding Regime Native range 1st record Invaded countries in invaded areas 1981, PT- ES-CAN, FR, PT-MAD MAD Habitat Hosts A phytophagous AsiaTropical I2, J100 Palms Cerataphis lataniae (Boisduval, 1867) A phytophagous Asiatropical 1867, FR CZ, ES-CAN, DE, FR, GB, IT, PL Cerataphis orchidearum (Westwood, 1879) A phytophagous AsiaTropical 1906, BE BE, ES, FI, FR, GB, HU, PT- J100 MAD, RU, SE Chaetosiphon Pentatrichopus fragaefolii (Cockerell, 1901) A phytophagous North America 1912, GB I1, J100 Fragaria Chaitophorus populifolii (Essig, 1912) A phytophagous North America 1956, DE AT, BE, BG, CH, CZ, ES, ES-CAN, FR, DE, GB, HR, HU, IE, IL, IT-SIC, IT, LV, MK, NL, NO, PT, PT-AZO, PT-MAD, RO, RS, SI DE, RS I2 Populus Chaitophorus saliapterus quinquemaculatus Bozhko 1976 A phytophagous Asia 1953,UA IT, UA F9 Salix I2, J100 Areca, Musa Orchids Chapin and Germain (2005), Germain and Chapin (2004), Ilharco (1984), Pérez Hidalgo et al. (2000) Boisduval (1867), Chapin and Germain (2005), Pérez Hidalgo et al. (2000) Germain and Chapin (2004), Heie (1980), Ilharco (1973), Ilharco (1974), Schouteden (1906) Balachowsky (1933), Theobald (1912) Pintera (1987), Poljaković-Pajnik and Petrović-Obradović (2009) Binazzi and Barbagallo (1991), Bozhko (1976), Pintera (1987) Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) Cerataphis brasiliensis (Hempel, 1901) References 462 Species Species Status Feeding Regime Native range Chromaphis juglandicola (Kaltenbach, 1843) A phytophagous Cinara Cedrobium laportei (Remaudière, 1954) A phytophagous Cinara Cinara cedri Mimeur, 1936 A phytophagous Africa 1974,IT Cinara Cinara curvipes (Patch, 1912) A phytophagous North America Drepanaphis acerifoliae (Thomas, 1878) A phytophagous Ericaphis scammelli Mason 1940 A Ericaphis wakibae (Hottes, 1934) A Habitat I2, G5 Hosts Juglans G3, G5, Cedrus I2 Cedrus. 1999, GB BE, CH, DK, ES, FR, GB, I2, G5 HR, HU, IL, IT, IT-SAR, ITSIC, RS, SI CZ, CH, DE, GB, RS, SK, SL I2 North America 1992, IT IT , ES I2 Acer phytophagous North America 1964, GB FR, GB, IT, NL I1, I2 Vaccinium phytophagous North America 1963, GB CZ, GB I1, B3 Fragaria Abies References Balachowsky and Mesnil (1935), Heie (1982), Kaltenbach (1843), Schouteden (1906), Theobald (1927) Covassi (1971), Emonnot et al. (1967), Leclant (1978) Covassi and Binazzi (1974), Fabre (1976) Angst et al. (2007), Jurc et al. (2009), Martin (2000), Poljaković-Pajnik and Petrović-Obradović (2002), Scheurer and Binazzi (2004) Lozzia and Binaghi (1992), Pérez Hidalgo et al. (2008) Barbagallo et al. (1999), Barbagallo et al. (1998), Prior (1971) Stroyan (1972) Aphids (Hemiptera, Aphididae). Chapter 9.2 1st record Invaded countries in invaded areas Asia< 1758 AT, BE, BG, CH, CZ, DE, Temperate Unknown DK, ES, ES-CAN, FR, FRCOR, GB, HR, HU, IL, IT, IT-SAR, IT-SIC, MD, MK, PL, PT-AZO, PT-MAD, PT, RO, RS, SE, SI, SK, UA Africa 1967, FR ES, FR, GB, IT, IT-SAR, ITSIC, NL, PT, SI 463 Status Feeding Regime Native range Eriosoma lanigerum (Hausmann, 1802) A phytophagous North America Essigella Essigella californica (Essig, 1909) A phytophagous North America Greenidea Greenidea ficicola Takahashi 1921 Hysteroneura setariae (Thomas, 1878) A phytophagous phytophagous AsiaTropical North America 2004, IT ES, IT, IT-SIC I2 Ficus 1982, PTMAD ES, PT-MAD E, I Prunus, fruit trees, Graminae Idiopterus nephrelepidis Davis, 1909 A phytophagous Tropical, subtropical 1915, GB I2, J1, J100 Tropical ferns indoors Illinoia Illinoia andromedae (MacGillivray, 1958)] Illinoia Illinoia azaleae Mason, 1925 A phytophagous phytophagous North America North America 1960, GB BE, CH, CZ, DE, DK, ES, ES-CAN, FR, GB, GR, IE, IL, IT, IT-SIC, NL, PL, PT, PT-AZO, PT-MAD, PT, RU, SE, SI, SK GB I2 Asteraceae A A 1950, GB References Balachowsky and Mesnil (1935), Marchal (1928) Aguiar and Ilharco (2001), Turpeau and Remaudière (1990) Barbagallo et al. (2005a), Mifsud (1998) Blackman and Eastop (2006), Meliá Masiá (1995), Van Harten (1982) Heie (1994), Laing (1923), Theobald (1926), Tsitsipis et al. (2007) Eastop (1962), Stroyan (1964) AT, CH, CZ, DK, ES, FI, FR, I2, J100 Rhododendron; Biurrun and Nieto Ericaceae DE, GB, HU, IT, IT-SIC, Nafría (1987), Heie NL, PL, PT, PT-AZO, PT(1995), Ilharco (1968b), MAD, RO, RU, SE, SI Stroyan (1950) Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) 1st record Invaded countries Habitat Hosts in invaded areas I, I1 Malus; 1787, GB AL, AT, BE, BG, CH, CY, orchard trees CZ, DE, DK, ES, ES-CAN, FR, DE, GB, GR, HR, HU, IE, IL, IT, IT-SAR, IT-SIC, LT, LV, MD, NO, PL, PT, PT-AZO, PT-MAD, RO, RU, RS, SE, SI, SK, UA 1988, FR ES,FR, IT, IT-SAR, IT-SIC, G5, I2 Pinus radiata, PT-MAD P. pinaster 464 Species Species Status Feeding Regime Native range A phytophagous North America Illinoia Illinoia liriodendri (Monell, 1879) Illinoia Illinoia morrisoni (Swain, 1918) A phytophagous phytophagous North America North America A 1998, FR 1960, GB Illinoia Masonaphis lambersi (MacGillivray, 1960) A phytophagous North America Illinoia Masonaphis rhododendri (Wilson, 1918)] Impatientinum Impatientinum asiaticum Nevsky 1929 A phytophagous phytophagous North 1939, GB America Asia1967, RU Temperate phytophagous phytophagous North 1954, DE America Asia1907, PT Temperate Iziphya flabella (Sanborn, 1904) Macrosiphoniella Macrosiphoniella sanborni (Gillette, 1908) A A A 1971, NL Habitat Hosts References Eastop (1962), Stroyan I2, J100 Astereacae (Aster, Erigero., (1964), Ward (1961) Solidago) DE, FR, GB, IT, SI G5, I2 Liriodendron Limonta (2001), Rabasse et al. (2005b) FR, GB I2 Cupressus Eastop (1962), Prior (1975), Rabasse et al. (2005b) Stroyan (1964) BE, CH, CZ, DK, GB, NL, I2 Rhododendron, Aguiar and Ilharco NO, PT-MAD, SK Kalmia (2001), Heie (1995), Hille Ris Lambers (1973), Stroyan (1971), Stroyan (1972) GB, NL, SK I2, J100 Rhododendron Eastop (1956), Heie (1994), Stroyan (1950) AT, CH, CZ, DE, DK, EE, G, I2, Impatiens Heie (1994), Holman FI, FR, GB, LV, PL, RO, RU, X25 (1971), Ilharco (1968b), SE, SI, SK Tambs-Lyche and Heie (1973) DE, UA I2 Carex Quednau (1954) AL, AT, BE, BG, CH, CY, I2, J100 ChrysanCZ, DK, ES, ES-CAN, FI, themum FR, DE, GB, GR, HR, IE, IL, IT, IT-SIC, LT, LV, MD, NO, PL, PT, PT-AZO, PT-MAD, RO, RS, RU,SE, UA Aphids (Hemiptera, Aphididae). Chapter 9.2 Illinoia Illinoia goldamaryae (Knowlton, 1938) 1st record Invaded countries in invaded areas 1960, GB GB Balachowsky and Mesnil (1935), Del Guercio (1911), Del Guercio (1913), Holman (2009), Ilharco (1968b), Ilharco (1974), Theobald (1926) 465 Status Feeding Regime Native range 1st record Invaded countries in invaded areas 1981, GB AT, BE, CH, DE, FR, GB, GR, IE, IT, IT-SIC, SE A phytophagous North America Macrosiphum Macrosiphum euphorbiae (Thomas, 1878) A phytophagous North America 1917, GB Macrosiphum Macrosiphum ptericolens Patch, 1919 A phytophagous North America 1972, GB Megoura lespedezae (Essig & Kuwana, 1918) A phytophagous Melanaphis bambusae (Fullaway, 1910) A Melaphis rhois (Fitch, 1866) A I1, I2 Hosts Lupinus, Fragaria AL, AT, BE, BG, CH, CZ, DK, EE, ES, ES-CAN, FI, FR, FR-COR, DE, GB, GR, HR, HU, IS, IE, IL, IT, ITSAR, IT-SIC, LT, LV, MD, MK, MT, NO, PL, PT, PTAZO, PT-MAD, RO, RS, RU, SE, SI, SK, UA GB, PL E, F, I, J, Polyphagous J100 (vegetables, Fragaria) Asia1994, CH Temperate CH I1 phytophagous Asia1961, IT Temperate ES, FR, GR, IT-SIC, IT, PT, PT-MAD, RS I2 Pteridium aquilinum (bracken) Polyphagous (vegetables; Lespedeza, Japanese clover) Bambusa phytophagous North America GB, SE I2 Rhus 1902, GB G References Carter et al. (1984), Hullé et al. (1998), Meier and Schweizer (1987), Piron (1987), Stroyan (1981) Blackman and Eastop (2000), Eastop (1958) Holman (2009), Lawton and Eastop (1975) Giacalone and Lampel (1996) Hille Ris Lambers (1966), Nieto Nafria et al. (2007) Blackman and Eastop (1994), Theobald (1918), Theobald (1929) Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) Macrosiphum Macrosiphum albifrons Essig, 1911 Habitat 466 Species Species Status Feeding Regime Native range 1st record Invaded countries in invaded areas 1985, ES IL, ES Habitat G5 A phytophagous North America Monelliopsis caryae (Monell ex Riley & Monell, 1879) A phytophagous North America 1984, FR ES, FR, HU, IL, IT, PT G5 Monelliopsis pecanis Bissell, 1983 A phytophagous North America 1995, PTMAD IT-SIC, PT-MAD G5 Myzaphis turanica Nevsky, 1929 Myzocallis Lineomyzocallis walshii (Monell ex Riley & Monell, 1879) C phytophagous phytophagous Cryptogenic North America 1976, ES ES,FR, GB, IT-SIC, SE I2 1988, FR BE, CH, CZ, DE, ES, FR, HU, IT, IT-SIC, RS G, I2 Myzus Myzus hemerocallis Takahashi, 1921 A phytophagous Asia1990, FR Temperate FR, PT-MAD I2 Myzus Myzus ornatus Laing, 1932 A phytophagous Asia1932 GB Temperate AL, AT, BE, BG, CH, CZ, DE, DK, EE, ES, ES-CAN, FI, FR, FR-COR, GB, GR, HR, HU, IE, IT, IT-SAR, IT-SIC, LV, NO, PL, PT, PT-AZO, PT-MAD, RO, RS, RU, SE, SI, SK I, J100, X8 A References Juglans, Carya Hermoso de Mendoza (1988), Nieto Nafría and Mier Durante (1998) Juglans, Carya Hullé et al. (1998), Mier Durante and Pérez Hidalgo (2002) Carya Aguiar and Ilharco (1997), Barbagallo and Suma (1999) Rosa rugosa Meliá Masiá (1998), Patti (1983) Quercus rubra Hullé et al. (1998), Petrović-Obradović et al. (2007), Remaudière (1989) Hemerocallis Aguiar and Ilharco (1997), Remaudière and Munoz Viveros (1992) Polyphagous Blackman and Eastop (Prunus (2000), Ilharco (1969), cornutaLaing (1932) primary host); many herbaceous plants and vegetablessecondary host) Aphids (Hemiptera, Aphididae). Chapter 9.2 Monellia caryella (Fitch, 1855) Hosts 467 Status Feeding Regime A phytophagous Myzus Nectarosiphon ascalonicus Doncaster, 1946 A phytophagous Myzus Nectarosiphon persicae Sulzer 1776 C phytophagous Myzus Sciamyzus cymbalariae Stroyan, 1954 C phytophagous Nearctaphis bakeri (Cowen ex Gillette & Baker, 1895) A phytophagous 1st record Invaded countries in invaded areas Asia1946, CH AL, AT, BA, BE, BG, CH, Temperate CZ, DE, ES, FR, FR-COR, MK, DE, GB, GR, HR, HU, IT, IT-SIC, PL, RO, RS, RU, SI, SK Asia1941, GB AL, AT, BE, BG, CH, CZ, Temperate DE, DK, ES, ES-CAN, FI, FR, MK, DE, GB, GR, HR, IE, IS, IT, LT, LV, NL, NO, PL, PT, PT-AZO, RO, RS, RU, SE, SK Crypto<1758 AL, AT, BE, BG, CH, CY, genic Unknown CZ, DK, EE, ES, ES-BAL, ES-CAN, FI, FR, FR-COR, MK, DE, GB, GR, GR-CRE, HR, HU, IE, IT, IT-SAR, IT-SIC, LT, LV, ME, MD, MT, NO, PL, PT, PT-AZO, PT-MAD, RO, RU, RS, SE, SI, SK, UA Crypto1950, GB BE, CH, CZ, DE, ES, FR, genic GB, GR, IT, PT-AZO, PTMAD North 1964,FR AL, CH, ES, ES-BAL, FR, America DE, GB, GR, IT, IT-SIC, PT, PT-AZO, SK UA Habitat Hosts References I2, G5 Prunus persicae, Clematis Blackman and Eastop (2000), Börner (1952), Hille Ris Lambers (1947) I2, E Fragaria, Allium Börner (1952), Doncaster (1946) G5 Polyphagous Balachowsky and Mesnil (1935), Blackman and Eastop (2000), Boisduval (1867), Buckton (1876), Koch (1855), Macchiati (1883), Schouteden (1906), Theobald (1926) I Polyphagous I, E Maloideae (primary hosts) and Fabaceae (secondary hosts; e.g. Trifolium) Blackman and Eastop (2000), Ilharco (1974), Stroyan (1954) Heie (1992), Leclant (1967), Stroyan (1972) Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) Myzus Myzus varians Davidson, 1912 Native range 468 Species Species Status Feeding Regime Native range Neomyzus circumflexus Buckton 1876 A phytophagous Asia Neophyllaphis podocarpi Takahashi, 1920 Neotoxoptera formosana (Takahashi, 1921) A phytophagous phytophagous AsiaTemperate Asia 1994, FI Neotoxoptera oliveri (Essig, 1935) Neotoxoptera violae (Pergande, 1900) A A A phytophagous phytophagous Panaphis juglandis (Goeze, 1778) A phytophagous Paoliella eastopi Hille Ris Lambers, 1973 A phytophagous DE, FI, FR, GB, IT, NL, PT-MAD Habitat Hosts References I2, J100 Polyphagous flower crops Blackman and Eastop (2000), Buckton (1876), Ilharco (1969) I2 Podocarpus Limonta (2001) I1, J1, J100 Allium 469 Aguiar and Ilharco (2001), Barbagallo Ciampolini (2000), Blackman and Eastop (2000) Asia1959, PT ES, FR, IT-SIC, PT-MAD, I1, J100 Viola, Allium Ilharco (1960), Ilharco Temperate PT, RS (1968b) Asia1939, IT ES, ES-CAN, FR, IT IT-SIC I2 Viola Barbagallo and Coccuzza Temperate (1998), Germain and Deogratias (2008) Silvestri (1939) Asia <1758 AL, AT, BE, BA, BG, CH, I2, G5 Juglans Blanchard (1840), unknown CZ, DK, ES, ES-CAN, FR, Goeze (1778), Ilharco FR-COR, DK, GB, GR, HR, (1968a), Kaltenbach HU, IL, IT-SIC, IT, MD, PL, (1843), Malkov (1908), PT, RO, RS, SE, SI, SK, UA Schouteden (1906), Walker (1848) Africa <2004, GB GB U Passionfruit in Nieto Nafria et al. native range (2007) (Kenya) Aphids (Hemiptera, Aphididae). Chapter 9.2 1st record Invaded countries in invaded areas 1876, GB AL, AT, BE, BG, CH, CZ, DE, DK, EE, ES, ES-CAN, FI, FR, FR-COR, GB, HR, HU, IE, IT, IT-SIC, LT, LV, MD, NL, NO, PL, PT, PTAZO, PT-MAD, RO, RU, SE, UA 1990, IT IT Status Feeding Regime Native range 1st record Invaded countries in invaded areas 1966, PT- GB, PT-AZO, PT-MAD MAD BG, HU, RS G, G5 Fraxinus Asia1975, IT Temperate AL, BG, CY, ES, FR, GR, IT, IT-SIC, RO, RS, UA I2, G5 Prunus; fruit trees (peach) North <2004, UA America Asia1998, PT Temperate Tropical, 1934, ES subtropical EE, UA G Populus ES, PT I2, G5 Ficus AL, AT, BA, BG, CH, CZ, DE, DK, ES, ES-CAN, FI, FR, GB, GR, HU, IL, IT, IT-SIC, LV, NL, PL, PT, PTMAD, RO, RS, SE, SI, SK I2, J100 Fragaria, Rosa (in greenhouses in Central Europe) A phytophagous Tropical, subtropical Periphyllus californiensis (Shinji, 1917) A phytophagous Asia1932, GB Temperate HR, DK, DE, GB, IT, NL, CH Prociphilus Meliarhizophagus fraxinifolii Riley ex Riley & Monell, 1879 Pterochloroides persicae (Cholodkovsky, 1899) A phytophagous North America A phytophagous A phytophagous phytophagous phytophagous A A 1922, GB 2003, HU DK, DE, GB, IL, IT, NL, PT- J100 AZO, ES-CAN Blackman and Eastop (1994), Doncaster (1954), Eastop (1956), Petrović-Obradović et al. (2007), Remaudière and Ripka (2003) Ciampolini and Martelli (1977), Petrović and Milanović (1999), Roberti (1975), Velimirovic (1976) Nieto Nafria et al. (2007) Barbagallo et al. (2005b) Ilharco (1969), Mimeur (1936), Tashev (1964) Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) Cairaschi (1942), Sűss (1972–73) I2, G5 Musa (preferred); Polyphagous on tropical and subtropical ornamental plants Acer North America Pterocomma pseudopopuleum Palmer, 1952 Reticulaphis distylii vand der Goot 1917 Rhodobium porosum (Sanderson, 1900) References Blackman and Eastop (1994), Ilharco (1974) phytophagous I2, F Hosts Populus A Pemphigus Pemphigus populitransversus Riley ex Riley & Monell, 1879 Pentalonia nigronervosa Coquerel, 1859 Habitat 470 Species Species Status Feeding Regime Native range 1st record Invaded countries in invaded areas 1921, IT AL, AT, BE, BG, CH, CZ, DE, ES, FR, GB, GR, HR, IT, IT-SIC, NL, PL, PT, PTAZO, PT-MAD, RO, RU A phytophagous North America Rhopalosiphum insertum (Walker, 1849) A phytophagous North America 1848 GB Rhopalosiphum maidis (Fitch, 1856) A phytophagous Asia 1903, IT Rhopalosiphum rufiabdominale (Sasaki, 1899) A phytophagous Asia1960 PT Temperate Sipha Sipha flava (Forbes, 1884) A phytophagous North America 1979, PTAZO I1 AL, AT, BY, BE, BG, CH, I1, E CZ, DE, DK, EE, ES, ESCAN, FI, YU, FR, FR-COR, DE, GB, GR, HU, IE, IT, LT, LV, MD, NL, NO, PL, PT, PT-AZO, PT-MAD, RO, RU, RS, SE, SI, SK, UA I1, E AL, BE, BG, CH, CY, CZ, DE, DK, ES, ES-CAN, FI, ,FR, FR-COR, GB, GR, GRCRE, HU, IT-SAR, IT-SIC, IT, LV, MD, NL, NO, PL, PT, PT-AZO, PT-MAD, RO, RS, RU, SE, ES, SK, UA BG, DK, ES, FI, FR, GR, IT, I1 IT-SIC, PT, PT-AZO, PTMAD, RU, UA AL, PT-AZO I1 Hosts Solanum; polyphagous on vegetables (Beta, Fragaria, Ipomea) and flowers (Gladiolus) Graminae (Poa, Festuca, Juncus) Maize, sorghum; other crops Rice roots, Gramineae Sugarcane References Blackman and Eastop (2000), Remaudière (1952), Tashev (1961) Blackman and Eastop (2000), Dospevski (1910), Ilharco (1968a), Walker (1849) Blackman and Eastop (2000), Del Guercio (1913), Del Guercio (1917), Dospevski (1910), Eastop (1956), Heie (1986), Ilharco (1961) Blackman and Eastop (2006), Heie (1986), Ilharco (1968a), Ilharco (1973) Sousa-Silva and Ilharco (1995) Aphids (Hemiptera, Aphididae). Chapter 9.2 Rhopalosiphoninus Rhopalosiphoninus latysiphon (Davidson, 1912) Habitat 471 Status Feeding Regime A Sitobion Sitobion luteum (Buckton, 1876) 1st record Invaded countries in invaded areas 1999, FR FR, IT Habitat Hosts phytophagous phytophagous North America Asia<2004, GB GB, NL Temperate G5, I2, FA I2, E C phytophagous Cryptogenic BE, DE, FR, GB, PT-MAD J100 Stomaphis mordvilkoi Hille Ris Lambers, 1933 Takecallis arundicolens (Clarke, 1903) A phytophagous phytophagous Asia1980, IT Tropical Asia1923, GB Temperate IT G Blackman and Eastop (2006), Nieto Nafria ei al. (2007) Orchidaceae, Blackman and Eastop Bromeliaceae, (2006), Buckton (1876), Araceae Del Guercio (1911) Schouteden (1906) Juglans Colombo (1981) CH, DE, ES, FR, GB, IE, IT, PT I2 Bamboos Takecallis arundinariae (Essig, 1917) A phytophagous Asia1961, GB Temperate CH, DE, ES, GB, GR, IT, IT-SIC, PT-MAD I2 Takecallis taiwana (Takahashi, 1926) A phytophagous Asia1923, GB Temperate CH, DE, ES, FR, GB, HR, IT, IT-SIC I2 Tinocallis Sappocallis nevskyi Remaudière, Quednau & Heie, 1988 A phytophagous Asia1978, PL Temperate AT, BE, CH, CZ, DE, DK, FI, GB, HU, IT, NL, PL, SE G, G5, I2, FA A A 1875 GB Cupressus References Rabasse et al. (2005a) Graminae Hille Ris Lambers (1947), Ilharco (1969), Laing (1923), Stroyan (1964), Stroyan (1977), Theobald (1927) Bamboos Giacalone and Lampel (1996), Pati and Tomatore (1988), Stroyan (1964), Stroyan (1977) Bamboos Giacalone and Lampel (Phyllostachys) (1996), Limonta (1990), Stroyan (1964) Ulmus Remaudière et al. (1988), Szelegiewicz (1978), Van Harten and Coceano (1981) Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) Siphonatrophia cupressi Swain, 1918 Sitobion Sitobion alopecuri (Takahashi, 1921) Native range 472 Species Status Feeding Regime A phytophagous Tinocallis Sappocallis takachihoensis Higuchi 1972 A phytophagous Asia1985, FR Temperate ES, FR, IT, IT-SIC G, G5, I2 Ulmus Tinocallis Sarucallis kahawaluokalani (Kirkaldy, 1906) A phytophagous Asia1984, IT Temperate DE, ES, FR, GR, IT, IT-SIC, ME I2, G5 Lagerstroemia indica Tinocallis Tinocallis ulmiparvifoliae Matsumura, 1919 A phytophagous Asia1973, GB Temperate ES, GB, IT I2, J100 Ulmus Tinocallis Tinocallis zelkowae (Takahashi, 1919) Toxoptera aurantii Boyer de Fonscolombe 1841 A phytophagous phytophagous Asia1973, GB Temperate Tropical, 1841 FR subtropical FR, GB I2, J100 Zelkova A Native range AL, BE, CH, CY, DE, ES, ES- I, G5, BAL, FR, FR-COR, GB, GR, J100 HR, IL, IT, IT-SAR, IT-SIC, ME, MT, PT-AZO, PT-MAD, PT, RO Polyphagous (mainly Citrus) References Holman and Pintera (1981), Hullé et al. (1998), Remaudière et al. (1988), Van Harten and Coceano (1981) Hullé et al. (1998), Leclant and Renoust (1986), Leclant and Remaudière (1986) Arzone and Vidano (1990), Leclant and Renoust (1986), Ossiannilsson (1959), Pati (1984), PetrovićObradović et al. (in press) Lucchi and Pollini (1995), Pérez Hidalgo and Nieto Nafria (2005), Prior (1971), Stroyan (1977) Prior (1971), Stroyan (1977) Boyer de Foscolombe (1841), Del Guercio (1917), Passerini (1861), Stroyan (1984), Tavares (1900) Aphids (Hemiptera, Aphididae). Chapter 9.2 Tinocallis Sappocallis saltans (Nevsky, 1929) 1st record Invaded countries Habitat Hosts in invaded areas Asia1976,RO ES, FR, HU, IT, IT-SIC, MD, G, G5, Ulmus Temperate NL, PL, RO, RU, UA I2 Species 473 Status Feeding Regime Native range 1st record Invaded countries in invaded areas 1994, PT- ES, PT, PT-MAD MAD A phytophagous Trichosiphonaphis Xenomyzus polygonifoliae (Shinji, 1944) A phytophagous Tropical, subtropical Asia1990, FR Temperate Tuberculatus Nippocallis kuricola (Matsumura, 1917) Uroleucon Lambersius erigeronense (Thomas, 1878) A phytophagous phytophagous Asia1981, PTTemperate MAD North 1952, FR America A Hosts References I, G5 Citrus Aguiar et al. (1994), Ilharco et al. (2005) FR, GB, HU, IT, RS, UA I2 Lonicera, Polygonum ES, PT, PT-AZO, PT-MAD G1, I2 Castanea, Quercus Asteraceae (Erigeron, Coniza) Coceano and PetrovicObradovic (2006), Petrović-Obradović et al. (in press), Remaudière et al. (1992) Ilharco (1984), Pedro Mansilla et al. (2001) Blackman and Eastop (2006), Heie (1995), Remaudière (1954) Uroleucon Uroleucon pseudoambrosiae (Olive, 1963) A phytophagous North America <2004 AT, BE, CH, CZ, DE, DK, J, J6 ES, FI, FR, GB, GR, HU, IT, IT-SIC, LV, MD, NL, PL, PTMAD, RO, RS, SE, SI, RK PL I Utamphorophora humboldti (Essig, 1941) A phytophagous North America 1974, GB FR, GB, GR, IE Wahlgreniella arbuti (Davidson, 1910) A phytophagous North America 1905, PT Wahlgreniella nervata (Gillette, 1908) A phytophagous North America 1973, GB ES, ES-BAL, FR, FR-COR, I2, F6 GB, GR, IT, IT-SAR, IT-SIC, NL, PT, PT-MAD AT, BE, ES, ES-CAN, FR, I2 GB, GR, IT-SIC I2 Asteraceae (Mainly Lactuca spp.) Physocarpus, Poaceae Arbutus, Arctostaphylos Rosa Blackman and Eastop (2000), Blackman and Eastop (2006), Nieto Nafria et al. (2007) Hullé et al. (1998), Prior (1975), Tsitsipis et al. (2007) Heie (1995), Ilharco (1969), Tavares (1905), Tsitsipis et al. (2007) Blackman and Eastop (2006), Prior (1975), Tsitsipis et al. (2007) Armelle Cœur d’Acier et al. / BioRisk 4(1): 435–474 (2010) Toxoptera citricidus Kirkaldy 1906 Habitat 474 Species A peer reviewed open access journal BioRisk 4(1): 475–510 (2010) RESEARCH ARTICLE doi: 10.3897/biorisk.4.45 BioRisk www.pensoftonline.net/biorisk Scales (Hemiptera, Superfamily Coccoidea) Chapter 9.3 Giuseppina Pellizzari1, Jean-François Germain2 1 Università di Padova - Dipartimento Agronomia Ambientale e Produzioni Vegetali, Agripolis - Viale dell’Università 16, 35020 Legnaro Padova, Italia 2 Laboratoire National de la Protection des Végétaux, Station de Montpellier, CBGP Campus international de Baillarguet CS 30016 34988 Montferrier-sur-Lez Cedex, France. Corresponding authors: Giuseppina Pellizzari (giuseppina.pellizzari@unipd.it), Jean-François Germain (ger- main@supagro.inra.fr) Academic editor: David Roy | Received 1 February 2010 | Accepted 24 May 2010 | Published 6 July 2010 Citation: Pellizzari G, Germain J-F (2010) Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 475–510. doi: 10.3897/biorisk.4.45 Abstract Scale insects are frequent invaders. With 129 established species, they numerically represent one of the major group of insects alien to Europe. Scales are usually small insects with wingless females. Due to this small size and concealment, many species, mainly belonging to the families Diaspididae, Pseudococcidae and Pseudococcidae, have been accidentally introduced to Europe, mostly originating from tropical regions and essentially from Asia. The trade of fruit trees and ornamentals appears to be the usual pathway of introduction. At present, alien scales represent an important component of the European entomofauna, accounting for about 30% of the total scale fauna. Keywords Europe, Alien, scale insects 9.3.1 Introduction Coccoidea or scale insects is a large superfamily in the order Hemiptera with a worldwide distribution. They are unusually small insects, highly specialized for plant parasitism, that have evolved different kinds of metamorphosis depending on sex and family. Scale insects are characterized by sexual dimorphism: females are wingless, usually small (from 0.5 – 10mm), with an oval or round but flat to fairly convex body Copyright G. Pellizzari, J-F. Germain. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 476 Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) form, sometimes bud shaped, and often protected by waxy secretions or covers. The adult females may exhibit reduction or loss of appendages, depending on family and instar, and are often sedentary or sessile. Adult males are usually winged and inconspicuous, do not feed and live a few days. Scale insect identification is mainly based upon the morphology of adult females that persist on the host plant longer than the other stages. Females usually take three or four developmental stages to reach maturity, males usually five. Parthenogenesis is quite common. Eggs are usually laid under the female body, under the scale cover, or in waxy egg-sacs. Dispersal is carried out by first instars. Scale insects feed on various parts of the host plant (leaves, fruits, stems, branches and roots) and are frequently introduced and acclimatized in different parts of the world. This is due to their small size (first instars are about 0.2–0.3mm; adult females usually are from 0.5 to10mm long) and their concealment using waxy secretions; beside many species live in hidden habitats (under leaf sheaths, in bark crevices or on roots) so that they can easily escape visual quarantine inspections. Once in a new territory, parthenogenesis and high fecundity favour quick colonization starting from a few females: for example, a single female Neopulvinaria innumerabilis may lay up to 8000 eggs (Canard 1968). 9.3.2 Taxonomy of the scale species alien to Europe According to Ben-Dov et al. (2006) the superfamily Coccoidea comprises 22 families, with more than 7300 described species. In Europe, native representatives of 12 families have so far been recognized. On the basis of the best known western and central European coccoid faunas (France, Italy, Hungary) (Ben-Dov et al. 2006, Foldi 2001, Pellizzari and Russo 2004), the total number of scale insects present in Europe is likely to reach about 400–450 species. Aliens recorded in Europe up until 2007 account for 129 species which include the following eight families: Diaspididae (60 species), Pseudococcidae (37), Coccidae (23), Eriococcidae (3), Margarodidae (2), Asterolecanidae, Ortheziidae, and the alien family Phoenicococcidae, each with one species (Table 9.3.1). Unlike for other taxa, aliens represent an important component of the scale fauna currently present in Europe, i.e. near 30% (Fig. 9.3.1). The remaining five native families (Aclerdidae, Cerococcidae, Kermesidae, Lecanodiaspididae, Micrococcidae) each have one or two species in Europe: none of them is a pest, with the exception of the family Kermesidae (8 species in Europe), in which Kermes vermilio and Nidularia pulvinata exhibit outbreaks in urban environments only. One species, Dactylopius coccus Costa, representing the alien family Dactylopiidae, has been included among aliens to Europe, even though it is present only in Canary islands, Madeira and Azores, where it was intentionally introduced. These islands belong politically to Europe (Spain, Portugal) but biogeographically they belong to Macaronesia, a biogeographic Atlantic region quite distinct from the European continent and with a unique flora and fauna. Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 477 Figure 9.3.1 Taxonomic overview of the scale species alien to Europe compared to the native fauna. Species alien to Europe include cryptogenics. Diaspididae Armoured scale insects are the commonest alien scales incidentally introduced all over the world: this is probably due to their small dimension and camouflage. The 60 alien species account for nearly half (44.6%) of an estimated 130 species in Europe. Many notorious pests of fruit trees such as Pseudaulacaspis pentagona (the white peach scale- see factsheet 14.45)) and Diaspidiotus perniciosus (San José scale - see factsheet 14.44)) belong to this family: these species are still pests of fruit trees in spite of the introduction of specific parasitoids from their native area. The Asiatic armoured scales of Citrus are largely found in European Citrus groves and presently number 10 species. Their ”invasion” started around 1850 with Parlatoria ziziphi and Lepidosaphes becki and is still going on with the arrival and establishment of Unaspis yanonensis (1969), Aonidiella citrina (1994), Chrysomphalus aonidum (2000). Several armoured scales commonly occur throughout European greenhouses (e.g. Diaspis echinocacti, Chrysomphalus dictyospermi, Diaspis bromeliae, Abgrallaspis cyanophylli), even if they cannot be considered as established. In some cases, species recorded only in greenhouses in northern and central Europe are established outdoors in southern countries (i.e. Furchadaspis zamiae, Chrysomphalus aonidum). Some armoured scales thought to be of Afrotropical origin or cryptogenic (e.g. Aspidiotus nerii, Hemiberlesia lataniae, H. rapax) are very common in natural habitats of the Mediterranean countries (including small islands). Pseudococcidae Mealybugs are covered with mealy or cottony wax, have a distinct segmentation and are mobile. The 37 alien mealybugs account for roughly one fourth (25.7%) of the ca. 140 European species and most of them are polyphagous. Planococcus citri, Pseu- 478 Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) dococcus longispinus, P. viburni and P. calceolariae arrived and established during the 19th century and are presently the most common species on ornamental plants, both outdoors and indoors. P. citri, first recorded in 1813, is still a pest of Citrus and ornamental plants. Several mealybugs have been recorded in only one or two countries to date (e.g. Palmicultor palmarum, Phenacoccus madeirensis, Rhizoecus americanus, Trochiscococcus speciosus), both outdoors and in greenhouses, on ornamental plants. Coccidae About 70 species of soft scales are recorded in Europe. Of these, there are 23 aliens to Europe representing 32.8% of the fauna, and are mainly pests of fruit trees and ornamentals. Among them, the polyphagous Coccus hesperidum and Saissetia oleae, the well-known Mediterranean Black Scale, are probably the most ancient arrivals which established in the countries surrounding the Mediterranean Basin. Most recent arrivals are Pulvinaria hydrangeae, P. regalis (see factsheet 14.41), Ceroplastes japonicus and, in warmer places, Protopulvinaria pyriformis, invasive on trees and ornamental plants in urban environments. Some species, such as Coccus pseudomagnoliarum, after first spreading in Mediterranean Citrus groves, later became more localised and less common. On the other hand, the American Pulvinaria innumerabilis is still considered a pest of vine, more than 40 years after its arrival in European vineyards. Several species (e.g. Saissetia coffeae, S. oleae, C. hesperidum, Eucalymnatus tessellatus, Parasaissetia nigra) are rather common in greenhouses of central and northern Europe, while in southern Europe are outdoors pests. Eriococcidae European felt scales number about 50 species. Among them, only three alien felt scales have been so far recorded. The Australian Eriococcus araucariae is widespread on Araucaria trees growing in Mediterranean countries, the American E. coccineus is recorded on succulent plants and Ovaticoccus agavium is quite common on Agave sp. growing outdoors. Margarodidae European margarodids recorded up until now number 15 species. Two alien margarodids, Icerya purchasi (the cottony cushion scale) and I. formicarum, invaded Europe at very different times. The latter species is known from a single record in 2001 in Corsica and its establishment is unknown. On the other hand, the Australian I. purchasi has both established and caused an agricultural and environmental impact. It arrived and established in many Mediterranean countries between the end of 1800 and the first Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 479 decades of 1900 and was very destructive to Citrus groves. The high infestations led to the introduction of the Australian coccinellid Rodolia cardinalis, for biological control. Presently, the cottony cushion scale is mainly a pest of ornamental plants such as Pittosporum, Acacia and Mimosa. It is also a very common species in semi-natural habitats (i.e. the Mediterranean maquis), far away from cultivated areas, where it develops on autochthonous wild plants such as Cistus, Genista, Smilax and Rosmarinus. Two other margarodids, Marchalina hellenica and Matsucoccus feytaudi, are alien in Europe, entirely due to deliberate introduction. Asterolecanidae About 10 species of asterolecanids are present in Europe. Of these, the only alien pit scale is the Asiatic Bambusaspis bambusae, a species associated with bamboos. Ortheziidae Ortheziids consist of 10 species in Europe. Among these, Insignorthezia insignis, a polyphagous Neotropical species, has been reported in European greenhouses since the end of 19th century. Apparently I. insignis is established outdoors only in Portugal and France. Phoenicococcidae Phoenicococcus marlatti, the Red Date Palm Scale, thought to originate in the Middle East or North Africa, is the only species currently placed in the family Phoenicococcidae. It is considered a minor pest of commercial dates, whereas in Spain, France and Italy, it infests ornamental palms (mainly Phoenix canariensis). 9.3.3 Temporal trends of introduction in Europe of alien scale species Fig. 9.3.2. presents the temporal variation in the mean number of new alien species recorded per year since 1492. Serious studies of the Coccoidea began in mid 19th century. From that time, to the mid-1970s, the introduction of alien species was relatively constant, averaging 0.66 species per year. Since then, there is an apparent increase in alien introductions, up to an average of 1.15 species per year. In interpreting this chart, account should be taken of “old” alien species, found and described in Europe, (i.e. Aspidiotus nerii, Planococcus citri, Coccus hesperidum, Saissetia oleae) for which the introduction date is based only on the date of their first description. In the case of the most harmful alien scales, the date of first introduc- 480 Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) Figure 9.3.2 Temporal trends in the mean number of new records per year of scale species alien to Europe from 1492 to 2007. The number above the bar indicates the absolute number of species in this time period. tion to Europe and the chronology of their invasion is known more precisely (i.e. for Pseudaulacaspis pentagona, Icerya purchasi, Diaspidiotus perniciosus). Moreover, records of alien scales depend on the presence of specialists in a given country. For instance, during the 1970–80s, advances in systematic knowledge and the increasing number of active coccidologists led to the “discovery” of several species which have probably been introduced a long time before. The great rise in the global exchanges of plants and quarantine inspections can explain the increases in subsequent years up until the present. Among the scale insects introduced to Europe from the end of 19th century to 1960s there are several pests of fruit trees and Citrus (i.e. Diaspidiotus perniciosus, Lepidosaphes gloverii, Pseudaulacaspis pentagona, Ceroplastes sinensis, Icerya purchasi), whereas in the last 40 years the most numerous introduced scales are pest of ornamental plants, both outdoors and indoors (i.e. Pulvinaria regalis, P. hydrangeae, Ceroplastes japonicus, Protopulvinaria pyriformis, Parassaisetia nigra, Trochiscococcus speciosus), the main scale of agricultural importance being Neopulvinaria innumerabilis, a pest of vine. Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 481 9.3.4 Biogeographic patterns of the scale species alien to Europe 9.3.4.1 Origin of the alien species The geographical origin of introduced scale insects shows a large dominance of species from tropical areas, essentially Asia, followed by southern American species (Fig. 9.3.3). The precise origin remains unknown for about one fourth of alien scales. Among the most widespread aliens to Europe are Diaspidiotus perniciosus of temperate Asian, Planococcus citri from tropical Asia, Ceroplastes sinensis from Central-America, Parthenolecanium fletcheri from Northern-America, Saissetia oleae from the Afrotropics, Icerya purchasi from Australasia, and Lepidosaphes beckii as cryptogenic species. 9.3.4.2 Distribution of the alien species in Europe It should be borne in mind that, as for the other arthropod groups, the number of records of alien scales in European countries, reflects, in part, differences of study intensity and the number of local taxonomists. Moreover, the geographic position of some countries such as France, Italy and Spain, whose climatic conditions vary from high montane, continental to Mediterranean, allows establishment of species from very different geographical areas. Two countries present a particularly high number of alien species: France with 90 species and Italy with 92 species (Fig. 9.3.4). Lagging far behind are Spain, Great Britain and Portugal with 50, 43 and 41 species, respectively. The islands of the Atlantic, not represented in the figure, have respectively 51 aliens in the Canaries, 44 in Madeira and 22 in the Azores. There are 12 alien species recorded in at least 20 countries, namely Coccus hesperidum (28 countries), Pulvinaria floccifera (21), Saissetia coffeae (24), S. oleae (26), Aspidiotus nerii (26), Diaspidiotus perniciosus (26), Pinnaspis aspidistrae (20), Pseudaulacaspis pentagona (21), Planococcus citri (22), Pseudococcus longispinus (22) and P. viburni (26). These are all polyphagous species, with the exception of Unaspis euonymi, monophagous on Euonymus spp., recorded in 22 countries. A total of 20 species (15%) are present only in one country. 9.3.4.3 Scale species alien in Europe With regard to scale insects alien in Europe, that is originating from another European area where native and introduced through human activity, only very few certain cases are known. Marchalina hellenica is native to Turkey and Greece and presently invasive in the small island of Ischia (Italy). It was introduced there in 1960 to study endosymbiosis, but unfortunately escaped from laboratory breeding and presently is a pest of pines (Tranfaglia and Tremblay 1984). Matsucoccus 482 Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) Figure 9.3.3 Geographic origin of the scale species alien to Europe. Figure 9.3.4 Numbers of established alien scale species in the European countries and main islands according to Table 9.3.1. Archipelago: 1 Azores 2 Madeira 3 Canary islands. Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 483 Figure 9.3.5 Ceroplastes ceriferus (Coccidae). Credit: Giuseppina Pellizzari Figure 9.3.6 Coccus hesperidum (Coccidae). Credit: Giuseppina Pellizzari feytaudi lives on Pinus pinaster and is native to the Atlantic regions of France, Spain and Portugal. It was introduced with its host plant in South-eastern France and from there spread towards Italy (Arzone and Vidano 1981). Both Aonidiella lauretorum and A. tinerfensis are endemic to the Atlantic islands of Canary (Spain) and Madeira (Portugal). They were introduced incidentally with their host plants 484 Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) Figure 9.3.7 Parasaissetia nigra (Coccidae). Credit: Giuseppina. Figure 9.3.8 Protopulvinaria pyriformis (Coccidae). Credit: Giuseppina Pellizzari. in the Botanic gardens of Sintra and Lisbon (Portugal), where they still persist (Balachowsky 1948). 9.3.6 Pathways of introduction in Europe of alien scale species Scale insects are highly specialized, sedentary, plant-parasitic insects and the only pathway of introduction is the horticultural and ornamental trade: importation and trade Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 485 Figure 9.3.9 Pulvinaria hydrangeae (Coccidae). Credit: Nico Schneider Figure 9.3.10 Pulvinaria floccifera (Coccidae). Credit: Nico Schneider of fruit and Citrus trees, ornamental trees and bushes, bulbs and corms, has led to incidental introduction and subsequent spread of scale insects. More recently, the “fashion” of succulent plant cultivation and the subsequent increase in plant importation and plant exchanges among collectors is responsible for the introduction and spread of several species such as Delottococcus euphorbiae, Hypogeococcus pungens, Trochiscococcus speciosus, Vryburgia rimariae, Spilococcus mamillariae and Eriococcus coccineus. Importation of bonsais from Asia could allow the introduction and spread of Rhizoecus hibisci, a mealybug living on roots and recently intercepted several times by European quarantine services. 486 Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) Figure 9.3.11 Chrysomphalus aonidum (Diaspididae). Credit: Giuseppina Pellizzari. Figure 9.3.12 Unaspis yanonensis (Diaspididae). Credit: Giuseppina Pellizzari. 9.3.7 Ecosystems and habitats invaded in Europe by alien scale species Alien, established scale insects colonize strongly anthropogenic habitats such as cultivated agricultural lands, horticultural and domestic habitats, urban environments, gardens and parks, botanic gardens, nurseries and greenhouses, but they have also spread to natural habitats. Mediterranean Citrus groves host a large community of alien scales: 18 different species have been so far recorded. These are: Icerya purchasi Planococcus citri, Pseudococcus calceolariae, P. longispinus, Ceroplastes sinensis, Coccus hesperidum, C. pseudomagnoliarum, Saissetia oleae, Aonidiella aurantii, A. citrina, As- Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 487 Figure 9.3.13 Comstockiella sabalis (Diaspididae). Credit: Jean Francois Germain Figure 9.3.14 Ovaticoccus agavium (Eriococcidae). Credit: Giuseppina Pellizzari pidiotus nerii, Chrysomphalus dictyospermi, C. aonidum, Lepidosaphes beckii, L. gloverii, Parlatoria pergandii, P. ziziphi and Unaspis yanonensis. Some polyphagous scales are urban pests, largely distributed in urban parks and gardens, on trees and ornamentals (i.e. Pulvinaria regalis, P. hydrangeae, Ceroplastes japonicus), whereas they are absent or very rare in the countryside. A few monophagous species are only known in Botanical gardens, where they persist outdoors, at a low population levels, on exotic plants 488 Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) Figure 9.3.15 Pseudococcus comstocki (Pseudococcidae). Credit: Giuseppina Pellizzari Figure 9.3.16 Pseudococcus longispinus (Pseudococcidae). Credit: Giuseppina Pellizzari introduced over there a long before (i.e. Aonidiella tinerfensis, Pseudaonidia paeoniae or Bambusaspis bambusae). Several other monophagous species remain strictly associated to their original, exotic ornamental plants, and have a correspondingly wide distribution in Europe (i.e. Parthenolecanium fletcheri, Pulvinaria mesembryanthemi, Eriococcus araucariae). On the other hand, some polyphagous species (i.e. Diaspidiotus perniciosus, Pseudaulacaspis pentagona, Pulvinaria floccifera) have spread from cultivated areas to natural woodland and forest habitats (Balachowsky 1932b, Balachowsky 1936). Others (Antonina graminis, Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 489 Figure 9.3.17 Pseudococcus calceolariae (Pseudococcidae). Credit: Jean Francois Germain Chorizococcus rostellum and Trionymus angustifrons) can be found in grasslands. In natural habitats of Mediterranean countries (including small islands), species such as the armoured scales Aspidiotus nerii (see factsheet 14.43), Hemiberlesia lataniae, H. rapax, the mealybug Planococcus citri, the wax scale Ceroplastes sinensis and the Australian I. purchasi are quite common on wild autochthonous plants, growing far away from cultivated plants. Their transfer from cultivated plants to authochtonous ones in natural environments confirms that they have fully acclimatized. 9.3.8 Impact of alien scale species Scale insects are plant pests, especially of fruit trees, woody ornamentals, forest trees and greenhouse plants. They cause damage to plants by sap sucking. Moreover, except for Diaspididae and Asterolecaniidae, they excrete honeydew that covers leaves and fruits and allows the development of sooty mould. This black sooty mould can reduce photosynthesis by 70%, leading to early senescence, with smaller and premature fruits, and loss of aesthetic value (Mibey 1997). Moreover, Coccidae and Pseudococcidae are vectors of closteroviruses. For example, Planococcus citri and Pulvinaria innumerabilis may transmit the Grapevine Leafroller-associated Virus (GLRaV-1, GLRaV-3) and the Corky Bark disease (GVA, GVB) (Sforza et al. 2003, Zorloni et al. 2006). Diaspididae 490 Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) cause discolouration on leaves, red or black spots on fruits, and twig dieback. Pesticides are commonly applied to control scale insects in fruit orchards and Citrus groves. Infestations of alien scales in orchards have led to the introduction to Europe, from their native area, of many natural enemies for biological control purposes. 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Bollettino di Zoologia agraria e di bachicoltura Ser II 24: 215–221. Russo A, Mazzeo G (1996) Dactylopius coccus Costa (Homoptera, Coccoidea): la cocciniglia del carminio. Informatore Fitopatologico 4: 10–13. Russo A, Mazzeo G, Suma P (1999) Sulla presenza di Entaspidiotus lounsburyi (Marlatt, 1908) su Mesembryanthemaceae in Italia (Homoptera, Coccoidea). Redia 82: 83–87. Seabra AF de (1942) Contribuiçoes para o inventario da fauna Lusitânica Insecta: Homoptera (Coccidae). Aditamento Memorias y Estudos do Museo Zoologico da Universidade de Coimbra 128: 1–2. Sforza R, Boudon-Padieu E, Greif C (2003) New mealybug species vectoring Grapevine leafroll-associated viruses-1 and-3 (GLRaV-1 and –3). European Journal of Plant Pathology 109: 975–981. Signoret V (1869a) Essai sur les cochenilles ou gallinsectes (Homoptères - Coccides), 4ème partie. Annales de la Société Entomologique de France 9: 109–138. Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 495 Signoret V (1869b) Essai sur les cochenilles ou gallinsectes (Homoptères - Coccides), 5ème partie. Annales de la Société Entomologique de France 9: 431–452. Signoret V (1875) Essai sur les cochenilles ou gallinsectes (Homoptères - Coccides), 15e partie. Annales de la Société Entomologique de France (série 5) 5: 305–352. Süss L, Trematerra P (1986) Hypogeococcus festerianus (Lizer y Trelles), nocivo alle Cactacee ormamentali in Liguria. Informatore Fitopatologico 10: 43–46. Targioni Tozzetti A (1884) Relazione intorno ai lavori della R Stazione di Entomologia Agraria di Firenze per gli anni 1879–80 article V-omotteri. Annali di Agricoltura (Ministero di Agricoltura, Industria e commercio) Firenze, Roma: 383–414. Targioni Tozzetti A (1867) Studii sulle Cocciniglie. Memorie della Società Italiana di Scineze Naturali Milano 3: 1–87. Targioni Tozzetti A (1886) (1885) Sull’insetto che danneggia I gelsi. Rivista di bachicoltura 18: 1–3 Tranfaglia A (1976) Studi sugli Homoptera Coccoidea IV Su alcune Cocciniglie nuovo o poco conosciute per l’Italia (Coccidae, Eriococcidae, Pseudococcidae). Bollettino del laboratorio di entomologia Agraria ‘Filippo Silvestri’ Portici 33: 128–143. Tranfaglia A (1981) Studi sugli Homoptera Coccoidea V Notizie morfo-sistematiche su alcune specie di Cocciniglie con descrizione di tre nuove specie di Pseudococcidi. Bollettini del Laboratorio di Entomologia Agraria ‘Filippo Sivestri’ Portici 38: 3–28. Tranfaglia A, Tremblay E (1984) Faunistic and systematic studies on Italian scale insects. In Verhandlungen des Zehnten Internationalen Symposiums über Entomofaunistik Mitteleuropas (SIEEC X), August 1983, Budapest (1984), 372–374. Tranfaglia A, Viggiani G (1988) Cocciniglie di importanza economica in Italia e loro controllo. Benevento, Regione Campania, Italy: Assessorato Agricoltura, Servizio Sperimentazione Abete Grafica 30pp. Williams DJ (1985) Scale insects (Homoptera: Coccoidea) of Tresco, isles of Sicily. Entomologist’s Gazette 36: 135–144. Williams DJ, Pellizzari G (1997) Two species of mealybugs (Homoptera Pseudococcidae) on the roots of Aloaceae in greenhouses in England and Italy. Bollettino di Zoologia agraria e di Bachicoltura ser II 29: 157–166. Zahradnik J (1990) Die Schildläuse (Coccinea) auf Gewächshaus- un Zimmerpflanzen in den Tschechischen Ländern. Acta Universitatis Carolinae Biologica 34: 1–160. Zorloni A, Prati S, Chiesa S, Bianco PA (2006) Transmission of grapevine leafroll associated virus 3 by the soft scale insect Neopulvinaria innumerabilis Rathvon. In Abstracts of the 13th Congresso Nazionale della Società Italiana di Patologia Vegetale September, 2006, Foggia, 17. Pellizzari G, Danzig E (2007) The bamboo mealybugs Balanococcus kwoni n. sp. and Palmicultor lumpurensis (Takahashi) (Hemiptera, Pseudococcidae) Zootaxa 1583: 65–68. Status Regime Native range 1st record in Europe Invaded countries Habitat Hosts References A Phytophagous Asia-Tropical 1941, ITSIC DK, ES, FR, GB, IT, ITSIC, PT, PT-MAD I2, J100 Bambusa Russell (1941) A Phytophagous Phytophagous Phytophagous Phytophagous Central1921, IT America South1930, FR America Asia-Tropical 1930, FR IT, ES-CAN, GB I2 Polyphagous CY, FR, GR, IL, MT, PT- I2 AZO, PT-MAD FR, IT, HR, SI I2 Polyphagous Green (1921b), Mori et al. (2001) Balachowsky (1930) CentralAmerica 1890, IT AL, ES, ES-CAN, FR, I2 FR-COR, GR,HR, IT, IT-SIC, MT, ME, PT, PT-AZO, PTMAD, RO I2 BE, BG, CH, CY, DE, DK, ES, ES-CAN, FR, FR-COR, GB, GR, HU, HR, IT, IT-SIC, IT-SAR, LV, NL, ME, MT, PT, PT-AZO, PT-MAD, SK, SI, RS, RO, UA FR, ES-CAN I2 Polyphagous Pellizzari and Camporese (1994) Del Guercio (1900) Polyphagous Costa (1829) Polyphagous Foldi (2001) Polyphagous Green (1921a) A A A Coccus hesperidum Linnaeus, 1758 A Phytophagous Tropical/ subtropical 1829, IT Coccus longulus (Douglas, 1887) Coccus pseudohesperidum (Cockerell, 1895) A Phytophagous Phytophagous Tropical/ subtropical SouthernAmerica 2001, FR A 1920, GB GB, LV, UA I2 Polyphagous Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) Family Species Asterolecaniidae Bambusaspis bambusae (Boisduval, 1869) Coccidae Ceroplastes ceriferus (Fabricius, 1798) Ceroplastes floridensis Comstock, 1881 Ceroplastes japonicus Green, 1921 Ceroplastes sinensis Del Guercio 1900 496 Table 9.3.1. List and main characteristics of the scale species alien to Europe. Status: A: Alien to Europe; C: cryptogenic species. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Only selected references are given. Last update 29/05/200 Status Eulecanium excresens Ferris, 1920 Cryptinglisia lounsburyi (Cockerell, 1900) Neopulvinaria innumerabilis (Rathvon, 1880) Parasaissetia nigra (Nietner, 1861) Parthenolecanium fletcheri (Cockerell, 1893) Protopulvinaria pyriformis (Cockerell, 1894) Pulvinaria floccifera (Westwood, 1870) A Phytophagous Native range 1st record Invaded countries in Europe Asia-Tropical 1974, GR, FR, GR, HR, IT, ITIT SIC,ME, SI A Phytophagous SouthernAmerica A Phytophagous Phytophagous Phytophagous NorthernAmerica Afrotropical A A Regime Habitat I2 Hosts Citrus References Barbagallo (1974) I2, J100 Livistona ., Palms Balachowsky (1954) I2 Malumphy (2005) 1982, IT IT I2 NorthernAmerica 1961, FR FR, HR, IT, SI I2 Vitis, polyphagous Hodgson (1994) AL, ES, ES-CAN,FR, FR-COR, IT, IT-SIC, MT, PT, PT-AZO, PT-MAD, PL, RO AT, BG, CH, CZ, DE, FR, HU, HR, LV, NL, PL, RO, S AL, ES, ES-CAN, FR, GR, IT, IT-SIC, PT, PTAZO, PT-MAD CH, CY, CZ, DE, ES, ES-CAN, FR, FR-COR, GB, GR, HU, HR, IT, IT-SAR, IT-SIC, NL,PT, PT-MAD, SE, SI, RO, RU I2 Polyphagous Marotta (1987) I2 Cupressus, Thuya Kawecki (1935) I2 Polyphagous Marotta and Tranfaglia (1990) I2 Ilex aquifolium, Taxus baccata Marchal (1907) A Phytophagous Afrotropical 1900, IT A Phytophagous NorthernAmerica 1935, PL A Phytophagous Asia-Tropical 1991, IT A Phytophagous AsiaTemperate 1889, FR Juglans, Wisteria Pelargonium Marotta (1987) 497 BG, DK, DE, ES, ESCAN, FR, IL, PT-MAD, PL, UA 1998, GB GB 1932, FR Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 Family Species Coccus pseudomagnoliarum (Kuwana, 1914) Eucalymnatus tessellatus (Signoret, 1873) Regime A Phytophagous A Phytophagous Phytophagous Phytophagous Phytophagous A A A Native range 1st record in Europe Asia2001, FR, Temperate/ GR Japan Northern2001, FR America Tropical/ 1928, GB subtropical Asia1968, FR temperate Afrotropical/ 1829, FR South Africa Saissetia coffeae (Walker, 1852) A Phytophagous Afrotropical Saissetia oleae (Olivier, 1791) A Phytophagous Afrotropical A Phytophagous CentralAmerica Dactylopiidae Dactylopius coccus Costa, 1829 Invaded countries FR, GR, HR, IT, ITSIC,ME, SI Habitat F, G CH, DE, FR, GB, HR, IT, I2 LU, NL, SI GB, ES-CAN I2 AT, BE, CH, DE, FR, GB, IRL, LU, NL ES, ES-CAN,FR, GB, GR, IT, IT-SAR, IT-SIC, MT, PT-MAD, SI 1867, IT BG, CH, DK, ES, ESCAN, FR, FR-COR, GB, GR, HR, HU, IT, IT-SAR, IT-SIC, LV, MT, NL, PT, PT-AZO, PT-MAD, PL, RO, S, UA 1791, FR, AL, AT, BG, CH, CY,DK, IT ES, ES-CAN, FR, FRCOR, GB, GR, HR, IT, IT-SAR, IT-SIC, ME, PT, PT-AZO, PT-MAD, NL, RO, SK, SI, RS, UA 1827, ES- ES-CAN, PT-AZO, PTCAN MAD Hosts References Aesculus, Acer, Foldi (2001) Ficus Polyphagous Foldi (2001) Polyphagous Green (1928) I2 Polyphagous Canard (1968) I2 Aizoaceae Balachowsky (1932a) I2, J100 Polyphagous Leonardi (1920) I, I2 Olea europea, Nerium oleander, polyphagous Olivier (1791) I Cactaceae Russo and Mazzeo (1996) Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) Pulvinaria hydrangeae (Steinweden, 1946) Pulvinaria psidii Maskell, 1893 Pulvinaria regalis Canard, 1968 Pulvinariella mesembryanthemi (Vallot, 1830) Status 498 Family Species Pulvinaria horii Kuwana, 1902 Family Species Diaspididae Abgrallaspis cyanophylli (Signoret, 1869) Status Regime Native range 1st record in Europe Habitat Phytophagous Cryptogenic Aonidiella aurantii (Maskell, 1879) A Phytophagous Asia-Tropical/ 1881, IT China Aonidiella citrina (Coquillett, 1891) Aonidiella taxus Leonardi 1906 Aonidiella tinerfensis (Lindinger, 1911) A Phytophagous Phytophagous Phytophagous Asia-tropical 1994, IT BG, CZ, DK, ES-CAN, I2 FR, FR-COR, IT, IT-SAR, IT-SIC, PL CY, ES, ES-CAN, FR, FR- I, I2 COR, GR, IT, IT-SAR, IT-SIC, PT-MAD CY, FR, FR-COR, IT I, I2 Asia-tropical 1906, IT ES, FR, IT, IT-SIC Africa/ Canary Islands cryptogenic 1936, PT A Aspidiotus destructor Signoret 1869 Aspidiotus nerii (Bouché, 1833) C Aulacaspis tubercularis Newstead, 1906 Chrysomphalus aonidum (Linnaeus, 1758) C A A Phytophagous Phytophagous Phytophagous Phytophagous Hosts References Polyphagous Signoret (1869a) Citrus, Polyphagous Leonardi (1918) Longo et al. (1994) I2 Citrus, Polyphagous Taxus Leonardi (1906) PT I2 Dracaena Fernandes (1992) 1898, IT FR, IT J100 Leonardi (1898) Afrotropical 1829, IT I, I2 Leonardi (1920) Cryptogenic 1990, IT AL, CH, CY, CZ, DE, DK, ES, ES-CAN, FR, FR-COR, GB, GR, HU, HR, IT, IT-SAR, IT-SIC, MT, PT, PT-AZO, PTMAD, PL, RO, RS, SE, SI IT, IT-SIC, PT Palms, Polyphagous Nerium oleander, Polyphagous I2 Mangifera Porcelli (1990) SouthernAmerica 1895, IT Citrus, Polyphagous Leonardi (1920) 499 BE, DE, DK, ES, ESI2 CAN, FR, FR-COR, GB, GR, HR, IT, PT-MAD, PL, RS Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 C A 1868, FR Invaded countries A Phytophagous Chrysomphalus pinnulifer (Maskell, 1891) Comstockiella sabalis (Comstock, 1883) Diaspidiotus osborni (Newell & Cockerell, 1898) Diaspidiotus perniciosus (Comstock, 1881) C Phytophagous A Phytophagous Phytophagous NorthernAmerica NorthernAmerica A Phytophagous Asiatemperate/ China Diaspidiotus uvae (Comstock 1881) Diaspis boisduvalii Signoret 1869 A Phytophagous Phytophagous NorthernAmerica SouthernAmerica Phytophagous SouthernAmerica Diaspis bromeliae (Kerner, 1778) A A A Regime Native range 1st record Invaded countries in Europe Asia-Tropical 1896, IT CZ,DK, ES, ES-CAN, FR, FR-COR, GB, GR, HR, IT, PT, PL,PT-AZO, PT-MAD, PL, RO, RS Cryptogenic 1957, ES ES, ES-CAN, PT-MAD Habitat Hosts References I2, J100 Citrus, Polyphagous Berlese and Leonardi (1896) I2 Polyphagous Gómez-Menor Ortega (1957) I2 Palms 1979, BG BG, CH, IT, IT-SIC I2 Platanus Germain and Matile-Ferrero (2006) Kozár et al. (1979) 1928, HU AT, BG, CH, CZ, DE, DK, ES, ES-CAN, FR, FR-COR, GB, GR, HU, HR, IT, IT-SAR, IT-SIC, MD, NL, PT, PT-MAD, PL, RO, SE, SI, UA 1944, ES ES, ES-CAN G, I Fruit trees, Polyphagous Melis (1943) I Polyphagous Ruiz Castro (1944) 2005, FR 1868, FR 1868, FR FR BG, DE, DK, ES, ESI2, J100 Polyphagous CAN, FR, FR-COR, GB, GR, IT, IT-SIC, PT, PTMAD, SE A, B, BG, CH, CZ, DE, I2, J100 Bromeliaceae DK, ES, ES-CAN FR, GB, HU, IT, IT-SIC, MT, NL, PT-AZO, PT-MAD, PL, SE Signoret (1869b) Signoret (1869b) Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) Status 500 Family Species Chrysomphalus dictyopsermi (Morgan, 1889) Family Species Diaspis echinocacti (Bouché, 1833) Regime A Phytophagous A Phytophagous Phytophagous Phytophagous A A A A Gymnaspis aechmeae Newstead, 1898 Hemiberlesia lataniae (Signoret, 1869) C Hemiberlesia palmae (Cockerell, 1892) A C Phytophagous Phytophagous Native range 1st record Invaded countries in Europe Central1827, IT DE, DK, ES, ES-CAN, America FR, FR-COR, GB, GR, HU, HR, IT, IT-SAR, IT-SIC, LU, LT, PT, PTMAD Afrotropical/ 1999, IT, IT, IT-SIC South Africa IT-SIC Afrotropical/ 1985, GB GB South Africa Asia-Tropical 1867, IT ES-CAN, FR, GR, IT, ITSIC, MT, PT-MAD Asia-Tropical 1952, PT Afrotropical Phytophagous Phytophagous Cryptogenic Phytophagous SouthernAmerica Cryptogenic 1895, IT IT, PT CH, CZ, DE, DK, ES, ES-CAN, FR, GB, IT, IT-SAR, IT-SIC, PT, PTAZO, PT-MAD, PL, SE, UA 1898, GB BE, BG, CH, CZ, DE, ES, FR, IT, IRL, Pl, RO, S 1869, FR AT, BE, BG, CY, CZ, DE, ES, ES-CAN, FR, FR-COR, GB, GR, IT, IT-SIC, PT, PT-MAD, PL, RO 1920, GB CY, GB, PT, PT-MAD Habitat Hosts References I2, J100 Cactaceae Leonardi (1920) I2 Aizoaceae Russo et al. (1999) I2 Polyphagous Williams (1985) I2 Polyphagous Targioni Tozzetti (1886), (1885) I2 Polyphagous Baeta Neves (1954) I2, J100 Cycadaceae, Zamiaceae Berlese and Leonardi (1896) I2, J100 Bromeliaceae Newstead (1898) I2, J100 Polyphagous Signoret (1869a) I2, J100 Palms Green (1920) Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 Entaspidiotus lounsburyi (Marlatt, 1908) Eulepidosaphes pyriformis (Maskell, 1897) Fiorinia fioriniae (Targioni Tozzetti, 1867) Fiorinia pinicola Maskell, 1897 Furchadaspis zamiae (Morgan, 1890) Status 501 Lepidosaphes gloverii (Packard, 1869) Leucaspis podocarpi (Green, 1929) Lindingaspis rossi (Maskell, 1891) Lopholeucaspis cockerelli (Grandpré & Charmoy, 1899) Oceanaspidiotus spinosus (Comstock, 1883) Odonaspis greenii (Cockerell, 1902) Regime C Phytophagous C Phytophagous Phytophagous Phytophagous C A C Phytophagous C Phytophagous Phytophagous Phytophagous Phytophagous A A C C Phytophagous A Phytophagous Native range 1st record Invaded countries in Europe Cryptogenic 1881, IT CZ, ES, ES-CAN, FR, GR, IT, IT-SAR, IT-SIC, MA, PT, PT-AZO, PTMAD, PL Cryptogenic 1896, IT FR, IT Habitat Hosts References I2, J100 Polyphagous Leonardi (1920) I2, J100 Polyphagous Berlese and Leonardi (1896) Cryptogenic 1954, FR CZ, DK, ES-CAN, F I2, J100 Polyphagous Balachowsky (1954) Asiatemperate/ China Japan Cryptogenic 1900, IT AL,FR, HR, IT, PL, SI, UA I2 Bamboos Lupo (1938) Polyphagous Bouché (1851) Citrus, Polyphagous Podocarpus Targioni Tozzetti (1884) Cryptogenic 1850, DE BG, CY, ES, ES-CAN, FR, I2 FR-COR, GB, GR, HR, IT, IT-SAR, IT-SIC, MA, PT, PT-AZO, PT-MAD 1884, IT ES, FR, FR-COR, HR, IT, I2 IT-SAR, IT-SIC, GR, P 1985, GB GB I2 Australasia/ New-Zealand Australasia/ 1942, PT ES, FR, IT, IT-SIC, PT, Australia PT-MAD Cryptogenic 1908, DE DE, GB, GR F, G, I2 Polyphagous Seabra de (1942) J100 Lindinger (1908) Cryptogenic I2, J100 Polyphagous Leonardi (1897) I2, J100 Bamboos Zahradnik (1990) 1890, ITSIC ES, ES-CAN, GB, IT, IT-SIC, PT, PT-AZO, PT-MAD Asia-Tropical 1963, CZ CZ, ES, IT Orchidaceae Williams (1985) Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) Howardia biclavis (Comstock, 1883) Ischnaspis longirostris (Signoret, 1882) Kuwanaspis pseudoleucaspis Kuwana, 1923 Lepidosaphes beckii (Newman, 1869) Status 502 Family Species Hemiberlesia rapax (Comstock, 1881) Status A A A A C C Regime Phytophagous Phytophagous Phytophagous Phytophagous Phytophagous Phytophagous Native range 1st record Invaded countries in Europe Asia-Tropical 1929, FR FR Cryptogenic C Phytophagous Cryptogenic Parlatoria theae Cockerell, 1896 Parlatoria ziziphi (Lucas, 1853) C Phytophagous Phytophagous Cryptogenic Pinnaspis aspidistrae (Signoret, 1869) A Phytophagous Pinnaspis buxi (Bouché, 1851) C Phytophagous A References Bamboos Balachowsky (1930) FR I2 Opuntia Balachowsky (1932a) ES, FR, IT, PT I2 Palms Lupo (1948) ES, FR, IT, IT-SIC, PT, PT-MAD 1887, GB FR, GB, IT, HU I2 Camellia, Polyphagous Croton Leonardi (1903) Last 1899, CY, DE, ES, ES-CAN, IT FR, FR-COR, GR, HR, IT, IT-SAR, IT-SIC, MT, PT, PT-MAD 1939, FR BG, CZ, DE, DK, FR, PL, UA I2, J100 Citrus, Polyphagous 1953, FR Asia-Tropical 1853, FR I2 I2, J100 Palms, orchids, Polyphagous ES, FR, PT-MAD, PL, UA I2 Polyphagous BG, CY, ES, ES-CAN, FR, I2 FR-COR, GR, HR, IT, IT-SAR, IT-SIC, PT, UA Asia-Tropical 1868, FR B, BG, CZ, DE, ES, ES- J100 CAN, FR, FR-COR, GB, HU, IT,IT-SIC, IE, MT, NL, PT, PT-MAD, PL, S, UA Cryptogenic 1851, DE DE, DK, FR, IT J100 Douglas (1887) Berlese and Leonardi (1899) Morrison (1939) Balachowsky (1953) Citrus, Rutaceae Lucas (1853) Polyphagous Signoret (1869b) Polyphagous Balachowsky (1938) 503 Parlatoria proteus (Curtis, 1843) Hosts I2 Southern1929, FR America Arabian 1947, IT peninsula Asia-Tropical 1903, IT Cryptogenic Habitat Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 Family Species Odonaspis secreta (Cockerell, 1896) Opuntaspis philococcus (Cockerell, 1893) Parlatoria blanchardi Targioni Tozzetti, 1883 Parlatoria camelliae Comstock, 1883 Parlatoria crotonis Douglas, 1867 Parlatoria pergandii Comstock 1881 J100 Polyphagous Tranfaglia and Viggiani (1988) J100 Asia-Tropical 1949, IT IT J100 Cycadaceae, Ericaceae Camellia Anagnou–Veroniki et al. (2008) Pegazzano (1949) Asia-Tropical 1992, IT FR, FR-COR, IT, IT-SIC, J100 SI AT, BG, CH, DE, ES, ES- G, J, I CAN, FR, FR-COR, GB, GR, HU, HR, IT, IT-SAR, IT-SIC, MA, NL, PT, PTMAD, SI, UA CZ, DE, ES, FR, IT, PT- I2, J100 MAD Polyphagous Russo and Mazzeo (1992) Fruit trees, Polyphagous Targioni Tozzetti (1867) Polyphagous Leonardi (1918) Polyphagous Balachowsky (1954) Asia-Tropical? 1886, IT Habitat Hosts References C Phytophagous Cryptogenic 1918, IT C Phytophagous Phytophagous Phytophagous Phytophagous Phytophagous Cryptogenic 1954, FR FR I2 Cryptogenic 2002, FR FR I2, J100 Polyphagous Afrotropical/ 1991, IT South Africa Afrotropical 1990, IT IT I2 Euphorbiaceae Marotta and Garonna (1991) IT I2 Polyphagous Pellizzari (1993) AsiaTemperate/ Eastern Asia AT, BG, CH, DE, ES, ES- I2 CAN, FR, FR-COR, GB, GR, HU, HR, IT, IT-SAR, IT-SIC, MT, NL, PL, PT, RO, SI, UA Euonymus Targioni Tozzetti (1884) C A A A 1884, IT Germain et al. (2002) Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) Pseudoparlatoria parlatorioides (Comstock, 1883) Pseudoparlatoria ostreata Cockerell, 1892 Rutherfordia major (Cockerell, 1894) Selenaspidus albus McKenzie, 1953 Umbaspis regularis (Newstead, 1911) Unaspis euonymi (Comstock, 1881) Native range 1st record Invaded countries in Europe Cryptogenic 1988, IT DE, ES-CAN, FR, GB, IT, PL Asia-tropical 2007, GR GR 504 Family Status Regime Species Pinnaspis strachani C Phyto(Cooley, 1899) phagous Poliaspis cycadis PhytoComstock, 1833 phagous Pseudaonidia paeoniae A Phyto(Cockerell, 1899) phagous Pseudaulacaspis cockerelli A Phyto(Cooley, 1897) phagous Pseudaulacaspis A Phytopentagona (Targioni phagous Tozzetti, 1886) Family Species Unaspis yanonensis (Kuwana, 1923) Eriococcidae Eriococcus araucariae Maskell, 1879 Ortheziidae Insignorthezia insignis (Browne, 1997) Native range 1st record Invaded countries in Europe Asia-Tropical 1969, FR ES, FR, FR-COR, IT Habitat A Phytophagous A Phytophagous Australasia/ Australia A Phytophagous Phytophagous NorthernAmerica NorthernAmerica Phytophagous Phytophagous Asia-Tropical 2001, FR FR Australasia/ Australia 1900, IT AL, CH, CY, ES, ESI, I2 CAN, FR, FR-COR, GR, HR, IT, IT-SAR, IT-SIC, MT, PT, PT-AZO, PTMAD, RO, SI A Phytophagous SoutherrnAmerica 1887, GB AT, CH, CZ, DE, DK, ES-CAN, FR, GB, HU, HR, PT, PT-AZO, PTMAD A Phytophagous North Africa 1930, FR A Phytophagous Asia-Tropical 1937, FR A A A I, I2 Hosts Citrus 1895?, IT ES, ES-CAN, FR, FRI2 Araucaria COR, GR, HR, IT, IT-SAR, IT-SIC, PT, PTAZO, PT-MAD 1930, FR FR,FR-COR, GR, HR, I2, J100 Cactaceae IT, IT-SIC 1888, GB FR, FR-COR, IT, IT-SIC, I2, J100 Agavaceae UA I2 References Bénassy (1969) Leonardi (1899) Balachowsky (1932a) Green (1915) Polyphagous Foldi (2001) Polyphagous Leonardi (1920) I2, J100 Polyphagous Douglas (1889) ES, FR, FR-COR, IT, ITSIC, PT-MAD I2 Palms Balachowsky (1930) ES, FR, GB, HR, UA I2 Poaceae Goux (1937) 505 Phoenicoccocidae Phoenicococcus marlatti (Cockerell, 1899) Pseudococcidae Antonina crawi Cockerell, 1900 Regime Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 Eriococcus coccineus Cockerell, 1894 Ovaticoccus agavium (Douglas, 1888) Margarodidae Icerya formicarum Newsteadt, 1897 Icerya purchasi (Maskell, 1879) Status A A A Phytophagous Phytophagous Phytophagous Australasia/ Australia Asia Habitat Hosts References E, I2 Poaceae Marotta (1992) 1918, IT FR, GB, IT, UA J100 Phormium Leonardi (1918) Last 2007,IT IT I2 Bamboos Pellizzari and Danzig (2007) Asia-Tropical 1990, IT IT I2 Bamboos Porcelli (1990) E, I Tranfaglia (1981) I2 Agavaceae, Gramineae Polyphagous I Polyphagous Balachowsky (1938) J100 Polyphagous Jansen (1995) I, J100 Polyphagous Leonardi (1913) I2 Bromeliaceae Marotta (1992) I2 Polyphagous Jansen (1995) I2, J100 Polyphagous Jansen (1995) I2, J100 Polyphagous Jansen (1995) Northern1979, GR FR, GR, HU, IT, IT-SAR America Afrotropical/ 1977, IT FR, IT, IT-SIC South Africa Cryptogenic Last 1938 PT-MAD PT-MAD 1933, NL ES-CAN, IT, IT-SIC,NL, PT-AZO, PT-MAD Last 1913 ES-CAN, FR ES-CAN 1989, IT IT CentralAmerica CentralAmerica SouthernAmerica/ Mexico Southern1988, NL IT, IT-SIC, NL America Southern1994, NL FR, NL America Asia-Tropical 1967, NL DK, FR, NL Tranfaglia (1981) Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) Dysmicoccus neobrevipes Beardsley 1959 Ferrisia virgata (Cockerell, 1893) Geococcus coffeae Green, 1933 Native range 1st record Invaded countries in Europe Asia-Tropical 1992, IT FR, IT 506 Family Status Regime Species Antonina graminis A Phyto(Maskell, 1897) phagous Balanococcus diminutus A Phyto(Leonardi, 1918) phagous Balanococcus kwoni A PhytoPellizzari & Danzig phagous 2007 Chaetococcus bambusae A Phyto(Maskell, 1892) phagous Chorizococcus rostellum A Phyto(Lobdell, 1930) phagous Delottococcus euphorbiae A Phyto(Ezzat & McConnell, phagous 1956) Dysmicoccus boninsis C Phyto(Kuwana, 1909) phagous Dysmicoccus brevipes A Phyto(Cockerell, 1893) phagous Dysmicoccus grassii A Phyto(Leonardi, 1913) phagous Dysmicoccus mackienziei A PhytoBeardsley 1965 phagous Native range 1st record Invaded countries in Europe Southern1986, IT FR, FR-COR, GR, IT, America IT-SIC Habitat I2 Hosts Cactaceae References Süss and Trematerra (1986) CentralAmerica Cryptogenic 1917, GB ES, ES-CAN, GB, IT-SIC, I2, J100 Polyphagous PT-MAD 2004, FR ES-CAN, F J100 Palms Chapin and Germain (2005) NorthernAmerica NorthernAmerica 1976, IT IT G, I2 Corylus Tranfaglia (1976) 1946, ES ES, ES-CAN I2 Polyphagous Gómez-Menor Ortega (1946) 1923, PT- FR, IT, IT-SIC, PT-MAD MAD 1999, IT, AL, IT, IT-SIC IT-SIC 1813, FR BG, CH, CY, CZ, ES, ES-CAN, FR, FR-COR, GB,GR, HU, HR, IT, IT-SAR, IT-SIC, NL, PL, PT, PT-AZO, PT-MAD, SI, UA 1989, IT IT I2 Polyphagous Green (1923) I2 Polyphagous Mazzeo et al. (1999) SouthernAmerica NorthernAmerica Asia-Tropical Planococcus halli Ezzat & McConnel, 1956 C Phytophagous Cryptogenic Pseudococcus calceolariae (Maskell, 1879) A Phytophagous Australasia/ Australia I2, J100 Polyphagous Risso (1813) I2 Marotta (1992) Nerium oleander, Polyphagous I2, J100 Polyphagous Green (1915) 507 1914, GB BG, CZ, ES, ES-CAN, FR, FR-COR, GB, HR, IT, IT-SAR, IT-SIC, PT, PT-AZO, UA Green (1917) Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 Family Status Regime Species Hypogeococcus pungens A PhytoGranara de Willink, phagous 1981 Nipaecoccus nipae A Phyto(Maskell, 1893) phagous Palmicultor palmarum C Phyto(Ehrhorn, 1916) phagous Peliococcus serratus A Phyto(Ferris, 1925) phagous Phenacoccus gossypii A PhytoTownsend & Cockerell, phagous 1898 Phenacoccus madeirensis A PhytoGreen, 1923 phagous Phenacoccus solani A PhytoFerris, 1918 phagous Planococcus citri (Risso, A Phyto1813) phagous Status A Phytophagous Phytophagous Native range 1st record in Europe Asialast 1989, Temperate MD Australasia/ 1867, IT Australia Invaded countries ES-CAN,FR, IT, MD, PT-MAD BG, CZ, DK, ES, ESCAN, FR, FR-COR, GB, GR, HU, HR, IT, IT-SAR, IT-SIC, LV, MT, PT, PTAZO, PT-MAD, PL, SI, UA B, BG, DE, DK, CZ, ES, ES-CAN, FR, FR-COR, GB, GR, HU, HR, IT, IT-SAR, IT-SIC, MT, NL, PT, PT-AZO, PT-MAD, PL, SI, SK, RS, UA IT, IT-SIC Habitat I, I2 Hosts Polyphagous References Ben-Dov (1994) I2, J100 Polyphagous Targioni Tozzetti (1886), (1885) I, I2 Polyphagous Signoret (1875) Polyphagous Russo and Mazzeo (1992) Pseudococcus viburni (Signoret, 1875) A Phytophagous NorthernAmerica 1875, FR Rhizoecus americanus (Hambleton, 1946) Rhizoecus cacticans (Hambleton, 1946) Rhizoecus dianthi Green, 1926 Rhizoecus latus (Hambleton, 1946) Spilococcus mamillariae (Bouché, 1844) Trionymus angustifrons Hall, 1926 A Phytophagous Phytophagous Phytophagous Phytophagous Phytophagous Phytophagous NorthernAmerica SouthernAmerica Australasia/ Australia SouthernAmerica NorthernAmerica Arabian peninsula 1992, IT, I2 IT-SIC 1961, NL BY, CZ, DK, ES-CAN, IT, I2 IT-SIC, NL, PL 1961, NL CZ, DK, FR, IT, NL, PL I2 Polyphagous Jansen (1995) Polyphagous Jansen (1995) 1995, IT IT I2 Polyphagous Marotta (1995) 1979, IT I2, J100 Cactaceae Tranfaglia (1981) 1966, PL CZ, DE, DK, FR, GB, HU, IT, IT-SIC CH, FR, PL E, I2 Koteja and Zak-Ogaza (1966) Phytophagous Afrotropical 1990, IT FR,IT J100 Trochiscococcus speciosus (De Lotto, 1961) A A A A A A Compositae, Tamarix, Urtica Liliaceae Williams and Pellizzari (1997) Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) A Regime 508 Family Species Pseudococcus cosmtocki (Kuwana, 1902) Pseudococcus longispinus (Targioni TozzettIT, 1868) Status A A A Regime Phytophagous Phytophagous Phytophagous Native range 1st record Invaded countries in Europe Afrotropical 1933, IT BG, DE, ES, FR,GR, IT, NL, P Afrotropical 1975, DK BE, DK, GB Afrotropical/ 1975, IT South Africa FR, IT, IT-SIC Habitat Hosts References I2 Polyphagous Menozzi (1933) I2 Polyphagous I2 Crassulaceae Kozarzhevskaya and Reitzel (1975) Tranfaglia (1981) Scales (Hemiptera, Superfamily Coccoidea). Chapter 9.3 Family Species Vryburgia amaryllidis (Bouché, 1837) Vryburgia brevicruris (McKenzie, 1960) Vryburgia rimariae Tranfaglia, 1981 509 510 Giuseppina Pellizzari & Jean-François Germain / BioRisk 4(1): 475–510 (2010) Table 9.3.2. List and main characteristics of the scale species alien in Europe. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Only selected references are given. Last update 29/05/2009 Family Species Diaspididae Aonidiella tinerfensis Lindinger (1911) Aonidiella lauretorum (Lindinger, 1911) Margarodidae Marchalina hellenica (Gennadius, 1883) Matsucoccus feytaudi Ducasse 1941 Regime Native range Invaded Habitat countries Hosts References Phytophagous Canary Islands PT I2 Dracaena Balachowsky (1948), Fernandes (1992), (1990) Phytophagous Canary Islands, Madeira PT I2 Polyphagous Balachowsky (1948) Phytophagous Greece, Turkey IT G Pinus Tranfaglia and Tremblay (1984) Phytophagous France, Spain, Portugal IT, FRCOR G Pinus pinaster Arzone and Vidano (1981), Jactel et al. (1996) A peer reviewed open access journal BioRisk 4(1): 511–552 (2010) doi: 10.3897/biorisk.4.63 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea) and Hemiptera Auchenorrhyncha Chapter 9.4 David Mifsud1, Christian Cocquempot2, Roland Mühlethaler3, Mike Wilson4, Jean-Claude Streito5 1 Junior College, Department of Biology, University of Malta, Msida MSD 1252, Malta 2 INRA UMR Centre de Biologie et de Gestion des Populations, CBGP, Campus international de Baillarguet, CS 30016, 34988 Montferrier-sur Lez, France 3 Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity, Humboldt University Berlin, Invalidenstrasse 43, 10115 Berlin, Germany 4 Department of Biodiversity & Systematic Biology, National Museum Wales, Cathays Park, Cardiff CF10 3NP, United Kingdom 5 Laboratoire national de la protection des végétaux, CBGP, Campus international de Baillarguet, CS 30016, 34988 Montferrier-sur Lez, France Corresponding author: David Mifsud (david.a.mifsud@um.edu.mt), Christian Cocquempot (cocquemp@ supagro.inra.fr), Roland Mühlethaler (roland.muehlethaler@mfn-berlin.de), Mike Wilson (mike.wilson@ museumwales.ac.uk), Jean-Claude Streito (streito@supagro.inra.fr) Academic editor: Wolfgang Rabitsch | Received 19 May 2010 | Accepted 24 May 2010 | Published 6 July 2010 Citation: Mifsud D et al. (2010) Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea) and Hemiptera Auchenorrhyncha. Chapter 9.4. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 511–552. doi: 10.3897/biorisk.4.63 Abstract Apart from aphids and scales, 52 additional Sternorrhyncha hemipteran species alien to Europe have been identified within Aleyrodidae (27 whitefly species), Phylloxeroidea (9 adelgids, 2 phylloxerans) and Psylloidea (14 species of jumping plant-lice) in addition to 12 Auchenorrhyncha species (mostly Cicadellidae- 8 species). At present, the alien species represent 39% of the total whitefly fauna and 36% of the total adelgid fauna occuring in Europe. The proportion is insignificant in the other groups. The arrival of alien phylloxerans and adelgids appeared to peak during the first part of the 20th century. In contrast, the mean number of new records per year of alien aleyrodids, psylloids and Auchenorrhyncha increased regularly after the 1950s. For these three groups, an average of 0.5–0.6 new alien species has been recorded per year in Europe since 2000. The region of origin of the alien species largely differs between the different groups. Alien aleyrodids and psylloids mainly originated from tropical regions whilst the adelgids and phylloxerans came equally from North America and Asia. A major part of the alien Auchenorrhyncha originated Copyright D. Mifsud et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 512 David Mifsud et al. / BioRisk 4(1): 511–552 (2010) from North American. Most of these alien species are presently observed in man-made habitats, especially in parks and gardens but alien adelgids are mainly observed in forests because of their association with conifer trees used for afforestation. Keywords alien, Europe, Adelgidae, Aleyrodidae, Cicadellidae, Psyllidae, Phylloxeridae, Auchenorrhyncha 9.4.1. Introduction This chapter will consider the hemipteran species alien to Europe belonging to the Sternorrhyncha superfamilies other than Aphidoidea and Coccoidea (i.e., Aleyrodoidea, and superfamilies Phylloxeroidea and Psylloidea) and to the Auchenorrhyncha (Cicadomorpha and Fulgoromorpha suborders). We will mainly follow the higher classification used in Fauna Europaea (Asche and Hoch 2004, Nieto Nafria and Binazzi 2005). Both Aleyrodoidae (whiteflies) and Psylloidea (jumping plant-lice or psylloids) are distributed throughout the major zoogeographical regions of the World, with their greatest diversity in tropical and south temperate regions. They are all sap-sucking insects and most of them are narrowly host-plant specific. This is particularly true for the psylloids were such specificity may also be present at higher taxonomic levels and not just at species level. Both adult whiteflies and psylloids possess a feeding rostrum, two pairs of flying wings and are fully mobile. Reproduction in both groups is generally sexual with some rare cases of parthenogenetic* development. The eggs in both groups are laid directly onto the host-plant surface. Whiteflies comprise a relatively small group of insects in a unique family Aleyrodidae, and we will later us only this family name. Whiteflies are the least speciose among the four groups of sternorrhynchous Hemiptera (whiteflies, aphids, jumping plant-lice and scale insects) with only 1,556 described species accommodated in 161 genera (Martin and Mound 2008). Adult whiteflies are very small insects, most measuring between 1–3 mm in body length. Life-cycles of whiteflies are somewhat unusual. The first-instar larvae are able to walk around (crawler) short distances on the host plant until a suitable feeding site is found; then, the remaining three larval instars are sessile. The final whitefly larval stage is usually termed as a puparium* where feeding goes on during the first part of this stage. It is also this stage which is used for almost all whitefly taxonomy and systematic with adults being identified only rarely. All whitefly species are free living during their larval stages. Jumping-plant lice (Psylloidea) comprise some 3,000 described species accommodated in the six currently recognized families. Adults range from 1–12 mm in body length. Life-cycles of psylloids are very straightforward with eggs laid singly or in clusters on the host plant, the immatures undergoing five larval instars (being all mobile unless gall-dwelling) and after these adults emerge. In jumping-plant lice, both adults and nymphal stages are used for species identifications. More than three-quarters of Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... 513 psylloid species are free-living during their larval stages, but some are gall-inducing and others live under protective scales or lerps (waxy constructions covering the body). The feeding activity of whiteflies and psylloids may negatively affect the host-plant by rendering weakness and thus more susceptibility to other diseases. The feeding activity of these insects (especially in whiteflies) may produce copious honeydew which may cover underlying leaves and fruits/flowers of the host-plant. Usually, this honeydew is immediately covered by black sooty mould which impairs photosynthesis and/ or renders unmarketable plant parts such as flowers and fruits. Notorious pest species in both groups (adults) are vectors of a number of plant pathogens such as viruses and phytoplasmas. Phylloxeroidea (adelgids and phylloxerans) is a closely related superfamily, which include some of the most destructive introduced plant pests in the World. They include minute insects (1–2 mm in body length), which are highly host specific but with a simple morphology. The two groups are distinguished from typical aphids (Aphididae) by the complete absence of siphunculi* and the retention of the ancestral trait of oviparity in all generations. Phylloxerans feed on angiosperms, particularly hickories and ashes (Juglandaceae), oaks and beeches (Fagaceae) and grapes (Vitaceae) but adelgids only develop on certain genera of the Pinaceae family, retaining their ancestral relationships with gymnosperms. Such as their host plants, adelgids are endemic to the Northern Hemisphere in boreal and temperate habitats. Despite the broad geographical distribution of these host plants, there are less than 70 and ca. 75 species of known adelgids and phylloxerans, respectively (Havill and Foottit 2007). However, there is considerable taxonomic uncertainty in both groups since several described species may not represent unique taxa but are actually different morphological forms of the same species found on different host plants. Both groups exhibit cyclical parthenogenesis and possess complex, multigenerational, polymorphic life cycles. Five generations make up the typical two- year adelgid holocycle*, three produced on the primary host, Picea spp. (noticed as -I- in Table 9.4.1) where sexual reproduction and gall formation occurs, and the last two are produced on a secondary host (Abies, Larix, Pseudotsuga, Tsuga, or Pinus, noticed as -II- in Table 9.4.1) which supports a series of asexual generations. Adelgids that are anholocyclic* complete their entire life cycle on either Picea or on a secondary host genus. Some anholocyclic species may in fact be holocyclic, but forms on the alternate host have not been described. Typically, sexual reproduction and host alternation nymphs and galls are formed in spring. Winged gallicolae* can disperse or can stay to lay eggs near the gall from which they emerged. Auchenorrhyncha, with some 42,000 described species worldwide is probably paraphyletic but composed of two well supported monophyletic groups, Fulgoromorpha (planthoppers) and Cicadomorpha (leafhoppers, froghoppers, treehoppers and cicadas). Hemipteran phylogeny is still controversial (Cryan 2005, Yoshizawa and Saigusa 2001) although Sternorrhyncha, Fulgoromorpha, Cicadomorpha, Coleorrhyncha and Heteroptera are considered monophyletic by most authors (Bourgoin and Campbell 2002, 514 David Mifsud et al. / BioRisk 4(1): 511–552 (2010) Dietrich 2002, Nielson 1985). Auchenorrhyncha usually feed on plant sap, either on phloem, xylem or parenchyma, and they occur therefore in almost all habitats colonized by vascular plants. Many are of economic importance due to the transmission of phytopathogenic organisms causing plant diseases such as phytoplasmas and virus diseases (Bourgoin and Campbell 2002, Carver et al. 1991, Dietrich 2005, Kristensen 1991, Nielson 1985). Most Auchenorrhyncha have a bisexual reproduction. Eggs are usually laid into plant tissue and there are 5 nymphal instars. While some species are good flyers and can be carried by wind over relatively long distances (Della Giustina and Balasse 1999), most of the translocations are considered due to anthropogenic causes. All the species introduced from North American and east Asiatic are assumed to have been imported with plants, either as eggs in the tissue or as nymphs or adults feeding on the host plants. Planthoppers (Fulgoromorpha) with 21 families and some 12,000 described species occur worldwide but are most diverse in the tropics. Only the widely distributed families Cixiidae and Delphacidae occur also in colder regions such as Northern Europe. In Europe, ca. 750 species of Fulgoromorpha are expected to occur (Asche and Hoch 2004). They can be distinguished by the following characters: pedicel of antenna bulbous or enlarged; presence of tegulae* on the mesothorax; bases of mid-coxae widely separated. The body size varies from 2–114 mm but most species are small (O’Brien and Wilson 1985). Cicadomorpha are characterised by following characters: antennal pedicel small; tegulae absent; meso-coxae small and narrowly separated. To date, 30,000 species of Cicadomorpha have been described in over than 5,000 genera and 13 families. Dietrich (Dietrich 2002) estimated that about 6–10% of plant-feeding insects belong to the Cicadomorpha. Despite their economic importance, there are surprisingly still many gaps in the knowledge on the taxonomy, phylogeny, life history and biology of Auchenorrhyncha. 9.4.2. Taxonomy and invasion history of the Aleyrodidae, Psylloidea, Phylloxeroidea, and Auchenorrhyncha alien to Europe The literature about alien species of Aleyrodidae, Psylloidea, Phylloxeroidea, and Auchenorrhyncha in Europe is relatively scattered, most of the studies dealing with alien pests of economic importance such as Bemisia tabaci and Trialeurodes vaporarium (Bedford et al. 1994, Martin et al. 2000) for Aleyrodidae or Metcalfa pruinosa and Scaphoideus titanus (Arzone et al. 1987, Dlabola 1981) for Auchenorrhyncha. Indeed, comprehensive data on alien species were available for only a few European countries. i.e., Albania, Bulgaria and Macedonia (Tomov et al. 2009), Austria (Essl and Rabitsch 2002), the Czech Republic (Šefrová and Laštùvka 2005), Germany (Geiter et al. 2002), Great Britain (Hill et al. 2005), Slovenia (Seljak 2002) and Switzerland (Kenis 2005). The ‘Handbook of alien species in Europe’ (DAISIE 2009), generated by the DAISIE project, listed a number of species alien to Europe (i.e, of exotic origin or cryptogenic) and alien in Europe (introduced by man from a European region to another where the species is not native) but the status of some of these species also Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... 515 needed to be reviewed. At the end of each group, we provide information on the species of this group we excluded from the alien list either because of confusion in their actual status or of misidentifications. Apart from the established species, the alien lists of Aleyrodidae, Phylloxeroidea and Psylloidea will also include species which were observed only in greenhouses and for which no data is available on their establishment in the wild in the mentioned territory. In contrast, the list of alien Auchenorrhyncha will only include established species in the wild. 9.4.2.1 Aleyrodidae A total of 27 species alien to Europe were recorded. Although the family Aleyrodidae include three subfamilies only two of these are represented in both the alien and the native European fauna. At present, the alien species represent 39% of the total whitefly fauna observed in Europe (Figure 9.4.1). Twenty alien species belong to Aleyrodinae, which is the most widespread and largest subfamily with over 1,400 described species. Seven species belong to the subfamily Aleurodicinae, which is mainly confined to South America, plus very few species in South-Eastern Asia and other geographical regions (121 described species) (Martin 1996). It is usually regarded as being more primitive than Aleyrodinae. In general, Aleurodicinae represent much larger species than typical whitefly, their additional wing venation being possibly a functional necessity associated with their large size. The pupal cases of the Aleurodicinae are generally more complex than those of the Aleyrodinae, bearing large compound wax-secreting pores on the dorsal surface. Species of whiteflies intercepted in greenhouses (occasionally or once) are rather few. Such species were included in the list because additional introductions as well as establishment in the wild are not to be excluded especially under global change conditions. These species include Filicaleyrodes williamsi, a species whose origin remains obscure; Aleuropteridis filicicola, an African species found on ferns; Aleurotulus nephrolepidis, a specialist fern feeder often found in greenhouses which is already known to occur in the wild in Macaronesia (Martin et al. 2000); Ceraleurodicus varus, an Aleurodicinae species which was found to colonize orchids in 1939- 1940 in an orchid house at the Budapest Botanical Garden, but was never intercepted again or recorded in other European countries; Aleurodicus destructor of which a single specimen was collected from Olea at a Garden Festival in Liverpool, UK, but which is occasionally intercepted by quarantine inspections in Europe (Martin 1996); a neotropical whitefly, Aleurotrachelus trachoides was intercepted in Great Britain on sweet potato leaves imported from Gambia (Malumphy 2005); and, Pealius azaleae. This latter species is often regarded as a minor pest of ornamental azaleas (Rhododendron spp.). It was originally described from Belgian material intercepted by quarantine officials in the United States but its origin is likely Eastern Asia. The occurrence of this species in Europe is very sporadic and records often reflect newly introduced populations with azalea hosts being kept indoors, in greenhouses or in very sheltered places. 516 David Mifsud et al. / BioRisk 4(1): 511–552 (2010) Figure 9.4.1. Comparison of the relative importance of Aleyrodidae, Psylloidea, Phylloxeroidea, and Auchenorrhyncha in the alien and native entomofauna in Europe. The number right to the bar indicates the number of species per family. An emergent whitefly pest in Europe is Alerocanthus spiniferus, commonly known as the Orange Spiny Whitefly. This species is listed as a quarantine threat to Europe and is included in the EPPO A1-List of species recommended for regulation as quarantine pests and in the EU Annex II/A1 under: “Pests known not to occur in the EU, whose introduction into, and/or whose spread within, all EU Member States is prohibited, with reference to specific plants or plant products”. The accidental introduction, acclimatization and spreading of this species in southern Italy (Porcelli 2008) is thus of concern to all the European Union. As pointed out by Porcelli (Porcelli 2008), the origin of the infestation of this species is still unknown, and the species has already spread in the Apulia Region to make its eradication impossible. A. spiniferus is a widespread tropical species, occasionally a pest on Annona and Citrus, but it is also recorded from woody hosts of more than 15 plant families (Martin 1996). Aleuroclava aucubae, a species described from Japan and most likely of Oriental origin, was recently recorded from Italy (Pellizari and Šimala 2007) and may also prove to be a potential pest in Europe. It is known to occur on more than 15 plant families (Mound and Halsey 1978) and in the Veneto region, the species was found on both greenhouse plants (Citrus x limon (L.) Osb., Ficus sycomorus L.) and outdoor host plants (Pittosporum tobira (Thunb.) Aiton , Prunus armeniaca L., Photinia). Some whitefly species not native to Europe have been found in Macaronesia and some of these are also penetrating into Europe. Aleuroplatus perseaphagus is a species of Neotropical origin, but was first described from Madeira. The species is common on avocado. Aleurotrachelus atratus is also a species of Neotropical origin, but was found in the Canary Islands (Martin et al. 2000) and is now being recorded on several endangered palm species on various islands in the south-western Indian Ocean and in glasshouses in Paris (Borowiec et al. 2010). Acaudaleyrodes rachipora was described from India and is probably native to Asia but the species is also known from the Ca- Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... 517 nary islands (Martin et al. 2000). Crenidorsum aroidephagus, introduced in Madeira, is a native of New World, colonising several plant species of the Araceae family in Central and South America, southern USA, and the Pacific Region. It is is also reported as a minor pest for growers of ornamental-foliage plants (Martin et al. 2001). Massilieurodes chittendeni is most probably a species originating from northern Asia, from where its host plant, rhododendrons, mainly originate. This species was described on material collected in England in 1928 (Laing 1928). Klasa et al. (2003) reported the introduction of this species to central Poland, the Czech Republic, Germany and the Netherlands. Two whitefly species with an uncertain area of origin include Dialeurodes kirkaldyi and Singiella citrifolii both potential pests of Citrus-plantations. D. kirkaldyi was originally described from Hawaii and later reported in several states in North America (Russell 1964). The species is also known from Africa and Asia. In Europe it was so far found in Cyprus and Portugal. S. citrifolii was originally described from the United States. It is known from the Oriental Regions and from the Neotropics and the Nearctic Region. In Europe the species is known from Madeira (Aguiar 1998) and recently it was reported from the Mediterranean Region (Lebanon) (Martin 2000). Parabemisia myricae, commonly known as the Japanese bayberry whitefly, is probably native to Japan. It arrived in the Mediterranean Basin and Southern Europe in the mid 1980s and in a very short time it invaded most of the Mediterranean countries with considerable damage to citrus plantations (Rapisarda et al. 1990). Some alien whitefly species show little dispersion in Europe. Trialeurodes packardi, a species native to the Nearctic Region where it is extremely polyphagous, was only noted in Hungary (Kozár et al. 1987) as a pest on strawberries. T. packardi is closely related to T. vaporariorum, and the two species can only be distinguished via microscopic examination of pupal cases, and this may also be a reason why the species was not recorded elsewhere in Europe. A highly polyphagous Neotropical species is Aleurodicus dispersus, commonly known as the Spiralling Whitefly. This species is occasionally detected in northern Europe on plants imported from the Far East (Martin 1996). In the 1970s this species began a rapid expansion of its range, westwards from the New World, and crossed the Pacific to the Philippines by 1983, and in 1990 its arrival in the Malay Peninsula was noted. Since then its spread continued into Thailand, Sri Lanka, southern India, the Maldive Islands, and Western Africa (Martin 1996). Its establishment in the Canaries dates back to the early 1960s, but the species is also known from Macaronesia where it is common on trees and shrubs in the open and seems to be a well established species. A species which co-exists with A. dispersus in the Canary Island is Lecanoides floccissimus, a second Neotropical species which is particularly damaging to numerous unrelated host-plants due to direct feeding and by the enormous populations depriving plants of sap and thus inhibiting growth. The species is also known to secrete copious honeydew on which sooty mould immediately grows and a final effect to people living in the area where this species is abundant, is the fluffy white “wool” secreted by the larval stages, which blows from trees, sticks to clothing and garden furniture, and even causes allergic reactions (Martin et al. 1997). The genus Paraleyrodes, also native to the 518 David Mifsud et al. / BioRisk 4(1): 511–552 (2010) Neotropical Region, is represented in the West Palaearctic by three species. Paraleyrodes species are all very small, comparable in size to members of the Aleyrodinae, and similarly having their fore wing venation reduced to a single unbranched main vein. However, the larval instars all possess wax-producing pores of compound structure, claws on the puparial legs and a quadrisetose ligula*, all being diagnostic characteristics for the Aleurodicinae. P. bondari, is well established in Madeira with material collected on several host-plants since 1995 and likelwise, P. citricolus, established on the same island at least since 1994 and is common on both Citrus spp. and Persea americana Miller (Martin 1996). P. minei, although originally described from Syria, is native to the Neotropics. This species has been established in Spain since the early 1990s where it provokes substantial damage on citrus plantations (Garcia Garcia et al. 1992). A fourth species, P. pseudonaranjae Martin has become established in Florida, Hawaii, Bermuda and Hong Kong and seems to be rapidly extending its native geographical range (Martin 2001). This species is polyphagous with Citrus included in its host-plant records and Europe should be alerted with respect to the high risk of introducing this species. With regard to the DAISIE list of alien Aleyrodidae published in the ‘Handbook of alien species in Europe’(DAISIE 2009), the identification of Aleuroclava guyavae by Pellizari and Šimala (Pellizari and Šimala 2007) was incorrect and should refer to A. aucubae, a closely related species (Martin, J. pers. comm., 2010). Bemisis afer (Priesner & Hosny) was not included as an alien species to Europe in this work as this group is in need of taxonomic revision. Several samples from Britain do however come from glasshouses and its status in Britain was reviewed by Malumphy (2003). Besides, several forms are known from Macaronesia, and before a proper revision of the group is done to define species boundaries no account on European material is included. Aleurolobus marlatti (Quaintance) was also removed from the list of alien species in Europe. The species has a very wide geographical distribution with native records from Southern Europe (Sicily and Malta). We also excluded Aleurolobus olivinus (Silvestri), a species which is widely found in Europe and wherever its preferred host-plant (olive tree) grows. Finally, Dialeurodes formosensis Takahashi was also excluded because the unconfirmed record to species level of Iaccarino (1985) was incorrect and should refer to Dialeurodes setiger (Goux), a species native of the Mediterranean area. 9.4.2.2 Psylloidea Jumping plant-lice alien to Europe include 14 species belonging to two families, Psyllidae (11 species) and Triozidae (3 species) (Figure 9.4.1). The Psyllidae family is the largest family of jumping lant-lice with a cosmopolitan distribution and some 1,800 described species accommodated in more than 150 genera. As presently constituted this family is difficult to define as, effectively, it comprises all those species that do not belong in any other of the five psylloid families. The family has a wide range of hostplants with many species utilising woody legumes. Some species are gall-inducers and all of the solitary lerp-forming species belong to this family. The genus Acizzia currently Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... 519 accommodates more than 30 described species of psylloids mainly found in Australia, New Zealand, the Old World tropics and extending through North Africa and the Middle East to the Mediterranean Basin (Hodkinson and Hollis 1987). Among other characteristics, male adults of this genus have a proctiger* with a conspicuous posterior lobe, forewing with a tapered pterostigma and distinct costal break, basal metatarsus with 1 or 2 black spures and apical segment of aedeagus often complex. Species feed on mimosoid legumes, particularly Acacia and Albizia. In Europe, four species are considered alien introductions. Acizzia hollisi was described from Saudi Arabia and Israel (Burckhardt 1981) on Acacia raddiana Savi and was found on the island of Lampedusa in 1987 (Conci and Tamanini 1989). Acizzia acaciaebaileyanae and A. uncatoides were originally described from Australia and New Zealand, respectively. Both species have been introduced and established in several European locations; A. acaciaebaileyanae in France (Malausa et al. 1997), Italy (Fauna Italia, Rapisarda 1985) and Slovenia (Seljak et al. 2004) whereas A. uncatoides in France, Italy, Portugal (Hodkinson and Hollis 1987), Montenegro (Lauterer 1993), Malta (Mifsud 2010) and the Canary Islands. Within this psylloid group, the latest arrival in Europe was Acizzia jamatonica, originally described from Asia. This species was first noted in Italy (Zandigiacomo et al. 2002), and it was later recorded from a number of European countries including France and Corsica (Chapin and Cocquempot 2005), Slovenia (Seljak 2003), Switzerland (Kenis 2005), Croatia (Seljak et al. 2004), and Hungary (Redel and Penzes 2006). Since 2006, this species was also introduced in the Nearctic Region and its occurrence in the south-eastern United States was surveyed (Wheeler Jr and Richard Hoebeke 2009). Another group of psylloids which are being accidentally introduced and established in Europe are those associated with eucalyptus plantations. The psylloid subfamily Spondyliaspidinae represents a group of insects associated with Myrtaceae, in particular with eucalyptus. Eucalypts, native to Australia, are planted for a variety of uses in many warmer regions throughout the Old and the New World. The commercial value of selected species for the production of ornamental foliage used in the cut flower industry and/or for pulp timber production has resulted in the widespread planting of Eucalyptus trees. Psylloids associated with such host-plants, have become established outside their native range and are sometimes responsible for severe damage to such plantations (Burckhardt and Elgueta 2000). One such psylloid is Blastopsylla occidentalis described from Australia, New Zealand and California, and subsequently reported from Mexico, Brazil and Chile (Burckhardt and Lauterer 1997). The species was recently reported in Italy (Laudonia 2006) and most likely this psylloid is already established in other Mediterranean countries. Glycaspis brimblecombei, commonly known as the Redgum Lerp Psyllid, originally described from Brisbane in Australia, is also expanding its range with records from Mauritius and California (late 1990s), and it has recently been intercepted in Spain and Portugal (Valente and Hodkinson 2008). The Redgum Lerp Psyllid is becoming a major ornamental pest of Red Gum Eucalyptus, but also occurs on Sugar Gum, Glue Gum and other Eucalyptus spp. Three species of Ctenarytaina also established in Europe, the first being C. eucalypti, com- 520 David Mifsud et al. / BioRisk 4(1): 511–552 (2010) monly known as the Eucalyptus psyllid. Originally described from specimens collected on blue gum in New Zealand, this species was first introduced into southern England, northern France and South Africa as early as the 1920s (Laing 1922, Mercier and Poisson 1926, Pettey 1925). This psylloid pest expanded and its current distribution includes France, Germany, Italy, Portugal, Madeira, the Azores, Spain, the Canary Islands, Switzerland and Great Britain (Hodkinson 1999, Wittenberg 2005). The two other species of Ctenarytaina have been introduced more recently. C. spatulata was first reported from France and Italy (Costanzi et al. 2003) and later from Portugal (Valente et al. 2004) and Spain (Mansilla et al. 2004), whereas C. peregrina was first intercepted and described from England (Hodkinson 2007) and recently reported from France and Italy (Cocquempot and Constanzi (Unpubl.)). The genus Cacopsylla includes more than 100 described species distributed mainly in the Holarctic Region, with species that penetrate the Oriental, Afrotropical and Neotropical Regions. Cocquempot and Germain (Cocquempot and Germain 2000) recorded Cacopsylla fulguralis, a species native to western Asia, for the first time from France and subsequently the species was found in Belgium (Baugnée 2003), Italy (Süss and Salvodelli 2003), Spain (Cocquempot 2008), Switzerland (Cantiani 1968) and the United Kingdom (Malumphy and Halstead 2003). Cacopsylla pulchella, a species strictly associated with the Juda’s tree (Cercis siliquastrum L.) is probably native to the Eastern Mediterranean basin but since the 1960s the species was found in various localities in Central and Northern Europe (Cantiani 1968, Hodkinson and White 1979b). The family Triozidae is the second largest family of Psylloidea with some 1,000 described species accommodated in 50 poorly diagnosed genera (Hollis 1984) with a worldwide tropical/temperate distribution. Species utilise host plants in a wide variety of families but never on legumes and many species produce characteristic galls on their host-plants. Four species are recorded as alien for Europe. Trioza neglecta was introduced to Europe from south-western and Central Asia, the area of its origin, with its host plant, Elaeagnus angustifolia L. grown as an ornamental shrub in parks and along roads. It is now widely distributed from Georgia, Armenia, Azerbaijan, Iran and Anatolia through Russia, Ukraine, Moldavia, Bulgaria, the former Yugoslavia and Romania to Central Europe (Hungary, Slovakia, the Czech Republic, Austria) (Lauterer and Malenovský 2002b). The other two introduced triozid psylloids include T. erytreae and T. vitreoradiata, both of economic importance and which are treated in detail under section 9.4.8. An additional triozid species, Bactericera tremblayi (Wagner), was included in the list of aliens of the DAISIE ‘Handbook of alien species in Europe’ (DAISIE 2009) but was removed from the present list. This species was abundant in Southern Italy and caused problems on onions since the late 1950s. However, around 1980 the populations of this species declined and now the species seems to be rare and localised. According to Tremblay (1988) the species could have been a recent introduction in Italy from the former USSR. There is not much to sustain such a statement given the fact that apart from Italy, the species is known to occur in Switzerland, France, Turkey, Iran and questionably from Syria and also because the species is polyphagous on herbaceous plants (Burckhardt and Mühlethaler 2003, Lauterer et al. in prep). Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... 521 In addition, several other psylloid species can be considered as alien in Europe. One is a species from the small Homotomidae family, which includes 80 described species in the world, accommodated in 11 genera. Host plants all belong to the Moraceae family, and mainly to the genus Ficus. Most known larvae are free-living, although some live in colonies under communal lerps and very few species are gall-inducers. Most species have a pan-tropical distribution but Homotoma ficus (L.), a native of Central-Southern Europe and the Middle East feeding on Ficus carica L., has been introduced in Southern England where it seems to be confined (Hodkinson and White 1979a). It is alien to North America (Hollis and Broomfield 1989). In the same category of alien in Europe are two Psyllidae species. Calophya rhois (Löw), a southern-European species, was reported as introduced in Britain on the basis of a single record from Scalpay in the Hebrides (Hodkinson and White 1979a). The genus Calophya is species-poor and distributed in the Neotropical, Holarctic and Oriental Regions with jumping plant-lice associated mainly with Anacardiaceae. Livilla variegata (Löw), is probably native to Eastern Europe. The species is known from France, Italy, Switzerland, Bosnia, Romania, Spain, Great Britain, Hungary, Germany, Austria and the Czech Republic (Hodkinson and White 1979b, Lauterer and Malenovský 2002b). This species is strictly oligophagous on Laburnum anagyroides Medik. and L.. alpinum (Mill.) Bercht. & Presl., and it is already a widespread element in Central Europe, where it colonises its host plant, L. anagyroides, an introduced Mediterranean ornamental tree commonly planted in parks and gardens, towns and villages and on roadsides. The introduction and spread of L. variegata in Central Europe escaped the notice of entomologists, similar to what happened in England, where it was collected for the first time in 1978 (Hollis 1978), but by which time it was already widespread in that country. A last species, Trioza alacris Flor, is most likely of Mediterranean origin but was introduced throughout central and Northern Europe (only in greenhouses or on laurels placed temporarily outside during summer) on cultivated bay laurel. It mostly develops on Laurel (Laurus nobilis L.) but is also reported on L. azoricus Seub., producing characteristic large leaf galls by rolling the leaf margins down to the lower leaf surface. Most probably the earliest record in Central Europe was that of Schaefer (1949) with material collected from Switzerland in 1917. The species was also introduced in USA (California and New Jersey), Brazil, Chile and Argentina (Conci and Tamanini 1985). 9.4.2.3. Phylloxeroidea – Adelgidae Following the 2007 revision by Havill and Footit (2007), a total of 9 adelgid species were identified as alien to Europe, including 6 species in the genus Adelges (subgenera Cholodovskaya, Dreyfusia, and Gilletteella) and 3 species in the genus Pineus (subgenera Pineus and Eopineus). At present, these alien species represent 36% of the total adelgid 522 David Mifsud et al. / BioRisk 4(1): 511–552 (2010) fauna observed in Europe (Figure 9.4.1). Most of them were introduced during the late 19th century- early 20th century alongside with their exotic conifer host trees which were massively used at that time for afforestation in Europe, e.g. Douglas-fir (Pseudotsuga menziesii Mirb. (Franco)) for Adelges cooleyi (Chrystal 1922) and A. coweni (Roversi and Binazzi 1996), Caucasian fir (Abies nordmanianna Spach.) for Adelges (Dreyfusia) nordmanianna (Marchal 1913) , A. prelli (Eichhorn 1967) and A. merkeri (Binazzi and Covassi 1988), and oriental spruce, Picea orientalis (L.) Link., for Pineus orientalis. Some other species were introduced along with ornamental trees originating from North America such as Pineus (Eopineus) strobi with the eastern white pine, Pinus strobus (Steffan 1972), and Pineus similis with Sitka spruce, Picea sitchensis (Bong.) Carrière (Carter 1975, Carter 1975). A majority (five out of nine) of the alien species are holocyclic, one is anholocyclic of first type developing entirely on Picea (Pineus similis) and three anholocyclic of second type developing entirely on Pseudotsuga (Adelges coweni), Larix (A. viridula) or Pinus strobus (Pineus strobi). In addition, several adelgid species native of the Alps and/or Central Europe can be considered as alien in Europe. Their primary host is mostly spruce (Picea), and then larch (Larix), fir (Abies), or pine (Pinus). They include Adelges (Adelges) laricis Vallot, which accompanied the plantations of larch in the lowlands (Glavendekić et al. 2007, Hill et al. 2005), and several species introduced from continental Europe to Great Britain, i.e. Adelges (Adelges) piceae Ratzeburg, A. (Sacchiphantes) abietis L. , A. (Sacchiphantes) viridis Ratzeburg , and Pineus pineoides Cholodkovsky (Hill et al. 2005). Similarly, the alpine Pineus cembrae (Cholodokovsky) colonized the Faroe islands with Swiss stone pine, Pinus cembra L. Adelges (Aphrastasia) pectinatae (Cholodkovsky), a species which develops on spruce and fir was first considered as an alien in Europe (DAISIE 2009) having established in Central and Northern Europe, including the Baltic countries (Gederaas et al. 2007, Holman and Pintera 1977). However, its origin is difficult to be ascertained since Havill and Footit (2007) indicated ‘Europe, China and Japan’. – Phylloxeridae There are two species of phylloxerans alien to Europe with regard to 15 native species (Figure 9.4.1). Moritziella corticalis is of unknown origin (cryptogenic) and was first reported as introduced in Britain (Barson and Carter 1972). The genus Moritziella accommodates two species living on Fagaceae. They are distinguished from Palaearctic species of Phylloxera by the absence of abdominal spiracles on segment 2–5 and by the presence of numerous well-developed, pigmented dorsal tubercles. Generic distinction between North American species of Phylloxera and Moritziella is however not satisfactory. The other species is the well-known ‘Phylloxera’, Viteus vitifoliae (=Dactylosphaera vitifoliae) which has devastated the European vineyards at the end of 19th century. The genus Viteus is a monotypic genus, the alatae* of which have paler abdominal stigmal* plates and a shorter distal sensorium* on the third antennal segment than the common Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... 523 European Quercus-feeding Phylloxera. Viteus vitifoliae typically goes through a two-year cycle involving a sexual phase and leaf-galling and root-feeding stages on American vines. On European vines it normally lives continuously on the roots, reproducing parthenogenetically. Leaf-galls occur in Europe on cultivars derived from hybrids between Vitis vinifera L. and American vines. The economic significance of this species is discussed in some detail under section 9.4.8. 9.4.2.4. Auchenorrhyncha A total of 12 species alien to Europe have been considered (Figure 9.4.1). Not surprisingly most of them belong to the species- rich family of Cicadellidae (17,000–20,000 worldwide; 1,236 species in Europe). Other families are represented only by a single species in each. Within Cicadomorpha, the Cicadellidae (leafhoppers) is the largest family with 50 subfamilies and 17,000–20,000 described species. Leafhoppers live in all zoogeographical regions and feed on a wide range of host plants, though individual species have often trophically and geographically restricted ranges (Dolling 1991, Nielson 1985). Cicadellidae varies in body length from 2–30 mm. Leafhoppers feed on a large range of plants (grasses, herbaceous plants, trees and shrubs). The majority of leafhoppers feed on phloem, some on xylem (especially the subfamily Cicadellinae), and only members of the subfamiliy Typhlocybinae are specialised parenchyma-feeder. Leafhoppers are well known vectors of plant diseases and of economic importance worldwide. For some leafhopper species migratory behaviour is documented (Della Giustina 2002). Eight leafhopper species are certainly alien to Europe. Probably most famous is the Rhododendron leafhopper, Graphocephala fennahi, a native to North America. The species was first reported from southern England in the 1930s but it crossed the Channel only after 1960, to the Netherlands from where it spread rapidly within continental Europe. Two other North American species, Scaphoideus titanus and Erythroneura vulnerata, are pest species on grapes. Especially Scaphoideus titanus has become an important pest since it is the vector of ‘flavescence dorée’ phytoplasma to grapevine. The Neartic leafhopper Kyboasca maligna does not seem to be problematic as an alien species to Europe for the time being. From Eastern Asia four cicadellid species have been introduced: Japananus hyalinus, Macropsis elaeagni, Orientus ishidae and Igutettix oculatus. None of them have yet been found to transmit plant diseases in Europe and are therefore not of economic importance. O. ishidae was only recently reported new to Europe (Günthart et al. 2004) but is spreading rapidly in Europe (Switzerland, Italy, Germany, Slovenia, France, Austria, Czech Republic). I. oculatus (=Vilbasteana oculata (Lindberg)) is originally an eastern Palaearctic species which was first found in Moscow in 1984 and is now spreading to the west (Finland (Söderman 2005)). It lives on Syringa. With around 3,200 described species Membracidae is the largest family of treehoppers. Membracids are widespread worldwide but only few species occur in Eu- 524 David Mifsud et al. / BioRisk 4(1): 511–552 (2010) rope. This family is most diverse in the Neotropics and North America. Characteristic is the enlarged pronotum with sometimes bizarre shaped extensions and elongations. They are medium sized with a body length of 2–24 mm. As with other members of Cicadomorpha, Membracidae lay their eggs into living plant tissue. If populations are too big this can cause serious damages to the host plant and therefore can be regarded as crop pests (e.g. apple trees, see e.g. (Arzone et al. 1987)). Only four species are native to Europe. One species (Stictocephala bisonia) was introduced from North America. The Fulgoromorpha group yet contributed for only three species alien to Europe, with one per family Delphacidae, Flatidae and Acanaloniidae, to be compared to 727 species native in Europe. Delphacidae are characterized by a moveable spur on the hind tibia. Species are generally small (2–6 mm) and are widely distributed also in colder regions. Worldwide around 1,500 delphacid species are described. They feed on monocotyledons and are economically important as pest species on rice, maize, wheat and sugarcane. Nilaparvata lugens (Stål) for example is a serious pest of rice in Asia (O’Brien 2002, Wilson and Claridge 1991). In Europe there are some 260 species. Only one alien delphacid has established in Europe, Prokelisia marginata, which was first found on the Algarve (Portugal) in 1994 and in Spain in 1998 (unpublished data M.R. Wilson). In Slovenia a well established population was found in 2004 (Seljak 2004). New, unpublished records are from southern England (2008) and France (2009). It is very likely that this planthopper is expanding its range rapidly along the European coasts. Species of the family Flatidae have often colourful opaque wings and can be distinguished from other Fulgoromorpha by the numerous parallel crossveins along the costal margin of the forewing and a single spine at each side of the second tarsomere of the hind leg. The body size varies between 4.5–32.0 mm. Flatids feed on different shrubs, trees and herbs (O’Brien 2002). The North American Metcalfa pruinosa has been introduced to Europe probably in plant material and was first recorded in Italy in 1983. From there it is spreading rapidly to the rest of southern Europe (France, Slovenia, Switzerland, Austria, the Czech Republic) causing damages on grapes (Della Giustina 1986, Dlabola 1981, Holzinger et al. 1996, Lauterer and Malenovský 2002a, Mani and Baroffio 1997, Seljak 2002). The Acanaloniidae is a small family of Fulgoromorpha with c. 80 described species accommodated in 14 genera. In general they resemble flatid planthoppers. This family is not native to Europe and the north American species Acanalonia conica was only recently introduced into northern Italy (D’Urso and Uliana 2006). A. conica has a similar biology to Metcalfa pruinosa and can often be found in mixed nymphal feeding groupings with the latter (Wilson and MacPherson 1981). Therefore this species could potentially be another pest insect for European vineyards. Tropiduchidae is a small family within the Fulgoromorpha with some 400 described species worldwide. Body size varies between 5–13 mm; the mesonotum with its apical angle is separated by a transverse groove. They feed on ferns, palms, grasses and Dicotyledonae (O’Brien 2002). Ommatissus lybicus Bergevin, the dubas bug, was Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... 525 for a long time regarded as a variety of O. binotatus Fieber (but see Asche and Wilson 1989). O. lybicus is a severe pest of date palms in the Middle East causing the death of trees. O. binotatus was described from Spain and feeds on Chamaerops humilis L. It was also found in Sicily and Portugal and is a native European species and should be deleted from the DAISIE list. Species with an Eurosiberian or a Holarctic distribution, Edwardsiana ishidai Matsumura and Kyboasca bipunctata (Oshanin), have been excluded from Table 9.4.1. Other leafhopper species with a doubtful alien status include: Cicadulina bipunctata (Melichar), a North African species which occurs also in the eastern Mediterranean; Empoasca punjabensis Singh-Pruthi, originally described from India but is also reported from the southern parts of European Russia , Ukraine, Bulgaria, Serbia and Greece; Jacobiasca lybica (Bergevin & Zanon), another North African species which is reported from other Mediterranean regions (Sicily, Sardinia and Greece); Melillaia desbrochersi (Lethierry), a North African species also reported from Greece, Sicily and Corsica; Psammotettix saxatilis Emeljanov, described from Kazakhstan and found in France but possibly conspecific with P. sierranevadae Dlabola from Spain. There are some papers reporting mainly records of Mediterranean Auchenorrhyncha new to Northern European regions (Maczey and Wilson 2004, Nickel and Holzinger 2006, Wilson 1981). Due to lack of sufficient historical information on the distribution of most Auchenorrhyncha species it is difficult to determine if anthropogenic factors and/or climatic influence are the main causes of range extension. There are for example some southern European Eupteryx species, which appear to have become in the last decades more common in central Europe or even extended their range to northern latitudes such as Denmark and the UK. These species may exploit certain man made habitats, e.g. in greenhouses where herbal plants are cultivated (such as Lamiaceae e.g. Melissa, Oreganum, etc.) but may also build up localised ‘wild’ populations. Such populations may be stable over years under good environmental conditions but can also easily break down depending on several conditions including weather, pressure of predators, parasites and others. Continental European Auchenorrhyncha species introduced to European islands are also excluded of this overview. Thus, five Cicadellidae species (Empoasca pteridis (Dahlbom), Grypotes puncticollis (Herrich-Schaffer), Iassus scutellaris (Fieber), Placotettix taeniatifrons (Kirschbaum) and Wagneripteryx germari (Zetterstedt)) are reported to be alien in the UK (Stewart 1993). On the other hand it is very likely that Philaenus spumarius L. (Aphrophoridae) was introduced into Iceland in the late 1970s. 9.4.3 Temporal trends of introduction in Europe of alien species of Aleyrodidae, Psylloidea, Phylloxeroidea, and Auchenorrhyncha The first records in Europe are approximately known for 60 of the 64 species considered here. Dates given are relatively imprecise, as most of these tiny species have probably been introduced several years before they were reported. 526 David Mifsud et al. / BioRisk 4(1): 511–552 (2010) The number of new records per time period largely differed among Aleyrodidae, Psylloidea, Phylloxeroidea and Auchenorrhyncha (Figure 9.4.2.). The arrival of alien phylloxerans and adelgids appeared to peak during the first part of the 20th century. Some species such as the Grape Phylloxera, Viteus vinifoliae, and the silver fir adelgid, Adelges nordmannianae, arrived earlier in the 19th century but most species, especially the ones associated with Douglas-fir (Adelges cooleyi and A. coweni) were probably introduced in the early 1900s. Only one new species having been introduced later (Pineus similis in 1971), and apparently none during the last ten years. In contrast, the mean number of new records per year of Aleyrodids, Psylloids and Auchenorrhyncha increased regularly from the 1950s. For these three groups, an average of 0.5–0.6 new alien species has been recorded per year in Europe since 2000. The first documented introduced alien Auchenorrhycha to Europe was Stictocephala bisonia (at that time under the name Ceresa bubalus) in eastern Europe (former Austro-Hungarian Empire) in 1912 (Horvaáth 1912). This treehopper was probably introduced with fruit tree cuttings and is now widespread all over Europe except the northern regions. It was followed by another North American species, Graphocephala fennahi, which was first found on rhododendrons in southern England in 1933. Since then other Auchenorrhyncha species from North America or East Asia have been introduced mainly to Central or Southern Europe benefiting from international trade of plants. In the case of Scaphoideus titanus it seems that this species had a first ancient introduction followed by multiple colonization events (Bertin et al. 2007). Figure 9.4.2. Temporal changes in the mean number of new records per year of Aleyrodidae, Psylloidea, Phylloxeroidea (Adelgidae/ Phylloxeridae) and Auchenorrhyncha alien to Europe from 1800 to 2009. The number right to the bar indicates the total number of species recorded per time period. Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... 527 9.4.4 Biogeographic patterns of the Aleyrodidae, Psylloidea, Phylloxeroidea, and Auchenorrhyncha alien to Europe 9.4.4.1 Origin of alien species The region of origin of the alien species largely differs between groups (Figure 9.4.3). Aleyrodids and psylloids mainly originated from tropical regions, the Neotropics and Australasia, respectively. Adelgids and phylloxerans came equally from North America and Asia, mostly because a number of adelgids were introduced from the Caucasus Mountains together with their conifer hosts. In contrast, most of the alien Auchenorrhyncha have a North American origin. For a few species, the area of origin remains uncertain. 9.4.4.2 Distribution of alien species in the European countries For whiteflies and psylloids, the distribution of alien species to Europe or to certain parts of Europe has been highlighted and documented in 9.4.2 and is also presented in Table 9.4.1. Most of the alien species of aleyrodids, psylloids, phylloxerans and adelgids did not spread largely within Europe yet. Indeed, 31 species out of 52 (i.e., 60%) have colonized less than five European countries. Only 4 species, two aleyrodids (Bemisia ta- Figure 9.4.3. Comparative origin of the Aleyrodidae, Psylloidea, Phylloxeroidea (Adelgidae/ Phylloxeridae) and Auchenorrhyncha species alien to Europe 528 David Mifsud et al. / BioRisk 4(1): 511–552 (2010) baci and Trialeurodes vaporariorum), one phyloxeran (Viteus vinifoliae) and one adelgid (Adelges nordmannianae) have colonized more than 20 countries (Table 9.4.1). Due to the lack of comprehensive data we cannot give appropriate information on the distribution of alien Auchenorrhyncha in Europe. However three species (Scaphoideus titanus, Metcalfa pruinosa and recently Acanalonia conica) could have first established in the Mediterranean region from where they spread northbound. Other species expanded their range from eastern Europe (Stictocephala bisonia, Macropsis eleagni) or central Europe (Japananus hyalinus, Orientus ishidae), one species started from the UK (Graphocephala fennahi). It is also possible that some of the alien species had multiple introductions (Scaphoideus titanus, Prokelisia marginata). Generally the introduced species could spread easily as long as the environmental conditions are appropriate for them (climate, host plants, etc.). Five out of the 12 alien species spread in more than 10 countries, with Stictocephala bisonia having expanded in 26 countries and islands (Table 9.4.1). 9.4.5 Pathways of introduction to Europe of the alien species of Aleyrodidae, Psylloidea, Phylloxeroidea, and Auchenorrhyncha Most alien species of whiteflies, psylloids, phylloxerids and adelgids were accidentally introduced with their host plant. In most circumstances such introductions occurred via trade of the host plant or of parts of the host plants such as fruit or cut flowers. It is reported that Auchenorrhyncha can migrate. Usually they are short-distance migrants to leave non-permanent habitats but some species are able to migrate over long distances (Della Giustina 2002). The probably most amazing example is the cicadellid Balclutha pauxilla Lindberg which invaded in swarms the Ascension Island in the Atlantic Ocean (about half way between South America and Africa) in 1976. The specimens must have flown more than 2,000 km over the sea probably coming from Africa (Ghauri 1983). Despite of the fact of possible migration, alien Auchenorrhyncha certainly profit of the worldwide trade of fruit trees, vine cuttings and ornamental plants. Especially eggs in the plant tissue can survive the transport even over long distances and time. Once arrived, the nymphs hatch and without their specific parasites they can build up strong populations. Not surprisingly some alien Auchenorrhyncha were first found around harbours (e.g. Prokelisia marginata) or cities (Orientus ishidae), an unmistakable trace of their pathway of introduction. 9.4.6 Ecosystems and habitats invaded in Europe by the alien species of Aleyrodidae, Psylloidea, Phylloxeroidea, and Auchenorrhyncha Apart from those species so far intercepted only in greenhouses and of which no reports exist of their establishment in Europe, the other introduced species of the five groups treated in this account are often confined to few related host plants. For exam- Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... 529 ple, several species of whiteflies which in their area of origin are highly polyphagous have shown to be strictly oligophagous in their new territories, occurring mainly on Citrus and some other woody hosts. Thus, the major part of these alien species is presently observed in man-made habitats, especially in parks and gardens where a number of exotic plants have been planted (Figure 9.4.4). Natural and semi-natural habitats are yet little colonized by alien Auchenorrhyncha and psylloids (<20%) and quite none by aleyrodids. A noticeable exception concerns adelgids because of their association with conifer trees used for afforestation. More than 60% of the alien adelgids are thus found in forest habitats together with fir, spruce and larch trees. Interestingly so far only one grassland species (Prokelisia marginata) was introduced to Europe. This species lives originally in salt marshes along the East-Coast of North America and is associated with Spartina grasses. All other alien Auchenorrhyncha colonize mainly anthropogenic habitats (vine yards, orchards, gardens, parks). Some of them are polyphagous and can therefore also be found in natural environments (e.g. Stictocephala bisonia in dry habitats or Orientus ishidae on willows and birch trees). 9.4.7 Ecological and economic impact of the alien species of Aleyrodidae, Psylloidea, Phylloxeroidea, and Auchenorrhyncha In terms of economic losses, the two most important whiteflies in Europe are Trialeurodes vaporariorum, commonly known as the glasshouse or greenhouse whitefly and Bemisia tabaci, commonly known as the Cotton Whitefly. T. vaporariorum is a member of a North American species-group. It was however described in 1856 from England, at which time the species was an already widespread and established pest. B. tabaci, Figure 9.4.4. Main European habitats colonized by the established alien species of Aleyrodidae, Psylloidea, Phylloxeroidea (Adelgidae/ Phylloxeridae) and Auchenorrhyncha. The number over each bar indicates the absolute number of alien species recorded per habitat. Note that a species may have colonized several habitats. 530 David Mifsud et al. / BioRisk 4(1): 511–552 (2010) a b d e c g f h j i k Figure 9.4.5. Aleyrodid species alien to Europe. a Aleurocanthus spiniferus adult b Aleurocanthus spiniferus puparium c Aleurocanthus spiniferus puparium from palm leaf (East-Timor) d Acaudaleyrodes rachipora puparium on leaf of Argania (Agadir, Morocco) e Aleurothrixus floccosus puparium on leaf of Citrus reticulata (France) f Aleurodicus dispersus puparium from leaf of Psidium gajava (Martinique) g Aleurodicus dispersus puparium on leaf of Psidium gajava (Martinique) h Aleurodicus dispersus damage on palm leaf i Aleurodicus dispersus damage on leaf j Bemisia tabaci from Thailand intercepted at Roissy airport, France on leaf of Eryngium foetidum k Trialeurodes vaporariorum adults and puparium on leaf of Fragaria (France). (Credit: a, b, h, i - Francesco Porcelli; c, d, e, f, g, j, k - LNPV Montpellier). Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... a c 531 b d Figure 9.4.6. Psylloid species alien to Europe. a Acizzia jamatonica adult on leaf of Albizia (Bordeaux, France) b Acizzia jamatonica immature on leaf of Albizia (Bordeaux, France) d Trioza vitreoradiata male under a leaf of Pittosporum tobira e Trioza vitreoradiata female. (Credits: a, b - LNPV Montpellier; c, d - Jean-Marie Ramel and Christian Cocquempot). probably of Asian origin, is now virtually cosmopolitan, usually found under glass in areas with continental climates. Several biotopes of this species are known (De Barro et al. 1998) and this taxon is known to transmit geminiviruses to cultivated plants of various unrelated groups (Bedford et al. 1994) and is a serious pest of both open-air and protected cropping. Some of the “emerging” whitefly pests in Europe may also prove to be of high economic impact to European agriculture and within this group the most promising species seems to be Aleurocanthus spiniferus. One of the most important species of psylloid in terms of economic losses is Trioza erytrea, a native to the Afrotropical Region. This species is a major pest of citrus plantations, but in its native range is also known to develop on Vepris undulata (Thunb.) Verdoorn & C.A. Sm. Zanthoxylum (=Fagara) capense (Thunb.) Harvey and Clausena anisata (Willd.) Hook. f. ex Benth. (Hollis 1984). The main economic importance of T. erytreae is as vector of the citrus disease caused by citrus greening bacterium (also transmitted by the psylloid, Diaphorina citri Kuwayana). Both psylloids are listed as A1 quarantine pests by EPPO and other phytosanitary organisations. Isolated outbreaks of this species were first noted in Europe in Madeira in 1994 and it seems that the species is now established on both the Canary Islands and Madeira (Borges et al. 2008, Gonzalez 532 David Mifsud et al. / BioRisk 4(1): 511–552 (2010) b a c d Figure 9.4.7. Adelgid and phylloxeran species alien alien to Europe. a, b - Viteus vitifoliae on roots of Vitis vinifera (France) c V. vitifoliae from galls on leaf of V.vinifera (France) (Credit: LNPV Montpellier) d Adelges cooleyi on needles of Douglas-fir (France) (Credit: A. Roques). 2003). T. erytrea is also a species of considerable taxonomic interest as it is part of a complex of species, all of which are difficult to define morphologically, but which have discrete host plant preferences (Hollis 1984). Another important psylloid of economic significance is Trioza vitreoradiata, a species native to New Zealand but recently established in Britain (Martin and Malumphy 1995), Ireland (O’Connor et al. 2004), and France (Cocquempot 2008). This psylloid is specific to Pittosporum where apart from direct loss by the plant in the form of sap depletion caused by the feeding activity of the psylloid, shallow pit galls are formed on young leaves, which remain visible for the life of the leaf. Sooty mould is also very common due to the large amounts of honeydew droppings on underlying leafs. The galling and presence of such sooty moulds make unmarketable ornamental plants of Pittosporum tenuifolium Gaertner, which are often grown for the cut-flower industry and also harvested for its foliage (Martin and Malumphy 1995). Two of the introduced Auchenorrhyncha are of high economical importance. Both are regarded as pest species of vine. Scaphoideus titanus is a vector of ‘flavescence dorée’, a phytoplasma disease (grape vine yellows), which can cause big crop losses. Metcalfa pruinosa affects the plants directly. Strong populations can weaken the plant by sucking and the excreted honeydew is medium for fungi, which can cause reduction in the quality of the fruits. The only phylloxerid of devastating economic significance and which was the cause of much trouble for the wine industry in Europe was the Grape Phylloxera, Viteus vitifoliae. This serious pest of grapes originated in North America where the local Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... a 533 b c d e f Figure 9.4.8. Auchenorrhyncha species alien to Europe. a Metcalfa pruinosa larvae b Metcalfa pruinosa adult c Graphocephala fennahi adult d Orientus ishidae adult e Scaphoideus titanus adult f Stictocephala bisonia adult. (Credit: a - LNPV Montpellier; b–f - Gernot Kunz) vines evolved with it and are not severely damaged by its feeding activity. The species was accidentally introduced to Europe around 1860. In Italy, the species was first reported in 1879 and one year later it was also found in Sicily. In certain countries, possibly due to strict quarantine notices of this new pest, several years passed by before its introduction (e.g. in Malta, Grape Phylloxera was introduced in 1919 (Mifsud and Watson 1999)) but eventually the species was introduced everywhere. It invaded the Mediterranean Region, the Middle East, Africa, Korea, Australia, New Zealand and parts of South America. Grape Phylloxera feeds on species of Vitis including grape vines. Foliar attack does not seem to be unduly damaging, but asexual forms attacking roots all year round can kill plants that did not originate from North America. Grafting European vines onto North American rootstocks has successfully solved this problem in the past, but concern has increased in recent years because this resistance 534 David Mifsud et al. / BioRisk 4(1): 511–552 (2010) is being broken in some parts of the World as new biotypes of Grape Phylloxera are evolving (King and Rilling 1985). 9.4.8. Conclusion Only few European countries produced comprehensive lists of alien Aleyrodidae, Psylloidea, Phylloxeroidea and Auchenorrhyncha. Most of these alien insects were probably introduced by plant material and once established could spread quickly into other European countries. Fortunately, only few species (Trioza erythrea, Trioza vitreoradiata, Scaphoideus titanus, Metcalfa pruinosas and Stictocephala bisonia) have to be regarded as pest or potential pest species so far. However, recent introductions (Acanalonia conica, Orientus ishidae, Prokelisia marginata) show that trade is the main factor of introduction and that at any time new problematic species can occur. On the other hand we have still not sufficient information on the migration of Auchenorrhyncha within European regions. Several observations indicate that in the last decades Mediterranean species expanded their distribution to the North but it is not clear if they can establish wild populations or not. Usually these species profit from anthropogenic habitats (e.g. agricultural areas and parks) and can cause problems. Therefore we need to monitor species migration carefully. Acknowledgements We thank Wolfgang Rabitsch and Alain Roques for their comments and corrections on the chapter draft, and Francesco Porcelli and Jon Martin for information on certain whitefly species, and Gernot Kunz for supplying photos. 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Notizia/ERSA 2002: 18–20 Suborder Status Regime Superfamily Family (Subfamily) Sternorrhyncha Aleyrodidae (Aleyrodinae) Acaudaleyrodes rachipora A Phyto(Singh, 1931) phagous Aleurocanthus spiniferus A Phyto(Quaintance, 1903) phagous Oriental Region Oriental Region 2000, ESCAN 2008, IT ES-CAN I2 Polyphagous IT I Aleuroclava aucubae (Kuwana, 1911) A Phytophagous Oriental Region 2007, IT IT I2, J100 Aleuroplatus perseaphagus Martin et al., 1996 Aleuropteridis filicicola (Newstead, 1911) A Phytophagous Phytophagous Neotropical 1991, ESRegion MAD Africa 1961, GB PT-MAD I2 Polyphagous; Porcelli (2008) occasionally a pest on Annona and Citrus Psidium, Pellizari and Šimala (2007) Cinnamomum, Citrus, Ficus, Pittosporum, Prunus, Photinia Avocado mainly Martin et al. (1996) GB J100 Aleurothrixus floccosus Maskell, 1895 A Phytophagous Neotropical 1968, Region ES-CAN; 1969, FR Aleurotrachelus atratus Hempel, 1922 A Phytophagous Neotropical 2000, ESRegion CAN AL, ES-CAN, FR, I2, J100 FR-COR, GR, IL, IT, IT-SAR, ITSIC, MT, PT, GB ES-CAN, FR I2, J100 1st record Invaded countries in Europe Habitat Hosts Pteris togoensis, Cyclosorus dentatus, Oleandra articulata Polyphagous; a preference for Citrus where established Cocus spp. References Martin et al. (2000) Mound (1961)1 Martin et al. (2000) Borowiec et al. (2010) Mound (Mound 1961) redescribed this species under the name of A. douglasi from material collected on ferns in Kew Gardens, UK. David Mifsud et al. / BioRisk 4(1): 511–552 (2010) 1 A Native range 544 Table 9.4.1. List and main characteristics of Aleyrodidae, Psylloidea, Phylloxeroidea, and Auchenorrhyncha species alien to Europe. Country codes abbreviations refer to ISO 3166 (see Appendix I). Habitat abbreviations refer to EUNIS (see Appendix II). Only selected references are given Status Crenidorsum aroidephagus Martin & Aguiar, 2001 Dialeurodes citri (Ashmead, 1885) A Dialeurodes kirkaldy (Kotinsky, 1907) Filicaleyrodes williamsi (Trehan, 1938) Massilieurodes chittendeni (Laing, 1928) A Regime Native range 1st record Invaded countries in Europe Neotropical 2005, GB Region C&S 1938, GB America ? Asia ? Phytophagous Phytophagous C&S America Oriental Region 1998, PTMAD 1945 ? ?A Phytophagous ? New World ? CY, IL, PT I2 A Phytophagous Phytophagous ? Tropical Africa Northern Asia 1938, GB GB, HU J100 1928, GB BE, CH, CZ, DE, I2 DK, FI, FR, GB, IT, NL, PL, SE A A A ? J100 ES, ES-CAN, GB, J100 HU AL, AT, BE, BG, I1, J100 CH, CY, CZ, DE, ES, ES-BAL, ES-CAN, FRCOR, FR, DE, GR-CRE, GR, HU, HR, IL, IT, IT-SAR, IT-SIC, MT, NL, NO, PL, PT, RO, RU DE, FR, PT-MAD J100, I2 AL, FR, FR-COR, I2 IL, IT, IT-SAR, IT-SIC, MT, SI Hosts Sweet potato leaves Ferns References Malumphy (2005) Trehan (1938) Polyphagous crops & greenhouses Martin et al. (2000) Araceae Martin et al. (2001), Streito (2004) Priore (1969) Polyphagous; a preference for Citrus where established Polyphagous; a preference for Jasminum and Morinda citrifolia Ferns Trehan (1938) Rhododendron Laing (1928) Russell (1964) 545 Phytophagous Phytophagous Phytophagous A GB Habitat Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... Suborder Superfamily Family (Subfamily) Aleurotrachelus trachoides (Back, 1912) Aleurotulus nephrolepidis (Quaintance, 1900) Bemisia tabaci (Gennadius, 1889) Status Regime A Habitat References Rapisarda et al. (1990) Phytophagous Phytophagous Phytophagous Phytophagous Polyphagous; a preference for citrus and avocados (in Europe) Rhododendron ? New World Nearctic Region North America Martin (2000) Aleyrodidae (Aleurodicinae) Aleurodicus destructor A Mackie, 1912 Aleurodicus dispersus A Russell, 1965 Phytophagous Phytophagous Neotropical ?, GB Region Neotropical 1962, ESRegion CAN Ceraleurodicus varus (Bondar, 1928) Lecanoideus floccissimus Martin et al., 1997 Paraleyrodes bondari Peracchi, 1971 Phytophagous Phytophagous Phytophagous Neotropical Region Neotropical Region Neotropical Region ?A A A A A A Asia Hosts Martin et al. (2000) 1998, PTMAD 1987, HU PT-MAD I2 Citrus mainly HU I2 1856, GB AL, AT, BG, CH, I2, J100 CZ, DE, DK, EE, FR, HU, IT, IT-SAR, IT-SIC, LT, MT, PT, RO, RS, SI Strawberries (in Kozár et al. (1987) Europe) Polyphagous Martin et al. (2000) GB J100 ES, ES-CAN, PT- I2 MAD Polyphagous Martin (1996) Martin (1996) Visnya (1941) Martin et al. (1997) 1939 HU J100 Polyphagous; a preference for Citrus where introduced Orchids 1994, ESCAN 1995, PTMAD ES-CAN I2 Polyphagous PT-MAD I2 Polypagous, also Martin (1996) on Citrus spp. David Mifsud et al. / BioRisk 4(1): 511–552 (2010) Phytophagous 1st record Invaded countries in Europe mid 1980’s CY, ES, ES-CAN, I2 FR, FR-COR, GR-CRE, IL, IT, IT-SAR, IT-SIC, PT Eastern Asia 1920, BE BE, GB, IT, NL I2, J100 Pealius azaleae (Baker & Moles, 1920) Singiella citrifolii (Morgan, 1893) Trialeurodes packardi (Morrill, 1903) Trialeurodes vaporariorum (Westwood, 1856) A Native range 546 Suborder Superfamily Family (Subfamily) Parabemisia myricae (Kuwana, 1927) Status Native range 1st record Invaded countries in Europe Habitat Hosts Neotropical 1994, PTRegion MAD Neotropical 1990, ES Region PT-MAD I2 ES I2 A Phytophagous Australia 1981, FR FR, IT, IT-SIC, SI I2, F Acizzia hollisi Burckhardt, 1981 A Phytophagous Africa 1987, IT IT (Lampedusa) I2, F Acizzia jamatonica (Kuwayama, 1908) A Phytophagous Western Asia 2002, IT CH, FR, FRCOR, HR, HU, IT I2, F Acizzia uncatoides (Ferris & Klyver, 1932) A Phytophagous Australia 1974, FR ES-CAN, FR, IL, I2, F IT, IT-SIC, ME, MT, PT Blastopsylla occidentalis Taylor, 1985 Cacopsylla fulguralis (Kuwayama, 1908) A Phytophagous Phytophagous Australia 2006, IT IT I2 Western Asia 1999, FR BE, CH, ES, FR, GB, IT I2 Elaeagnus x ebbingei Eastern Mediterranean 1964, FR FR, GB, CH, IT, IT-SIC I2 Cercis siliquastrum A A A Phytophagous References Citrus spp., Martin (1996) Persea Americana Mainly on Garcia Garcia et al. (1992) Citrus spp. Acacia baileyana Malausa et al. (1997), Rapisarda (1985),Stoch (2003), Seljak et al. (2004) Acacia raddiana, Conci and Tamanini (1989) cultivated Acacia spp. Albizzia Chapin and Cocquempot (2005), julibrissima Seljak et al. (2004), Seljak (2003), Wittenberg (2005), Rédei and Pénzes (2006), Zandigiacomo Acacia floribunda Hodkinson and Hollis (1987), Lauterer (1993), Malausa et al. (1997), Stoch (2003), Seljak et al. (2004) Eucalyptus spp. Laudonia (2006) Baugnée (2003), Cocquempot (2008), Cocquempot and Germain (2000), Malumphy and Halstead (2003), Süss and Salvodelli (2003), Wittenberg (2005) Klimaszewski (1973), Hodkinson and White (1979a), Hodkinson and White (1979b), Burckhardt (1983), Stoch (2003) 547 Phytophagous Phytophagous Cacopsylla pulchella (Löw, 1877) A Regime Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... Suborder Superfamily Family (Subfamily) Paraleyrodes citricolus Costa Lima, 1928 Paraleyrodes minei Iccarino, 1990 Psylloidea Psyllidae Acizzia acaciaebaileyanae (Froggatt, 1901) Status Regime Native range 1st record Invaded countries in Europe Habitat Hosts Phytophagous Australia 1922, GB CH, DE, ES, FR, I2, G5 GB, IE, IT, PT Eucalyptus spp. Ctenarytaina peregrina Hodkinson, 2007 Ctenarytaina spatulata Taylor, 1967 Glycaspis brimblecombei (Moore, 1964) Triozidae Trioza erythreae (Del Gercio, 1918) Trioza neglecta (Loginova, 1978) A Phytophagous Phytophagous Phytophagous Australia 2006, GB FR, GB, IT I2 Australia 2002, PT ES, FR, IT, PT I2, G5 Eucalyptus parvula Eucalyptus spp. Australia 2008, ES, PT ES, PT I2, G5 Eucalyptus spp. Phytophagous Phytophagous 1994, MAD 1982, CZ ES-CAN, PTI2 MAD AT, BG, CZ, HU, I2 SK, RO, YU Trioza vitreoradiata (Maskell, 1879) A Phytophagous Western Africa Southwestern and Central Asia New Zealand 1993, GB FR, GB, IE A phytophagous 1913, GB AT, CH, CZ, DE, G3, I2 DK, FR, GB, IE, IT, NL, PL, PT, RO, RS, SE, SK, UA Phylloxeroidea Adelgidae Adelges (Gilletteella) cooleyi (Gillette, 1907) A A A A Western North America I2 Citrus trees Elaeagnus angustifolia Burckhardt (1998), Cavalcaselle (1982), Hodkinson (1999), Hodkinson and White (1979a), Laing (1922), Mercier and Poisson (1926), Nogueira (1971), Rupérez and Cadahia (1973), Wittenberg (2005) Hodkinson (2007) Costanzi et al. (2003), Mansilla et al. (2004), Valente et al. (2004) Valente and Hodkinson (2008) Borges et al. (2008), Gonzalez (2003) Lauterer (1993), Lauterer and Malenovský (2002b) Pittosporum spp. Cocquempot (2008), Malumphy et al. (1994), O'Connor et al. (2004) Picea (I), Pseudotsuga (II) Chrystal (1922), Covassi and Binazzi (1981), Essl and Rabitsch (2002), Forster (2002), Glavendekić et al. (2007), Nieto Nafria and Binazzi (2005), Pašek (1954) David Mifsud et al. / BioRisk 4(1): 511–552 (2010) A References 548 Suborder Superfamily Family (Subfamily) Ctenarytaina eucalypti (Maskell, 1890) 1st record Invaded countries in Europe >1900, IT Habitat Hosts AT, IT, PT G3, I2 Pseudotsuga (anholocyclic) Adelges (Dreyfusia) merkeri Eichhorn 1957 A phytophagous Asia Minor >1900, IT AT, CZ, DE, IT, SE G3 Picea (I), Abies (II) Adelges (Dreyfusia) nordmannianae (Eckstein, 1890) A phytophagous Caucasus Mountains 1840, DE AT, BG, CH, CZ, G3 DE, DK, EE, FR, GB, HU, IE, IT, LV, NL, PL, PT, RS, SE, SI, SK, UA Picea (I), Abies (II) Adelges (Dreyfusia) prelli Grosmann, 1935 A phytophagous Caucasus mountains <1900, IT AT, CH, CZ, DE, G3 IT, SE, SK Picea, Picea orientalis (I), Abies (II) Adelges (Cholodkovskya) viridula (Cholodkowsky, 1911) Pineus (Pineus) orientalis (Dreyfuss, 1889) A phytophagous ?, CZ CZ, DK, ES, GB, G3 SE, SI, SK, YU Larix (anholocyclic) A phytophagous Northwestern Russia Caucasus mountains 1913, CZ CZ, DE, DK, GB, G3, I2 IT, NL, SK, UA Picea orientalis (I), Pinus (II) References Carter (1983), Essl and Rabitsch (2002), Louro and Cabrita (1989), Nieto Nafria and Binazzi (2005), Roversi and Binazzi (1996), Steffan (1972) Binazzi and Covassi (1988), Fauna Italia, Nieto Nafria and Binazzi (2005) Binazzi and Covassi (1988), Dimitrov and Ruskov (1927), Eichhorn (1967), Eichhorn (1991), Essl and Rabitsch (2002), Fauna Italia, Glavendekić et al. (2007), Marchal (1913), Nieto Nafria and Binazzi (2005), Pašek (1954), Varty (1956) Binazzi and Covassi (1988), Eichhorn (1967), FranckeGrossmann (1937a), FranckeGrossmann (1937b), Nieto Nafria and Binazzi (2005), Šefrová and Laštùvka (2005) Nieto Nafria and Binazzi (2005), Šefrová and Laštùvka (2005), Steffan (1972) Bayer (1914), Covassi and Binazzi (1981), Hill et al. (2005), Marchal (1913), Nieto Nafria and Binazzi (2005) Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... Suborder Status Regime Native Superfamily range Family (Subfamily) Adelges (Gilletteella) coweni A phytoNorth (Gillette, 1907) phagous America 549 Viteus vitifoliae (Fitch, 1855) Auchenorrhyncha Cicadomorpha Cicadellidae Erythroneura vulnerata (Fitch, 1851) Graphocephala fennahi Young, 1977 Igutettix oculatus (Lindberg, 1929) A Regime Native range 1st record Invaded countries in Europe Habitat North America Eastern North America C phytophagous Cryptogenic 1970, GB AT, DE, GB, IT, MD, NL, UA A phytophagous North America 1860, FR AL, AT, BG, CH, I CZ, DE, ES, FR, GR, HR, HU, IE, IL, IT, IT-SAR, IT-SIC, MD, MT, PT, PT-MAD, RO, RS, SI, UA Vitis A Phytophagous Phytophagous North America North America 2004, IT IT Vitis Duso et al. (2005) 1933, GB Rhododendron Sergel (1987) Phytophagous East Asia 1984, RU AT, BE, CH, CZ, FB, G, I2, DE, DK, FR, GB, X11, F IT, NL, SI FI, RU I2 Syringa Söderman (2005) A A GB G3 1900, CZ AT, BG, CH, CZ, G3, I2 DE, DK, GB, IT, LV, NL, PL, RO, RS, SE, SK, UA References phytophagous phytophagous A 1971, GB Hosts G,I2 I Picea sitchensis (anholocyclic) Pinus strobus (anholocyclic) Carter (1975), Carter (1975) Quercus petreae Barson and Carter (1972), Fauna Italia, Nieto Nafria and Binazzi (2005) Aloi (1898), Anonymous (1894), Baudyš (1935), Essl and Rabitsch (2002), Fauna Italia, Glavendekić et al. (2007), Nieto Nafria and Binazzi (2005), Roll et al. (2007), Stani et al. (1974), Teodorescu et al. (2005), Tremblay (1981), Tsitsipis et al. (2007), Wittenberg (2005) Bayer (1920), Essl and Rabitsch (2002), Glavendekić et al. (2007), Martelli (1960), Nieto Nafria and Binazzi (2005), Steffan (1972) David Mifsud et al. / BioRisk 4(1): 511–552 (2010) Phylloxeridae Moritziella corticalis (Kaltenbach, 1867) Status 550 Suborder Superfamily Family (Subfamily) Pineus (Pineus) similis (Gillette 1907) Pineus (Eopineus) strobi (Hartig, 1837) Kyboasca maligna (Walsh, 1862) Macropsis elaeagni Emeljanov, 1964 Orientus ishidae (Matsumura, 1902) Scaphoideus titanus Ball, 1932 Membracidae Stictocephala bisonia Kopp & Yonke, 1977 Fulgoromorpha Acanaloniidae Acanalonia conica (Say, 1830) Delphacidae Prokelisia marginata (Van Duzee,1897) Status Regime Native range A Phytophagous East Asia 1942, AT A Phytophagous Phytophagous Phytophagous Phytophagous North America Asia (Caucasus) East Asia 1997, FR North America 1958, FR A Phytophagous North America < 1912, HU A Phytophagous North America A Phytophagous North America A A A 1st record Invaded countries in Europe 1982, CZ 2002, CH Habitat AT, BG, CH, CZ, I2, G1 DE, ES, FR, HU, IT, ME, RO, RS, RU, SI, SK BE, FR I AT, BG, CZ, DE, I2, G5 HU, RO, SI, UA AT, CH, CZ, DE, I2 FR, IT, SI AL, AT, BG, CH, I1 ES, FR, HU, IT, PT, RS, SI Hosts Acer References Seljak (2002) Pyrus, Crataegus Della Giustina and Remane (2001) Elaeagnus Holzinger and Remane (1994) Salix, Betula, fruit tress Vitis Guglielmino (2005), Günthart et al. (2004) Arzone et al. (1987) AL, AT, BA, BE, I2 BG, CH, CZ, DE, ES, FR, HR, HU, IT, IT-SAR, IT-SIC, MD, ME, MK, NL, PL, RO, RS, SI, SK, UA Polyphagous Arzone et al. (1987), Seljak (2002) 2003, IT IT I, J Polyphagous D’Urso and Uliana (2006) 2003, SI ES, FR, GB, PT, SI D6 Spartina maritima Seljak (2004) Other Hemiptera Sternorrhyncha (Aleyrodidae, Phylloxeroidea, and Psylloidea)... Suborder Superfamily Family (Subfamily) Japananus hyalinus (Osbom, 1900) 551 Status A Regime Phytophagous Native range North America 1st record Invaded countries in Europe 1970, FR Hosts Polyphagous References Dlabola (1981), Lauterer and Malenovský (2002a) David Mifsud et al. / BioRisk 4(1): 511–552 (2010) AL, AT, BA, BG, I CH, CZ, FR, FRCOR, GR, HR, HU, IT, IT-SAR, IT-SIC, RS, SI, SK Habitat 552 Suborder Superfamily Family (Subfamily) Flatidae Metcalfa pruinosa (Say, 1830) Glossary of the technical terms used in the book Glossary of the technical terms used in the book (marked by *) Alatae: winged forms in aphids, adelgids, and other hemipterans. Ampelophagous: related to the grapevine. Anholocyclic: in cyclically parthenogenetic organisms, life cycles that do not include a sexual generation (e.g., in adelgids). Archegonia: female multicellular egg-producing organ occurring in mosses, ferns, and most gymnosperms. Archeozooan: an alien animal introduced to Europe since the beginning of the Neolithic agriculture but before the discovery of America by Columbus in 1492 (Daisie 2009). Arrhenotoky: a common form of sex-determination in Hymenoptera and some other invertebrates, in which progeny are produced by mated or unmated females, but fertilized eggs produce diploid female offspring, whereas unfertilized eggs produce haploid male offspring by parthenogenesis (only the females are biparental). Carina (sg.), Carinae (pl.): a ridgelike structure (e.g. antennal longitudinal ridge). Cercus (sg.), Cerci (pl.): paired sensory structures at the posterior end of some arthropods. Clava: apically differentiated region (sometimes club-like) of the antennal flagellum. Dealate: having lost its wings; used for ants and other insects that shed their wings after the mating flight. Declivity: posterior portion of the elytra that descends to its apex. Domestic: living in human habitats. Endofurca: the internal skeleton of the meso-and metathorax, that provides important muscle insertion points. In some thrips, the metasternal endofurca provides the insertion for powerful muscles that are associated with a remarkable jumping ability of adults. Endophytic (adj): living inside a plant. Endopterygote: insect that undergoes complete metamorphosis, with the larval and adult stages differing considerably in their structure and behaviour. Epigyne: the external female sex organ in arachnids. Exarate: for a pupa, having the appendages free and not attached to the body (as opposed to Obtect). Exopterygote: insect that undergoes incomplete metamorphosis. The young (called nymphs) resemble the adults but lack wings; these develop gradually and externally in a series of stages or instars until the final moult produces the adult insect. There is no pupal stage. Flagellum: the part of the antenna beyond the pedicel, which is differentiated into three regions, the anellus, funicle and clava. Frass: waste material produced by feeding insects, including excrement and partially chewed vegetation. Funicle: region of the antennal flagellum between the anellus and clava. Gallicolae: leaf gall making forms; e.g., in phylloxerans. Gnathosoma: anterior body region in mites. Halobiont: an organism that lives in a salty environment. Alain Roques et al. (Eds) / BioRisk $$: @@–@@ (2010) Hemimetabolous: the type of insect development in which there is incomplete or partial metamorphosis, typically with successive immature stages increasingly resembling the adult; see Exopterygote. Holocyclic: in cyclically parthenogenetic organisms, life cycles that include a sexual generation (e.g., in adelgids). Holoptic: as in flies, with compound eyes meeting along the dorsal midline of the head. Hyperparasitoid: a parasitoid living on or in another parasitoid. Idiobiont parasitoid: a parasitoid which prevents further development of the host after initial parasitization. Idiosoma: abdomen of mites and ticks. Kleptoparasitoid: a parasitoid which preferentially attacks a host that is already parasitized by another species. Koinobiont parasitoid: a parasitoid which allows the host to continue its development and often does not kill or consume the host until the host is about to either pupate or become an adult. Ligula: the apical lobe of the labium. Megagametophyte: female haploid, gamete-producing tissue in conifers. Mesothorax: the second, and usually the largest, of the three primary subdivisions of the thorax in insects. Mesonotum: the dorsal part of the mesothorax. Metathorax: the third of the three primary subdivisions of the thorax in insects. Metanotum: the dorsal part of the metathorax. Moniliform: bead-like (as in antennae). Mycangium (sg.), mycangia (pl.): usually complex structures on the insect body that are adapted for the transport of symbiotic fungi, usually spores. Neozooan: an alien animal introduced to Europe after the discovery of America by Columbus in 1492 (Daisie 2009) . Notaulix (sg.), Notaulices (pl.): one of a pair of grooves on the mesoscutum, from the front margin to one side of the midline and extending backward; divides the mesoscutum into three parts. Obtect: for a pupa, having the legs and other appendages fused to the body. Oniscomorph: the state as in ‘pill’ millipedes of being able to roll up in a ball. Opisthosoma: posterior part of the body in spiders and mites. Paranota: lateral wings. Parthenogenesis, parthenogenetic (adj.): the production of offspring from unfertilized eggs. Special cases of this state are arrhenotoky, pseudo-arrhenotoky, and thelytoky. Phytoplasma: prokaryotes that are characterized by the lack of a cell wall, associated with plant diseases. Phytotelmatum (sg.), Phytotelmata (pl.): a small, water-filled cavity in a tree or any similar environment. Podosoma: anterior section of idiosoma in ticks; serving as connecting area for the four pairs of legs. Porrect: extended, especially forward; e.g., porrect mandibles. Glossary of the technical terms used in the book Proctiger: the reduced terminal segment of the abdomen which contains the anus. Prognathous: with the head more or less in the same horizontal plane as the body, and the mouthparts directed anteriorly. Pronotum: the dorsal part of the prothorax. Propodeum: the first abdominal segment. Prosoma: anterior part of the body in spiders and mites; also called cephalothorax. Prothorax: The first of the three primary subdivisions of the thorax in insects. Pseudo-arrhenotoky: A form of sex-determination (especially in some scale insects and mites) in which males and females arise from fertilized eggs and are diploid. However, males become haploid by inactivation of the paternal genomic complement. Puparium (sg.), puparia (pl.): the enclosing case of a pupa. Reticulate: net-like, anastomosing. Rostrum: beak-shaped projection on the head; e.g., in weevils. Scutellum: the middle region of the mesonotum or metanotum, behind the scutum. Scutum: the anterior part of the mesonotum or metanotum. Secondary pest: a pest that attacks only weakened plants. Sensorium: sensory structure present on antenna. Siphunculi, siphuncular (adj.): pair of protruding horn-shaped dorsal tubes in aphids which secrete a waxy fluid. Spatula sternalis: median cuticular sclerite, often bilobed, on the ventral side of the prothoracic segment of the last instars of some midge larvae; plays a role in larval locomotion. Stigma: conspicuous, usually melanised area at the apex of a vein of the forewing, generally at the leading wing edge. Sulcate: having narrow, deep furrows or grooves. Synanthropic: ecologically associated with humans. Tegula: Small, typically oval sclerite that covers the region of the mesothorax where the forewing and thorax articulate. Thelitoky: A form of sex-determination (especially in Hymenoptera Symphyta and Cynipidae) in which only diploid female progeny are produced by parthenogenesis. Termen: distalmost edge of wing. Transhumance: in the case of hives, moving to new environments, according to the change in season. Xylophagous (adj.): feeding on wood. Alien terrestrial arthropods of Europe Edited by Alain ROQUES, Marc KENIS, David LEES, Carlos LOPEZ-VAAMONDE, Wolfgang RABITSCH, Jean-Yves RASPLUS and David B. ROY Sofia–Moscow 2010 BioRisk 4(2) (Special Issue) Alien terrestrial arthropods of Europe Edited by Alain Roques, Marc Kenis, David Lees, Carlos Lopez-Vaamonde, Wolfgang Rabitsch, Jean-Yves Rasplus And David B. Roy This work was supported by a grant from the Sixth Framework Programme of the European Commission under the project DAISIE (Delivering Alien Species Inventories in Europe), contract SSPI-CT-2003-511202. We thank very much Jean-Marc Guehl (INRA department of «Ecologie des Forêts, Prairies et Milieux Aquatiques») and Olivier Le Gall (INRA department of «Santé des Plantes et Environnement ») for their financial help which allowed to publish this book. We are also very grateful to all colleagues who gently supplied us photos to illustrate the alien species: Henri-Pierre Aberlenc, C. van Achterberg, Daniel Adam, G. Allegro, J.J. Argoud, Margarita Auer, Juan Antonio Ávalos, Ab Baas, Antony Barber, Claude Bénassy, Christoph Benisch, C. van den Berg, Mark Bond, Nicasio Brotons, Gert Brovad, Peter J. Bryant, David Capaert, Jérôme Carletto, Rémi Coutin, David Crossley, Györgi Csóka, Massimiliano Di Giovanni, Joyce Gross, L. Goudzwaard, Jan Havelka, Jean Haxaire, Franck Hérard, R. Hoare, R. Kleukers, Zoltán Korsós, Gernot Kunz, Jørgen Lissner, Jean-Pierre Lyon, Mike Majerus†, Kiril Makarov, Chris Malumphy, Erwin Mani, Paolo Mazzei, Tom Murray, Louis-Michel Nageilesen, Laurence Ollivier, Jean-Pierre Onillon, Claude Pilon, Francesco Porcelli, Jean-Paul Raimbault, Urs Rindlisbacher, Gaëlle Rouault, Gilles San Martin, R.H. Scheffrahn, Vaclav Skuhravý, John I. Spicer, Massimo Vollaro, Jordan Wagenknecht, Beate Wermelinger, Alex Wild, Vassily Zakhartchenko, and the Montpellier Station of the Laboratoire National de Protection des Végétaux, France. Olivier Denux did a great job in realizing all the distribution maps. First published 2010 ISBN 978-954-642-555-3 (paperback) Pensoft Publishers Geo Milev Str. 13a, Sofia 1111, Bulgaria Fax: +359-2-870-42-82 info@pensoft.net www.pensoft.net Printed in Bulgaria, July 2010 Contents 553 Chapter 10. Diptera Marcela Skuhravá, Michel Martinez & Alain Roques 603 Chapter 11. Lepidoptera Carlos Lopez-Vaamonde, David Agassiz, Sylvie Augustin, Jurate De Prins, Willy De Prins, Stanislav Gomboc, Povilas Ivinskis, Ole Karsholt, Athanasios Koutroumpas, Fotini Koutroumpa, Zdeněk Laštůvka, Eduardo Marabuto, Elisenda Olivella, Lukasz Przybylowicz, Alain Roques, Nils Ryrholm, Hana Šefrová, Peter Šima, Ian Sims, Sergey Sinev, Bjarne Skulev, Rumen Tomov, Alberto Zilli & David Lees 669 Chapter 12. Hymenoptera Jean-Yves Rasplus, Claire Villemant, Maria Rosa Santos Paiva, Gérard Delvare & Alain Roques 767 Chapter 13.1. Thrips (Thysanoptera) Philippe Reynaud 793 Chapter 13.2. Psocids (Psocoptera) Nico Schneider 807 Chapter 13.3. Dictyoptera (Blattodea, Isoptera), Orthoptera, Phasmatodea and Dermaptera Jean-Yves Rasplus & Alain Roques 833 Chapter 13.4. Lice and Fleas (Phthiraptera and Siphonaptera) Marc Kenis & Alain Roques 851 Chapter 13.5. Springtails and Silverfishes (Apterygota) Jürg Zettel 855 Chapter 14. Factsheets for 80 representative alien species Alain Roques & David Lees (Eds) 1023 Abbreviations and glossary of technical terms used in the book Alain Roques & David Lees 1029 Index of the latin names of the arthropod species mentioned in the book Alain Roques & David Lees A peer reviewed open access journal BioRisk 4(2): 553–602 (2010) doi: 10.3897/biorisk.4.53 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Diptera Chapter 10 Marcela Skuhravá1, Michel Martinez2, Alain Roques3 1 Bítovská 1227/9, 140 00 Praha 4, Czech Republic 2 INRA Centre de Biologie pour la Gestion des Populations (CBGP), Campus International de Baillarguet, 34988 Montferrier-sur-Lez, France 3 INRA UR633 Zoologie Forestière, 2163 Av. Pomme de pin, 45075 Orléans, France Corresponding authors: Marcela Skuhravá (skuhrava@quick.cz), Michel Martinez (martinez@supagro.inra. fr), Alain Roques (alain.roques@orleans.inra.fr) Academic editor: David Roy | Received 4 February 2010 | Accepted 24 May 2010 | Published 6 July 2010 Citation: Skuhravá M et al. (2010) Diptera. Chapter 10. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(2): 553–602. doi: 10.3897/biorisk.4.53 Abstract Of the 19,400 native species and 125 families forming the European diptera fauna, 98 species (less than 0.5%) in 22 families are alien to Europe. These aliens constitute 66 species (18 families) of the suborder Brachycera and 32 species (4 families) of the suborder Nematocera. By family in this category, there are 23 Cecidomyiidae species, 18 Drosophilidae, nine Phoridae, eight Tachinidae and seven Culicidae. Another 32 fly species belonging to five families are considered to be alien in Europe. These invasives native to other European countries are composed of 14 species of Cecidomyiidae, seven Syrphidae, five Culicidae and three species each of Anthomyiidae and Tephritidae. The date of the first record in Europe is known for 84 alien species. Arrivals of alien species of Diptera have accelerated rapidly since the second half of the 20th century. North America appears to be the dominant contributor of the alien flies. The majority of alien Diptera were introduced into or within Europe unintentionally, with only three predators released intentionally for biological control. Alien Diptera are predominantly phytophagous (35.6%), while a lesser portion are zoophagous (28.6%) or detrivorous /mycetophagous (29.6%). Ecological impacts on native fauna and flora have not been documented for any of the alien species established in Europe. However, 14 alien species have economic impacts on crops. Keywords alien, Europe, Diptera Copyright Skuhravá M et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 554 Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) 10.1 Introduction Diptera is one of the largest insect orders, with a worldwide distribution. The order includes 172 to 179 families (depending on authors) with about 132,000 species described which probably underestimates the actual fauna by at least a half. About 19,400 native species and 125 families have been recorded in Europe (Fauna Europaea). The alien entomofauna is comparatively very limited with only 98 species observed to date, i.e. less than 0.5% of the total dipteran fauna in Europe. Commonly called true flies, mosquitoes, midges, deer- and horseflies and houseflies feature among the most familiar Diptera. Flies are not only abundant in popular perception but also have particular veterinary and medical importance for vectoring diseases and as pests of agriculture, forestry and husbandry. However, some species are useful to man as parasitoids and predators of insect pests and as plant pollinators. Generally, adults are minute to small, soft-bodied insects with a highly mobile head, large compound eyes, antennae of variable size and structure, and sucking mouthparts. They have only one pair of functional wings, the second pair being changed into small head-like bodies called halteres. Legs are usually long, with five-segmented tarsi. Adults are usually very active and are found in all major habitats. They are often associated with flowers and with decaying organic matter, but females of some groups are blood-sucking. Larvae are eruciform and legless in most species. They develop mainly in moist or wet habitats such as soil, mud, decaying organic matter, and in plant or animal tissues. Only a small proportion of larvae is truly aquatic. The majority are liquid-feeders or microphagous. 10.2.Taxonomy of the Diptera species alien to Europe The 98 species of Diptera alien to Europe belong to 22 different families (Table 10.1), which all have native representatives. A larger number of aliens belong to the suborder Brachycera (66 species and 18 families) than to the suborder Nematocera (32 species and 4 families). However, this apparently large diversity is confusing. More than 40% of the alien species are either midges (Cecidomyiidae- 23 species) or fruit flies and their relatives (Drosophilidae- 18 species). The other 20 families show less than 10 species each (Figure 10.1). The arrival of these alien species has largely modified the composition of some families such as Braulidae and Drosophilidae where at present aliens respectively account for 33.3% and 14.8% of the total fauna observed in Europe. However, the native entomofauna includes 103 additional families for which no alien species has yet been recorded in Europe, especially for some ecologically and economically- important groups such as Chironomidae, Syrphidae, Asilidae, Tipulidae and Anthomyiidae. The alien dipterans belong to the following families: Diptera. Chapter 10 555 Suborder Brachycera Agromyzidae. All species in the family are phytophagous, including a number of serious pests of cultivated plants. Larvae live in plant tissues, usually forming characteristic galleries as mines. Most larvae live in the parenchyma of leaves, or mine stems, few attack fruits and seeds. The majority of the species are monophagous, some of them are widely polyphagous, attacking different plants of several families. To date, only five alien species have been observed in Europe relatively to 903 recorded native species (Fauna Europaea). However, the alien fauna includes three species of Liriomyza (L. chinensis, L. huidobrensis - see factsheet 14.23, and L. trifolii) which are highly damaging to vegetable crops (Arzone 1979, Martinez 1982, Trouvé et al. 1991). Braulidae. Larvae live as commensals within cells of honey-bee nests (Apis species). They feed on pollen, honey and organic debris. Adults are “food-parasites” of adult bees, attaching themselves to the body of the queen or rarely to a worker. They feed on liquids from the mouth of the bees. There is only one genus present in Europe, Braula, which includes one alien species, B. schmitzi (Dobson 1999), and two native species. Calliphoridae. This is a key family for human health. Adults are potential vectors of bacteria, viruses, protozoaires and helminthes because they actively search for and sit on feces, fresh and cooked meat, fish, dairy products, and wounds. Larvae are parasitoids or predators of living snails, or feed on blood of nestling birds. A few species are obligate producers of myiasis in various animals. Only one alien species, Chrysomya albiceps (Mercier 1927), has been observed in Europe compared to 112 native species. Canacidae (=Tethinidae). Most species are strictly associated with salty habitats (halobionts), e.g. coastal salt marshes, seashore wrack, sandy beaches, shores of inland salt lakes, alkaline springs etc, and only a few species are also known from habitats that are apparently without increased salinity (forests, meadows, deserts). Some species have been reared from deposits of seaweed. There is only one alien species, Pelomyia occidentalis (Irwin et al. 2001), compared to a total of 39 native species. Ceratopogonidae. Biting adults of this family are potential vectors of major animal diseases. In particular, Culicoides species transmit bluetongue orbivirus between ruminant hosts. A species of Afro-Asian origin, C. imicola Kieffer, has been considered as the main agent of the recent outbreaks of bluetongue disease in Europe although some native species could also be involved (e.g., C. pulicarius L. and C. newsteadi Austen complexes (Purse et al. 2007)). However, it seems that the most likely mode of incursion of C. imicola in Europe was via passive transport on the wind as aerial plankton“ (Mellor et al. 2008, Purse et al. 2007). Thus, this species was not considered in this chapter. Dolichopodidae. Adults and larvae of most species are predaceous and feed on softbodied invertebrates. They occupy all terrestrial habitats from coastal beaches to high elevations, but they generally prefer humid areas. Larvae are mostly found in moist soils or in the litter layer while a few others depend on sap runs and tree rot holes for their development. There is only one alien species, Micropygus vagans (Chandler 2004), in comparison to 790 native species in Europe. 556 Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) Figure 10.1. Relative importance of the families of Diptera in the alien and native entomofauna in Europe. Families are presented in a decreasing order based on the number of alien species. Species alien to Europe include cryptogenic species. Only the most important families of native species (> 50 spp.) have been considered. The number over each bar indicates the number of species observed per family. Drosophilidae. Species in this family show very diverse biological habits. The larvae of most species develop in fermenting substrates, but some mine living plants. Some species are used as important laboratory animals. Drosophilids occur in all terrestrial habitats, from lowlands up to alpine meadows. They may be found near the habitats of their insect hosts or preys (mealybugs, bees, wood-boring beetles), around toadstools (Polyporales) and in the flower heads of thistles. Aliens include 18 species in the genera Drosophila (8 species) (Bächli et al. 2002, Grassi et al. 2009), Chymomyza (4 species) (Band 1994, Carles-Tolra and Andersen 2002, Perju 1959, Trent Band et al. 2005), Zaprionus (3 species) (Chassagnard and Kraaijeveld 1991, Monclus 1976, Tsacas et al. 1977), Scaptomyza (2 species) (Nicoli Aldini 2005, Nicoli Aldini and Baviera 2002) and Dettopsomyia (1 species) (Prevosti 1976) compared to 104 native species. Ephydridae. Adults are usually associated with moist substrates, especially shores, marshes and wet meadows. Some develop in decomposing matter or excrement, other are leaf miners or parasitoids. Aquatic and semiaquatic habitats are typical of the family. A total of 335 native species occur in Europe with only three alien species - in the genera Elephantinosoma, Placopsidella and Psilopa (Gatt and Ebejer 2003). Diptera. Chapter 10 557 Fanniidae. Species inhabit forests, rarely open landscape and wetlands. Larvae are generally saprophagous and mostly feed on decaying organic matter as human or animal faeces, decaying material in gardens, and rotting leaf litter. Some species have been reared from fungi, others occur in bird nests, burrows of vertebrates, and nests of social Hymenoptera. There is only one alien species, Fannia pusio (Carles-Tolra and Andersen 2002), compared to 82 native species. Heleomyzidae. Larvae develop in sporocarps of fungi or live in association with mycelia in forest soil, some are necrophagous or saprophagous. There is only one alien species, Prosopantrum flavifrons (Ismay and Smith 1994) compared to 145 native species Hippoboscidae. Adults are bloodsucking ectoparasites of birds and mammals. Females of all species are macrolarviparous, i.e. retaining the larva in the uterus to the end of the third instar. There is only one alien species, Crataerina melbae (Popov 1995), compared to 29 native species. Milichiidae. Larvae are saprophagous and develop in decaying vegetation, wood detritus, in nests of birds, ants (myrmecophilous species) and of other social insects, but also in excrements, carrion, dead insects and snails. Adults of some species are commensals or kleptoparasites of predatory insects and spiders. There are two alien species, in the genus Desmometopa (Roháček (2006b)), compared to 41 native species. Muscidae. Larvae develop in various kinds of decaying organic matter, often showing facultative or even obligatory carnivorous behaviour. Larvae of some species appear to be predaceous during their entire larval life. Adults feed on nectar or plant sap, sometimes also on decaying liquids and some species are predaceous. Some species are adapted to anthropogenically-altered ecosystems. Blood-sucking species are of medical and veterinary importance, being vectors of some diseases. There are two alien species, the sorghum pest Athrerigona soccata (Vercambre et al. 2000), and a predator of house flies, Hydrotaea aenescens (Rozkošný 2006, Saccà 1964), compared to 585 native species. Phoridae. Adults are found in all types of terrestrial habitats, particularly in forests and meadows but also in steppe-like and xerothermic sites. Food preferences of larvae appear to be remarkably different. Most species are polysaprophagous with different degrees of specialisation. Parasitic species are often found in the nests of ants and termites. Some fungus breeders feed on the fungi but others are obligate predators or parasitoids of other fungus feeders such as larval Sciaridae. There are nine alien species in the genera Megaselia (three species) (Campobasso et al. 2004, Disney 2008, Disney and Durska 1999), Chonocephalus (two species) (Disney 1980, Disney 2002), Dohrniphora (two species) (Disney 2002, Disney 2004), Hypocerides (one species) (Disney 2004), and Puliciphora (one species) (Disney 1983) in comparison to a total of 596 native species. Sphaeroceridae. Larvae and adults are saprophagous. Larvae develop in diverse organic matter and feed as saprophages on microorganisms destroying rotting plants, dung, carrion or fungi and also on the decomposed liquid substances. Adults occur in all habitats that contain the breeding media of the larvae, preferably in damp places. A few polyphagous species are synanthropic, living near human habitats. Many copro- 558 Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) phagous species develop in dung heaps near stables or in pastures. There are four alien species, belonging to the genera Thoracochaeta (two species feeding on seaweeds) (Roháček and Marshall 2000), Coproica (one species) (Carles-Tolra and Andersen 2002), and Trachyopella (one species) (Roháček (2006a)), in comparison to a total of 253 native species. Stratiomyidae. Terrestrial and aquatic larvae of this family live as scavengers. Adults feed on nectar of flowers, exploiting a wide range of flowering plants, especially umbels alongside water margins but also in open sunny places. There are two alien species, the scavenger Hermetia illucens (Venturi 1956), which has been used to control house fly, and a soldier fly, Exaireta spinigera (Lapeyre and Dauphin 2008), compared to a total of 138 native species. Tachinidae. Larvae live as endoparasitoids of arthropod larvae. Many species are parasitoids of important pests of agricultural crops and forest trees and are regarded as economically beneficial. Aliens include 8 species of different genera (Blepharipa, Catharosia, Clytiomya, Phasia, Leucostoma, Sturmia, Trichopoda and Zeuxia) (Carles-Tolra and Andersen 2002, Cerretti 2001, Clemons 2001, Colazza et al. 1996, Vaňhara et al. Tschorsnig 2006) in comparison to a total of ca. 870 native species. Tephritidae. So called “fruit flies” because larvae of most species inhabit the fruits or other seed-bearing organs of flowering plants. Larvae are phytophagous, some being leaf miners and stem-borers and others developing in roots. Many species are associated with Asteraceae. Adults feed on pollen and nectar. Some species are pests but others are used as biological control agents of weeds. Aliens include 4 species in the genus Rhagoletis (3 species) (Duso 1991, Lampe et al. 2005, Merz 1991) and the major fruit pest Ceratitis capitata (see factsheet 14.28) in comparison to a total of 264 native species. Ulidiidae. The biology and immature stages are largely unknown. Adults occur in dry, sunny habitats, such as steppe meadows, and thin steppe forests. Larvae are mostly saprophagous and develop in rotting matter, under bark or in dung but a few seem to be phytophagous. Adults live in marshland habitats, woodland areas, sandy, salty or steppe meadows. They are often observed on flowers, shrub leaves, tree trunks, and on excrement and manure heaps. There are only two aliens, compared to a total of 106 native species, Euxesta pechumani, living on carrion and dung (Delage 1969) and Euxesta notata living on bulbs (such as onions) and sometimes considered as a pest (Martinez, unpublished). Suborder Nematocera Cecidomyiidae. Larvae of gall midges are either phytophagous, zoophagous or mycophagous. Phytophagous species cause galls on various parts of their host plants (hence the common name “gall midges“) but some larvae live free in flower heads or in the stems without making galls, or in conifer cones, or are associated with cambium layers of various trees. Some gall-causing species are serious pests of cultivated plants and forest trees. The zoophagous larvae are predators of the larvae of other gall Diptera. Chapter 10 559 midges, aphids, mites, coccids, and other arthropods and some of them are used for biological control of pests. Larvae of several species are endoparasites of aphids, psyllids and tingids. This is the dominant group of aliens in Diptera with 23 species (see Table 10.1 for references) but altogether 1800 native midge species are known to occur in Europe. Culicidae. Larvae develop in water. Females of most species are haematophagous and feed by sucking the blood of vertebrates, whereas males may feed on flower nectar. Adults may transmit various disease pathogens, viz. viruses, malaria and filarioses. Most Culicidae are distributed in tropical and subtropical areas of the world. Whereas the European native fauna only includes 93 species within this family, seven alien species have established in Europe: two species belonging to the genus Aedes (the Asian tiger mosquito, A. albopictus- see factsheet 14.27, and the Asian rock pool mosquito, A. japonicus (Schaffner et al. 2009)); three Asian species of the genus Culex (Adhami 1987, Ramos et al. 1998, Samanidou and Harbach 2003) and two species of Ochlerotatus (Romi et al. 1999, Schaffner et al. 2001). Aedes aegypti, the vector of yellow fever which has been present in Europe for a long time, now seems to be extinct; no exotic species of Anopheles has yet established (Schaffner et al. 2001). Mycetophilidae. Larvae are mycophagous, feeding on the mycelia or fruit bodies of various fungi or myxomycetes. Adults fly in the undergrowth of forests, on meadows and steppe habitats. There is only one alien species, Leia arsona (Halstead 2004) compared to a total of ca. 950 native species. Sciaridae. Larvae are mostly free living in the upper soil layer of nearly all terrestrial habitats. Some species develop inside plant stems, leaves or decaying wood. They feed on fungal mycelia or decomposing plant tissue. There is only one alien species, Bradysia difformis (White et al. 2000), compared to a total of 629 native species. 10. 3.Temporal trends of introduction in Europe of alien Diptera The date of the first record in Europe is more or less precisely known for 84 (ie., 86%) of the alien species of Diptera, whilst it remains unknown for the other 14 species (Table 10.1). Considering, cautiously, this first record in Europe as a proxy, the arrival of alien dipterans showed a significant, exponential acceleration since the second half of the 20th century (Figure 10.2). The mean number of new records per year increased from 0.25 during 1900–1950 to 2.2 during 2000–2008. In parallel, an increasing diversification of the dipteran families involved in the arrivals was observed. Only a few aliens, mostly Cecidomyiidae, were newly recorded during the 19th century. Probably originating from the subtropics, the midge Feltiella acarisuga was first found and described in France in 1827 (Vallot 1827) . It was subsequently discovered in several other European countries to be finally introduced intentionnally in a large part of the world as a biological control agent for red spider mites in greenhouses. Four more alien dipterans, of which three midges and one fruit fly, were subsequently recorded during the second half of the 19th century, each showing different patterns of 560 Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) Figure 10.2. Temporal changes in the mean number of records per year of dipteran species alien to Europe from 1800 to 2009. The number over each bar indicates the absolute number of species newly recorded per time period. expansion in Europe. Contarinia quinquenota (Cecidomyiidae), developing in flower buds of Hemerocallis fulva (Liliaceae), was first found in Austria in 1885 (Löw 1888 and subsequently in 11 other countries. Clinodiplosis cattleyae (Cecidomyiidae), which forms conspicuous swellings on the aerial roots of Cattleya species (Orchidaceae), was first observed in England in 1885 but later only in France (Molliard 1902). Orseolia cynodontis (Cecidomyiidae) was first observed in 1892 in Italy (Massalongo 1892) and then in three other countries. The fruit fly Ceratitis capitata (Tephritidae) was discovered in Italy in 1873 and subsequently in 15 other European countries. The first half of the 20th century saw the arrival of 13 more alien dipterans of which six are Cecidomyiidae, five Drosophilidae, one Calliphoridae and one Stratiomyidae. Two of these species have not shown any expansion in Europe. A cecidomyiid from tropical Asia, Procontarinia matteiana, was only first observed in 1906 within the Botanical Garden of Palermo (Sicily), galling leafs of a plant imported from India, Mangifera indica (Anacardiaceae) (Kieffer and Cecconi 1906). According to recent information, the host plant has subsequently died out; this alien midge may be considered as extinct in Europe. Discovered in England in 1913, a North American midge, Rhopalomyia grossulariae, causing galls on Ribes grossularia (Grossulariaceae), has not been found anywhere else since that time (Theobald 1913). On the contrary, an other North American midge, Janetiella siskiyou (=Craneiobia lawsonianae), which develops in cones of Chamaecyperis lawsoniana (Cupressaceae), was first observed in the Netherlands Diptera. Chapter 10 561 in 1931 (Meijere 1935) and subsequently in 10 further countries. A gall midge of Asian origin, Rhopalomyia chrysanthemi, damaging leaves of cultivated Chrysanthemum (Asteraceae), was observed in France and Denmark in 1935 (Bovien 1935) and subsequently found in greenhouses of eight more countries. An other Asian midge, Stenodiplosis panici, developing in inflorencesces of Panicum miliaceum (Poaceae), was discovered in southern Russia in 1926 (Dombrovskaja 1936) and then in four other countries. The African predatory midge, Dicrodiplosis pseudococci, attacking the scale Planococcus citri (Pseudococcidae) was found in Italy in 1914 (Felt 1914) and then in Spain. Five Drosophila species of unknown origin were first found in Great Britain in 1900 and then in several countries of northern and central Europe. The cryptogenic Chrysomyia albiceps (Calliphoridae) was recorded in 1927 in France (Mercier 1927) and later expanded to most of southwestern and central Europe. Finally, a Stratiomyidae, Hermetia illucens, was first discovered in Malta in 1936 but subsequently spread to 6 more countries (Venturi 1956). The second half of the 20th century consisted of two distinct periods of invasion of alien dipteran species. From 1950 to 1974, only seven new alien species (i.e. 0.2 species per year on the average) were recorded. They belong to families Cecidomyiidae (Contarinia citri (Genduso 1963) and Stenodiplosis sorghicola (Starostin et al. 1987), both of African origin), Dolichopodidae (Micropygus vagans found in Great Britain in 1970 (Chandler 2004)), Muscidae (a north American predator of house fly, Hydrotaea aenesecens (Saccà 1964)), and Sciaridae (Bradysia difformis recorded from Great Britain in 1965 (White et al. 2000) and subsequently found in Northern Europe). In contrast, a total of 39 alien species were subsequently observed in Europe from 1975 to 1999 (i.e. 1.6 species per year on the average). These later invasions involved a much larger number of dipteran families than previously. By order of importance, families include Drosophilidae (eight species), Cecidomyiidae (six species), Culicidae (six species among which the tiger mosquito, Aedes albopictus, arrived in 1979 in Albania (Adhami 1987)), Phoridae (five species, including the mushroom pest Megaselia tamilnaduensis in 1999 (Disney and Durska 1999), Tachinidae (three species), Tephritidae (three species of Rhagoletis fruit pests), Agromyzidae (three species among which the crop pests Liriomyza trifolii in 1979 (Aguilar & Martínez 1979) and L. huidobrensis in 1989 (Trouvé et al. 1991)), and one species in the families Braulidae, Heleomyzidae, Hippoboscidae, Muscidae, and Mycetophilidae. Since 2000, alien dipterans were observed in Europe at a proportionally higher rate, with 20 species newly recorded from 2000 to 2009, i.e. an average of 2.2 species per year. In addition to families already represented by alien species such as Phoridae (four species) (Disney 2002, Disney 2004), Cecidomyiidae (four species among which the quickly spreading Obolodiplosis robiniae galling Robinia pseudoacacia (Duso C and Skuhrava 2003) - see factsheet 14.26) (Calvo et al. 2006, Gagné 2004, Harris and Goffau 2003), Drosophilidae (three species), Agromyzidae (two species) (Bella et al 2007, Süss 2001), Culicidae (Schaffner et al. 2003), Stratiomyidae (Lapeyre and Dauphin 2008) and Ulidiidae (one species each) (Martinez, unpublished), representatives of two new families were observed: Ephydridae shore flies (three species mostly linked to poultry dung) (Gatt and Ebejer 2003) and Canacidae (one species) (Irwin et al. 2001). 562 Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) 10. 4. Biogeographic patterns of the dipteran species alien to Europe 10.4.1. Origin of alien species A region, or more simply a continent, of origin could be traced for only 78 of the 98 dipteran species alien to Europe, i.e. in ca. 80% of the species. However, in a number of cases, the origin of the dipteran species could only be assumed from that of its host. Several species of Cecidomyiidae illustrate the difficulties and uncertainties in assigning origins. Some species were found and described for the first time in Europe but it is likely that they are non-native and introduced together with their host. For example, the Asian origin of a gall midge Procontarinia matteiana, first described in Sicily (Kieffer and Cecconi 1906), and the African origin of Orseolia cynodontis, another gall maker on Cynodon dactylon (Poaceae), first discovered at Verona (Italy) (Massalongo 1892), were assumed from the source of their host plants, imported from India and North Africa, respectively. Similarly, that of Dicrodiplosis pseudococci, a predator midge of a scale, Planococcus citri (Pseudococcidae), also discovered in Sicily (Felt 1914), was assumed from the subtropical and tropical origin of its insect prey. The cases of Rhopalomyia grossulariae and Dasineura gibsoni are even more complex. The larvae of Rhopalomyia grossulariae which develop in enlarged, deformed leaf buds of Ribes uva-crispa (Grossulariaceae) were first discovered in Ohio (USA) and were later found in Great Britain (Theobald 1913); specimens of Dasineura gibsoni were described developing in flower heads of Cirsium arvense (Asteraceae) in Ottawa, Canada (Gagné 1989), before being also found in Great Britain (Harris 1976). Both species were thus considered to be native of the Nearctic, and then introduced to Europe. However, both host plants are not Nearctic species but archaeophytes of Eurasian origin. Therefore, R. grossulariae as well as D. gibsoni might also be of such origin. However, neither larvae nor adults of these two species have been discovered in continental Europe until now. Further genetic studies may contribute to tracking the exact origin of such species. In contrast to the general trend observed for arthropods and insects, North America appears to be the dominant contributor of the alien dipteran fauna, with almost one-third of the species originating from this continent, far beyond Asia whilst a significant percentage of species came from Africa (Figure 10.3). The 30 alien species originating from North America consists of Cecidomyiidae (10 species), Drosophilidae (6 species), Sphaeroceridae (3 species), Tephritidae (3 species; the fruit fly pests Rhagoletis completa, R. cingulata and R. indifferens), Ulidiidae (2 species), and Agromyzidae, Canacidae, Culicidae, Muscidae, Stratiomyidae, and Tachinidae (one species each). The insects originate from various part of this large continent; for example Janetiella siskiyou (Gagné 1972) and Resseliella conicola (Gagné 1989, Skuhrava et al. 2006) developing in cones of Abies and other conifers (Pinaceae) from the northwestern region whereas Obolodiplosis robiniae and Dasineura gleditchiae (Gagné 1989) developing in leaflet galls on Gleditsia triacanthos (Fabaceae) arrived from the northeast. Diptera. Chapter 10 563 Figure 10.3. Origin of the 98 species of Diptera alien to Europe. The 19 dipteran species coming from Asia consists of six species of Cecidomyiidae, five species of Culicidae, two species of Agromyzidae, Phoridae and Tachinidae, and one species of Drosophilidae and Ephydridae. Most species originate from the temperate, eastern Asia such as Contarinia quinquenotata damaging flower buds of Hemerocallis fulva (Liliaceae), Epidiplosis filifera, a predator of the coccid scale Ceratoplates floridensis on citrus fruits (Nijveldt 1965), and probably Rhopalomyia chrysanthemi (Cecidomyiidae) (Barnes 1948) whilst Cerodontha unisetiorbita (Agromyzidae) (Süss 2001), Aedes japonicus (Culicidae) (Schaffner et al. 2009) and Drosophila curvispina (Drosophilidae) (Bächli et al. 2002) originate from Japan. However, tropical Asia, mainly India, has also contributed to the alien entomofauna, having supplied Aedes albopictus (Eritja et al. 2005), Culex tritaebiorhynchus (Samanidou and Harbach 2003), C. vishnui (Culicidae) (Adhami 1987), Placopsidella phaenota (Ephydridae) (Gatt and Ebejer 2003), Procontarinia matteiana (Kieffer and Cecconi 1906), Horidiplosis ficifolii (Cecidomyiidae), causing leaf galls on Ficus benjamina (Moraceae) (Harris and Goffau 2003), and Megaselia tamilnaduensis (Phoridae) (Disney and Durska 1999). A few species came from Middle East (Leucostoma edentata; Tachinidae) (Chassagnard and Kraaijeveld 1991) and Western Asia (Ochlerotatus subdiversus; Culicidae) (Schaffner et al. 2001). The 16 species coming from Africa consist of Cecidomyiidae (five species), Drosophilidae (three Zaprionus species), Phoridae (three species), Ephydriidae (two species), and one species of Tephritidae (Ceratitis capitata), Culicidae and Mycetophilidae. In addition to the species mentionned above (D. pseudococci and O. cynodontis), midges include Stenodiplosis sorghicola associated with Sorghum (Poaceae), and Contarinia citri developing in flower buds of Citrus sp. (Rutaceae), which probably originates from Mauritius. The Phoridae species came from tropical Africa. 564 Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) Five alien dipteran species of different families are known to originate from Central and South America. They include Clinodiplosis cattleyae (Cecidomyiidae) from Brazil (Gagné 1994), Liriomyza huidobrensis (Agromyzidae) from South America (Trouvé et al. 1991) before having been introduced in Central America, Asia and Africa, Fannia pusio (Fanniidae) (Hill et al. 2005), Prosopantrum flavifrons (Heleomyzidae) (Ismay and Smith 1994), and the recently- arrived, Phytoliriomyza jacarandae (Agromyzidae) (Bella et al 2007). Another 5 dipteran species originate from Australasia, viz. Micropygus vagans (Dolichopodidae) from New Zealand (Chandler 2004), Megaselia gregaria (Phoridae) from Tasmania (Disney 2002), Coproica rufifrons (Sphaeroceridae) from Papua-New Guinea (Carles-Tolra and Andersen 2002), Exaireta spinigera (Stratiomyidae) from Australia (Lapeyre and Dauphin 2008), and Dohrniphora cornuta (Phoridae) from Australasia (Disney 2002). Three other dipteran species are only known to originate from the tropical and subtropical parts of the world. They include Dettopsomyia nigrovittata (Drosophilidae), which has been found only once in Canary islands (Prevosti 1976), Puliciphora borinquenensis (Phoridae), found only once in Great Britain (Disney 1983) and Megaselia scalaris (Phoridae), a saprophagous species which may be dangerous to human health and has largely spread in western and central Europe (Disney 2008). 10.4.2. Distribution of alien species in Europe Alien dipteran species and families are not evenly distributed throughout Europe. Large differences exist between countries in the number of alien species present within each territory (Figure 10.4). As for the other arthropod groups, it may reflect differences in sampling intensity and in the number of local taxonomists specialized in these families. The number of alien dipterans is significantly and positively correlated with the country surface area (after log-transformation; P= 0.0282). Indeed, Great Britain hosts the largest number of aliens (36 species of 11 families), followed by continental Spain (33 species; 17 families), continental France (29 species; 13 families) and continental Italy (28 species; 11 families). However, the family diversity is similar in three countries of Central Europe of much smaller size, the Czech Republic, Switzerland, and Slovakia which host each 11 families of alien dipterans for ca. 20 species. Although the western and southern countries seem to host more aliens (Figure 10. 4), the number of species per country relatively to their size is not correlated with longitude (P= 0.4106) nor with latitude (P= 0.3896). The European islands host proportionally more alien dipterans than continental countries relatively to their size (Kruskall- Wallis test on the number of aliens per km2; P=0.0098). Thus, 14 alien species of 10 families were found in the small island of Malta occupying 316 km2 in the Mediterranean Sea. Most alien dipterans still have a very restricted distribution. More than 30% of the species (30 species) have been observed in only one country such as Culex deserticola (Culicidae) and Dohrniphora papuana (Phoridae) as yet only recorded from Spain Diptera. Chapter 10 565 Figure 10.4. Comparative colonization of continental European countries and islands by dipteran species alien to Europe. Archipelago: 1 Azores 2 Madeira 3 Canary islands. (Disney 2004, Eritja et al. 2000, Ramos et al. 1998), Chymomyza wirthi (Drosophilidae) in Great Britain (Gibbs 1994), Placopsidella phaenota (Ephydridae) in Malta (Gatt and Ebejer 2003), and Exaireta spinigera (Stratiomyidae) in France (Lapeyre and Dauphin 2008). Another 17 species are present in only two, often nearby, countries such as Cerodontha unisetiorbita (Agromyzidae) found in Italy and Albania (Süss 2001), Drosophila suzukii (Drosophilidae) in Spain and Italy (EPPO 2010) and Culex tritaeniorhynchus (Culicidae) in Albania (Adhami 1987) and Greece (Samanidou and Harbach 2003). No alien Diptera is present in more than 24 of the 65 countries and large islands of Europe. Only 9 species have been introduced or have expanded in 15 countries or more. Most are plant pests such as the agromyzid leaf miners Liriomyza huidobrensis (24 countries) (EPPO 2006, Fauna Europaea) and L. trifolii (22 countries) (Fauna Europaea), a midge Obolodiplosis robiniae (20 countries) (Glavendekić et al. 2009), and a fruit fly Ceratitis capitata (20 countries) (Fauna Europaea). The Tiger mosquito, Aedes albopictus, and the predator midge, Feltiella acarisuga are also 566 Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) present in 13 and 21 countries, respectively. In most cases, it is not known whether the species has expanded naturally once established in a country or if the extended distribution corresponds to repeated introductions from abroad. However, very patchy distributions probably result from independent introductions. Thus, Hypocerides nearcticus (Phoridae) was found in Spain and Sweden (Disney 2004), and Coproica rufifrons (Sphaeroceridae) in Malta and in the Canary islands (Carles-Tolra and Andersen 2002). In contrast, the occurence of an alien species within a whole geographic region is likely to proceed, at least partly, from natural dispersion such as for Pelomyia occidentalis (Canacidae) which is currently present throughout Central Europe (Czech Republic, Germany, Hungary, Poland and Slovakia) (Roháček (2006a), Roháček (2006c)). Some other species are known to combine both methods of dispersal. Aedes albopictus was introduced independently by human activity in Albania, France, Italy, Netherlands but probably spread naturally along the Adriatic coast (see map on factsheet 14.27). The honeylocust gall midge, Obolodiplosis robiniae, also spread very rapidly throughout Europe (Glavendekić et al. 2009). Four years after its discovery in Italy in 2003, it occupied a large area from southern England in the west to eastern Ukraine in the east and from northern Germany to southern Italy (see map on factsheet 14.26). Dipterans alien in Europe, i.e. originating from one part of Europe and introduced through human activity in an other part, are a matter of debate because it is often difficult to discriminate between a natural expansion, an introduction, or simply a lack of previous information regarding the actual species‘ native range. Table 10.2 present some of these species. They include species of Mediterranean origin, likely to have been introduced with their Mediterranean hosts in more northern countries, for example Monarthropalpus flavus, a gall-maker of common box (Buxus sempervirens) in Central-European countries. In addition, the date of first record is likely to differ largely from the date of arrival for a few species specifically associated with archaeophyte plants. For instance, two gall midges, Contarinia pisi and C. lentis, specifically galling plants in the family Fabaceae, Pisum sativum and Lens culinaris respectively, have been recorded in Europe only rather recently, although their host plants have been introduced for cultivation since the prehistoric times, probably from the Mediterranean region or the Middle East. Other species followed their host plant introduced from continental Europe to islands on which the plant was absent. Dipterans specifically related to larch such as several species of Strobilomyia larch cone flies (Anthomyiidae) (Ackland 1965; Roques, unpubl.) and a larch gall midge, Dasineura kellneri (Hill et al. 2005) or to spruce (a spruce cone gall midge, Kaltenbachiola strobi) (Hill et al. 2005) are thus considered to be alien in Great Britain. 10.5. Main pathways of introduction to Europe of alien dipteran species Intentional introductions represent a much smaller proportion of alien arrivals in Diptera than the average in arthropods in general (3.1% vs. 10%). Only three dipteran Diptera. Chapter 10 567 predators of different families were introduced intentionally for biological control and have subsequently become established. Two of them, Hydrotaea aenescens (Muscidae) and Hermetia illucens (Stratiomyidae), were released from North America to control houses flies in poultry farms and stables (Saccà 1964). The third species, Feltiella acarisuga (Cecidomyiidae), is a cryptogenic species of cosmopolitan distribution preying exclusively on tetranychid red spider mites. Larvae and adults were found in several countries of Europe, in northern Africa, Asia, North America, Australia and New Zealand. It has been intentionally released, mostly in glasshouses, in Italy, Denmark and Poland, to protect crops. Similarly, as for the other taxa, trying to identify pathways for the remaining 97% of accidental introductions is not a straightforward task. In a number of cases, it however could be inferred from the species biology, for that of the plant/animal host or from repeated interceptions with merchandise at borders. Thus, eggs and larvae of the Asian tiger mosquito, Aedes albopictus, and those of the Asian rock pool mosquito, A. japonicus, have been shown to be imported as stowaway through the trade of secondhand tyres (Reiter 1998, Schaffner et al. 2009). Larvae of A. albopictus were also found inside bags watering “lucky bamboos” (Dracaena senderiana) for horticultural markets. Larvae, such as these of Liriomyza spp., that are leaf-miners of vegetable crops, are regularly intercepted at borders along with agriculture imports, as well as fruits infested by larvae of Ceratitis capitata and Rhagoletis spp. More generally, pathways can be hypothesized for about a half of the 95 alien Diptera which were accidentally introduced. Horticultural and ornamental trade is probably the most significant pathway, with a total of 30 species more or less closely associated. Horidiplosis ficifolii, a midge causing leaf galls on Ficus benjamina (Moraceae) was probably imported with infected fig plants in containers from South-eastern Asia (Taiwan) as well as the midge Asphondylia buddleia, developing in swollen aborted flowers of Buddleia racemosa (Scrophulariaceae), from El Salvador to southern France (Beguinot 1999). A similar process is likely to have occurred for the agromyzids Cerodontha unisetiorbita with Phyllostachys bamboos imported from south Asia (Süss 2001), and Phytoliriomyza jacarandae developing on ornamental blue Jacaranda trees (Jacaranda mimosifolia) introduced to Sicily and mainland Italy (Bella et al. 2007). Some other gall midges are assumed to have been transported to Europe with seedlings of plants for planting as very small larvae hidden in undeveloped plant organs, as for example Obolodiplosis robiniae, Dasineura gleditchiae, Dasineura oxycoccana and Prodiplosis vaccinii, the two last species developing in bud galls of cultivated species of Vaccinium (Ericaceae) in North America (Gagné 1989). Orchid trade was probably responsible for the transport of the midge Clinodiplosis cattleyae whereas cone and seed trade can be assumed as the vector of a seed midge, Janetiella siskiyou, infesting Chamaecyprais lawsonniana (A. Murr.) Parl. and a cone midge, Resseliella conicola on Picea sitchensis (Bong.) Carrière. Comparatively few species (10) have larvae that appear to be associated with the trade of vegetable crops (the agromyzids L. huidobrensis and L. trifolii with a large number of different crops; L. chinensis with Allium; the cecidomyiids Stenodiplosis pa- 568 Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) nici with Panicum and S. sorghicola with Sorghum) and fruit crops (the midge Contarinia citri with Citrus, and the tephritids Ceratitis capitata, Rhagoletis completa, R. cingulata and R. indifferens). The movement of stored products seems responsible for the introduction of another 10 species, mostly drosophilds but also several species associated with the mushroom trade such as the phorids Megaselia tamilnaduensis (Disney and Durska 1999) and M. scalaris (Disney 2008) and the mycetophilid Leia arsona (Halstead 2004). Movement of compost is the problable pathway for two species of Stratiomyidae, Exaireta spinigera (Lapeyre and Dauphin 2008) and Hermetia illuscens (Venturi 1956). Finally, three species are associated with animal husbandry such as Crataerina melbae (Hippoboscidae) (Popov 1995) and Chonocephalus depressus (Phoridae) (Disney 2002). 10.6. Ecosystems and habitats invaded in Europe by alien dipteran species Alien dipterans predominantly exhibit phytophagous habits (35 species- 35.6%). However, zoophagous and detrivorous/mycetophagous species each represent nearly one-third of aliens (28.6% and 29.6%, respectively) whilst the feeding habits remains unknown for ca. 2% of the species. Leaves constitute the most important feeding niche for the alien phytophagous species (24 species), far beyond fruits (10 species including cones and seeds). Leaves are exploited by “true” leaf miners (agromyzids and cecidomyids) and by gall-makers (cecidomyids) but not by external feeders. About 85 % of the alien Diptera seem to have firmly established in their new European environment and its habitats. However, there is little evidence of the establishment status of the 15 % remaining species which have been recorded only once or twice. Nearly 65% (64.1%) of the alien Dipteran species established in Europe are only present in man-made habitats, essentially around and in buildings, in agricultural lands, parks and gardens and glasshouses (Figure 10.5). This proportion is not significantly different from the average value observed for all arthropods. In addition, 16 of the 35 phytophagous aliens (45.7%) remain strictly related to their original, exotic plants used as ornamentals at the vicinity of human habitations such as Cerodontha unisetiorbita on bamboo, Dasineura gleditchiae on Gleditsia, Asphondylia buddleia on Buddleia, Obolodiplosis robiniae on honey locust Robinia pseudoacacia. Woodlands and forests have been colonized by a few alien species (11.7 %). The remaining species occur quite equally in diverse natural and semi-natural habitats, viz. in coastal areas, inland surface waters, mires and bogs, grasslands, and heathlands. 10.7. Ecological and economic impact of alien dipteran species Like most insects, alien dipteran species are better known for their economic and sanitary impact than for their ecological impact. Indeed, ecological impacts on native fauna and flora are not documented at all for any of the species established in Europe. Nega- Diptera. Chapter 10 569 Figure 10.5. Main European habitats colonized by the established alien species of Diptera. The number over each bar indicates the absolute number of alien dipterans recorded per habitat. Note that a species may have colonized several habitats. tive economic impacts on crops have been recorded for a total of 14 species. They include the agromyzid leaf miners Liriomyza trifolii and, more especially, L huidobrensis, whose larvae mine a wide range of vegetables and ornamental plants in glasshouses in a large part of Europe but also outdoors in the Mediterranean basin (see factsheet 14.23, 14.24). Of economic importance are also the tephritid fruit flies. Ceratitis capitata damage fruits of many host plants and has a large impact on fruit crops, especially citrus fruits and peach, all over the Mediterranean basin but also in some countries of central Europe (see factsheet 14.28). Other fruit fllies in the genus Rhagoletis, affect cherry (R. cingulata and R. indifferens) (Lampe et al. 2005, Merz 1991) and walnut crops (R. completa) (Duso 1991, Merz 1991) in Western Europe. The recently introduced Drosophila suzukii is also a fruit pest (EPPO 2010). Some mycetophagous species have a local impact on cultivated mushrooms (Megaselia tamilnaduensis, Megaselia gregaria, and Bradysia difformis) (Disney 2008, Disney and Durska 1999, White et al. 2000). Two other species of midges, Stenodiplosis panici and Stenodiplosis sorghicola developing in inflorescences of Panicum and Sorghum, respectively, may become economic pests in the future if the development conditions become more suitable for outbreaks. Positive impacts are considered for the 3 dipteran species deliberately introduced to Europe for biological control of house flies and tetranychid mites (see 10.5). However, their possible ecological impact on the native, non-target fauna is not documented. 570 Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) a f g b h c d e Figure 10.6. Some alien midges and their damage. a Unopened and swollen flower bud (right) of Hemerocallis fulva caused by larvae of Contarinia quinquenotata b leaflets of Gleditsia triacanthos changed into galls by larvae of Dasineura gleditchiae c Leaf bud gall on Pisum sativum caused by larvae of Contarinia pisi d Fruits of Pyrus communis heavily deformed by larvae of Contarinia pyrivora e female of Dasineura kellneri sitting on the bud of Larix decidua and laying eggs f Swollen buds of Larix decidua capped with resin caused by larvae of Dasineura kellneri g Galls in form of indistinct shallow blisters apparent on both sides of the leaf of Buxus sempervirens, caused by larvae of Monarthropalpus flavus h Rolled leaf margins of Pyrus communis caused by galls of Dasineura pyri. Diptera. Chapter 10 571 Some other alien predators which have been accidentally introduced such as Dicrodiplosis pseudococci and Epidiplosis filifera, may be used for biological control of coccids in the future. A total of 7 alien dipterans may have a sanitary impact on human and animal health. Six of the 7 introduced species of mosquitoes in the family Culicidae are capable of transmitting diseases through female bites (Taylor et al. 2006). The most important one, Aedes albopictus, is now established along the Mediterranean coast from south eastern France to northern Greece and is the vector of Chykungunya disease as well as many arboviruses, avian plasmodia and dog heartworm filariasis (see factsheet 14.27). Other alien culicids may be vectors of the West Nile virus (Aedes japonicus (Schaffner et al. 2009), Culex tritaeniorhynchus, C. vishnui, O. atropalpus), Japanese encephalitis (A. japonicus, C. tritaeniorhynchus) and Sindbis virus (C. tritaeniorhynchus). In addition, a detrivorous phorid, the scuttle fly Megaselia scalaris, may be a cause of allergies whilst it is reported in tropical areas to cause wound and intestinal myiasis in humans (Disney 2008). 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Family Species Native range 1st record in Invaded Europe and countries and country islands Habitat* Hosts References A Phytophagous Asia 2001, IT AL, IT I2 Bamboos Süss (2001) A Phytophagous Phytophagous Asia 1982, FR FR I1 Allium spp. Martinez (1982) C. & S. America 1989, FR AL, AT, BE, BG, CH, CY, CZ, ES, ESCAN, FR, GR, GR-CRE, HU, HR, IL, IT, ITSIC, MT, NL, PL, NO, PL, PT, RS AL, AT, BA, BE, CH, CY, ES, ES-CAN, FR, GR, HR, IT, IT-SAR, ITSIC, IL, MT, NL, NO, PT, RO, RS, RU IT-SIC, IT I1, I2, J100 Polyphagous Beschovski and Karadjova (1996), Carlesleaf miner Tolra and Andersen (2002), Cerný (2006), Cerný and Vála (2006), EPPO (2006), Gederaas et al. (red.) (2007), Glavendekić et al. (2007), Roll et al. (2007), Süss (1991), Trouvé et al. (1991) I1 Polyphagous Aguilar & Martínez (1979), Arzone (1979), leaf miner Carles-Tolra and Andersen (2002), Cerný and Vála (2006), EPPO (2006), Gederaas et al. (red.) (2007), Glavendekić et al. (2007), Máca (2006), Roll et al. (2007) I2 Jacaranda mimosifolia A Liriomyza trifolii (Burgess, 1880) A Phytophagous North America 1976, FR Phytoliriomyza jacarandae Steyskal & Spencer, 1978 A Phytophagous South America 2006, ITSIC Diptera. Chapter 10 Agromyzidae Cerodontha unisetiorbita Zlobin, 1993 Liriomyza chinensis Kato, 1949 Liriomyza huidobrensis (Blanchard, 1926) Status Regime Bella et al (2007) 585 Status Regime Native range 1st record in Invaded Europe and countries and country islands Habitat* Hosts References C Parasitic/ Cryptogenic 1998, GB Predator BG, ES, FR, GB, GR, IT, PT E, J Bees Carles-Tolra and Andersen (2002), Dobson (1999) Calliphoridae Chrysomyia albiceps (Wiedemann, 1819) C Parasitic/ Cryptogenic 1927, FR Predator AT, BG, CH, ES, ES-BAL, ES-CAN, FR, HR, MT, PTMAD, PTAZO, SK E Cadavers Carles-Tolra and Andersen (2002), Fauna Europaea, Kubík (2006), Mercier (1927) Canacidae (=Tethinidae) Pelomyia occidentalis A Williston, 1893 Cecidomyiidae Asphondylia buddleia A Felt, 1935 Clinodiplosis cattleyae A (Molliard, 1903) ? North America 2001 CZ, DE, HU, PL, SK U unknown Irwin et al. (2001), Roháček (2006a), Roháček (2006c) Phytophagous Phytophagous North America C. & S. America 1999, FR FR I2 1885, GB FR, GB J, J100 Beguinot (1999), Gagné (1989), Skuhravá et al. (2005) Barnes (1948), Gagné (1994), Molliard (1902), Skuhravá et al. (2005) Contarinia citri Barnes, 1944 A Phytophagous Africa 1957, CY AL, CY, IL, IT, I IT-SIC Buddleia racemosa Cattleia and other Orchidaceae Citrus spp. Contarinia quinquenotata (F. Löw, 1888) A Phytophagous Asia 1885, AT (Temperate) AT, BG, CZ, DE, F- COR, GB, HU, LV, NL, NO, PL, SE I2, J6 Genduso (1963), Georghiou (1977), Sinacori and Mineo (1997), Skuhravá and Skuhravý (2004a) Hemerocallis Balas (1943), Dittrich (1913), Docters fulva van Leeuwen (1957), Halstead and Harris (1990), Löw (1888), Miller (1956), Prell (1916), Skuhravá (1975), Skuhravá and Skuhravý (in prep.), Skuhravá et al. (1991), Spungis (1988), Wahlgren (1944) Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) Braulidae Braula schmitzi Orosi Pal, 1939 586 Family Species Family Species Dasineura gibsoni Felt, 1911 Dasineura gleditchiae (Osten Sacken, 1866) Status Regime North America North America A Phytophagous Predator North America Africa Epidiplosis filifera (Nijveldt, 1965) A Predator Feltiella acarisuga (Vallot, 1827) C Predator A A 1st record in Invaded Habitat* Hosts Europe and countries and country islands 1976, GB GB, F4, I Cirsium References Gagné (1989), Harris (1976) AL, AT, BG, CH, CZ, DE, DK, ES, FR, FR-COR, GB, GR,HU, IT, LU, NL, PL, RS, SK I2 Gleditchia triacanthos Bolchi Serini and Volonté (1985), Dauphin (1991), Dimitrova and Pencheva (2004), Dini-Papanastasi and Skarmoutsos (2001), EPPO (2008), Estal et al. (1998), Halstead (2004), Hrubík (1999), Labanowski and Soika (1997), Lambinon et al. (2001), Meyer and Jaschhof (1999), Nijveldt (1980), Simova-Tošić (2008), Simova-Tošić et al. (2000), Skuhravá (2004), Skuhravá M (Unpublished), Skuhrava et al. (2006), Steyrer et al. (2002) Bosio et al. (1998), Gagné (1989), Seljak (2004) Carles-Tolra and Andersen (2002), Felt (1914), Solinas (1971) 1997, IT AL, FR, IT, SI FB 1914, ITSIC ES, IT-SIC I Eastern Asia 1965, IL GB, IL, LV, PT-MAD U Cryptogenic 1827, ? AL, AT, BE, I, J100 CH, CZ, DE, DK, ES, FI, FR, GB, IE, IT, IT-SIC, LT, LV, NL, NO, PL, RU, SE Vaccinium spp. Scale, Planococcus citri Scale Harris (2004), Nijveldt (1965), Skuhravá (Ceratoplates (2008), Skuhravá et al. (2006), Spungis floridensis) (2003) Mites EPPO (2002), Fiedler (2005), Kahrer and (TetraSkuhravá (2000), Mamaev and Krivosheina nychidae) (1965), Meijere (1939), Roberti (1955), Skuhrava et al. (2006), Spungis (2003), Vacante and Firullo (1985), Vallot (1827), Vimmer (1931) 587 1975, NL Diptera. Chapter 10 Phytophagous Phytophagous Dasineura oxycoccana (Smith, 1890) Dicrodiplosis pseudococci (Felt, 1914) A Native range A A Native range Phytophagous Phytophagous Asia (Tropical) North America 1st record in Invaded Habitat* Hosts Europe and countries and country islands 2001, NL? DK, GB, ITX24 Ficus SIC, NL benjamina 1931, NL AL, CZ, DE, I2 ChamaeDK, FR, GB, cyparis IT, NL, PL, SK lawsoniana Lestodiplosis aonidiellae Harris, 1968 A Predator Africa 1999, ITSIC Obolodiplosis robiniae (Haldeman, 1847) A Phytophagous North America 2003, IT Orseolia cynodontis Kieffer & Massalongo, 1902 A Phytophagous Africa 1892, IT Procontarinia matteiana Kieffer & Cecconi, 1906 A Phytophagous Asia (Tropical) 1906, ITSIC IT-SIC J100 References Harris and Goffau (2003), Skuhrava et al. (2007), Suma et al. (2007) Coutin (1976), Gagné (1972), Harris (2004), Juhásová and Hrubík (1984), Kapuscinski (1948), Meijere (1935), Skuhravá (1979), Skuhrava et al. (2006), Stelter (1978) Siscaro et al. (1999), Skuhrava et al. (2007) Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) Horidiplosis ficifolii Harris, (2003 Janetiella siskiyou Felt, 1917 Status Regime 588 Family Species Scale (Aonidiella aurantii) AL, AT, BA, G, I2, H1 Robinia Bathon (2007), Berest and Titar (2007), CH, CZ, DE, pseudoacacia Csóka (2006), Duso C et al. (2005), Duso FR, FR-COR, C and Skuhrava (2003), Glavendekić GB, GR-ION, et al. (2009), Laguerre and Dauphin HR, HU, IT, (2007), Roskam et al. (2008), SimovaMK, NL, PL, RS, SI, SK, UA Tošić and Skuhravá (1995), Skuhravá M (Unpublished), Skuhravá and Skuhravý (2004b), Skuhravá (2007), Skuhrava et al. (2008), Wehrmaker (2007), Wermelinger and Skuhravá (2007), Zúbrik et al. (2007) FR, HU, IT, E1 Cynodon Houard (1902), Massalongo (1892), RO dactylon Moesz (1938), Roman and Ionescu (1967), Skuhrava et al. (1972) IT-SIC I2 Mangifera Kieffer and Cecconi (1906), Skuhrava et indica al. (2007) Family Species Prodiplosis vaccinii (Felt, 1926) Prodiplosis violicola (Coquillett, 1900) Resseliella conicola (Foote, 1956) Rhopalomyia chrysanthemi (Ahlberg, 1939) Stenodiplosis sorghicola (Coquillett, 1899) Culicidae Aedes albopictus (Skuse, 1894) A A A A A Phytophagous Phytophagous Phytophagous Phytophagous Native range 1st record in Invaded Habitat* Hosts Europe and countries and country islands North 2001, ES ES I2 Vaccinium America spp. North 2004, NL/ NL, SE I Viola spp. America SE North 1999, DK DK I2 Picea America sitchensis Asia 1935, FR BE, CH, DE, X24, I2 Chrysan(Temperate) DK, FI, FR, themum GB, NO, PL, (cultivated) SE Phytophagous Phytophagous North 1913, GB America Asia 1926, RU (Temperate) A Phytophagous Africa A Parasitic/ Asia Predator (Tropical) A G1 Ribes grossularia Panicum spp. BG, RS, RU, SI, UA I 1964, IT FR, GR, IT, RU I Sorghum spp. 1979, AL AL, CH, ES, FR, GR, GRION, HR, IL, IT, IT-SIC, ME, RS, SI J6 Humans (biting) Calvo et al. (2006), Skuhravá et al. (2006) Gagné (2004) Skuhrava et al. (2006) Barnes (1948), Behr (1949), Blauvelt (1939), Bovien (1935), Gjaerum (1949), Häflinger (1945), Skuhrava et al. (2006), Suire (1935), Szadziewski (1991), Vappula (1965), Wahlgren (1951) Barnes (1948), Theobald (1913) Dombrovskaja (1936), Janežič (1972), Krištal (1959), Martinovic and Bjegovic (1949), Simova-Tošić et al. (1996), Simova-Tošić et al. (2000), Skuhravá et al. (1991) Coutin (1969), Mariani and Beccari (1964), Skuhravá et al. (2005), Starostin et al. (1987) Adhami and Murati (1987), Dalla Pozza and Majori (1992), Eritja et al. (2005), Klobučar et al. (2006), Patsoula et al. (2006), Reiter (1998), Romi (1995), Romi et al. (1999), Sabatini et al. (1990), Šuligoj (2005), Urbanelli et al. (2000) 589 GB, References Diptera. Chapter 10 Rhopalomyia grossulariae Felt, 1911 Stenodiplosis panici Plotnikov, 1926 Status Regime Chymomyza procnemis (Williston, 1896) Chymomyza procnemoides Wheeler, 1952 Chymomyza wirthi Wheeler, 1954 1st record in Invaded Habitat* Europe and countries and country islands Parasitic/ Asia 2000, FR BE, CH, DE, J6 Predator (Temperate) FR Parasitic/ Africa 1993, ES ES C1, D Predator Parasitic/ Asia 1987, AL AL, GR D6, C1 Predator (Tropical) Parasitic/ Asia 1987, AL AL C1, D Predator (Tropical) Parasitic/ North 1996, IT IT J6 Predator America Parasitic/ Asia 1987, RS RS D Predator (Temperate) Humans (biting) Humans (biting) Humans (biting) Humans (biting) A Parasitic/ Australasia Predator 1970, IE GB, IE G, I2 Broadleaved Chandler (2004) woodlands A PhytoNorth phagous, America Detrivorous 1975, CZ G, I2, J1 Apple, fruits, nuts Clemons (2009), Máca (2006), Máca (2006), Pakalniškis et al. (2006), Trent Band et al. (2005) A Detrivorous Detrivorous North America North America 2000, ESCAN 1992, HU AT, CH, CZ, DE, ES, FR, GB, HU, LT, MO, NL, RO, RS, RU, SK, ES-CAN Unknown Carles-Tolra and Andersen (2002) HU G Unknown Band (1994), Perju (1959) Detrivorous North America 1994, GB GB B Unknown Gibbs (1994) A A A A A A A A Native range ? Hosts References Humans Andreadis et al. (2001), Schaffner et al. (biting) (2003), Schaffner et al. (2009) Wild rabbits Eritja et al. (2000), Ramos et al. (1998) Adhami (1987), Samanidou and Harbach (2003) Adhami (1987) Romi et al. (1999) Bozicic (1987), Schaffner et al. (2001) Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) Aedes japonicus (Theobald, 1901) Culex deserticola Kirkpatrick, 1925 Culex tritaeniorhynchus Giles, 1901 Culex vishnui (Theobald, 1901) Ochleroratus atropalpus (Coquillett, 1902) Ochleroratus subdiversus (Martini, 1926) Dolichopodidae Micropygus vagans Parent, 1933 Drosophilidae Chymomyza amoena (Loew, 1862) Status Regime 590 Family Species Family Species Dettopsomyia nigrovittata (Malloch, 1924) Drosophila curvispina Watabe & Toda, 1984 Drosophila busckii Coquillett, 1901 Drosophila hydei Sturtevant, 1921 Status Regime A Detrivorous Tropical Subtropical 1st record in Invaded Europe and countries and country islands <1976, ES- ES-CAN CAN A Detrivorous Detrivorous Detrivorous Asia 2002, CH C C Native range Cryptogenic 1900, GB Cryptogenic 1900, GB C Detrivorous Cryptogenic 1900, GB Drosophila repleta Wollaston, 1858 C Detrivorous Cryptogenic 1900, GB Drosophila melanogaster Meigen, 1830 Drosophila suzukii (Matsamura, 1931) Drosophila tsigana Burla & Gloor, 1952 Scaptomyza adusta (Loew, 1862) C Detrivorous Phytophagous Detrivorous Phytophagous Cryptogenic 1900, GB C C A Cryptogenic 2009, IT Cryptogenic ? North America 1996 G CZ, GB, LT, SK CZ, ES, ESBAL, GB, LT, PT, PT-AZO, PT-MAD, SK CZ, ES, ESCAN, GB, LT, PT, PT-AZO, PT-MAD, SK CZ, ES, ESBAL, ES-CAN, GB, LT, PT, PT-AZO, PTMAD, SK CZ, GB, IT, LT, PT, SK IT, SP G AT, FR, HU, PT IT, GR, MT, ES-CAN, PTAZO Unknown G References Carles-Tolra and Andersen (2002), Prevosti (1976) Fungi (forest Bächli et al. (2002) floor) Unknown Hill et al. (2005), Máca (2006), Máca (2006), Pakalniškis et al. (2006) Unknown Carles-Tolra and Andersen (2002), Hill et al. (2005), Máca (2006), Pakalniškis et al. (2006) G Fruits Carles-Tolra and Andersen (2002), Hill et al. (2005), Máca (2006), Pakalniškis et al. (2006) G Unknown Carles-Tolra and Andersen (2002), Hill et al. (2005), Máca (2006), Pakalniškis et al. (2006) G Unknown G Fruits Bächli (2004), Hill et al. (2005), Máca (2006), Pakalniškis et al. (2006) EPPO (2010), Grassi et al. (2009) G Leaf miner Fauna Europaea I, J Nicoli Aldini (2005), Nicoli Aldini and Vegetables (Leaf miner) Baviera (2002) 591 CH Hosts Diptera. Chapter 10 Drosophila immigrans Sturtevant, 1921 Habitat* Heleomyzidae Prosopantrum flavifrons Tonnoir & Malloch, 1927 Hippoboscidae Crataerina melbae (Rondani, 1879) Milichiidae Desmometopa microps Lamb, 1914 A A A A A A A Phytophagous Phytophagous Phytophagous Phytophagous Native range North America Africa Africa Africa Parasitic/ Africa Predator Parasitic/ Asia Predator Parasitic/ Africa Predator 1st record in Invaded Habitat* Hosts Europe and countries and country islands ? GB, SP- CAN J100, I Fruits References Carles-Tolra and Andersen (2002), Roll et al. (2007) Chassagnard and Kraaijeveld (1991) 1991, CY CY Fruits 1976, ESCAN 1977, ESCAN AT, ES-CAN, IL, IT CY, ES-CAN, IL, MT Fruits 2003, MT MT B Shore fly Gatt and Ebejer (2003) 2003, MT MT B Shore fly Gatt and Ebejer (2003) 2002, ESCAN ES-CAN, MT B Shore fly Carles-Tolra and Andersen (2002), Gatt and Ebejer (2003) Poultry dung Carles-Tolra and Andersen (2002) Fruits Carles-Tolra and Andersen (2002), Monclus (1976), Roll et al. (2007) Carles-Tolra and Andersen (2002), Roll et al. (2007), Tsacas et al. (1977) A Detrivorous C. & S. America ? ES, FR, MT J A Detrivorous C. & S. America 1991, GB GB J A Parasitic/ Cryptogenic 1990, DE Predator BG, CH, DE, ES, IT J Haematophagous on birds Carles-Tolra and Andersen (2002), Popov (1995) A Parasitic/ Cryptogenic ? Predator CZ, SK, HU J Adults attack bees Roháček (2006a), Roháček (2006b) Ismay and Smith (1994) Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) Scaptomyza vittata (Coquillett, 1895) Zaprionus ghesquierei Collart, 1937 Zaprionus indianus Gupta, 1970 Zaprionus tuberculatus Malloch, 1932 Ephydridae Elephantinosoma chnumi Becker, 1903 Placopsidella phaenota Mathis, 1986 Psilopa fratella (Becker, 1903) Fanniidae Fannia pusio (Wiedemann, 1830) Status Regime 592 Family Species Family Species Desmometopa varipalpis Malloch, 1927 Mycetophilidae Leia arsona Hutson, 1978 Phoridae Chonocephalus depressus Meijere, 1912 Chonocephalus heymonsi Stobbe, 1913 Dohrniphora cornuta (Bigor in de la Sagra, 1857) Native range 1st record in Invaded Habitat* Hosts Europe and countries and country islands Cryptogenic ? FR, G, ES, CH J Biofilters, sewage filters, decaying vegetables and fruits References Carles-Tolra and Andersen (2002), Roháček (2006b) Detrivorous C PhytoCryptogenic 1998, FR phagous Parasitic/ North 1964, IT Predator America FR, GR, HU, I1 IT, IT-SIC, AT, CH, CZ, E, J DE, ES, ESCAN, FR, GB, IE, IT, IT-SAR, MT, PT, PTAZO, SK A Detrivorous Africa CH, ES-CAN, GB, MT, NL, PT-AZO, PTMAD I, J Fungus gnat Carles-Tolra and Andersen (2002), Halstead (2004) A Detrivorous Detrivorous Detrivorous Asia 2002, MT (Temperate) Africa 1981, GB MT J Ripe fruits Disney (2002) GB, J100 Ripe fruits Disney (1980), Disney (2002) Australasia AT, BG, BE, CY, CZ, DE, ES, ES-CAN, FR, GB, NL, PL, PT, PT-AZO, PTMAD, SI, SK, J saprophagous Beschovski and Langourov (1997), CarlesTolra and Andersen (2002), Disney (1991), Disney (2002), Mocek (2006) A A A 1978, GB 1997 Sorghum Vercambre et al.(2000) Predator of house fly Carles-Tolra and Andersen (2002), Gregor and Rozkošný (2006), Rozkošný (2006), Saccà (1964) 593 A Diptera. Chapter 10 Muscidae Athrerigona soccata Rondani, 1871 Hydrotaea aenescens (Wiedemann, 1830) Status Regime A A A Detrivorous Detrivorous Detrivorous Native range Africa 1st record in Invaded Habitat* Europe and countries and country islands 2004, ES ES J6 Hosts References Africa 2004, ES ES, SE U Australasia 2003 CZ, DE, DK, GB, NO, PL, PT, SE, SI, SK BE, BG, CH, DE, DK, ES, ES-CAN, FR, GB, IT, NL, PT-MAD J6, J1 Mushroom house J6, J1 Decaying Bourel et al (2004), Campobasso et al. material, (2004), Carles-Tolra and Andersen (2002), cadavers, Dewaele et al. (2000), Disney (1994), myasis agent Disney (2002), Disney (2008), Haenni Pers. comm. (2009), Langourov (2004), McCrae (1967), Miller (1979), Zwart et al. (2005) Cultivated Disney and Durska (1999) oyster mushrooms (Pleurotus) Disney (1983) Disney (2004) Disney (2004) Carles-Tolra and Andersen (2002), Disney (2002), Mocek (2006) Megaselia scalaris (Loew, 1866) A DetriTropical, vorous, Subtropical facultative predator/ parasite 1994, ES Megaselia tamilnaduensis Disney in Mohan, Mohan & Disney, 1996 Puliciphora borinquenensis Wheeler, 1900 Sciaridae Bradysia difformis Frey, 1948 A Detrivorous Asia (Tropical) 1999, PL CH, PL J A Detrivorous Tropical, Subtropical 1983, GB GB J C Detrivorous Cryptogenic 1965, GB ES, ES-CAN, GB, NO, SE, SK J100, J1 Mushrooms; Carles-Tolra and Andersen (2002), Heller ornamentals and Menzel (2006), Hellqvist (1994), in nurseries White et al. (2000) A Detrivorous Australasia ES-CAN, MT U Animal dung; leaf litter Sphaeroceridae Coproica rufifrons Hayashi, 1991 ? Carles-Tolra and Andersen (2002) Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) Dohrniphora papuana Brues, 1905 Hypocerides nearcticus (Borgmeier, 1966) Megaselia gregaria (Wood, 1910) Status Regime 594 Family Species Family Species Thoracochaeta johnsoni (Spuler, 1925) Thoracochaeta seticosta (Spuler, 1925) Trachyopella straminea Roháček & Marshall, 1986 A A A A C C C C A 1st record in Invaded Habitat* Hosts Europe and countries and country islands 1999, GB GB, IT B Seaweed Roháček and Marshall (2000) 1999, GB Seaweed Roháček and Marshall (2000) Saprophagous Carles-Tolra and Andersen (2002), Roháček (2006a) Compost, houses compost heaps, poutry, bee hives; used for control of house flies Lapeyre and Dauphin (2008) Detrivorous Detrivorous Detrivorous North America North America North America Detrivorous Detrivorous Australasia 2008, FR FR North America 1936 CH, ES, ESJ6 BAL, ES-CAN, FR, IT, MT, PT Parasitic/ Predator Parasitic/ Predator Parasitic/ Predator Parasitic/ Predator Parasitic/ Predator ? Cryptogenic ? DK, GB, NO, B SE AD, CY, CZ, U ES, ES-CAN, GR-CRE, HU, MT, SK I References Carles-Tolra and Andersen (2002), Venturi (1956) ? Unknown ? Unknown Carles-Tolra and Andersen (2002), Vaňhara et al. Tschorsnig (2006) Carles-Tolra and Andersen (2002) Cryptogenic ? AD, CZ, ES, GB, SK AD, ES, GB, PT ES, GB ? Unknown Carles-Tolra and Andersen (2002) Cryptogenic 2001, GB GB ? Unknown Clemons (2001) Asia IT ? Cryptogenic ? 1995, IT Cerretti (2001) 595 Tachinidae Blepharipa schineri (Mesnil, 1939) Catharosia pygmaea (Fallén, 1815) Clytiomya continua (Panzer, 1789) Phasia barbifrons (Girschner, 1887) Leucostoma edentata Kluger, 1978 A Native range Diptera. Chapter 10 Stratiomyidae Exaireta spinigera (Wiedemann, 1830) Hermetia illucens (Linnaeus, 1758) Status Regime Status Regime C Trichopoda pennipes (Fabricius, 1781) A Zeuxia zejana Kolomiets, 1971 Tephritidae Ceratitis capitata (Wiedemann, 1824) A 1st record in Invaded Habitat* Hosts Europe and countries and country islands Parasitic/ Cryptogenic ? ES, GB I2, E5, Danaid Predator FA butterflies (Ideopsis, Parantica) Parasitic/ North 1989 AL, ES, FR, IT, I Squash bug; Predator America IT-SIC southern green stinkbug Parasitic/ Asia 1995, IT IT ? Unknown Predator A Phytophagous Africa 1873, IT Rhagoletis cingulata Loew, 1862 A Phytophagous North America 1993, DE Rhagoletis completa Cresson, 1929 Rhagoletis indifferens Curran, 1932 A Phytophagous Phytophagous North America North America 1991, IT A 1983, CH AL, AT, BG, I CH, CZ, ES, ES-BAL, ESCAN, FR, G, IL, IT, IT-SAR, IT-SIC, ME, PT, PT-AZO, PT-MAD, SI, SR DE, HU, NL, G, I2 SI AL, CH, DE, I2 FR, HR, IT, SI CH I2 References Carles-Tolra and Andersen (2002) Carles-Tolra and Andersen (2002), Colazza et al. (1996) Cerretti (2001) Fruits (poly- Carles-Tolra and Andersen (2002), phagous) Kinkorová (2006), Peyrek (1960) Prunus van Aartsen (2001), EPPO (2007), Lampe fruits (wild et al. (2005), Szeőke (2006) P. avium, P. padus, P. serotina) Juglans fruits Duso (1991), EPPO (2004), Merz (1991), Seljak and Zežlina (1999) Prunus fruits Merz (1991) (cultivated). Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) Sturmia bella (Meigen, 1824) Native range 596 Family Species Family Species Ulidiidae Euxesta notata (Wiedemann, 1830) Euxesta pechumani Curran, 1938 Status Regime A A Detrivorous, Phytophagous ? Detrivorous Native range 1st record in Invaded Europe and countries and country islands Habitat* Hosts References North America 2009 FR ? ? Martinez (Unpublished) North America 1969, FR BG, CH, ES, FR, SK E, I Carrion; dung Carles-Tolra and Andersen (2002), Delage (1969), Fauna Europaea, Roháček (2006d) Diptera. Chapter 10 597 Asphondylia borzi (Stefani, 1898) Contarinia lentis Aczél, 1944 Regime Native range Invaded countries Habitat* and islands Phytophagous Phytophagous Phytophagous Alps BE, DK, GB, NL G Larix spp. Ackland (1965), Roques (Unpublished) Alps BE, GB, DK, NL G Larix spp. Ackland (1965), Roques (Unpublished) Alps BE, DK, NL, GB G Larix spp. Ackland (1965), Roques (Unpublished) Predator Central, South Europe Mediterranean Eastern Mediterranean Western Asia GB, LV, RU (?) I, J100 Harris (1976), Mamaev and Krivosheina (1965), Pakalniškis et al. (2006) GB I BG, CZ, FR, HU, SK I Adelges abietis (Adelgidae) Rhamnus alaternus Lens culinaris Pisum sativum Ambrus (1958), Buhr (1939), Forsius (1922), Kieffer (1898), Krištal (1947), Kutter and Winterhalter (1933), Loew (1850), Mamaeva (1969), Meijere (1911), Pakalniškis et al. (2006), Perju (1959), Pileckis and Vengeliauskaite (1977), Schøyen (1926), Simova-Tošić et al. (1996), Simova-Tošić et al. (2000), Skuhravá and Skuhravý (1960), Skuhravá and Skuhravý (2009), Skuhravá et al. (2005), Skuhravá et al. (1991), Skuhrava et al. (2006), Spungis (1977), Theobald (1911), Tullgren (1917) Phytophagous Phytophagous Contarinia pisi (Loew, Phyto1850) phagous AL, AT, BE, BG, I CH, CZ, DE, DK, FI, FR, GB, HU, LT, LV, NL, NO, PL, RO, RS, RU, SE, SI, UA Hosts References Harris (1976), Hill et al. (2005) Aczél (1944), Baudyš (1947), Coutin (1965), Skuhravá (1989), Skuhravá et al. (1991) Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) Family Species Anthomyiidae Strobilomyia infrequens (Ackland, 1965) Strobilomyia laricicola (Karl, 1928) Strobilomyia melania (Ackland, 1965) Cecidomyiidae Aphidoletes abietis (Kieffer, 1896) 598 Table 10.2. Diptera species alien in Europe. List and characteristics. Country codes abbreviations refer to ISO 3166 (see Appendix I). Habitat abbreviations refer to EUNIS (see Appendix II). Last update 05/02/2010. Regime Dasineura abietiperda (Henschel, 1880) Dasineura kellneri (Henschel, 1875) Phytophagous Phytophagous Dasineura pyri (Bouché, 1847) Phytophagous Phytophagous Dasineura rhododendri Phyto(Kieffer, 1909) phagous Kaltenbachiola strobi (Winnertz, 1853) Monarthropalpus flavus (Schrank, 1776) Phytophagous Phytophagous Invaded countries Habitat* Hosts and islands DK, GB, LV, NO, I Pyrus SE communis References Harris (1976), Máca (2006), Skuhravá and Skuhravý (in prep.), Skuhrava et al. (2006), Spungis (1977), Wahlgren (1944) GB, IT G3 Picea abies Harris (1976), Hill et al. (2005) GB G3 Larix decidua Harris (1976), Hill et al. (2005) DK, FI, GB, NO, SE I Pyrus communis Forsius (1922), Harris (1976), Hill et al. (2005), Skuhrava et al. (2006), Wahlgren (1944) GB I Rhododendron ferrugineum Chandler (Ed) 1 (1998), Harris (1976) GB, NL G3 Picea abies Harris (1976), Hill et al. (2005), Roques (Unpublished) AT, CH, CZ, DE, GB, HU, NL, PL, RO, SE, UA I Buxus Ambrus (1958), Docters van Leeuwen (1957), Harris (1976), sempervirens Meyer and Jaschhof (1999), Ryberg (1941), Skuhravá and Skuhravý (1960), Skuhravá and Skuhravý (2009), Skuhrava et al. (1972), Skuhrava et al. (2008), Wahlgren (1944) GB I Quercus ilex Chandler (Ed) 1 (1998), Harris (1976) 599 Phyllodiplosis cocciferae Phyto(Tavares, 1901) phagous Native range Central, eastern Europe, southwest Asia North-east Europe Central Europe, Alps, Carpathians Central, eastern Europe, southwest Asia Central, south Europe, mountains North-east Europe Western Asia, southern Europe, Mediterranean Mediterranean Diptera. Chapter 10 Family Species Contarinia pyrivora (Riley, 1886) Regime Culicidae Aedes vexans (Meigen, Parasitic/ Conti1830) Predator nental Europe Aedes cinereus (Meigen Parasitic/ Conti1818) Predator nental Europe Culex territans Walker, Parasitic/ Eastern 1856 Predator Europe Culex pipiens molestus Parasitic/ ContiL., 1758 Predator nental Europe Syrphidae Chamaesyrphus caledonicus Collin, 1940 Parasitic/ Predator Invaded countries Habitat* Hosts and islands DE I Lavandula angustifolia BE, DK, GB, NL G3 Larix decidua Meyer and Jaschhof (1999) Roques (Unpublished), Skrzypczynska et al. (1993), Skuhrava et al. (2006) GB C1, D Human (biting) GB D, J Human Taylor et al. (2006) (biting), dog GB D, J GB D, J Human Taylor et al. (2006) (biting), dog Human Taylor et al. (2006) (biting), hot-blooded animals GB Continental Europe References G3 Aphids larval predator (pine forests) Taylor et al. (2006) Sivell and Phillips (1999) Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) Phytophagous Phytophagous Native range Mediterranean Central Europe, Alps, Carpathians 600 Family Species Resseliella lavandulae (Barnes, 1953) Resseliella skuhravyorum Skrzypczynska, 1975 Family Species Dasysyrphus friuliensis (van der Goot, 1960) Regime Native range Parasitic/ Predator Continental Europe Parasitic/ Predator Eriozona erratica (Linnaeus, 1758) Continental Europe Parasitic/ ContiPredator nental Europe Parasitic/ Predator Eriozona syrphoides (FallÚn, 1817) Continental Europe References Stubbs and Falk (2002) Stubbs and Falk (2002) Ball and Morris (2000) Diptera. Chapter 10 Didea intermedia Loew, 1854 Invaded countries Habitat* Hosts and islands GB Aphid larval predator (spruce forests); Pollinator Ranunculus and Umbelliferae (Adult) G3 GB Aphid larval predator (Schizolachnus pineti; pine G3 forests) GB Aphid larval predator G3 (forests) GB Aphid larval predator (spruce and pine forestslarva); Pollinator of Hogweed (Heracleum sphondylium) G3 (adult) Ball and Morris (2000) 601 Invaded countries Habitat* Hosts and islands GB Narcissus Hill et al. (2005) and bluebell I2, G1 bulbs GB G3 Ball and Morris (2000) Aphid larval predator Tephritidae Bactrocera (Daculus) oleae (Rossi, 1790) Tephritis praecox (Loew, 1844) Phytophagous Phytophagous CH Terellia fuscicornis (Loew, 1844) Phytophagous Mediterranean I GB, NL Continental Europe Mediterranean GB I2 References Olea Neuenschwander (1984) Calendula arvensis (flower heads) Artichoke (flower head) Jones (2004), Kabos and van Aartsen (1984) Whittington (2002) Marcela Skuhravá, Michel Martinez & Alain Roques / BioRisk 4(2): 553–602 (2010) Native range Parasitic/ ContiPredator nental Europe Parasitic/ ContiPredator nental Europe Parasyrphus malinellus (Collin, 1952) Regime 602 Family Species Merodon equestris (Fabricius, 1794) A peer reviewed open access journal BioRisk 4(2): 603–668 (2010) doi: 10.3897/biorisk.4.50 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Lepidoptera Chapter 11 Carlos Lopez-Vaamonde1, David Agassiz2,3, Sylvie Augustin1, Jurate De Prins4, Willy De Prins4, Stanislav Gomboc5, Povilas Ivinskis6, Ole Karsholt7, Athanasios Koutroumpas8, Fotini Koutroumpa8, Zdeněk Laštůvka9, Eduardo Marabuto10, Elisenda Olivella11, Lukasz Przybylowicz12, Alain Roques1, Nils Ryrholm13, Hana Šefrová14, Peter Šima15, Ian Sims16, Sergey Sinev17, Bjarne Skulev18, Rumen Tomov19, Alberto Zilli20, David Lees1,2 1 INRA UR633 Zoologie Forestière, 2163 Av. Pomme de Pin, 45075 Orléans, France 2 Department of Entomology, The Natural History Museum, Cromwell Road, London, SW7 5BD, UK 3 The Garden House, Stafford Place, Weston-super-Mare, BS23 2QZ, UK 4 Entomology Section, Royal Museum for Central Africa, Leuvensesteenweg 13, B-3080 Tervuren, Belgium 5 Siskovo naselje 19, SI-4000 Kranj, Slovenia 6 Nature Research Centre, Institute of Ecology, Akademijos str. 2, Lt 08412 Vilnius, Lithuania 7 The Natural History Museum of Denmark, Zoologisk Museum, Universitetsparken 15, DK-2100 København Ø, Denmark 8 National Agricultural Research Foundation (NAGREF), Plant Protection Institute of Volos, PB 10303, Fytokou str, 38001 Volos, Greece 9 Department of Zoology, Fisheries, Hydrobiology and Apidology, Faculty of Agronomy, Mendel University in Brno, Zemědělská 1, CZ-613 00 Brno, Czech Republic 10 CBA - Centro de Biologia Ambiental, Faculdade Ciências Universidade de Lisboa, Campo Grande, edificio C2 - Lisboa, Portugal 11 Museu de Ciències Naturals de Barcelona (Zoologia), Passeig Picasso s/n, E-08003 Barcelona, Spain 12 Polish Academy of Sciences, Institute of Systematics and Evolution of Animals, Slawkowska 17, 31-016 Krakow, Poland 13 Department of Natural Sciences, University of Gävle, S-801 76 Gävle, Sweden 14 Department of Crop Science, Breeding and Plant Medicine, Faculty of Agronomy, Mendel University in Brno, Zemědělská 1, CZ-613 00 Brno, Czech Republic 15 Koppert Biological Systems, Komárňanská cesta 13, 940 01 Nové Zámky, Slovakia 16 Syngenta International Research Centre, Jealott’s Hill, Bracknell, Berkshire RG42 6EY, UK 17 Zoological Institute RAS, Universitetskaya nab. 1, 199034 St.Petersburg, Russia 18 Brøndsted 411, DK-3670 Veksø, Denmark 19 University of Forestry, 10 Kliment Ohridski blvd., 1756 Sofia, Bulgaria 20 Museo Civico di Zoologia, Via U. Aldrovandi 18, I-00197 Rome, Italy Corresponding author: Carlos Lopez-Vaamonde (carlos.lopez-vaamonde@orleans.inra.fr) Academic editor: David Roy | Received 31 August 2009 | Accepted 24 May 2010 | Published 6 July 2010 Citation: Lopez-Vaamonde C et al. (2010) Lepidoptera. Chapter 11. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(2): 603–668. doi: 10.3897/biorisk.4.50 Abstract We provide a comprehensive overview of those Lepidopteran invasions to Europe that result from increasing globalisation and also review expansion of species within Europe. A total of 97 non-native Lepidoptera species (about 1% of the known fauna), in 20 families and 11 superfamilies have established so far in Copyright C. Lopez-Vaamonde et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 604 Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Europe, of which 30 alone are Pyraloidea. In addition, 88 European species in 25 families have expanded their range within Europe and around 23% of these are of Mediterranean or Balkan origin, invading the north and west. Although a number of these alien species have been in Europe for hundreds of years, 74% have established during the 20th century and arrivals are accelerating, with an average of 1.9 alien Lepidoptera newly established per year between 2000–2007. For 78 aliens with a known area of origin, Asia has contributed 28.9%, Africa (including Macaronesian islands, Canaries, Madeira and Azores) 21.6%, North America 16.5%, Australasia 7.2% and the neotropics just 5.2%. The route for almost all aliens to Europe is via importation of plants or plant products. Most alien Lepidoptera established in Europe are also confined to man-made habitats, with 52.5% occuring in parks and gardens. We highlight four species in particular, Diaphania perspectalis, Cacyreus marshalli, Cameraria ohridella and Paysandisia archon, as the most important current economic threats. Keywords biological invasion, introduction, pest species, Europe, Lepidoptera, globalisation 11.1 Introduction Apart from the undoubted impact of climate change, various other facets of human activity, including the increasingly efficient means of transport in the last century, increased trade and globalisation, are having a dramatic effect on the composition of European faunas. Lepidoptera, as a mostly alate and largely phytophagous insect group, are particularly affected, not only by increased transport of the invasive species, but by increased trade in plants and stored plant products. In addition, many species are spreading to hostplants not used in their area of origin. Lists of naturalized non-native Lepidoptera are already available for a number of European countries (Agassiz 1996a, Essl and Rabitsch 2002, Geiter et al. 2001, Karsholt and Nielsen 1998, Kenis 2005, Šefrová and Laštůvka 2005). In addition, several detailed case studies have been published on the process of invasion to Europe of several non-native Lepidoptera species (Nash et al. 1995, Šefrová 2001, Šefrová 2002a, Šefrová 2002b, Šefrová and Laštůvka 2001, Whitebread 1990). The first list of terrestrial invertebrate species alien to and within Europe included 272 Lepidoptera species, of which 122 were alien to Europe, 139 alien to countries within Europe, and 11 of cryptogenic origin (DAISIE 2008). We substantially revise and update this list here, in the first comprehensive review of known naturalized non-native Lepidoptera known to Europe. We divided species into two categories: 1. Naturalized exotic species (originating from a continent other than Europe) whose first introduction into Europe appears to be a direct or indirect (deliberate or accidental) result of human activity (Table 11.1). This includes now well known alien lepidoptera such as the Neotropical castniid moth Paysandisia archon (Burmeister, Lepidoptera. Chapter 11 605 1880) or the South African lycaenid butterfly Cacyreus marshalli (Butler, 1898). We also considered in this category species of unknown origin (cryptogenics) such as the leaf-mining moth Phyllonorycter platani (Staudinger, 1870). It is worth noting that we also included here species introduced into confined environments like greenhouses which while not apparently spreading of their own accord, have been introduced with their hostplants, with the potential to spread due to horticultural trade. For instance, 11 species of aquatic Pyralidae have been introduced accidentally by man from Asia and North America into Europe, mostly as contaminants of plants. Current climate makes their establishment in the wild unlikely, but global warming could allow their establishment in the near future. 2. European species spreading throughout the continent as a result of human activity (Table 11.2). This category includes the invasive leaf-mining moth Cameraria ohridella Deschka and Dimić, 1986, now understood to be Balkan in origin (Valade et al. 2009). It is worth noticing that although many aliens are highly invasive our review also includes naturalised aliens that are not necessarily invasive such as the saturniid moth Samia cynthia (Drury, 1773). We excluded all the following cases, here giving examples: i) Species showing clear range expansions/contractions at a country level, which are known to follow global climate change trends (Warren et al. 2001). The butterfly Colotis evagore (Klug, 1829) in Spain (Fric, 2005), the processionary pine moth Thaumetopoea pityocampa (Denis & Schiffermüller, 1775), in central Europe (Battisti et al. 2005) and several British butterfly species (Asher et al. 2001) are classical examples of this phenomenon. However, it must be noted that T. pityocampa has apparently been introduced through human activity from continental Italy to Sardinia (Luciano et al. 2007). ii) Naturally-expanding species known as migrants which have established without clear human assistance, such as the choreutid Tebenna micalis (Mann, 1857) in Azores (Karsholt and Vieira 2005) and the geometrid Peribatodes secundaria (Denis & Schiffermüller, 1775) in Great Britain (Kimber, 2008) as well as rare vagrants that may or may not sporadically naturalize, such as Acontia crocata Guenée, 1852 in France (Letellier, 2004); Pardasena virgulana (Mabille, 1880) in Great Britain (Honey, 1994) and Gelechia sabinellus (Zeller, 1839), Eccopsis effractella Zeller, 1848 and Zophodia grossulariella (Hübner, 1809), all recently recorded from Great Britain (Agassiz 1978a, Agassiz 1996b, Roche 1982). iii) New records of species probably overlooked in particular countries for which there is no clear evidence of range expansion. For instance, in Great Britain the presence of Bucculatrix ulmifoliae Hering, 1931 and Ocnerostoma spp. (Heath and Emmet 1996, Langmaid et al. 2007). iv) Deliberate translocations of species between European countries, such as the introduction of the butterflies Araschnia levana (Linnaeus, 1758) (Frohawk, 1940), Ma- 606 Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) culinea arion (Linnaeus, 1758) (Thomas et al. 2009) and Lycaena dispar (Haworth, 1803) (Ford, 1945) into Great Britain. These translocations result from solitary enthusiasts or are for conservation management purposes including the reintroduction of extinct species, but have nothing to do with our subject of biological invasion, essentially the aspects associated with increased globalisation. v) Species once apparently established but now extinct (e.g. in Great Britain, the blastobasid Blastobasis phycidella (Zeller, 1839) and the oecophorid Euclemensia woodiella (Curtis, 1830) (Emmet 1988, Koster and Sinev 2003) vi) The large number of living display species (this is the case of butterfly houses), unless these species are either establishing in the wild, or there is evidence they have become greenhouse pests (we have no examples). Nevertheless, we highlight the risks involved in importation of butterflies for butterfly houses and for a new practice of wedding releases. Finally, the introduction of exotic host plants by man has indirectly allowed several lepidopteran species to expand their distribution range. We consider as alien species Stigmella speciosa Frey, 1857, Caloptilia rufipennella (Hübner, 1796) and Phyllonorycter geniculella (Ragonot, 1874), all feeding on Acer pseudoplatanus in northern Europe; Stigmella suberivora (Stainton, 1869) feeding on Quercus ilex in Great Britain; Eupithecia phoeniceata (Rambur, 1834) feeding on Juniperus and various Cupressaceae in Belgium and Great Britain, Cydia grunertiana (Ratzeburg, 1868) in Belgium, Denmark and Sweden; C. illutana (Maslov, 1988) and C. pactolana (Zeller, 1840) feeding on Larix, the last two in Great Britain; Thera cupressata (Geyer, 1831), feeding on imported Abies in Sweden and Cupressaceae in Great Britain, and Lithophane leautieri (Boisduval, 1829) on Cupressaceae cultivars in Great Britain. Polychrysia moneta (Fabricius, 1787) started to spread as early as 1891 in Europe, possibly as a result of rise in popularity of ornamental hostplants such as Delphinium in gardens (Agassiz, 1996a). Other well known examples of species which have followed the invasion of their host plants are the milkweed butterflies, Danaus plexippus (Linnaeus, 1758) and Danaus chrysippus (Linnaeus, 1758). The larvae of both species feed on ornamental and invasive milkweeds (Apocynaceae) which have been introduced in some Macaronesian islands and the Iberian Peninsula (Baez, 1998). We summarise the relative importance of naturalized alien invasives by family, in relation to their proportion in the relatively well known European fauna, finding great disparities in their prevalence. 11. 2 Diversity of alien lepidopteran species Lepidoptera is one of the largest insect orders, with around 175,000 described species in 128 families and 47 superfamilies (Kristensen and Skalski 1999, Mallet 2007). About 9,428 native species in 83 families and 31 superfamilies have been recorded in Lepidoptera. Chapter 11 607 Europe (Karsholt and Kristensen 2003). A total of 97 non-native Lepidoptera species, in 20 families and 11 superfamilies have established so far in Europe (Table 11.1). Our analysis reveals that there is a significant correlation between the number of alien species and the number of native species per family (Spearman’s rho correlation: r= 0.48, P < 0.001). In addition, 88 European species in 25 families have expanded their range within Europe and many of these are of Mediterranean origin, invading northern and western areas of Europe (Table 11.2). The 20 families which contain alien species to Europe are: Pyralidae (30 species), Tortricidae (10), Gracillariidae (8), Tineidae (7), Noctuidae (6), Gelechiidae (6), Blastobasidae (5), Yponomeutidae (4), Oecophoridae (4), Cosmopterigidae (3), Saturniidae (3), Pterophoridae (2), Nymphalidae (2) and Bucculatricidae, Agonoxenidae, Lycaenidae, Geometridae, Arctiidae, Nolidae and the alien family Castniidae, each with one species (Table 11.1). Agonoxenidae: Sixteen species of agonoxenids are native to Europe. The Asian species Haplochrois theae (Kusnezov, 1916) represents the only alien. During the 20th century this was a serious pest on tea plantations in Georgia and to a lesser degree, in the Krasnodar Territory of Russia (Sinev, 1994). Arctiidae: One hundred and one species of arctids are native to Europe but only one species, the North American Fall Webworm, Hyphantria cunea (Drury, 1773), is alien to the region. The larvae are highly polyphagous, feeding on hundreds of different species of deciduous trees on which they form conspicuous webbed nests in late summer and autumn. Blastobasidae: Only 41 species of native blastobasid moths have been recorded in Europe, a large evolutionary radiation of which 26 species occur in Madeira (Karsholt and Sinev 2004). However, the number of alien species in this family (five) is relatively high, mainly because the larvae feed usually on dead organic matter. Some species, such as Blastobasis lacticolella (Rebel, 1940) are pests of stored foodstuffs. Interestingly, all alien Blastobasidae appear to have colonized continental Europe (mostly Great Britain and/or mainland Portugal) from Madeira, presumably with the import of ornamental plants. The common species B. adustella Walsingham, 1894 (originally described as a form of B. lignea Walsingham) (Sinev, 2007) is another example. However, B. adustella has widely been treated, erroneously, as a synonym of the Madeira endemic species B. vittata Wollaston, 1858. Although there are records attributed to B. vittata on the internet, including from the British Isles, there are no unambigously identified instances of the introduction of this species outside Madeira at present. Bucculatricidae: There are 53 native bucculatricids known in Europe. One macaronesian species, Bucculatrix chrysanthemella (Rebel, 1896), was recently introduced from the Canaries into Italy and France, where it seems to have established populations. This species has also recently been recorded from Finland, at which latitudes it seems unlikely to become established (Siloaho, 2008). B. chrysanthemella attacks Paris Daisy (Argyranthemum frutescens), an economically important ornamental crop in some parts of Europe. Castniidae: This family has no native species in Europe. The majority of castniid moths are Neotropical, while some species are also found in Australia and South-east 608 Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Asia. The Neotropical Paysandisia archon is the only alien castniid known to occur in Europe and is currently spreading along the Mediterranean coast attacking a wide range of palm species. The castniid Riechia acraeoides (Guérin-Méneville, 1832) is one of numerous sporadic adventatives included in the previous list (DAISIE 2008) that we discount here. Cosmopterigidae: There are 79 species of cosmopterigids native to Europe, with three species considered as aliens. Two of these are African species that feed on Acacia in Malta. The larvae feed internally on the leaves, seeds and stems of their hostplants. There is no evidence that Cosmopterix pulchrimella Chambers, 1875, recently established in Cornwall, Great Britain, arrived there directly through human agency. Gelechiidae: There are 697 species of gelechiids known to occur in Europe. The larvae of most species are concealed feeders on plant tissues, many of them feeding internally in seed heads and fruits, some mining and even producing galls. Six alien gelechiids are known from Europe, among them major agricultural pests such as the Tomato Leafminer Tuta absoluta (Meyrick, 1917), the cosmopolitan Angoumois Grain Moth Sitotroga cerealella (Olivier, 1789), which attacks stored whole cereal grains, and the Pink Bollworm Pectinophora gossypiella (Saunders, 1844), whose larvae bore into the flowers and seeds of cotton. Geometridae: There are 1,024 species of geometrids native to Europe, but only one non-native species appears to have naturalized in Europe. This is Pseudocoremia suavis (Butler, 1879), an endemic geometrid to New Zealand (Berndt et al. 2004), which was recorded on five separate occasions in Cornwall in 2007 (James 2008, Skinner 2009), suggesting establishment in the wild. This species, polyphagous on various gymnosperms, represents a potential risk to European conifer forests. Gracillariidae: There are 249 species of native gracillariids known in Europe and eight alien species have been recorded. Among these are pests of economic importance, such as the Citrus Leafminer Phyllocnistis citrella Stainton, 1856. Lycaenidae: One hundred and thirty-six species of lycaenids are native to Europe. The South African Cacyreus marshalli is one of the few butterflies which are naturalised aliens in Europe (see also under Nymphalidae). This is a pest of cultivated Pelargonium plants, mainly in Mediterranean region but it was found to be breeding in Great Britain in 1997 (Lewes, East Sussex), where it became temporally established in greenhouses until May 1998 but was eradicated (Holloway, 1998). Noctuidae: This is the most species-rich family of Lepidoptera in Europe, with over 1,435 native species. Six alien noctuids have been recorded so far, including some major agricultural pests such as Chrysodeixis eriosoma (Doubleday, 1843) and Spodoptera litura (Linnaeus, 1758). However, on a cautionary note, these genera are known to have strong migratory tendencies. Indeed we may never know, due to lack of sufficient historical records, when or whether certain noctuids arrived as invasives to Europe or by artificial agency. One good example of this is Araeopteron ecphaea (Hampson, 1914) (type locality Nigeria). It is also interesting to note the African and Austral-Oriental fern-feeding species Callopistria maillardi (Guenée, 1862) seems to have been accidentally imported with Nephrolepis ornamental ferns, but this species has five subspecies and the precise origin Lepidoptera. Chapter 11 609 of the introduced individuals is unknown. Some records of Chrysodeixis acuta (Walker, 1858) could also represent misidentifications of C. chalcites (Esper, 1789). Following our exclusion criteria, we have not included singleton records, for example of Acontia crocata Guenée, 1852, a specimen of which was collected in Irais (Deux-Sevres), France (Letellier, 2004), possibly resulting passively from a plant import from SE Asia (Hacker et al. 2008). Nolidae: Thirty-five species of nolids are native to Europe, but only one exotic species has repeatedly been recorded within the region, the Spotted Bollworm, Earias vittella (Fabricius, 1794). The larva of this species feeds on several plants of the family Malvaceae, in particular Okra (Abelmoschus esculentus) pods, Gossypium (it is one of the most important pests of cotton) and Hibiscus. It has been found as a vagrant in Great Britain and seems to also be present in southern Spain (Nash, 2003). Its establishment needs to be confirmed. Nymphalidae: There are 239 species of nymphalid butterflies native to Europe. Two non-native danaine species, the Monarch butterfly Danaus plexippus and the Plain Tiger D. chrysippus have established themselves in the Macaronesian islands and Iberian Peninsula. We have included both species despite them being well known migrants because their introduction and establishment in Europe has followed the invasion and establishment in Europe of their Apocynaceae host plants (Asclepias curassavica, of Neotropical origin and Gomphocarpus fruticosus of Afrotropical origin). Thus, the Monarch’s range has greatly expanded during the 19th and 20th centuries from North America and now encompasses numerous Atlantic, Pacific and Indian Ocean islands and Australia. A number of hypotheses have been developed to explain this great range expansion (Vane-Wright 1993). Oecophoridae: There are 120 native species of oecophorids in Europe. Only four alien oecophorids are established in the region, three of which feed on dead plant material. Pterophoridae: There are 166 native pterophorids known to Europe. Two species, Megalorhipida leucodactylus (Fabricius, 1794) and Lantanophaga pusillidactylus (Walker, 1864) are known to be alien to Europe. M. leucodactylus has a circum-tropical distribution and has established populations in Sicily (Bella and Ferrauo 2005) and Israel. It has also been recorded in Spain, but its presence there needs confirmation (Gielis, pers. comm.). The larvae feed on Amaranthaceae, Cucurbitaceae, Goodeniaceae, Leguminosae, Nyctaginaceae, Rosaceae and Asteraceae (Vargas, 2007). The Lantana Plume Moth L. pusillidactylus is also a pantropical species whose origin, as for M. leucodactylus, is not clear. This species has been introduced with its Verbenaceae hostplant (which is of neotropical origin), Lantana camara, into Spain, Portugal and southern Italy (Aguiar and Karsholt 2006, Bella and Marchese 2007, King 2000). The moth is used as the biocontrol agent against this plant, itself an invasive in many parts of the world. Pyraloidea (Pyralidae and Crambidae): This superfamily has 898 native species known in Europe. Pyraloidea also has the highest number of species (30) alien to Europe. This is probably due to the high number of alien crambid pyrales that have larvae feeding on submerged and floating aquatic plants used in aquariums and ponds (11 species) as well as cosmopolitan pests that feed on stored products (seven species). These invasives include the North American wax moth Vitula edmandsii (Packard, 1865), whose larvae damage the combs of honeybee and bumblebee nests. 610 Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Saturniidae: Seven saturniids are native to Europe. Three Asian species have deliberately been introduced into Europe for silk production, but have naturalized from escapes. This family is also very popular among amateur breeders and sometimes there are reports of adult moths of a wide number of species in urban areas. Tineidae: There are 262 species of native tineids in Europe and seven alien species have also been recorded to the region. At least five of these feed on stored products, cloths, and detritus, such as the Common Clothes Moth (Tineola bisselliella (Hummel, 1823)), whose larvae feed on clothing and natural fibres. Tortricidae: About 977 species of tortrix moths have been recorded as native to Europe. Among the 10 alien species recorded to Europe, there are some economically important pests, in particular of apple trees, for example the oriental fruit moth (Grapholita molesta (Busck, 1916)) and the light brown apple moth Epiphyas postvittana (Walker, 1863). Larvae of the latter species are not easily distinguished from the larvae of other tortricid leafrollers; only DNA-based testing appears to work reliably for identification. Interestingly, half of the tortricids recorded as alien to Europe (five out of 10 species) are specialists on Cedrus and have been introduced into southern France, where plantations of these trees are common. Yponomeutidae: There are 113 species of ermine moths native to Europe, with four alien species having been recorded. The larvae tend to form communal webs, and some species are agricultural forestry pests, such as the Arborvitae Leafminer, Argyresthia thuiella (Packard, 1871) and Prays citri (Millière, 1873), a well-known Citrus pest in the Mediterranean region. Two North American leafminers of the genus Argyresthia attack Cupressaceae in Europe. In our analysis, it is interesting that we found a similar number of alien species to Europe (Table 11.1) as species that have expanded their range within Europe due to human activity (Table 11.2). Indeed, there is a significant correlation between the number of alien species per family to Europe and the number of alien species per family within Europe (Spearman’s rho correlation: r= 0.39, P = 0.044). However, several families exhibit some species which have expanded their range within Europe, yet have very few or no recorded aliens to Europe. For instance, strikingly, Geometridae features only one species alien to Europe within a fauna of 1,024 species, a number of which are known migrants, whereas as many as 11 species have been recorded invading other countries within Europe (Table 11.2). The North American sterrhine geometrid Idaea bonifata (Hulst, 1887) has been intercepted several times with imports of dried plant material but, as far as known, is not yet established in Europe (Martinez and Coutin 1985). The absence of alien species within other species-rich families, such as Coleophoridae (533 spp.), Nymphalidae (239 spp.), Psychidae (231 spp.) Nepticulidae (242 spp.) and Sphingidae (39 spp.) is also notable. In spite of the known high mobility of the last family, several exotic species (i.e. the American Sphinx drupiferarum Smith, 1797, Agrius cingulatus (Fabricius, 1775) and the African Polyptychus trisecta (Aurivillius, 1901)) have been recorded (sometimes repeatedly) within the region, with no confirmed establishment (Marabuto 2006, Pittaway 1993, Waring et al. 2003). Lepidoptera. Chapter 11 611 11.3 Temporal trends The precise date of arrival is not known for two species. An analysis of the 95 species for which the date of the first record in Europe is known shows that the arrival of alien Lepidoptera has dramatically accelerated during the second half of the 20th century (Figure 11.2). This trend is still increasing, with an average of 1.9 alien Lepidoptera newly established per year in Europe between 2000 and 2007 (Figure 11.2). This average is twice that during the period 1975 to 1999 (1.1 species per year). The same trend has been observed for all groups of alien terrestrial invertebrates analysed together (Roques et al. 2008). This temporal trend might be due to the acceleration of processes that happened in much wider time frames in the past, such as global climate change and human assisted transportation via the much faster and more efficient means of transport nowadays. Alien species have historically been introduced for centuries, so it should not be considered that invasive species are necessarily a 20th century phenomenon, although the poor documention of older cases inevitably also provides more scope for speculation. One such case is Euclemensia woodiella, belonging to a North American oecophorid lineage (Koster and Sinev 2003) found in numbers near Manchester in 1829 and not since. A much older potential example is the lasiocampid Pachypasa otus (Drury, 1773) with a scattered distribution in southern Italy, whose larva feeds mainly on Cupressus, could even have been introduced by the Romans for “Coan” silk production, as it possibly represents the “Assyrian Bombyx” mentioned in Naturalis Historia by Plinus (Good, 1995). 11.4 Biogeographic patterns For at least 19 alien species, the precise area of origin is not known and these we consider as cryptogenic. We have classified Phyllonoryctyer platani (Gracillariidae) as cryptogenic because there are some doubts regarding its origin (Šefrová, 2001). Thus, P. platani is either of North American origin and was introduced to Europe with American Platanus occidentalis, or it originated in Southeastern Europe and Southwestern and Central Asia, on Platanus orientalis. We have included C. ohridella as alien within Europe (Table 11.2) since recent genetic studies suggest a Balkan origin as most likely (Valade et al. 2009). An analysis of the 78 alien species for which the native area of origin is known, shows that Asia has contributed the most alien species with 28.9% (28 out of 97 species) (Figure 11.3). Africa (including Macaronesian islands, Canaries, Madeira and Azores) supplied 21.6% of alien species (21 out of 97 species) followed by North America with 16.5%, Australasia with 7.2%, and the Neotropics, surprisingly few with 5.2%. Large differences exist among European countries in the number of alien Lepidoptera recorded per country (Figure 11.4). With 42 species, the United Kingdom is the 612 Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Figure 11.1. Comparison of the number of species per family of Lepidoptera in the alien and native entomofauna in Europe. Families are presented in a decreasing order based on the number of alien species. Only the most important families of native species (> 50 spp.) have been considered. European country with the highest number of alien Lepidoptera, followed by France (mainland) with 41 and Spain (mainland) with 39 species. Both Moldavia and Luxemburg are the European countries with the lowest number (with one alien species each). These differences are very likely to result at least partly from variation in sampling effort and the availability of local taxonomic expertise, but the area and the geographical location of a country is also a very important factor, in this respect. 11.5 Main pathways and vectors to Europe As far as we know, most Lepidoptera alien to Europe have been introduced accidentally (96.9%). A clear exception is some saturniid species that were imported from Asia into Europe for silk production in the nineteenth century, and subsequently became naturalized, including in urban areas. On the other hand, the Silkmoth Bombyx mori Linnaeus, 1758. has not been included in the analysis, because although it is widespread in captivity throughout Europe, its flightlessness has prevented naturalisation. The import of ornamental plants (particularly palms, geraniums and azaleas) is most likely responsible for the introduction of several species such as Paysandisia archon, Cacyreus marshalli and Caloptilia azaleella. Transport also plays an important role in the dispersal of some species, including ones alien within Europe. For instance, Cameraria Lepidoptera. Chapter 11 613 Figure 11.2. Rate of established alien Lepidoptera in Europe since 1492 as mean number of alien Lepidoptera recorded per year. Calculations are made on 95 alien species for which the first record is precisely known. Numbers above bars indicate number of new species recorded per period. ohridella seems to feed almost exclusively on Aesculus hippocastanum trees planted in urban areas and parks. The main means of its spread is likely to be wind dispersal, but human assisted transportation played a major role in the long distance dispersal of this species (Gilbert et al. 2004). Since the advent of tropical butterfly houses in the 1980s, a potential new threat has emerged, the use of mass butterfly releases for weddings, a practice increasingly popular in countries such as Italy, where one of us (AZ) has recorded a number of exotic species flying freely in cities. Usually Monarch butterflies are used, but less scrupulous companies may be using a range of exotics, many of which are likely to find climate change and the availability of hostplants for some papilionid butterflies, such as Rutaceae planted in city gardens propitious for establishment of at least temporary populations. 11.6 Most invaded ecosystems and habitats Most alien Lepidoptera are phytophagous (78.3%), whereas detritivores represent only 21.6% (Table 11.1). The majority of alien Lepidoptera established in Europe are confined to man-made habitats, and only a few species have become established in a more or less natural environment, mostly in woodlands. Examples of the latter include the 614 Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Figure 11.3. Regions of origin of the Lepidoptera species alien to Europe Figure 11.4. Colonization of continental European countries and main European islands by Lepidoptera species alien to Europe. Lepidoptera. Chapter 11 615 Figure 11.5. Main habitats of 97 alien Lepidoptera species established in Europe (note that a species may live in several habitats). arctiid Hyphantria cunea, the gracillariid Phyllonorycter issikii in Central Europe and the saturniid Antheraea yamamai in the Balkans. In Europe, most alien Lepidoptera species feed on their original hostplants. However, some species seem to have been able to switch to other hostplants that are often closely related. For instance, Paysandisia archon specializes on Trithrinax campestris (Arecaceae) and to a lesser extent on Cocos yalai in its native area (Argentina, Uruguay). However, in Europe this moth has expanded its host range to many ornamental exotic palms (Phoenix canariensis, Latania sp.) as well as posing a threat to the native Chamaerops humilis (Montagud Alario 2004). About 50.5% of alien Lepidoptera live indoors in domestic, industrial and other artificial habitats such as 16.5% in greenhouses (Figure 11.5). Six out of the nine species that feed on stored products show a cosmopolitan distribution. Parks and gardens host 52.6% of alien species, where they are frequently introduced with their native hostplant, while 25.8% have colonized agricultural land (Figure 11.5). 11.7 Ecological and economic impact The impact of most alien Lepidoptera species has not been quantified in detail. However, negative economic impact has been recorded for 16 alien species. The Indian 616 Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) a b c d e f g h i j k l Figure 11.6. Adult habitus of some lepidopteran species alien to Europe: a Argyresthia thuiella b Parectopa robiniella c Phyllonorycter issikii winter form d Phyllonorycter issikii summer form e Phyllonorycter leucographella f Phyllonorycter platani g Phyllonorycter robiniella h Plodia interpunctella i Tineola bisselliella j Ephestia kuehniella k Hyphantria cunea male l Hyphantria cunea female (drawings by Aleš Laštůvka). Lepidoptera. Chapter 11 a b c d e f g h i j k l m n 617 Figure 11.7. Adult habitus of some lepidopteran species alien in Europe: a Coleophora laricella b Coleophora spiraeella c Cameraria ohridella d Caloptilia roscipennella e Leucoptera malifoliella f Acalyptris platani g Stigmella aurella h Stigmella atricapitella i Stigmella centifoliella j Stigmella pyri k Stigmella speciosa l Stigmella suberivora m Argyresthia trifasciata; n Ectoedemia heringella. (drawings by Aleš Laštůvka). 618 Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) a b Figure 11.8. Alien saturnid orginating from Asia, Antherea yamamai a adult b 2nd instar larva (credit Zdenek Laštůvka) Meal Moth Plodia interpunctella (Hübner, 1823) may severely affect grain and grain products, dried fruits and seeds in households and warehouses. The Common Clothes Moth Tineola bisselliella is another example of a major pest in houses where it feeds on clothes, carpets, rugs, and upholstered furniture. However, along with several other tineids, this species has become rare due to the increase in use of man-made fibres and the dry environment created by central heating (Kimber, 2008). The most serious alien lepidopteran pests in orchards in many parts of Europe include Grapholita molesta, Hyphantria cunea and Prays citri. Some species can also cause aesthetic impact. Thus, species causing severe infestations can lead to almost complete defoliation of the hostplants. For instance, C. ohridella causes premature defoliation of the white-flowered horse-chestnut, Aesculus hippocastanum. The trees do not die but the aesthetic impact is so severe that in some countries, heavily infested trees have been felled and removed. Lepidoptera. Chapter 11 619 a b c Figure 11.9. Clearwings (Sesiidae) alien in Europe. a Pennisetia hylaeiformis ♂ b Synanthedon andrenaeformis ♂ c Synynthedon myopaeformis ♂. (credit Zdenek Laštůvka) Little is known, however, about the ecological impact of alien Lepidoptera in natural areas of Europe (Kenis et al. 2009). Four alien Lepidoptera species seem to have a potentially important ecological impact: 1) the recently introduced pyralid Diaphania perspectalis that could represent a serious threat to topiary Box hedges and plants in nurseries, parks and gardens, and Buxus shrubs growing in the wild; 2) C. ohridella, that recent studies suggest could have a potential negative impact on native leafminers via apparent competition and could be adapting to Acer species in some areas (Péré et al. 2009); 3) the lycaenid Cacyreus marshalli, which threatens both native geraniums and Geranium-consuming lycaenids (Quacchia et al. 2008); 4) finally, as previously mentioned, Paysandisia archon represents a serious threat to the conservation of natural populations of Chamaerops humilis, the only native palm in Europe (Montagud Alario 2004, Sarto i Monteys 2002). Lastly, we recommend that in order to guarantee the well being of natural ecosystems and also to keep track of future additions to the European alien Lepidoptera list, natural areas of special conservation concern like those under the Natura-2000 framework should be monitored more intensively and regularly for the early detection of potential threats, which according to our results are expected to increase. 620 Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) a b c d e Figure 11.10. Damage by alien lepidopteran larvae. a mines of Parectopa robiniella on Robinia b 3rd instar larva of Cameraria ohridella extracted from its mine on Aesculus c damage of Hyphantria cunea on Acer negundo d mines of Phyllonorycter issikii on Tilia e mines of Phyllonorycter platani on Platanus (Credit: Hana Šefrová). Acknowledgements We would like to thank Leif Aarvik, Giorgio Baldizzone, Jarosław Buszcko, Martin Corley, Mirza Dautbasic, Willem Ellis, Eddie John, Natalia Kirichenko, Ferenc Lakatos, Phil Lambdon, Paul Sammut and Jaan Viidalepp for sending information about alien Lepidoptera from their respective countries. We also thank Cees Gielis and Klaus Sattler for comments on Pterophoridae and Gelechiidae respectively; Shipher Wu and Shen-Horn Yen for their comments on Callopistria maillardi and Juan Jose Pino Perez and Antonio Verdugo Paez for their comments on Danaus. David Lees was funded by a STUDIUM fellowship during preparation of this paper. Lepidoptera. Chapter 11 621 References Aastrup CH (1969) Phthorimaea operculella Zell. Index of Flora and Fauna 1890–1994 75: 63–64. Abafi-Aigner L, Pável J, Uhryk F (1896) Ordo Lepidoptera. Budapest: Regia Societas Scientarium Naturalium Hungarica, 82 pp. Adams RG (1979) Tinea murariella Staudinger in Britain (Lepidoptera: Tineidae). 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Weidner H (1971) Vorratsschädlinge, Bestimmungstabellen der Vorratsschädlinge und des Hausungeziefers Mitteleuropas. Stuttgart, New York: G. Fischer, 328 pp. Whitebread SE (1990) Phyllonorycter robiniella (Clemens, 1859) in Europe (Lepidoptera, Gracillariidae). Nota Lepidopterologica 12: 344–353. Wolff NL (1953) Sommerfuglelarver importeret med bananer. Entomologiske Meddelelser 26: 512–521. Wolff NL (1969) Sterrha inquinata Scop. (Lep., Geometridae) i Danmark. Indeks for Flora og Fauna 1890–1994, 75: 31–32. Wolff NL (1971) Lepidoptera. The Zoology of Iceland 3: 1–193. Yaroshenko VA (1972) Biologiya amerikanskoi beloi babochki i mery bor‘by s nei. Nauchnye trudy Kubanskogo Universiteta 151: 61–91. Zagulajev AK (1982) The potato moth - Phthorimaea operculella Zll. (Lepidoptera, Gelechiidae). Entomologicheskoe Obozrenie 61: 817–820. Zangheri S, Cavalloro R (1971) Sulla presenza in Italia di Epichoristodes acerbella (Walker) (Lepidoptera Tortricidae). Bolletino della Società Entomologica Italiana 103: 186–190. Zilli A (1997) Lepidoptera. In: Zapparoli M (Ed) Gli Insetti di Roma. Roma: Fratelli Palombi Editori, 294–311. Zolnir M (1977) Invertarizacija sladiscnih skodljivcev: porocilo za leto 1975 in koncno porocilo. Zalec: RSS, 27 pp. Zverezomb-Zubowsky EV (1918) Kratkij otchet o deyatel’nosti Donskogo byuro po bor’be s vreditelyami sel’skokhozyaistvennykh rastenij v 1917 godu. Rostov-na-Donu: 1–36. Table 11.1. List and characteristics of the lepidopteran species alien to Europe. Status: A Alien to Europe C cryptogenic species. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Last update 01/06/2009 Family Species Arctiidae Hyphantria cunea (Drury, 1773) Native range 1st record in Europe and country Invaded countries Alien Habitat Hosts References A Phyto- North phagous America 1949, YU AL, AT, BA, BG, CH, G5, I1, I2G CZ, DE, DK, EE, FR, DE, GR, HR, HU, IT, LT, MD, ME, MK, PL, RO, RS, RU, SI, SK, GB Polyphagous on deciduous trees (Acer negundo, Populus, Morus, Prunus, Juglans) Buszko and Nowacki (2000), Essl and Rabitsch (2002), Gaedike and Heinicke (1999), Huemer and Rabitsch (2002), Ippolito and Parenzan (1981), Janežič (1968), Karsholt and Nielsen (1998), Luig and Kesküla (1995), Montermini (1985), Rezbanyai-Reser (1991), Šefrová and Laštůvka (2005), Yaroshenko (1972), Surányi (1946), Torp (1987) A Phyto- Asia phagous 1915, RU RU I1 Thea, Camellia Demokidov (1916), Koster and Sinev (2003) A Detriti- Africa, 1902, IE vorous Macaronesia (PT-MAD) Detriti- Africa, 1946, PT vorous Macaronesia (PT-MAD) BE, FR, GB, IE, NL G5, I2, Decaying vegetal J6 material PT F5, G5, Wide variety of I2, J1, foodstuffs, including J6 leaf-litter, vegetation, and stored products G5, I2, Decaying vegetal J6 material A Blastobasis lacticolella Rebel, 1940 A Detriti- Africa, 1946, GB vorous Macaronesia (PT-MAD) GB, IE Aguiar and Karsholt (2006), De Prins et al. (2009), Karsholt and Sinev (2004) Corley et al. (2006), Karsholt and Sinev (2004) Aguiar and Karsholt (2006), Karsholt and Sinev (2004) 641 Blastobasis decolorella (Wollaston, 1858) Lepidoptera. Chapter 11 Agonoxenidae Haplochrois theae (Kusnezov, 1916) Blastobasidae Blastobasis adustella Walsingham, 1894 Status Regime Cosmopterigidae Anatrachyntis simplex (Walsingham, 1891) Ascalenia acaciella Chrétien, 1915 Bifascioides leucomelanellus (Rebel, 1917) Gelechiidae Coleotechnites piceaella (Kearfott, 1903) A A Native range 1st record in Invaded countries Europe and country 1990, PT ES, FR, PT Detriti- Africa, vorous Macaronesia (PT-MAD) Detriti- Africa, 1998, GB vorous Macaronesia (PT-MAD) GB Alien Habitat B, F5, G5, I2, J6 G5, I2, J6 References Decaying vegetal material Passos de Carvalho and Corley (1995) Decaying vegetal material Aguiar and Karsholt (2006), Karsholt and Sinev (2004) Argyranthemum frutescens Cocquempot and Nel (2009), Constanzi et al. (2008), Klimesch (1979) A Phyto- Africa phagous (Macaronesia) 2007, IT FI, FR, IT A Phyto- Neotropics phagous (South America) c.1995, ES CY, DK, ES, ES-BAL, I2, J100 Palm trees (Phoenix spp, Aguilar et al. (2001), Colazza et FR, GR, GR-CRE, Thritrinax, Chamaerops, al. (2005), Espinosa et al. (2003), IT, IT-SIC, SI Livistona, Trachycarpus, Hollingsworth (2004) Washingtonia). A Phyto- Asiaphagous Tropical 1999, PT CY, ES, GB, PT J1 Polyphagous, cotton, pomegranate fruits Heckford (2004), Koster and Sinev (2003) A Phyto- Africa phagous Phyto- Africa phagous 2001, MT MT I2 Acacia Koster and Sammut (2006) 2004, MT MT I2 Acacia Koster and Sammut (2006) 1952, GB AT, CZ, DE, FR, GB, G3, G5, Picea HU, IT, SK I2 A A Phyto- North phagous America I2 Hosts Essl and Rabitsch (2002), Hill et al. (2005), Huemer and Rabitsch (2002), Reiprich (1991), Šumpich et al. (2007) Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Blastobasis maroccanella Amsel, 1952 Blastobasis rebeli Karsholt & Sinev, 2004 Bucculatricidae Bucculatrix chrysanthemella (Rebel, 1896) Castniidae Paysandisia archon (Burmeister, 1879) Status Regime 642 Family Species Family Species Pectinophora gossypiella (Saunders, 1844) Phthorimaea operculella (Zeller, 1873) Status Regime Native range 1st record in Invaded countries Alien Hosts Europe and Habitat country 1935, IT AL, BG, ES, GR, GR- I1 Cotton CRE, IL, IT, IT-SIC Phyto- Australasia phagous A Phyto- Neotropics phagous (South America) 1899, MT AL, AT, BE, BG, DK, I1, J1 ES, GB, GR, HU, IL, IT, IT-SAR, IT-SIC, MK, MT, NL, PT, PT-AZO, PT-MAD, RU, SE, SI Sitotroga cerealella (Olivier, 1789) A Grain feeder Australasia 1790, DE? AL, AT, BE, BG, BY, J1 CH, CZ, DE, DK, ES, FR, GB, GR, HR, HU, IS, IT, IT-SAR, IT-SIC, LT, MK, MT, NL, NO, PL, PT, PTAZO), RO, RU, RS, SE, SI, SK, GB Tecia solanivora (Povolny, 1973) Tuta absoluta (Meyrick, 1917) A Phyto- Neotropics phagous Phyto- Neotropics phagous 1999, ESCAN 2006, ES ES-CAN A I1, J1 AL, BG, CH, ES, ES- I1, J1 BAL, FR, FR-COR, IT, IT-SIC, LT, MT, SI Karsholt and Nielsen (1986), Povolny (1996), Roll et al. (2007), Russo (1939) Aastrup (1969), Bentinck (1963), Potato, tobacco and other Solanaceae, stored Borg (1899), García Mercet (1926), Huemer and Rabitsch products and fields (2002), Janežič (1951), Karsholt and Sinev (2004), Mendes (1910), Petralia (1949), Roll et al. (2007), Stanev and Kaitazov (1962), Zagulajev (1982) Stored products Borg (1932), Dei (1871), Glavendekić et al. (2005), Hrubý (1964), Huemer and Rabitsch (2002), Ivinskis (1993), Janežič (1951), Karsholt and Nielsen (1976), Karsholt and Vieira (2005), Lindeman (1880), Mehl (1977), Ostrauskas and Taluntyte (2004), Šefrová and Laštůvka (2005), Snellen (1898), Tschorbadjiew (1930) Potato OEPP/EPPO (2005) Tomato Lepidoptera. Chapter 11 A References Harizanova et al. (2009), Ostrauskas and Ivinskis (2010), Urbaneja et al. (2007) 643 Status Regime Native range 1st record in Europe and country Invaded countries Alien Habitat Hosts A Phyto- Australasia phagous (New Zealand) 2007, GB GB Gracillariidae Caloptilia azaleella (Brants, 1913) A Phyto- E Asia phagous 1920, NE AT, BE, CH, CZ, I2, J100 Rhododendron DE, DK, ES, FR, GB, IT, NL, NO, PL, PT, PT-MAD, RU, SE, SI, SK Parectopa robiniella Clemens, 1863 A Phyto- North phagous America 1970, IT AT, BG, CH, CZ, I2, FA, Robinia DE, ES, FR, HR, G1, G5 HU, IT, LT, MK, PL, RO, RS, SI, SK, GB Phyllocnistis citrella (Stainton, 1856) A Phyto- Asia phagous 1993, ES AL, CY, ES, FR, I2 GR, IL, IT, IT-SAR, IT-SIC, MT, PT, PTAZO, PT-MAD, RS Citrus Phyllocnistis vitegenella Clemens, 1859 A Phyto- North phagous America 1997, IT AL, IT, SI Vitis G3, X25 I1 Nothofagus spp., Podocarpus, Kunzea ericoides, Pinus spp. (mainly P. radiata) and Pseudotsuga menziesii James (2008), Skinner (2009) Aguiar and Karsholt (2006), Brants (1913), Della Beffa (1931), Emmet et al. (1985), Gomboc (2003), Huemer and Rabitsch (2002), Jørgensen (1982), Lhomme (1946–1963), Opheim and Fjeldså (1983), Šefrová and Laštůvka (2005), Starý (1936) Buszko and Nowacki (2000), Huemer and Rabitsch (2002), Ivinskis and Rimsaite (2008), Maček (1982), Marek et al. (1991), Olivella (2001), Vidano (1970) de Carvalho and Aguiar (1997), Corley et al. (2000), Garijo and Garcia (1994), Karsholt and Vieira (2005), Mihelakis (1997), Ortu and Delrio (1995), Roll et al. (2007) Posenato et al. (1997), Seljak (2005) Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Geometridae Pseudocoremia suavis Butler, 1879 References 644 Family Species Family Species Status Regime Native range 1st record in Invaded countries Alien Europe and Habitat country 1985, RU AT, BG, BY, CZ, DE, I2, FA, Tilia EE, FI, HR, HU, LT, G1, G5 PL, RO, RU, SI, SK, GB A Phyto- E Asia phagous Phyllonorycter leucographella (Zeller, 1850) A Phyto- SW Asia phagous Phyllonorycter platani (Staudinger, 1870) C Phyto- Cryptogenic 1870, IT phagous Phyllonorycter robiniella (Clemens, 1859) A Phyto- North phagous America 1850, IT 1983, CH AT, BE, BG, CH, DE, CZ, DK, FR, GB, GR, HR, HU, IT, NL, PL, RS, SE, SI, SK I2, FB AL, AT, BE, BG, CH, CZ, DE, DK, ES, FR, FR-COR, GB, GR, HR, HU, IL, IT, IT-SAR, IT-SIC, MK, NL, PL, PT, PTMAD, SE, SI, SK, GB AL, AT, BE, BG, CH, CZ, DE, DK, ES, FR, HR, HU, IT, LT, NL, PL, RO, RS, SI, SK, GB I2, X11, Platanus FA, G5 Rosaceous bushes, mainly firethorn (Pyracantha) I2, X11, Robinia FA, G1, G5 References Bednova and Belov (1999), Buszko and Nowacki (2000), Ermolaev and Motoshkova (2008), Gomboc et al. (in press), Huemer and Rabitsch (2002), Noreika (1998), Šefrová (2002a), Tokár et al. (2002) Baraniak and Walczak (2000), Buhl et al. (1994), Csoka (2001), De Prins (1994), Glavendekić et al. (2005), Huemer and Rabitsch (2002), Maček (1976), Šefrová (1998), Šefrová (1999), Stigter and Frankenhuyzen (1991) Aguiar and Karsholt (2006), BaetaNeves (1945), Frankenhuyzen (1983), Huemer and Rabitsch (2002), Janmoulle (1954), Maček (1968), Roll et al. (2007), Šefrová (2001), Skala (1936), Skala (1937) 645 Bolchi Serini and Trematerra (1989), Buhl et al. (2005), Buszko and Nowacki (2000), De Prins and Groenen (2001), Glavendekić et al. (2005), Huemer and Rabitsch (2002), Huisman et al. (2003), Ivinskis and Rimsaite (2008), Olivella (2001), Šefrová (2002b), Seljak (1995), Tomov (2003), Whitebread (1990) Lepidoptera. Chapter 11 Phyllonorycter issikii (Kumata, 1963) Hosts Lycaenidae Cacyreus marshalli Butler, 1898 Status Regime Native range 1st record in Europe and country Invaded countries Alien Habitat Hosts References Phyto- Africa phagous 1987, ESBAL BE, CH, DE, ES, ES- I2, J1 BAL, FR, FR-COR, GB, IT, IT-SAR, ITSIC, MT, PT Pelargonium Aistleitner (2003), Fuentes Garcia (1997), Sammut (2007), Sarto i Monteys (1992), Trematerra et al. (1997), Troukens (1991), Zilli (1997) A Phyto- North phagous America 1967, RU, RU, GB Ragweed (Ambrosia spp.) Poltavsky and Artokhin (2006), Rezbanyai-Reser et al. (2005), Shchurov (2004) A Phyto- Africa phagous 1987, GR/ ES FR, FR-COR, GR, ES, BAL Rezbanyai-Reser et al. (2004), Robinson et al. (2010), Tautel (2008) Callopistria maillardi (Guenée, 1862) C Chrysodeixis acuta (Walker, 1858) C Chrysodeixis eriosoma (Doubleday, 1843) A Phyto- Cryptogenic 1983, DE, phagous (Oriental, DK Australasia, Pacific and Africa) Phyto- Cryptogenic 1998, AT phagous (Tropical/ Subtropical) Phyto- Australasia 2002, DE phagous Unknown, a New Guinea species of Ecphaea feeds on legume pods Ferns (Adiantum, Lygodium, Nephrolepis, Plleaea) Noctuidae Acontia (Emmelia) candefacta (Hübner, 1831) (according to Fauna Europaea) Araeopteron ecphaea (Hampson, 1914) DE, DK I1, J6 I1, I2 AT, ES, ES-CAN, FR, I1, I2 GB, IE, PT-MAD DE I1,I2 Bathon (1984), Buhl et al. (1985), Karsholt (1994) Polyphagous: Tomato, Aguiar and Karsholt (2006), cotton, soybean, Huemer and Rabitsch (2002) banana, tobacco, Citrus Highly polyphagous, Geiter et al. (2001) foliage and fruit of many field and vegetable crops, ornamentals and weeds: chickpeas, lucerne, maize, potato, sunflower, etc. Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) A 646 Family Species Family Species Spodoptera litura (Fabricius, 1775) Status Regime Native range 1st record in Invaded countries Europe and country 1978, GB AL, DE, GB Alien Habitat Hosts Phyto- Asiaphagous Tropical Nolidae Earias vittella (Fabricius, 1794) A Phyto- Asia phagous 2003, ES ES I1, J6 Okra (Abelmoschus Nash (2003) esculentus) pods, Gossypium and Hibiscus Nymphalidae Danaus chrysippus (Linnaeus, 1758) A Phyto- Africa phagous 1982, ES ES, ES-CAN, GR, HR, IT-SIC I1, I2, G, H A Phyto- North phagous America 1887, ESCAN ES, ES-CAN, PT, PT- I1, I2, AZO, PT-MAD G, H Asclepias, Cynanchum acutum, Gomphocarpus fruticosus, Gossypium arboreum, Caralluma burchardii Asclepias, Gomphocarpus fruticosus, Gossypium arboretum A Detritivorous Detritivorous 1961, PL ES, FR, NL, PL 1972, NL BE, DE, DK, NL, SE J1 Oecophoridae Borkhausenia nefrax Hodges, 1974 Eratophyes amasiella (Herrich-Schäffer, 1854) Neomariania rebeli (Walsingham, 1894) A A Phyto- Africa, 1986, PT phagous Macaronesia (PT-MAD and ESCAN) Detriti- Australasia 1908, GB vorous J1 PT B GB I2 Baez (1998), Gómez de Aizpúrua (2004), Tapia-Domínguez (1982) Baez (1998), Gómez de Aizpúrua (2004), Neves et al. (2001), TapiaDomínguez (1982) Decaying plant material Buszko and Vives Moreno (1992), Kuchlein and van Lettow (1999) Decaying wood Buhl et al. (1991), Buhl et al. (2004), De Prins (2007), Svensson (2007) Unknown Riedl (1990) Withered leaves, leaflitter Hind (2000) 647 Tachystola acroxantha (Meyrick, 1885) A North America Asia (Turkey) Seymour and Kilby (1978) Lepidoptera. Chapter 11 A Danaus plexippus (Linnaeus, 1758) F5, F6, Highly polyphagous, F8, I1, crops and ornamentals I2,J100 References Megalorhipida leucodactylus (Fabricius, 1794) C C Pyralidae + Crambidae Agassiziella A angulipennis (Hampson, 1891) Arenipses sabella A Hampson, 1901 Native range 1st record in Europe and country Invaded countries Alien Habitat Hosts References Phyto- Cryptogenic 1973, PTphagous (tropical, MAD type locality, Jamaica) Phyto- Cryptogenic 1967, ITSIC phagous (tropical , type locality Virgin Islands) ES, IT, PT-AZO, PT- I2 MAD Lantana camara IL, IT-SIC F5,F8, I2 Acacia neovernicosa, Bella and Ferrauo (2005), Gielis Mimosa tenuiflora (1996) (Fabaceae), Boerhavia diffusa, B. coccinea, B. chinensis, B. repens, Commicarpus tuberosus Okenia hypogaea (Nyctaginaceae), Amaranthus (Amaranthaceae), Scaevola frutescens (Goodeniaceae),Tessaria absinthioides (Asteraceae). Phyto- Asia phagous GB, NL J1, J100 Aquatic water plants ES, FR I2 1977, GB Phyto- Africa 1999, ES phagous (North Africa, Middle east) Aguiar and Karsholt (2006), Bella and Marchese (2007), Kimber (2008) Goater (1986), Goater et al. (2005) Palm trees (Phoenix spp) Asselbergs (1999), Streito and Martinez (2005) Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Pterophoridae Lantanophaga pusillidactylus (Walker, 1864) Status Regime 648 Family Species Family Species Status Regime C Phytophagous Cadra figulilella (Gregson, 1871) C Detritivorous Chilo suppressalis (Walker, 1863) Corcyra cephalonica (Stainton, 1866) A Phytophagous Detritivorous C 1st record in Invaded countries Europe and country Cryptogenic 1900,? AL, AT, BE, BG, CH, (type CY, CZ, DK, ES, FI, locality: FR, FR-COR, DE, India) GB, GR, GR-CRE, HU, IE, IT, IT-SAR, IT-SIC, LV, LT, MT, NL, NO, PL, PTAZO, PT-MAD, RO, RU, SE, SK Cryptogenic 1871, GB AL, AT, BA, BE, BG, (type BY, CH, CY, CZ, locality: DE, ES, FR, FRLiverpool, COR, GB, GR, GREngland) CRE, HR, HU, IE, IT, IT-SAR, IT-SIC, LT, LU, MK, MT, NO, PL, PT, PTMAD, RO, RS, RU, SE, SI, SK, GB Asia 1949, ES ES, FR, FR-COR, RU Cryptogenic 1866, GB AT, BE, BG, CH, (Tropical, CZ, DE, DK, ES, FR, subtropical, GB, GR, IT, IT-SIC, (type LV, PL, PT, PT-AZO, locality, RO, SE, GB Great Britain) Alien Habitat Hosts References J1 Stored Products: dried fruits, nuts, grain Aguiar and Karsholt (2006), von Andres (1916), Filipjev (1932), Huemer and Rabitsch (2002), Janmoulle (1965), Karsholt and Vieira (2005), Kenis (2005), Mehl (1977), Ostrauskas and Taluntyte (2004), Reiprich (1990), Šefrová and Laštůvka (2005), Paoli (1922) J1 Dried fruits, raisins, figs Carnelutti (1975), De Crombrugghe (1909), Goater (1986), Huemer and Rabitsch (2002), Kenis (2005), Reiprich and Okáli (1989), Roesler (1973), Šefrová and Laštůvka (2005) I1 Rice (leaves, stems) Feron (1973), Gerasimov (1949) J1 Stored grain (Poaceae: e.g. rice) Drensky (1930), Goater (1986), Huemer and Rabitsch (2002), Janmoulle (1938), Karsholt and Vieira (2005), Palm (1986), Šefrová and Laštůvka (2005), Silvestri (1943) Lepidoptera. Chapter 11 Cadra cautella (Walker, 1863) Native range 649 Native range A Phyto- Asia phagous A Phyto- Asia phagous A Phytophagous Phytophagous Phytophagous A A A C Phytophagous Detritivorous 1st record in Invaded countries Europe and country 2007, DE CH, DE, FR, NL Alien Habitat Hosts References I2 Buxus Brua (2008), Rennwald (2008) 2000, PT BE, DK, ES, ES-BAL, E3 MT, NL, PT Carex Buhl (in press), Muus and Wullaert (2008), Speidel et al. (2007) Asia 1978, DK CZ, DK, FI, GB, NL J1, J100 Aquatic plants AsiaTropical AsiaTropical 1978, DK/ GB 1978, GB CZ, DK, GB J1, J100 Aquatic plants GB J100, J1 Aquatic plants Buhl et al. (1982), Goater et al. (2005) Buhl et al. (1982), Vrabec and Heřman (2006) Agassiz (1978b) GB J100 Goater (1986) North 1968, GB America Cryptogenic 1796, DE (type locality, Germany) AL, AT, BE, BG, CH, J1 CY, CZ, DE, DK, EE, ES, FI, FR GB, GR, GR-CRE, HU, IE, IS, IT, IT-SAR, IT-SIC, LT, LV, MK, MT, NL, NO, PL, PT, PT-AZO, PTMAD, RO, RU, SE, SI, SK Waterlily Stored nuts, dried fruits, Abafi-Aigner et al. (1896), grain, etc. Aguiar and Karsholt (2006), Caruana Gatto (1905), De SélysLongchamps (1844), Filipjev (1932), Huemer and Rabitsch (2002), Karsholt and Vieira (2005), Kenis (2005), Mehl (1977), Petersen (1924), Reid (2008), Šefrová and Laštůvka (2005), Speiser (1903), Paoli (1922) Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Diaphania perspectalis (Walker, 1859) Diplopseustis perieresalis (Walker, 1859) Elophila difflualis (Snellen, 1880) Elophila manilensis Hampson, 1917 Elophila melagynalis (Agassiz, 1978) Elophila obliteralis (Walker, 1859) Ephestia elutella (Hübner, 1796) Status Regime 650 Family Species Family Species Status Regime C Eustixia pupula Hübner, 1823 Herpetogramma licarsisalis (Walker, 1859) A Leucinodes orbonalis (Guenée, 1854) Paralipsa gularis (Zeller, 1877) C A A 1st record in Invaded countries Alien Hosts Europe and Habitat country Detriti- Cryptogenic 1879,? AL, AT, BA, BE, BG, J1 Stored nuts, dried fruits, grain, etc. vorous (no type CH, CY, CZ, DE, locality) DK, EE, ES, FI, FR, GB, GR, GR-CRE, HU, IE, IS, IT, ITSAR, IT-SIC, LT, LV, ME, MT, NO, PL, PT, PT-AZO, PTMAD, RO, RS, RU, SE, SI, SK Phytophagous Phytophagous North 1997, GB America Cryptogenic 1994, CY (type locality: Malaysia: Sarawak, Old world tropics: Asia and Africa) Africa 2004, BE Phytophagous Detriti- SE Asia vorous (type locality: Japan) 1921 GB I1 CY, ES, MT, PT, PT- I2, E1 MAD,SE BE I1 AT, BE, CH, CZ, DE, DK, FR, GB, HU, IT, IT-SIC, LV, NL, NO, SE J1 Cabbage, Lepidium virginicum (Cruciferae) Monocots, turf grasses, pastures Solanum melongena (eggplant) Dry fruits, occasionally in imports of nuts for chocolate industry. References Aguiar and Karsholt (2006), Bolle (1921), Borg (1932), De Crombrugghe (1906), Glavendekić et al. (2005), Goater (1986), Hrubý (1964), Huemer and Rabitsch (2002), Janežič (1951), Karsholt and Vieira (2005), Kenis (2005), Mehl (1977), Palm (1986), Šefrová and Laštůvka (2005), Zverezomb-Zubowsky (1918) Budd and Goater (1998) Aguiar and Karsholt (2006), Karsholt and Vieira (2005), Sammut (2000) Lepidoptera. Chapter 11 Ephestia kuehniella Zeller, 1879 Native range Nyst (2004) De Prins (1983), Giunchi (1957), Goater (1986), Huemer and Rabitsch (2002), Mariani (1941– 1943), Mehl (1977), Palm (1986), Šefrová and Laštůvka (2005), 651 Native range A Phyto- North phagous America A Phytophagous Phytophagous Phytophagous A C Parapoynx fluctuosalis (Zeller, 1852) C Parapoynx obscuralis Grote 1881 Parapoynx polydectalis Walker, 1859 Phycita diaphana (Staudinger, 1870) A A C AsiaTropical AsiaTropical Cryptogenic (Old world tropics: Asia and Africa) Phyto- Cryptogenic phagous (ES, Asia and Africa, type locality, Natal) Phyto- North phagous America Phyto- Australasia phagous 1st record in Invaded countries Europe and country 1870, AT AT, IT I2 Juglans Huemer and Rabitsch (2002), Trematerra (1988) 1978, DK DK, GB, SE J100 Aquatic plants 1979, GB GB J100 Aquatic plants Hancock (1984), Karsholt and Nielsen (1998) Goater (1986) 1977, GB AT, CZ, DK, FI, GB J100 Nymphaea Buhl et al. (1982), Goater (1986), Goater et al. (2005), Huemer and Rabitsch (2002) 1979, GB GB J100 Aquatic plants Goater (1986) 1967, GB GB J100 Aquatic plants Goater (1986) 1979, GB GB, NL J100 Aquatic plants Goater et al. (2005) ES, GR, PT I2,J6 Ricinus communis Corley et al. (2000) Detriti- Cryptogenic 1870, ES, vorous (type (2002, PT) locality: Spain, Malaga) Alien Habitat Hosts References Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Paramyelois transitella (Walker, 1863) Parapoynx bilinealis Snellen, 1876 Parapoynx crisonalis (Walker, 1859) Parapoynx diminutalis Snellen, 1880 Status Regime 652 Family Species Family Species Status Regime Native range C Detritivorous Pseudarenipses insularum Speidel & Schmitz, 1991 C Phytophagous Spoladea recurvalis (Fabricus, 1775) A Phytophagous Vitula edmandsii (Packard, 1865) ssp. serratilineella Ragonot, 1887 Saturniidae Antheraea pernyi (Guérin-Méneville, 1855) A Detritivorous A Phyto- Asia phagous 1900, ES ES, ES-BAL, HU Hosts References J1 Stored plant products Aguiar and Karsholt (2006), Borg (1932), Goater (1986), Hrubý (1964), Huemer and Rabitsch (2002), Ivinskis (1976), Karsholt and Vieira (2005), Kenis (2005), Martin (1991), Mehl (1977), Palm (1986), Rebel (1901), Šefrová and Laštůvka (2005), Zolnir (1977) I2 Phoenix canariensis Reynaud et al. (2002), Sammut (2003), Sammut (2005) I1, I2 Beta vulgaris, Trianthema postulacastrum, Celosia sp., Chenopodium sp., Portulaca sp., Amaranthus sp. De Prins (2005), Karsholt and Vieira (2005), Nuss ( 2010) J Honey, pollen, broods in bee nests Kullberg and Mikkola (2001), Palm (1986), Svensson (1986), Weidner (1971) G1,G5, Quercus, Fagus, Betula, I2 Aesculus Pittaway (2008) 653 Plodia interpunctella (Hübner, 1813) Alien Habitat Lepidoptera. Chapter 11 1st record in Invaded countries Europe and country Cryptogenic 1813, DE? AL, AT, BE, BG, BY, (no type CH, CZ, DE, DK, locality) EE, ES, FI, FR, GB, GR, GR-CRE, HU, IE, IS, IT, LV, LT, ME, MK, MT, NL, NO, PL, PT, PTAZO, PT-MAD, RO, RU, SE, SI, SK, GB Cryptogenic 2002, FR, ES, ES-CAN, FR, (type 2003, MT MT locality: Tenerife, Santa Cruz) Tropics: 1968, BE, DK, IT, NL, PTAsia (type NL (from AZO, PT-MAD locality: Canaries) India Orientali) South America and Africa North late 1940’s, DE, DK, FI, NO, SE America DE Status Regime Native range 1st record in Invaded countries Europe and country 1866-1868, AT, BA, DE, HR, SI HU, IT, MK, RO, RS, SI Alien Habitat Hosts References A Phyto- Asia phagous Samia cynthia (Drury, 1773) A Phyto- Asia phagous 1854, IT G1,G5, Quercus, Aesculus, Fagus, Blažič et al. (1995), Casale (1973), I2 Castanea, etc. Glavendekić et al. (2005), Huemer and Rabitsch (2002), Pittaway (2008) AL, AT, CH, DE, ES, I2, X24 Ailanthus and other Huemer and Rabitsch (2002), FR, HR, IT, SI deciduous trees Kenis (2005), Kollar(1854), Koster and Sinev (2003), Lepidopterologen Arbeitsgruppe (2000), Quajat (1904) A Phyto- Africa phagous 1923, PTMAD DK, PT, PT-AZO, PT-MAD, SE A Phyto- C. Africa phagous 1910, PTMAD Tineidae Opogona omoscopa (Meyrick, 1893) Opogona sacchari (Bojer, 1856) I1, I2J1 Stored products (grain, fruits), plants with mosses AL, BE, BG, CH, I2, J1, Dracaena, Strelitzia, CZ, DK, ES, ESJ100 Yucca, Alpinia, CAN, GB, GR, HU, Begonia, Bougainvillea, Bromeliaceae, Palms IT, NL, PL, PT-AZO, (Chamaedorea etc.), PT-MAD Cordyline, Cycas, Hibiscus, Dieffenbachia, Poinsettia, Ficus, Gloxinia, Heliconia, Ippeastrum, Maranta, Philodendron, Sansevieria Saintpaulia, banana plantations (Musa acuminata) Buhl et al. (1997), Corley (2005), Gaedike and Karsholt (2001), Karsholt and Vieira (2005) Aguiar and Karsholt (2006), Ciampolini (1973), Gaedike and Karsholt (2001), Jannone (1966), Karsholt and Vieira (2005), Sitek (2003), Walsingham (1910), Wolff (1953) Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Antheraea yamamai (Guérin-Méneville, 1861) 654 Family Species Family Species Status Regime Native range 1st record in Invaded countries Europe and country Detriti- Cryptogenic before 1979, CY, MT, PT-AZO, PT-MAD PT-MAD vorous (type locality: Sierra Leone) Phyto- Asia (type 1909, IE GB, IE phagous locality: Ireland) C Psychoides filicivora (Meyrick, 1937) A Tinea pallescentella Stainton, 1851 A Detriti- Neotropics vorous (South America) Tinea translucens Meyrick, 1917 A Detriti- S Asia (type 1856, UK vorous locality: Pakistan) Tineola bisselliella (Hummel, 1823) C Detriti- Cryptogenic 1794, SE vorous (type locality, Europe) 1840, IE Hosts F5, F6, Pteridium F7, I1, I2 E5, F3, Ferns (Polystichum J100 setiferum, Dryopteris filix-mas, Phyllitis scolopendrium), often found indoors. AT, BE, CZ, DE, DK, J1, J2, Furs, dry fish FI, FR, GB, HU, IE, J6 IT, LV, NL, NO, RO, RU, SE, SK Aguiar and Karsholt (2006), Gaedike and Karsholt (2001), Karsholt and Vieira (2005) Beirne (1940), Gaedike and Karsholt (2001), Kimber (2008) Heath and Emmet (1985), Karsholt and Nielsen (1998), Mehl (1977), Šefrová and Laštůvka (2005), Tokár et al. (1999), Vives Moreno (2003) Stored products, clothes Buhl et al. (1987), Ivinskis (1993), Opheim and Fjeldså (1983), Pelham-Clinton (1985), Reiprich (1992), Šefrová and Laštůvka (2005), Tokár et al. (2002) Stored products, clothes Drenowsky (1909), Hrubý (1964), Karsholt and Nielsen (1998), Mehl (1977), Mendes (1904), Mendes (1905), Palionis (1932), Peterson and Nilssen (2004), Šefrová and Laštůvka (2005) 655 AL, AT, CY, CZ, DE, J1 DK, ES, FR, GB, GR, GR-CRE, HR, HU, IS, IT, IT-SAR, IT-SIC, LV, LT, NO, PT, RO, RU, RS, SK, GB AT, BE, BG, BY, CH, J1, J2 CZ, DK, EE, ES, FI, FR, FR-COR, DE, GB, HU, IS, IE, IT, LT, LV, NL, NO, PL, PT, RO, RU, SE, SI, SK, GB References Lepidoptera. Chapter 11 Praeacedes atomosella (Walker, 1863) Alien Habitat Cryptophlebia leucotreta (Meyrick, 1927) Dichelia cedricola (Diakonoff, 1974) Epichoristodes acerbella (Walker, 1864) Native range 1st record in Europe and country Invaded countries Alien Habitat Hosts References A Phyto- Asia phagous 1998, ES? ES, FR FA, G3, Cedrus I2, X11 Vives Moreno (2003) A Phyto- North phagous America 1979, DE DE,DK, GB, PTMAD I1, I2, J100 Buhl et al. (1997), Hill et al. (2005) A Phyto- Africa phagous 1965, FI IL,FI A Phyto- Asia phagous Phyto- Africa phagous POST-2001, FR FR 1960, DK DK, ES, FR, GB, IT, IT-SAR, IT-SIC, NO, RS Phyto- Africa phagous Phyto- Africa phagous Phyto- Australasia phagous A Epinotia algeriensis Chambon, 1990 Epinotia cedricida Diakonoff, 1969 A Epiphyas postvittana (Walker, 1863) A A Citrus, Euphorbia pulcherrima, strawberries, and low herbaceous plants I1, J100 Citrus, Macadamia terniflora, Ricinus communis, cotton G3, I2 Cedrus I2 Polyphagous, especially Dianthus POST-1990, FR FR 1968, FR AT, BG, FR G3 Cedrus G3, I2 Cedrus 1911, GB I1, I2 Polyphagous (Malus, etc.) GB, PT-AZO Bradley (1959), Hamburger et al. (2000), Karvonen (1983) Fabre et al. (2001) Costa Seglar and Vives Quadras (1976), Fjelddalen (1965), Glavendekić et al. (2005), Thygesen et al. (1965), Zangheri and Cavalloro (1971) Chambon et al. (1990) Du Merle (1988), Huemer and Rabitsch (2002), Leclant (1969), Vives Moreno (2003) Agassiz (1996a), Karsholt and Vieira (2005) Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Tortricidae Acleris undulana (Walsingham, 1900) Clepsis peritana (Clemens, 1860) Status Regime 656 Family Species Family Species Status Regime Native range 1st record in Invaded countries Alien Hosts Europe and Habitat country 1920, SI AL, AT, BA, BG, CH, I2 Rosaceae (Prunus, Pyrus, Malus) CZ, DE, DK, ES, FR GR, HU, IT, IT-SAR, IT-SIC, LT, ME, MK, PT-AZO, RO, RS, RU, SI, SK, GB Phyto- Asia phagous Lozotaenia cedrivora Chambon, 1990 Yponomeutidae Argyresthia cupressella Walsingham, 1890 A Phyto- Africa phagous 1968, FR FR G3, I2 Cedrus A Phyto- North phagous America 1997, GB GB I2 Argyresthia thuiella (Packard, 1871) A Phyto- North phagous America 1971, NL AT, BE, BG, CH, CZ, DE, HU, NL, PL, SI, SK I2, FA Cupressaceae (Chamaecyparis, Cupressocyparis , Juniperus) Thuja, occasionally other Cupressaceae Prays citri (Millière, 1873) A Phyto- Asia phagous 1877, IT Prays peregrina Agassiz, 2007 C Phyto- Cryptogenic 2003, GB phagous AL, DK, ES, FR, I2, J100 Citrus FR-COR, GR, GRCRE, IL, IT, IT-SAR, IT-SIC, NL, PT, PTAZO, PT-MAD GB I2 Unknown Domínguez García-Tejero (1943), Dufrane (1960), Glavendekić et al. (2005), Hrdý and Krampl (1977), Huemer and Rabitsch (2002), Ivinskis (1993), Janežič (1951), Karsholt and Vieira (2005), Katsogiannos and Koveos (2001), Kyparissoudas (1989), Paoli (1922), Strygina and Shutova (1966), Tzalev (1979) Fabre (1997) Agassiz (1999) De Prins (1983), Frankenhuyzen (1974), Huemer and Rabitsch (2002), Šefrová and Laštůvka (2005), Škerlavaj and Munda (1999), Tokár et al. (1999) Buhl et al. (2001), de Carvalho (1995), Franco et al. (2006), Karsholt and Vieira (2005), Liotta and Mineo (1963), Roll et al. (2007) Agassiz (2007) 657 A Lepidoptera. Chapter 11 Grapholita molesta (Busck, 1916) References Coleophoridae Coleophora coracipennella (Hübner, 1796) Coleophora laricella (Hübner, 1817) Coleophora spiraeella Rebel, 1916 Coleophora versurella Zeller, 1849 Epermeniidae Epermenia aequidentellus (Hoffmann, 1867) Ethmiidae Ethmia terminella Fletcher, 1938 Regime Native range Invaded countries Alien Habitat Hosts Refs Phyto- Europe, W Asia AT, BE, CH phagous & N Africa B3 Algae and lichens Fologne (1859), Huemer and Rabitsch (2002), Kenis (2005) Detrivorous AT, GR, HR, PT-AZO, SK, RU G, F4-9, FA,G, J6 Decaying plant material Gozmány (2008), Huemer and Rabitsch (2002), Karsholt and Vieira (2005), Tokár et al. (2002) PT-MAD I1, I2, X24 Malus Aguiar and Karsholt (2006) BE, DK, HR, EE, FI, GB, HR, IE, LT, LV, MK, NL, NO, RS, SE Phyto- C Europe (incl. DE, HU, IT, LT, SE, SK phagous CZ, AU) G3 Larix Bond et al. (2006), De Fré (1858) G, I2 Spiraea Phyto- Europe phagous PT-AZO E1 Atriplex, Chenopodium Baldizzone (pers. comm.), Huemer and Rabitsch (2002), Reiprich and Janovský (1981) Karsholt and Vieira (2005) Phyto- C & S Europe phagous PT-AZO U Daucus carota Karsholt and Vieira (2005) B2 Echium vulgare Svensson (1992) ?, described from North America Phyto- W Europe phagous Phyto- European Alps phagous Phyto- Europe to N SE phagous Africa and Asia Minor Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Family Species Arctiidae Eilema caniola (Hübner, 1808) Autostichidae Oegoconia novimundi Busck, 1915 658 Table 11.2. List and characteristics of the lepidopteran species expanding within Europe (alien in Europe). Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Last update 01/06/2009. Family Species Gelechiidae Athrips rancidella (Herrich-Schäffer, 1854) Chrysoesthia sexguttella (Thunberg, 1794) Gelechia senticetella (Staudinger, 1859) Regime Native range Invaded countries Alien Habitat Hosts Refs I2 Cotoneaster horizontalis Chalmers-Hunt (1985) Phyto- Europe and/or phagous N Africa Phyto- European Alps phagous PT-AZO D6 Chenopodium Karsholt and Vieira (2005) BE, DK, GB, NL, BU I2, G Juniperus, Cupressus Platyedra subcinerea (Haworth, 1828) Geometridae Bupalus piniaria (Linnaeus, 1758) Erannis defoliaria (Clerck, 1759) Eupithecia carpophagata Staudinger, 1871 Eupithecia indigata (Hübner, 1813) Phyto- Europe phagous PT-AZO I2 De Prins (1989), van Nieukerken et al. (1993), Buhl et al. (2007) Malva sylvestris, seeds, flowers Karsholt and Vieira (2005) Phytophagous Phytophagous Phytophagous Phytophagous Europe IE G3 Pinus Europe IS G1, I2 Mediterranean DE E4 G3 Polyphagous (Quercus, Betula, Wolff (1971) Ulmus, Acer, Tilia) Silene (S. saxifraga, S. Geiter et al. (2001) rupestris) Pinus sylvestris, Picea, Larix Skou (1986) Eupithecia phoeniceata (Rambur, 1834) Eurranthis plummistaria (De Villers, 1789) Idaea inquinata (Scopoli, 1763) Phytophagous Phyto- Mediterranean phagous Phyto- S Europe to phagous Asia Minor, N Africa Phyto- Europe to E phagous Asia I2 Juniperus phoenicea De Prins (2007) DE F6 Dorycnium Geiter et al. (2001) DK, FI, LV, NL, SE G1,G5,J1 Ever-lasting flowers/dry and withered petals Naves (1995), Skou (1986), Wolff (1969) IE G3 Pinus sylvestris Roques et al. (2006) Macaria liturata (Clerck, 1759) Europe from IE Urals W and S to Alps Atlantic Europe BE, GB Moffat (1897) 659 GB Lepidoptera. Chapter 11 Phyto- Europe phagous Regime Invaded countries Alien Habitat Hosts Refs Europe to Caucasus Europe IS G Deciduous trees Peterson and Nilssen (2004) DE F4 Juniperus Savela (2010) Europe SE G3 Abies, Pinus Skou (1986), Svensson (1977) Phyto- Southern phagous Balkans AT, BA, BE, BG, BY, CH, I2, X11, FA, Aesculus hippocastanum CZ, DE, DK, ES, FI, FR, G1 FR-COR, GB, HR, HU, IT, LV, LT, NL, PL, RO, RU, RS, SE, SI, SK, UK Caloptilia rufipennella (Hübner, 1796) Caloptilia roscipennella (Hübner, 1796) Phyto- Europe phagous Phyto- Europe or SW phagous Asia? LT, LV, NO, SE I2, G3 Acer pseudoplatanus Buhl et al. (2003), Butin and Führer (1994), De Prins and Puplesiene (2000), Hill et al. (2005), Huemer and Rabitsch (2002), Karsholt and Kristensen (2003), Łabanowski and Soika (1998), Laštůvka et al. (1994), Milevoj and Maček (1997), Šefrová and Laštůvka (2001), Stigter et al. (2000), Vives Moreno (2003) Kimber (2008) I2, G3 Juglans regia Šefrová and Laštůvka (2005) Phyllonorycter geniculella (Ragonot, 1874) Phyllonorycter joannisi (Le Marchand, 1936) Phyto- Europe phagous AT, BE, CH, CZ, DE, ES, FR, FR-COR, HU, IT, ITSIC, MD, PL, RO, RU, UK GB, LV, LT, SE I2, G5 Acer pseudoplatanus Emmet et al. (1985) GB I2, G5 Acer platanoides Emmet et al. (1985) Phyto- Europe phagous Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Phytophagous Phytophagous Phytophagous Native range 660 Family Species Operophtera brumata (Linnaeus, 1758) Peribatodes perversaria (Boisduval, 1840) Thera britannica (Turner, 1925) Gracillariidae Cameraria ohridella Deschka & Dimić, 1986 Family Species Phyllonorycter messaniella (Zeller, 1846) Phyllonorycter strigulatella (Zeller, 1846) Lasiocampidae Dendrolimus pini (Linnaeus, 1758) Regime Native range Refs Phyto- Europe phagous GB G,J1, H1 Alnus incana Hill et al. (2005) Phyto- Most of Europe GB phagous E to Urals and S to S. Italy and Sicily, NW North Africa and Asia Minor to Caucasus and Near East G3 Pinus spp. Kimber (2008), Mikkola and Ståhls (2008) Lyonetiidae Leucoptera malifoliella (O. Costa, 1836) Phyto- Mediterranean phagous PT-MAD I1, G1, G2 Polyphagous, mostly Rosaceae Aguiar and Karsholt (2006) (Malus, Pyrus, Sorbus, Crateagus, Prunus), Betula Nepticulidae Acalyptris platani (Müller-Rutz, 1934) Ectoedemia heringella (Mariani, 1939) Stigmella atricapitella (Haworth, 1828) Stigmella aurella (Fabricius, 1775) Stigmella centifoliella (Zeller, 1848) Phytophagous Phytophagous Phytophagous Phytophagous Phytophagous CH, ES, HR, FR, FR-COR, IT, PT, SI, GB FA, G, I2, X11 I2, G2 Platanus van Nieukerken et al. (2004) Quercus ilex leaf miner Hill et al. (2005) PT-MAD, ES G1,G4,X10 Quercus Aguiar and Karsholt (2006) Europe PT-AZO I1 Karsholt and Vieira (2005) Europe PT-MAD B1,X24,X25 Rosa Quercus, Fagus, Castanea Aguiar and Karsholt (2006), Karsholt and Vieira (2005) Rubus Aguiar and Karsholt (2006) 661 PT-AZO S. Europe (Adriatic) Europe Hosts Lepidoptera. Chapter 11 Phyto- Europe phagous Alien Habitat I2, G E Balkans Invaded countries Euplexia lucipara (Linnaeus, 1758) Lithophane leautieri (Boisduval, 1829) Polychrysia moneta (Fabricius, 1787) Sesamia nonagrioides (Lefèbvre, 1827) Spodoptera littoralis (Boisduval, 1833) Nolidae Native range Invaded countries Alien Habitat Hosts Refs Phyto- Europe phagous Phyto- Europe phagous Phyto- S Europe phagous SE I1 Pyrus Johansson et al. (1990) DK, GB G, I2 Acer pseudoplatanus Heath and Emmet (1983) GB G3,G4 Quercus ilex Heath and Emmet (1983) Phyto- E phagous Mediterranean, N & NE Africa Phyto- Mediterranean phagous &/or tropical Africa Phyto- Europe & W phagous Asia, N Africa Phyto- Mediterranean phagous expanding to C Europe, N Africa Phyto- C & SE Europe phagous to W Asia Phyto- S Europe, N, phagous W, and SW Africa Phyto- Subtropical phagous Africa Madagascar and S Europe CH I2, X11 CZ, PL, SE J100, I1 Vegetables in glasshouses Šefrová and Laštůvka (2005) PT-AZO G Ferns Karsholt and Vieira (2005) DK, GB, NL I2 Chamaecyparis, Cupressocyparis BE, DK, DE, GB I2 Delphinium Bednova and Belov (1999), Bech (2009), Heath and Emmet (1983), (Vanholder (2000), Vuure (1981) Kimber (2008) PT-AZO, PT-MAD I1 Corn, sugar cane AL, CH, DE, DK, ES, ESF5, F6, F8, CAN, FR, FR-COR, GB, IT, I1, I2 IT-SIC, PT, PT-MAD Rezbanyai-Reser (1983) Aguiar and Karsholt (2006), Karsholt and Vieira (2005) Hoffmeyer (1962), Roll et al. Polyphagous (vegetables, flowers, fruit trees, introduced (2007), Valletta (1949) with Chrysanthemum) Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Chrysodeixis chalcites (Esper, 1789) Regime 662 Family Species Stigmella pyri (Glitz, 1865) Stigmella speciosa (Frey, 1857) Stigmella suberivora (Stainton, 1869) Noctuidae Caradrina ingrata Staudinger, 1897 Family Species Earias vernana (Fabricius, 1787) Notodontidae Thaumetopoea pityocampa (Denis & Schiff., 1775) Nymphalidae Pararge aegeria (Linnaeus, 1758) Hofmannophila pseudospretella (Stainton, 1849) Native range Invaded countries Phyto- C&S Europe & SE phagous W Asia Alien Habitat G,FA Hosts Refs Populus alba Hyden et al. (2006) Phyto- S Europe & N phagous Africa IT-SAR G3 Pinus spp Mendes (1905) Phyto- Europe to E phagous Asia and N Africa PT-MAD G2,X10 Brachypodium sylvaticum (Poaceae) Aguiar and Karsholt (2006), Jones and Lace (1992) Carpets, corks of wine bottles, dried plant material, dried foodstuffs indoors. Occurs outdoors in dried bracket-fungi on trees Fabrics, including carpets, upholstery, leather and books, but more especially infesting dried foodstuffs Abafi-Aigner et al. (1896), Hrubý (1964), Martin (1991), Mehl (1977), Šefrová and Laštůvka (2005), Ulmer et al. (1918) Amsel (1959), Hill et al. (2005), Hrubý (1964), Jürivete et al. (2000), Mehl (1977), Šefrová and Laštůvka (2005) Brassica Aguiar and Karsholt (2006) Detrivorous Mediterranean? AT, BE, BY, CH, CZ, DE, J1, G DK, EE, FI, GB, IE, IS, LT, LV, NL, NO, PL, RO, SE, SK Detrivorous Mediterranean? AT, BE, BY, CH, CZ, DE, DK, EE, FI, GB, IS, IE, LT, LV, NL, NO, PL, PT, RO, SE, SK Pieridae Pieris rapae (Linnaeus, Phyto- Palaeartic and 1758) phagous N America, Australia Plutellidae PT-MAD J1, I2 X22, X23, X24, X25 Lepidoptera. Chapter 11 Oecophoridae Endrosis sarcitrella (Linnaeus, 1758) Regime 663 Regime Native range Pterophoridae Emmelina monodactyla Phyto- Europe, Africa, (Linnaeus, 1758) phagous Asia, and/or N America, Mexico Stenoptilia Phyto- Atlantic Europe millieridactylus phagous (Bruand, 1861) Pyralidae + Crambidae Aglossa caprealis Detri- Mediterranean (Hübner, 1809) vorous Apomyelois ceratoniae (Zeller, 1839) Detrivorous Mediterrranean? Cadra calidella (Guenée, 1845) Detrivorous Mediterranean SE Alien Habitat E5, I2 Hosts Refs Hesperis matronalis Gustaffson (2010) PT-AZO E, F, I2 Bindweeds (Convolvulus and Calystegia spp.), occasionally Morning glory (Ipomoea), Chenopodium and Atriplex Wild Mossy saxifrage (Saxifraga hypnoides) Karsholt and Vieira (2005) GB, IE I2 AT, BE, CZ, DE, DK, GB, NL, PL, PT-AZO, PT-MAD J1 Stored Products AT, BE, CH, CZ, DE, DK, GB, HU, NL, NO, PL, RO, RU, SE, UK AT, BE, CH, CZ, DE, DK, FI, GB, IE, NL, NO, RO, SE, SK J1 Stored products: dry fruits, dates, nuts, carob, pistachio J1 Dried fruits, nuts, figs Hill et al. (2005) Aguiar and Karsholt (2006), Buhl et al. (2007), Karsholt and Vieira (2005), Šefrová and Laštůvka (2005) Palm (1986), Sterneck and Zimmermann (1933) Hance (1991), Huemer and Rabitsch (2002), Mehl (1979), Palm (1986), Reiprich (1989), Vlach (1938) Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Phyto- Europe and phagous Asia Minor, expanding throughout N America Invaded countries 664 Family Species Plutella porrectella (Linnaeus, 1758) Family Species Duponchelia fovealis Zeller, 1847 Invaded countries Alien Habitat J100, J1 Phyto- Mediterranean phagous and Canary Islands BE, CZ, DE, DK, FI, GB, NL, NO, SE Phyto- SW Europe phagous (Spain) MT F6 Phyto- S & C Europe phagous to Asia (apparently expanding to Siberia, China and E USA) GB Hosts Refs Polyphagous in greenhouses (Begonia, Gerbera, Cyclamen, Anthurium, Kalanchoe, Poinsettia, Rosa, aquatic plants, maize, pepper and other vegetables), can reproduce outside, but surviving winters Palm trees (Phoenix canariensis and P. dactilifera) Buhl et al. (2006), Deurs (1958), Huisman and Koster (1995), Marek and Bártová (1998) B, E Grass stems used for thatching Wagner et al. (2003) Phyto- SW Europe CH phagous (Spain, France) G3 Pinus sylvestris Lepidopterologen Arbeitsgruppe (2000) Phytophagous Phytophagous GB I1, I2 Ribes Reiprich (1980) SE I2 Viburnum lantana Torstenius and Lindmark (2000) PT-MAD I2 Malus Aguiar and Karsholt (2006) Europe to W Asia Europe &/or Asia Minor to W Asia Phyto- SC Europe &/ phagous or Asia Minor and Egypt Sammut (2005) 665 Saturniidae Graellsia isabellae Graells, 1849 Sesiidae Pennisetia hylaeformis (Laspeyres, 1801) Synanthedon andrenaeformis (Laspeyres, 1801) Synanthedon myopaeformis (Borkhausen, 1789) Native range Lepidoptera. Chapter 11 Euclasta varii (Popescu-Gorj & Constantinescu, 1973) Sclerocona acutellus (Eversmann, 1842) Regime Regime Native range Invaded countries Alien Habitat Mediterranean AT, BE, CZ, DE, DK, FI, GB, LT, NL, NO, PL, RU, SE, SK Cereals Haplotinea insectella (Fabricius, 1794) Detrivorous Mediterranean AT, BE, CH, CZ, DE, DK, J1 FI, GB, IE, LT, NL, NO, PL, RU, SE, SK Tinea murariella Staudinger, 1859 Detrivorous Mediterranean? CH, ES, FR, GB, HR, IT, IT-SIC, NO, PT, PT-AZO, PT-MAD, RO J1 Stored products Trichophaga tapetzella (Linnaeus, 1758) Detrivorous J1 Mediterranean? AL, AT, BE, BG, BY, CH, CY, CZ, DE, DK, EE, FI, FR, FR-COR, GB, GR-CRE, HR, IE, IT-SAR, LU, LV, LT, NL, NO, PL, PT-AZO, SE, SI, SK, UK Stored products Stored products Refs Heath and Emmet (1985), Ivinskis (1988), Reiprich (1991), Šefrová and Laštůvka (2005) Heath and Emmet (1985), Hrubý (1964), Ivinskis and Mozūraitis (1995), Mehl (1977), Šefrová and Laštůvka (2005) Adams (1979), Gaedike and Karsholt (2001), Karsholt and Vieira (2005), Opheim and Fjeldså (1983) De Graaf (1851), Hrubý (1964), Karsholt and Vieira (2005), Lederer (1863), Palionis (1932), Robinson and Nielsen (1989), Šefrová and Laštůvka (2005) Phyto- Europe phagous PT-AZO FB Rosa Karsholt and Vieira (2005) Phyto- Europe phagous GB I1, I2 Polyphagous, fruit trees (Prunus, Malus, Rosa) and deciduous (Alnus, Betula, Populus, Salix) Bradley et al. (1973) Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Detrivorous Tortricidae Acleris variegana (Denis & Schiffermüller, 1775) Adoxophyes orana (Fischer von Röslerstamm, 1834) J1 Hosts 666 Family Species Tineidae Haplotinea ditella (Pierce & Metcalfe, 1938) Family Species Cacoecimorpha pronubana (Hübner, 1799) Regime Native range Phyto- S Europe phagous BE, CH, CZ, DE, DK, GB, HU, IE, LT, LU, NL Clavigesta sylvestrana (Curtis, 1850) Cydia grunertiana (Ratzeburg, 1868) Cydia illutana (Herrich-Schäffer, 1851) Cydia milleniana Adamczewski, 1967 Cydia pactolana (Zeller, 1840) Cydia pomonella (Linnaeus, 1758) Phyto- Europe phagous Phyto- E Europe phagous Phyto- Europe phagous PT-AZO, PT-MAD Alien Hosts Habitat FB, I2, X11, Polyphagous, especially G1, J100 on Dianthus but also on Acacia, Acer, Chrysanthemum, Citrus, Coriaria, Coronilla, Euphorbia, Ilex, Jasminum, Laurus, Mahonia, Malus, Olea, Pelargonium, Populus, Prunus, Rhododendron, Rosa, Rubus, Syringa G3 Pinus BE, DK, SE I2 Larix GB G3 Larix, Picea Phytophagous Phytophagous Phytophagous BE, DK, GB G3 Larix GB G3 Picea I1 Malus Cydia splendana (Hübner, 1799) Cydia strobilella (Linnaeus, 1758) Notocelia rosaecolana (Doubleday, 1850) Europe, PT-AZO, PT-MAD expanding to E USA Europe PT-AZO, PT-MAD Phytophagous Phyto- Europe phagous Phyto- Europe phagous G1 Castanea, Quercus but also Aguiar and Karsholt (2006), Fagus and Juglans, fruit borer Karsholt and Vieira (2005) Picea, cone borer Coldewey and Vári (1947) I2 Rosa NL IS Refs Billen (1999), de Carvalho (1995), Glavendekić et al. (2005), Ivinskis (2004), Janmoulle (1974), Thygesen (1963) Aguiar and Karsholt (2006), Karsholt and Vieira (2005) Falck and Karsholt (1993), Groenen and De Prins (2004) Hill et al. (2005) Hill et al. (2005), Buhl et al. (2004) Hill et al. (2005) Lepidoptera. Chapter 11 Europe and Asia Europe Invaded countries Aguiar and Karsholt (2006), Karsholt and Vieira (2005) 667 Yponomeutidae Argyresthia laevigatella (Heydenreich, 1851) Argyresthia trifasciata Staudinger, 1871 Prays oleae (Bernard, 1788) Zelleria oleastrella (Millière, 1864) Zygaenidae Theresimima ampellophaga (BayleBarelle, 1808) Native range Phyto- Europe phagous Invaded countries PT-AZO Phyto- Europe PT-MAD phagous expanding to N America Phyto- Mediterranean SE phagous and/or N Africa, Asia Minor Alien Habitat I2 Hosts Refs Karsholt and Vieira (2005) X15,X16 Holly (Ilex aquifolium) and blueberry (Vaccinium myrtillus) Pinus I1 Brassica Svensson (2006) Larix shoots Kimber (2008) Buhl et al. (1998), De Prins (1996), Gomboc (2003), Huemer and Rabitsch (2002), Šefrová and Laštůvka (2005) Karsholt and Vieira (2005) Aguiar and Karsholt (2006) Phyto- N or C Europe phagous &/or Japan Phyto- European Alps phagous DK, FI, GB, HU, IE, LT, LV, G3 NL, NO, SE AT, BE, CZ, DE, DK, GB, I2, FA HU, LV, NL, PL, SE, SI, SK Phyto- Mediterranean phagous Phyto- Mediterranean phagous PT-AZO I2, J100 Juniperus (not spiked species), very occasionally Cupressocyparis, Chamaecyparis Olea (240) trees GB, PT-MAD I2, J100 Olea (240) trees Aguiar and Karsholt (2006) AT I1 Vitis vinifera Huemer and Rabitsch (2002), Prinz (1907), Tarmann (1998) Phyto- Mediterranean phagous Carlos Lopez-Vaamonde et al. / BioRisk 4(2): 603–668 (2010) Rhyacionia buoliana (Denis & Schiffermüller, 1775) Selania leplastriana (Curtis, 1831) Regime 668 Family Species Rhopobota naevana (Hübner, 1817) A peer reviewed open access journal BioRisk 4(2): 669–776 (2010) doi: 10.3897/biorisk.4.55 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Hymenoptera Chapter 12 Jean-Yves Rasplus1, Claire Villemant2, Maria Rosa Paiva3, Gérard Delvare1, Alain Roques4 1 UMR Centre de Biologie et de Gestion des Populations, CBGP, (INRA/IRD/CIRAD/Montpellier SupAgro), Campus international de Baillarguet, CS 30016, 34988 Montferrier-sur Lez, France 2 UMR Origine, Structure et Evolution de la Biodiversité, OSEB, (MNHN/CNRS) CP50, Muséum National d’Histoire Naturelle 45 rue Buffon, 75005 Paris, France 3 DCEA, Faculdade de Ciências e Tecnologia, Universidade Nova de Lisboa, 2829-516 Campus de Caparica, Lisbon, Portugal 4 INRA UR633, Zoologie Forestière. Centre de recherche d’Orléans, 2163 Avenue de la Pomme de Pin, CS 40001 Ardon, 45075 Orléans Cedex 2, France Corresponding author: Jean-Yves Rasplus (rasplus@supagro.inra.fr), Claire Villemant (villeman@mnhn. fr), Maria Rosa Paiva (mrp@fct.unl.pt), Gérard Delvare (delvare@supagro.inra.fr), Alain Roques (alain. roques@orleans.inra.fr) Academic editor: David Roy | Received 31 March 2010 | Accepted 26 May 2010 | Published 6 July 2010 Citation: Rasplus J-Y et al. (2010) Hymenoptera. Chapter 12. In: Roques A et al. (Eds) Arthropod invasions in Europe. BioRisk 4(2): 669–776. doi: 10.3897/biorisk.4.55 Abstract We present the first review of Hymenoptera alien to Europe. Our study revealed that nearly 300 species of Hymenoptera belonging to 30 families have been introduced to Europe. In terms of alien species diversity within invertebrate orders, this result ranks Hymenoptera third following Coleoptera and Hemiptera. Two third of alien Hymenoptera are parasitoids or hyperparasitoids that were mostly introduced for biological control purposes. Only 35 phytophagous species, 47 predator species and 3 species of pollinators have been introduced. Six families of wasps (Aphelinidae, Encyrtidae, Eulophidae, Braconidae, Torymidae, Pteromalidae) represent together with ants (Formicidae) about 80% of the alien Hymenoptera introduced to Europe. The three most diverse families are Aphelinidae (60 species representing 32% of the Aphelinid European fauna), Encyrtidae (55) and Formicidae (42) while the Chalcidoidea together represents 2/3 of the total Hymenoptera species introduced to Europe. The first two families are associated with mealybugs, a group that also included numerous aliens to Europe. In addition, they are numerous cases of Hymenoptera introduced from one part of Europe to another, especially from continental Europe to British Islands. These introductions mostly concerned phytophagous or gall-maker species (76 %), less frequently parasitoids. The number of new records of alien Hymenoptera per year has shown an exponential increase during the last 200 years. The number of alien species introduced by year reached a maximum of 5 species per year between 1975 and 2000. North America provided the greatest part of the hymenopteran species Copyright J-Y. Rasplus et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which ermits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 670 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) alien to Europe (96 species, 35.3%), followed by Asia (84 species, 30.9%) and Africa (49 species, 18%). Three Mediterranean countries (only continental parts) hosted the largest number of alien Hymenoptera: Italy (144 spp.), France (111 spp.) and Spain (90 spp.) but no correlation was found with the area of countries. Intentional introduction, mostly for biological control, has been the main pathway of introduction for Hymenoptera. Consequently, the most invaded habitats are agricultural and horticultural as well as greenhouses. To the contrary, Hymenoptera alien in Europe are mostly associated with woodland and forest habitats. Ecological and economic impacts of alien Hymenoptera have been poorly studied. Ants have probably displaced native species and this is also true for introduced parasitoids that are suspected to displace native parasitoids by competition, but reliable examples are still scarce. The cost of these impacts has never been estimated. Keywords Hymenoptera, alien, Europe, biological invasions 12.1. Introduction Hymenoptera is one of the four large insect orders exceeding 100 000 species in the world, the other major orders being Coleoptera, Lepidoptera and Diptera (Gauld and Bolton 1988, Goulet and Huber 1993). The Hymenoptera order contains about 115 000 described species and authors estimated that there are between 300,000 and 3,000,000 species of Hymenoptera (Gaston 1991), possibly around 1,000,000 (Sharkey 2007). These estimates mean that only 1/10 has been described so far and 9/10 awaits description. However, the number of Hymenoptera species is difficult to estimate with accuracy, as most of the mega diverse regions of the world have not been extensively studied and inventoried regarding this group (LaSalle and Gauld 1993). In Europe, about 15,000 species have been reported belonging to 73 families, but undoubtedly thousands of species remains to be discovered and described. From our recent review of the literature, the alien species of Hymenoptera comprise 286 species belonging to 30 families. The order ranks third just following the Coleoptera and the Hemiptera in terms of alien species diversity (Roques et al. 2008). Additionally, 71 European species have been translocated from one part of Europe to another (adding 5 more families) and 11 species are considered cryptogenetic. All together within Europe, at least 368 Hymenoptera species have been introduced in different parts of the continent. Hymenoptera have been traditionally subdivided into three assemblages (the paraphyletic sub-order Symphyta and the monophyletic Aculeata and Parasitica belonging to the sub-order Apocrita). Each group exhibits different biology. ‘Symphyta’ are mostly phytophagous and are the most primitive members of the order. Parasitica are mainly parasitic species but some of them have returned secondarily to phytophagy, while Aculeata encompass a larger spectrum (predators, pollinators, parasitoids); all eusocial hymenoptera belong to this last group. Members of the Hymenoptera are familiar to a general audience and common names exist for a large variety of groups: “wasps”, “bees”, “ants”, “bumblebees”, “saw- Hymenoptera. Chapter 12 671 flies”, “parasitic wasps”. Hymenoptera adult sizes range from the very small Mymaridae (0.5 mm) to the large aculeate wasps (up to 5 cm long in Europe). This group of mandibulate insects is well defined by the combination of several characters: they have two pairs of functional wings (with the exception of apterous species) bearing fewer veins than most other insect groups and rarely more than seven cross veins. The abdominal tergum 1 is fused to the metanotum and in most Hymenoptera the metasoma (apparent gaster) is joined to the mesosoma (apparent thorax) by a petiole. Hymenoptera have two main larval types. ‘Symphyta’ have larvae that are caterpillar-like, but true caterpillars (Lepidoptera) have at most four pairs of prolegs (abdominal segments 3–6) while sawflies larvae have at least five pairs of prolegs (abdominal segments 2–6). Furthermore the prolegs of Symphyta do not bear crochets, whereas those of Lepidoptera larvae do. ‘Apocrita’ have legless grub-like larvae that are nearly featureless unless they have a differentiated head (Goulet and Huber 1993). All Hymenoptera have haplodiploid sex determination (haploid males and diploid females). Arrhenotoky is the most common mode of reproduction in Hymenoptera (Heimpel and de Boer 2008). The males develop parthenogenetically from unfertilised eggs while the females develop from fertilised eggs. Females can control fertilisation by releasing sperm to an egg upon oviposition, and can thus adjust the sex-ratio of their progeny. Ecologically and economically few groups of insects are as important to mankind as the Hymenoptera. Bees provide the vital ecosystem service of pollination in both natural and managed systems (Gallai et al. 2009) while parasitic Hymenoptera control populations of phytophagous insects (Tscharntke et al. 2007) and can be effective agents for control of pest insects (Bale et al. 2008, Brodeur and Boivin 2004, Jonsson et al. 2008). Some of the phytophagous hymenoptera have an intimate association with their hostplants (Nyman et al. 2006) and can also be considered as major pests to forests (e.g. Diprionidae) (De Somviele et al. 2004, Lyytikainen-Saarenmaa and Tomppo 2002). Ant invasions cause huge economic and ecological costs (Holway 2002, Lach and Thomas2008) and Hymenoptera stings, specifically those of wasps, hornets and bees cause serious allergic reactions and anaphylaxis (Flabbee et al. 2008, Klotz et al. 2009). 12.2.Taxonomy of alien species The 286 species of Hymenoptera alien to Europe belong to 30 different families (Table 12.1), which also have native representatives. Among these alien species, 35 species are phytophagous, 1 detritivorous, 3 pollinators, 47 predators whilst 200 are parasitoids or hyperparasitoids. These results show that only 13.3% of the alien wasp and bee species are phytophagous (including pollinators), the great majority of which (86.4 %) are predators and parasitoids (respectively 16.4% and 70.0%). Most parasitoids were intentionally introduced to control pests. Interestingly, among the 71 Hymenoptera that have been introduced from one part of Europe to another (aliens in Europe - Table 12.2), an opposite proportion is observed. Fifty-four species (76.0 %) are phytopha- 672 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) gous and only 17 (23.9%) are parasitic or predatory. These species have mostly followed their host plants throughout Europe. Consequently, most alien Hymenoptera in Europe belong to the sub-order Parasitica (228 spp. and 20 families, 79.4% of the species), while Aculeata (51 spp. and 7 families, 17.8%) and Symphyta (8 spp. and 3 families, 2.8%) are less represented. Six families of wasps (Aphelinidae, Encyrtidae, Eulophidae, Braconidae, Torymidae, Pteromalidae) represent together with ants (Formicidae) about 80% of the alien Hymenoptera in Europe. Each of these families has more than 10 introduced species in Europe. The three most diverse families in terms of alien species are Aphelinidae (60 species), Encyrtidae (55) and Formicidae (42). By far the richest superfamily is the Chalcidoidea that includes 198 alien species (69.2% of the total alien Hymenoptera). Below we give a short synopsis for all Hymenoptera families containing introduced species to Europe (including cryptogenic and translocated species). Suborder Symphyta Argidae. The second largest family of ‘Symphyta’ with about 1000 species described, but only 60 in Europe. Alien species to Europe have not yet been found. One species only, Arge berberidis, is considered as introduced from one part of Europe to another,. Females deposit eggs in leafs of various angiosperms and the larvae are phytophagous, feeding mostly on woody plants (Salicaceae, Rosaceae, Betulaceae). Blasticotomidae. This is a very small family represented by one species only, Blasticotoma filiceti, in northern and central Europe. Larvae are stem borers, developing within the rachis of ferns (e.g., Athyrium filix-femina (L.) Roth) (Schedl 1974). B. filiceti has been infrequently introduced into Great Britain from continental Europe, mostly with horticultural plants. Diprionidae. A small family of ‘Symphyta’ that mostly occurs in northern Europe. It comprises about 100 species in the northern hemisphere, of which 20 occur in Europe. The larvae attack softwood trees (e.g. conifers) and are considered as major pests in forestry. Diprioninae develop on Pinaceae and Monocteninae on Cupressaceae, but only the first subfamily contains invaders. Alien species have not yet been recorded. However, five species are considered as alien in Europe. Neodiprion sertifer and Gilpinia hercyniae cause severe damage to pine and spruce plantations. Females of some species produce pheromones that attract males. The larvae consume needles, sometimes gregariously, and when mature drop to the ground, pupate and overwinter within a cocoon (rarely upon trees). Diapause can last for more than one winter (Pschorn Walcher 1991), the wasps emerging and dispersing in the early spring. Pamphiliidae. A small holarctic family containing about 60 species in Europe (van Achterberg and van Aartsen 1986, Viitasaari 2002). Only Cephalcia alashanica is an alien species introduced from temperate Asia. Six other species are alien in Europe, most of them having been introduced from the Alps to northern countries with their host trees. Some species attack conifers and are considered as forest pests. Females lay eggs Hymenoptera. Chapter 12 673 in a slit cut in a needle, the normally gregarious larvae either spin silk webs in which they develop (Cephalciinae) or roll the host plant leaves (Pamphiliinae). They overwinter as pupae within pupal chambers in the soil and adults emerge in early spring. Siricidae. A small Holarctic family (16 European species) of large and conspicuous wasps (woodwasps). Nine species are considered as alien in Europe, with only 5 alien species introduced from North America with imported timbers. The family is subdivided into two subfamilies, the Siricinae attacking conifers and the Tremecinae that attack angiosperm trees. The females, which do not feed, oviposit in recently fallen or dying trees and introduce spores of symbiotic fungus along with the eggs. The larvae develop in 2 or 4 years as woodborers and pupate in the bark. Tenthredinidae. This cosmopolitan family is the most diverse group of ‘Symphyta’ including 1050 species in Europe of which only two are alien to Europe , Nematus (Pteronidea) tibialis (a pest of black locust) and Pachynematus (Larinematus) itoi (a larch pest) and 23 alien in Europe. Some native European species are also considered serious pests in North America where they have been introduced. All species are phytophagous and the larvae are mostly external feeders on diverse species of angiosperms and conifers. The females embed their eggs in the tissue of the plant, using their ovipositor as a saw. The larvae feed singly on leaves, or are stem borers, gall makers or leaf miners. Tenthredinidae mostly overwinter as prepupae in the ground, sometimes as mature larvae or eggs, the adults emerge relatively early in the spring. Suborder Apocrita Parasitica Chalcidoidea Agaonidae. A small-sized family with only 6 species of wasps reported in Europe, four of which are introduced from tropical Asia, along with two ornamental trees Ficus microcarpa L.f. and F. religiosa L. Agaonidae are the pollinators of fig trees and are mutualistically associated with their host plant. Several groups of non-pollinating fig wasps are associated with figs, either as gall-makers, inquilines or parasitoids. Their taxonomic position has been discussed and they are here grouped within Agaonidae for convenience (Bouček 1988, Rasplus et al. 1998). Aphelinidae. This is a moderately sized family of wasps represented in Europe by less than 200 species of which sixty are aliens. Aphelinidae species have been introduced from diverse geographic areas as biological control agents. Along with encyrtid, the Aphelinidae is the most important family for biological control. Species are primarily endoparasitoids or ectoparasitoids, sometimes hyperparasitoids, of sternorrhynchous Hemiptera (mostly Aphidoidea, Coccoidea or Aleyrodoidea). Some species may have complicated ontogeny (Hunter and Woolley 2001) and males and females may attack different hosts either as parasitoids or hyperparasitoids. Chalcididae. A small family of chalcid wasps comprising about 80 species in Europe, including one alien species, introduced from North Africa to control fruit flies. 674 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) The hosts of these obligate parasitoids or hyperparasitoids are mostly Lepidoptera and Diptera, less frequently Coleoptera, Neuroptera or Hymenoptera (Delvare 1995, Delvare 2006). The females lay eggs within the host larva and the pupation take place in the host pupa. Encyrtidae. A large family of wasps represented by more than 700 species in Europe (Trjapitzin 1989), of which 55 are considered to be alien, introduced from different parts of the world for biological control of economically important pests (Noyes and Hayat 1994). Most of the Encyrtidae are endoparasitoids of scale insects. Some species also develop as endoparasitoids of other insect orders, mostly Lepidoptera, Coleoptera and Hymenoptera). The egg is laid inside the host and the larva develop as a parasitoid sometimes as an hyperparasitoid, and pupates within the host. Eulophidae. A large family of wasps that contains 1100 species in Europe (Gauld and Bolton 1988), including 29 alien species. Most alien species have been introduced for biological control but a few (3) are gall makers that develop at the expense of plant tissue of Eucalyptus (Branco et al. 2009). Eulophid are primarily solitary parasitoids of eggs, pupae or larvae of various endophagous insects (Diptera, Coleoptera, Thysanoptera, Lepidoptera or Hymenoptera). Some species attack economically important leaf miners or gall makers (e.g. Agromyzidae, Cecidomyiidae). Eupelmidae. A small family represented by about 100 native (Gibson 1995) and 5 alien species in Europe (Eupelmus and Anastatus spp.). Eupelmidae are primarily ectoparasitoids (idiobionts) of egg or larval stages of various insects and spiders (Askew et al. 2000). Some species within this family are generalist parasitoids. Eurytomidae. A medium-sized family with about 300 species in Europe (Zerova 1978), of which seven are alien. Interestingly, these alien species are not parasitoids but phytophagous and pests of crops or horticultural plants whilst most eurytomids are primarily ectoparasitoids or hyperparasitoids of extremely diverse groups of endophagous insects (Lotfalizadeh et al. 2007). Phytophagous species are either stem-borers or seed-feeders or gall-makers on different host-plant groups (e.g. Graminaceae, Leguminosae). Some species are both entomophagous then phytophagous during their larval development. Mymaridae. A medium-sized family including about 450 species in Europe, of which only two are alien, Anaphes nitens and Polynema striaticorne. All mymarids are internal, solitary (rarely gregarious) parasitoids of the eggs of various insects (Huber 1986). The most common hosts are eggs of Hemiptera Auchenorrhyncha (Cicallidae, Cixiidae) but mymarids also parasitize eggs of other insects (Coleoptera, Hemiptera). Female oviposit within concealed eggs, and there are 2 to 4 larval stages. Perilampidae. A small family of chalcid wasps that includes 40 European species. The only alien species in this family (Steffanolampus) originates from North America and is a parasitoid of wood-boring Coleoptera. Most perilampids are hyperparasitoids of Lepidoptera through Tachinidae (Diptera) or Ichneumonoidea (Steffan 1952). Females deposit their eggs away from the host, however the young larvae (planidium) are mobile, and may either attach themselves to the primary host, at any stage of larval development, or enter the host to attach to its endopara- Hymenoptera. Chapter 12 675 sitoids. In some species, an adult host carries the larva to a suitable location where host larvae occur (Darling 1999). Pteromalidae. A large, paraphyletic family including more than 1100 species in Europe (Graham 1969). Only ten are considered alien species, most of which were unintentionally introduced with their hosts, some (3) for biological control purposes. The diversity of the group is reflected by the diversity of the biology exhibited. Pteromalids are mostly ectoparasitoid idiobionts, but some species are koinobionts. Miscogasterinae are larvo-pupal endoparasitoids of dipteran leaf miners. Eunotinae (e.g. Moranila) are predators on Coccoidea eggs within the female body (Boucek and Rasplus 1991). Signiphoridae. A small family of tiny chalcids (0.5–2 mm) comprising only 8 European species, one of which is an introduced hyperparasitoid (Chartocerus) (Woolley 1988). Signiphoridae are known as parasitoids (sometimes hyperparasitoids) of cyclorrhaphous dipterans, scale-insects (Coccoidea) or white-flies (Aleyrodidae). Torymidae. A medium-sized family that includes about 350 European species (Grissell 1995, Grissell 1999), of which 13 are considered as alien to Europe. Most of the alien species (12) belong to the genus Megastigmus and are considered pest of conifer seeds (Roques and Skrzypczynska 2003). Most torymines are idiobiont ectoparasitoids of gall-makers (Cynipidae and Cecidomyiidae) and other endophytic insects but most Megastigminae are specialist phytophages. Megastigmus females lay their eggs in the ovules of conifers before fertilization has taken place (Roques and Skrzypczynska 2003) (Figure 12.9). Megastigmus biological habits have been shown to be particularly prone to invasion. Since most of their development takes place within seed, their presence is usually overlooked in traded seed lots, the infested seeds showing up only when X-rayed (Figure 12.10). In addition, insect are able to become dormant during the larval stage, for up to 5 years (prolonged diapause) following the annual size variations of the seed crop, thus broadening the chances that adult emergence will occur under favourable circumstances near a suitable new host. Moreover, some species such as the Douglas-fir seed chalcid, M. spermotrophus, appear capable of preventing the abortion of unfertilized seeds. The invasive insect larva may thus achieve its development in unpollinated, unfertilized seeds by altering the physiology of the ovule so that it allocates de novo resources to the larva (von Aderkas et al. 2005). Trichogrammatidae. A moderately-sized family containing about 150 European species. The nine alien species belong mostly to three genera: Trichogramma, Oligosota, Uscana and have been introduced to Europe for the control of agricultural pests (Lepidoptera and Coleoptera) (Pintureau 2008). Trichogrammatids are primarily solitary or gregarious endoparasitoids of insect eggs (mostly Lepidoptera, Hemiptera, Coleoptera) and can sometimes develop as hyperparasitoids. Ichneumonoidea Ichneumonidae. This is the first megadiverse Apocrita family in Europe with about 5500 species, six of them are considered as alien to Europe. These species have been in- 676 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) tentionally introduced for biological control. The family is divided into more than 30 subfamilies. Consequently, the biology of ichneumonids is extremely diverse. Ichneumonids mostly parasitize the immature stages of the Holometabola, and are frequently associated with Lepidoptera and sawflies (Hymenoptera). Ectoparasitism is considered the primitive condition and endoparasitism has evolved several times independently within the family. Braconidae. Braconids represent the second megadiverse family with nearly 3500 European species, 16 of which are considered as alien. Altogether, Ichneumonoidea may account for nearly 10000 species in Europe. Like ichneumonids, braconids exhibit a large range of biological characteristics. They are mostly parasitoids of other insects. Some of the braconid groups are larvo-nymphal koinobiont parasitoids; others are idiobiont ectoparasitoids. Introduced species are mostly koinobiont endoparasitoids and are associated with aphids (Aphidiinae), moths (Miscogasterinae), and fruit flies (Opiinae). Ceraphronoidea Ceraphronidae. A small family represented by 100 European species, only one of which is considered as alien, Aphanogmus bicolor. Their biology is poorly known but some species are endoparasitoids of nematocerous dipterans whilst others attack Thysanoptera or Neuroptera. Some species are considered as antagonists of biological control agents since they are parasitoids of predaceous midges or hymenopteran primary parasitoids. Cynipoidea Cynipidae. A medium-sized family confined to the Holarctic and containing 350 European species. Only the chesnut gall wasp, Dryocosmus kuriphilus, is alien to Europe (Figure 10.8). Six more species, mostly from the genus Andricus, are considered as aliens in Europe. Most Cynipinae are gall inducers on Quercus, Rosa and some Compositae but others (Synergini) are inquilines. Figitidae. This medium-sized family contains ca. 400 species in Europe, the family as presently understood includes the previous Eucoilidae, Charipidae and Anacharitidae (Ronquist 1995). Only one species (Aganaspis daci) is considered as alien and has been introduced to Europe for the control of fruitflies. Figitid larvae develop as internal parasitoids of other endophytic insect larvae. The hosts are mostly dipteran larvae but Charipinae Alloxystini are hyperparasitoids of aphids through Braconidae Aphidiinae and Aphelinidae. The egg is deposited inside a young host larva, which continues to develop normally (koinobionts), the parasitoid larvae emerges before the host death and can achieve its development as an ectoparasitoid. Hymenoptera. Chapter 12 677 Platygastroidea Platygastridae. A medium-sized family with about 500 species in Europe but only two (Amitus spp.) are considered as alien, having been introduced into Europe for the control of whiteflies. Many Platygastridae are endoparasitoids of gall-making dipterans whilst others attack immature hemipterans or ant larvae. The biology of most species remains largely unknown. Some species are thelytokous and very few polyembryonnic. The larvae have an uncommon appearance and superficially resemble cyclopoid copepods. Scelionidae. A medium-sized family that includes about 600 species in Europe, three of them considered as alien. Scelionids are primarily endoparasitoids in a wide variety of insect eggs (few on other arthropods), more rarely hyperparasitoids. Introduced species attack Hemiptera or Lepidoptera eggs and have been used for pest control. The family has been synonymized with Platygastridae but we still keep it apart for consistency (Murphy et al. 2007). Suborder Apocrita Aculeata Chrysidoidea Bethylidae. A medium-sized family represented by about 230 species in Europe. Four species are considered alien. Cephalonomia waterstoni, Holepyris sylvanidis and Plastanoxus laevis are cosmopolitan. They were introduced into Europe with stored products. Laelius utilis is a parasitoid of Anthrenus. Bethylidae mainly attack larvae of Lepidoptera and Coleoptera. The female stings and paralyses the host, and then lays several eggs on its skin. Larvae develop as ectoparasitoids. For a few species, females tend the eggs and developing larvae. Pupation occurs next to the host remains. Chrysididae. A medium- sized family that comprises 420 European species. Cukoo- wasps are parasitoids or kleptoparasitoids of Aculeate wasps. The nests of the host are sought out by the female chrysid that oviposits into the host cells. A true parasitoid larva develops as an ectoparasitoid on the host larva whilst a kleptoparasite larva kills the egg or the young larva of the host before consuming the stored food. One East European species introduced in western parts of Europe, Chrysis marginata, is considered as alien in Europe (Pagliano et al. 2000). Dryinidae. A medium-sized family that comprises about 100 species in Europe. All dryinids are parasitoids of immature and adult Hemiptera Auchenorrhyncha. The larva is rather endoparasitoid than ectoparasitoid during the last instars, forming a bag (thylacium) constituted by the exuviae of the parasitoid and bulging from the host abdomen. Only one species alien to Europe, Neodryinus typhlocybae, was introduced in northern Italy and subsequently in France for biological control of the Nearctic planthopper Metcalfa pruinosa (Hemiptera, Flatidae) (Malausa et al. 2003, Malausa et al. 2008). 678 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Apoidea Apoidea represents a superfamily including more than 2000 species in Europe. Depending on the classification used, the group comprises seven families (ancient subfamilies of the single family Apidae) to eleven families if sphecid wasps, the sister group of bees, are included (Sharkey 2007). Here we followed the more recent classification system and adopted a subdivision into several families. Bees are flower visitors and efficient pollinators of angiosperms. Their larvae are phytophagous and develop on a mixture of pollen and nectars. Bees are now recognized as an important group of ecosystem engineers that modulate resources availability (i.e. plants) to other organisms (Jones et al. 1994). Two families of bees contain alien species in Europe. Sphecid wasps comprise 4 families of wasps that feed their progeny with a wide range of preys (mainly insects or spiders), depending on genera. All alien species belong to the family Sphecidae. Apidae. This small family of eusocial bees includes social species, with colonies attaining large sizes. It comprises less than 70 species in Europe, all except one (Apis meliffera) belonging to the genus Bombus. Some of these pollinator species have been introduced from some parts of Europe into other European regions for crop pollination purposes and honey production. Megachilidae. This family comprises about 480 species in Europe, two are considered as alien. The alfalfa leafcutter bee, Megachile rotundata, is a west European species that has been used commercially for pollination of alfalfa, and introduced in Russia. Osmia cornifrons is an alien species that has been introduced from Japan into Denmark for pollination of fruit trees. Megachilidae nest in burrows in soil or in pithy stems. A few species build stony mud nests. Cells of Megachilidae are made of foreign materials (leaf pieces for Megachile species) brought into the nest. Sphecidae. This family in its narrow sense comprises about 70 species, four of which are alien species accidentally introduced into Western Europe from North America (Sceliphron caementarium and Isodontia mexicana) or from Asia (S. curvatum and S. deforme). Adults of most species (e.g., Isodontia) prey on orthopteroids but some of them, such as Sceliphron spp., catch Araneae. While S. deforme has possibly not established in the Balkans, both other species became established and threaten autochtonous species of Sceliphron (Cetkovic et al. 2004). While Isodontia puts its preys in pre-existing cavities, Sceliphron are mud-daubers that often built their nests in or around buildings (Bitsch and Barbier 2006, Bitsch et al. 1997). Vespoidea Formicidae. This family includes about 650 species in Europe, 42 of which are alien to Europe, one is cryptogenetic and seven are European species introduced into other areas of Europe. Ants exhibit a remarkable range of life histories. They have colonized most habitats and form colonies of variable sizes in the soil, plant debris, trees and infrastructures of human origin. The nest contains one to several reproductive females as well as workers and broods. Males are produced seasonally. Mating usually takes Hymenoptera. Chapter 12 679 place outside the nest but may occur inside the nest. In Europe, the argentine ant Linepithema humile (Mayr) is extremely abundant throughout the Mediterranean basin, causing economic damage by fostering some hemipteran pests and upsetting the action of natural enemies; However, it may occasionally act as a beneficial natural enemy in forest ecosystems (Way et al. 1997). Vespidae. This medium-sized family comprises 300 species in Europe classified into four subfamilies: Masarinae, Eumeninae, Polistinae and Vespinae (22 species). Vespinae are social wasps that built aerial or subterranean nests made of carton and composed of several combs protected by an envelope. Recently, a hornet species alien to Europe, Vespa velutina nigrithorax, was accidentally introduced from Asia into southern France (Haxaire et al. 2006, Villemant et al. 2006) (Figure 10.11). The European yellowjackets, Vespula germanica (Fabricius, 1793) and V. vulgaris (Linné, 1758) were introduced to Iceland from continental Europe, the last into Feroe Islands (Olafsson 1979). For nine families the number of alien species exceeds 5% of the species known in Europe (Figure 12.1). Four of these families are small (Agaonidae, Signiphoridae, Siricidae and Sphecidae) and consequently the number of alien species is marginal. However Aphelinidae, Encyrtidae, Trichogrammatidae and Formicidae are mediumsized families comprising between 150 and 700 species and consequently the number of alien taxa is relatively important. Interestingly, the number of alien Aphelinids introduced into Europe for biological control represents about one third of the specific diversity of the family in Europe. Aphelinidae, Encyrtidae and Trichogrammatidae, three families largely used for biological control, rank among the top five in terms of proportion of alien species in the European fauna. Aphelinidae and Encyrtidae are mostly biological control agents of the three mealybug families that include most of the pest species alien to Europe (Diaspididae, Pseudococcidae and Coccidae; see Chapter 9.3). Finally, Formicidae also include a large proportion of alien species to Europe and represent a major group of alien species to Europe. 12.3.Temporal trends First records in Europe are known for 262 of the 286 hymenopteran species alien to Europe (92%). Dates given here are relatively imprecise, as most species may have been introduced two to five years before they were reported. Furthermore, we did not try to check all literature and collections in order to report the dates of first interception within Europe. The number of new records per time period shows an exponential increase in the number of alien Hymenoptera to Europe during the last 200 years (Figure 12.2). The mean number of new records of alien hymenoptera varies from less than one species per year during the period (1800–1924) to about 5 species per year between 1975 and 2000. Interestingly, we observed a decrease in the number of Hymenoptera reported during the last 10 years. This overall increase in the number of introduced species also corresponded to an increase in the number of hymenopteran families newly found in Europe. 680 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Figure 12.1. Taxonomic overview of the alien Hymenoptera. Right- Relative importance of the hymenopteran families in the alien entomofauna. Families are presented in a decreasing order based on the number of alien species. Species alien to Europe include cryptogenic species. The number over each bar indicates the number of alien species observed per family. Left- Percentage of aliens vs. total species in each Hymenoptera family in Europe. The number over each bar indicates the total number of species observed per family in Europe. From 1800 to 1924 (125 years) only 35 species, representing 8 families, of alien hymenoptera were reported in Europe. Most of them are biological control agents or ants. Only one species of chalcid wasp (furthermore a hyperparasitoid) is reported from that period while Chalcidoidea is the most diverse group of alien Hymenoptera. However, during that period of time the European fauna was still poorly known and little studied (which is still the case for the majority of families) and the number of alien species is likely to have been underestimated. Nevertheless, over 1/3 of the alien ant species presently known in Europe were introduced between 1847 and 1929. About 79% of the alien Hymenoptera were introduced in Europe in the last 60 years. During that period of time, 61.5% of the phytophagous alien and only 38.3% of the predator alien were introduced into Europe. Among the three most diverse families of alien Hymenoptera (namely Formicidae, Aphelinidae and Encyrtidae), Formicidae exhibited a relatively stable pattern, regarding the number of introductions per year over time, varying between 0.08 and 0.36, with a maximum of introductions during the periods 1925–1949 and 1975–1999 (Figure 12. 3). Aphelinids and encyrtids both show a relatively similar pattern, but somewhat different to the pattern exhibited by ants. These two families, largely used in biological control, showed a peak of introduc- Hymenoptera. Chapter 12 681 Figure 12.2. Temporal trend in number of alien Hymenoptera to Europe per period of 25 years from 1492 to 2006. Cryptogenic species excluded. The number above the bar indicates the number of species introduced. tions during the period 1950–1999 (between 0.52 and 1.32 species per year), which roughly corresponds to the ‘golden years’ of biological control. More specifically, our analysis showed that 77.5% of the total number of parasitoids alien to Europe were introduced between 1950 and 1999. In the last 10 years, the rate of introduction drops to less than 0.1 species per year. This trend is probably due to both the decreasing interest in research on biological control and to the growing concern over possible nontarget effects of biological control. 12.4. Biogeographic patterns Origin of alien species We could ascertain a region of origin for 272 (95.1%) alien wasp species introduced to Europe. Overall there are no major difficulties in identifying the areas of origin of these wasps. The distribution of the genera of the hosts or the plant-hosts and also the origin of the taxonomists describing these species provide evidence of likely origins. However, for subsequent spread within Europe it is difficult, without genetic analyses, to separate spreading from adjacent countries from independent colonization events. 682 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Figure 12.3. Rates of introduction of the three most diverse families of invasive Hymenoptera during the two last centuries. North America provided the greatest part of alien Hymenoptera occurring in Europe (96 species, 35.3%), followed by Asia (84, 30.9%) and Africa (49, 18%) (Figure 12.4). This pattern is similar to the one found for Diptera (see Chapter 10) but differs from that observed in most other insect groups. Whatever the main areas of origin, trends of introduction are similar over time, and there is no evidence of a change in the origin of alien species through time (Figure 12.5). The only difference seemed to be a decrease of the afro-tropical species in the last 30 years, whereas rates of introduction still increased for both North America and Asia. However it must be noted that origins of alien species can differ from one country to another and general trends are not supported in all countries. Israel for example received more species from Asia and Africa than from North America (Roll et al. 2007). Interestingly, the composition of the introduced guilds originating from different continents differed taxonomically. The alien guilds introduced from North America contains several phytophagous species (Siricidae, Torymidae, Eurytomidae) and several species of Ichneumonoidea that are absent from oriental invader guilds. Overall, phytophagous aliens mostly originate from North America and temperate Asia. This is the case for xylophagous Siricidae, most Megastigmus seed-feeders (Torymidae), several Eurytomid species. Introduced plants (e.g. Ficus and Eucalyptus) came into Europe with species of their phytophagous guilds (Agaonid and Eulophidae gall-makers). Alien Formicidae originates from Africa (10 species), Asia (14) and South America (7) while only two were introduced from North America. South American ants mostly originated from areas with Mediterranean-like climate. Parasitoid wasps originated from all continents with no particular trends. Hymenoptera. Chapter 12 683 Figure 12.4. Origin of the 286 alien species of Hymenoptera established in Europe. Distribution of alien species in Europe Alien Hymenoptera species and families are not evenly distributed throughout Europe and large differences exist between countries (Figure 12.6, Table 12.3). However, results might have been influenced by large variations in the number of taxonomists involved, as well as by the intensity of the studies and of the samplings conducted in different regions. Little information is available for some countries of central and north-eastern Europe and consequently these areas appear to host comparatively few alien species of Hymenoptera. Continental Italy hosts the largest number of alien Hymenoptera (144 spp.), followed by continental France (111 spp.) and continental Spain (90 spp.). Bosnia, Andorra and Latvia are the countries from which the lowest number of invasive Hymenoptera has been reported so far, with only one alien species. No correlation with the country surface area has been found but there is a latitudinal trend of decreasing number of alien species to Europe from southern to northern Europe As most of the alien hymenopterans are biological control agents, they were mostly introduced in one or few countries by national research projects that attempted to control target pest. Large-scale European projects for biological control are rare and consequently wasps have been introduced on a local scale. About 150 alien species (i.e., more than 50% of the total species) have been reported from only one or two countries. In contrast, 31 species are reported from at least 10 countries, among them 13 of the 36 species were introduced before 1924. These aliens mostly belong to the three diverse families of alien Hymenoptera (namely Aphelinidae, Encyrtidae and Formicidae). Most of these widespread alien wasps were parasitoids introduced for biological control. For example, Aphelinus mali against the woolly apple aphid, Eriosoma lanigerum (Hausmann); Aphidius colemani and A. smithi as generalist 684 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Figure 12.5. Evolution of the rate of alien Hymenoptera from different origin through time. parasitoids used against several species of pest aphids, i.e., Acyrthosiphon pisum (Harris), Aphis gossypii Glover and Myzus persicae (Sulzer); Cales noacki against the aleyrodid Aleurothrixus floccosus (Maskell), a pest on Citrus; Encarsia formosa mostly as a biological control agent of greenhouse whitefly, Trialeurodes vaporariorum (Westwood); Leptomastix dactylopii Howard against Planococcus citri (Risso); Aphytis mytilaspidis as a parasitoid of the oystershell scale, Lepidosaphes ulmi (L.), and some other diaspidid scales; Eretmocerus eremicus as a parasitoid of the Bemisia complex (Hemiptera, Aleyrodidae) in the native range; and, Mesopolobus spermotrophus against the seed chalcid pest Megastigmus spermotrophus. Only three of the widespread alien Hymenoptera are phytophagous and were introduced during the 19th century (Megastigmus spermotrophus, Nematus tibialis, Sirex cyaneus). Seven species of Formicidae appear widely distributed in Europe: Hypoponera punctatissima (31 countries), Lasius neglectus (10), L. turcicus (15), Linepithema humile (17), Monomorium pharaonis (23), Paratrechina longicornis (13), Pheidole megacephala (14) 12.5. Main pathways to Europe Intentional introductions represent a large proportion of the introduced species in Europe (180 of 286, 63%) and this is mostly due to the high number of introduced Hymenoptera. Chapter 12 685 Figure 12.6. Colonization of continental European countries and main European islands by hymenopteran species alien to Europe. Archipelagos: 1 Azores 2 Madeira 3 Canary Islands. biological control agents. Among the 106 species clearly accidentally introduced in Europe, 32 (30.1%) are phytophagous species, only 24 (22.6%) parasitoids or hyperparasitoids that were sometimes unintentionally introduced with their parasitic hosts although the real status of some of these parasitoids is difficult to ascertain, while the majority (47 species; i.e., 44.3%), are social Hymenoptera and Sphecidae. Several species are cryptogenic and represent ancient introductions in Europe, mostly with stored products. Identifying the origin of accidental introductions is not easy but clearly introductions of plants for planting (e.g. cultivated conifers, ornamental trees) and plant seeds appeared to be the main pathways of introduction for phytophagous Hymenoptera. Thus, the lack of regulatory measures for seed imports in Europe probably resulted in the repeated establishment of alien species of Megastigmus seed chalcids since the beginning of the 20th century. Aliens presently represent 43% of the total fauna of tree seed chalcids in Europe (Roques and Skrzypczynska 2003). The development of trade in plant material through the Internet is likely to increase 686 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) this process because there is less control, especially for tree seeds which can be moved quite freely all over the world. 12.6. Most invaded ecosystems and habitats Most of the habitats colonized by Hymenoptera alien to Europe correspond to habitats strongly modified by humans (Figure 12.7). About half of the species occur in agricultural and horticultural habitats and this proportion reaches 2/3 of the species if greenhouses are considered. Only 20% of the aliens to Europe occur in woodland and forest habitats. However, the proportion is reversed if we consider Hymenoptera alien in Europe; in this case, half of the translocated species are phytophagous pests of trees. 12.7. Ecological and economic impact The ecological impacts of alien invertebrate species have been recently reviewed by Kenis et al. (2009) and Hymenoptera represent well all impact categories described in this review. Biological control programmes against pests, using introduced parasitoids, were initiated in Europe about 100 years ago. These programs using relatively hostspecific parasitoids are long supposed to decrease the risk to nontarget species, however there is increasing concern about the ecological costs of biological control (Louda et al. 2003, Simberloff and Stiling 1996). All introduced natural enemies present a certain Figure 12.7. Main European habitats colonized by the species of Hymenoptera alien to Europe and alien in Europe. The number over each bar indicates the absolute number of alien hymenopterans recorded per habitat. Note that a species may have colonized several habitats. Hymenoptera. Chapter 12 687 degree of risk to non-target species and there is clear evidence of non-target effects (Lynch and Thomas 2000). Indeed, some butterfly populations have suffered a range reduction likely due to parasitism from an introduced wasp (Benson et al. 2003a, Benson et al. 2003b). Recently, Babendreier et al. (2003) found in laboratory experiments that Trichogramma brassicae (a parasitoid largely used against Ostrinia nubilalis (Hübner) on maize) parasitizes eggs of 22 out of 23 lepidopteran species tested, including several which are listed on the Swiss red list of endangered species. Because researchers have not looked systematically for non-target effects, they are probably underestimated in Europe. Biological control is potentially a valuable control strategy against invasions of alien insect pest species in agricultural and forest ecosystems. Nevertheless, postrelease monitoring of biological control agents on target and nontarget species has yet to be developed. This is an ethical responsibility of scientists (Delfosse 2005) and it could help to resolve uncertainties in the impact of releases. One of the most pernicious effects of introduced ants is the elimination or displacement of native ants and potential cascading effects on other trophic levels. Indeed, invasive ant species have huge colonies that exploit local resources and therefore represent a considerable threat to native ants. This ecological advantage of invasive ant species is partly attributed to their unicoloniality that promotes high worker densities and to the presence of several queens that accelerate colony growth and propagation Figure 12.8. Chestnut gall induced by the chestnut gall wasp, Dryocosmus kuriphilus (Credit: Milka Glavendekić). 688 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Figure 12.9. Female of cedar seed chalcid, Megastigmus schimitscheki, ovipositing on a cedar cone. (Credit: Gaëlle Rouault). (Giraud et al. 2002), sometimes coupled with diet plasticity allowing them to exploit human residues. Introduced alien parasitoids have also been suspected to displace native parasitoids by competition; however, reliable examples are still rare. One reported case in Europe is the probable displacement of Encarsia margaritiventris (Mercet), a parasitoid of the whiteflies Aleurotuba jelineki (Frauenfeld) following the introduction of Cales noacki (Viggiani 1994b). There is still debate about the extent to which an introduced bee could alter native pollinator communities. Some studies clearly show that introduction of non-native bees may have strong impacts on local communities of bees (Goulson 2003), but their effects have been poorly documented in Europe. However, it is important to keep in mind that generalist polylectic bees (i.e. Apis, Bombus) may compete with native flower visitors (bees, wasps, butterflies, moths, beetles and flies) (Ings et al. 2006), as well as competing for nest sites. There is also evidence that introduced bees could bear pathogenic, commensal and mutualistic organisms, that could be co-introduced and transmitted to native Apidae (Goka et al. 2001). Exotic bees could also disrupt native pollinator services and could be the only pollinators of weeds, improving their seed set and spread. Genetic impacts of Hymenoptera are clearly underestimated and there is strong risk that introduced species may hybridize with localy adapted populations. This case has been reported for Bombus and Apis, and there is a strong risk that commercial and native subspecies will hybridize with alien ones (Goulson 2003, Ings et al. 2005, Hymenoptera. Chapter 12 689 Figure 12.10. X-ray picture of Douglas fir seeds showing seeds infested by larvae and pupae of the Douglas-fir seed chalcid, Megastigmus spermotrophus (Credit: Jean-Paul Raimbault). Figure 12.11. Nest of Asian Hornet, Vespa velutina nigrothorax (Credit: Claire Villemant) 690 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Kanbe et al. 2008). Introduction of Mediterranean subspecies of Apis mellifera, A. m. carnica and A.m. ligustica, in northern Europe has led to extended gene flow and introgression between these subspecies and the native black honeybee, A. m. mellifera in different parts of Europe (De La Rùa et al. 2002, Jensen et al. 2005). Introduced phytophagous Hymenoptera may also have strong economic and ecological impact. During mass-outbreaks they defoliate trees, reduce their growth and lead, sometimes, to their death. This is the case for diprionid outbreaks (De Somviele et al. 2004, Lyytikainen-Saarenmaa and Tomppo 2002) as well as for xylophagous siricids that threaten pine plantations (Yemshanov et al. 2009). Economic impacts of alien Hymenoptera have received little attention In Europe and consequently are clearly underestimated. However introduced alien ant species account for over $120 billion of annual costs in the United States alone (Gutrich et al. 2007, Pimentel et al. 2000, Pimentel et al. 2005, Vis and Lenteren 2008). Introduced siricids in the United States are considered as an economically serious threat with a total projected loss of more than $ 0.76 billion over 30 years (Yemshanov et al. 2009). The recent introduction in France of Vespa velutina would also have a significant impact on beekeeping because this hornet mainly preys on honeybees (see factsheet 14.62). 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Cotonello degli agrumi (Pseudococcus citri (Risso)) del isola di Procida mediante l’impiego di du parassiti esotica, Pauridia peregrina Timb. e Leptomastix dactylopii How. Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri 18: 257–284 Zinna G (1961) Specializzazione entomoparassiti negli Aphelinidae: studio morfologico, etologico e fisiologico del Coccophagus bivittatus Compere, nuovo parassita del Coccus hesperidum L. per l’Italia. Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri 19: 302–358. Table 12.1. Hymenoptera species alien to Europe. List and characteristics. Status: A: Alien to Europe; C: cryptogenic species. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Last update 01/03/2010 Status A phytophagous Asia 1968, IL IL, IT I2, G Ficus Eupristina verticillata Waterston, 1921 Josephiella microcarpae Beardsley & Rasplus, 2001 Odontofroggatia galili Wiebes, 1980 Aphelinidae Ablerus chionaspidis (Howard, 1914) A phytophagous phytophagous Asia 1991, ESCAN 1997, ESCAN ES-CAN, IT, IT-SIC I2, G Ficus ES-CAN, IT, IT-SIC I2, G Gall maker on Ficus leaves A phytophagous Asia 1979, GRSEG GR-SEG, IL, IT, IT-SIC I2, G Ficus A parasitic/ predator Asia 1972, IT ES, IL, IT, RS, G4 Ablerus clisiocampae (Ashmead, 1894) A parasitic/ predator Asia 1953, FR FR, IT G4 Ablerus perspeciosus Girault, 1916 A parasitic/ predator Asia 1972, FR FR, IL, IT, RS, YU G3, G4 Diaspidid scale insects (Hyperparasitoid and parasitoid) Diaspidid scale insects and lepidopteran eggs (Hyperparasitoid and parasitoid both of ) White peach scale, Pseudaulacaspis pentagona (parasite) A Regime Native range Asia First Record in Europe Invaded countries Habitat Host References Koponen and Askew (2002), Lo Verde et al. (1991) Beardsley and Rasplus (2001), Lo Verde (2002) Compton (1989), Lo Verde et al. (1991), Wiebes (1980) Galil and Eisikowitch (1968) Herting (1972), Herting (1977), Ofek et al. (1997) Peck (1963), Yasnosh (1978) Hymenoptera. Chapter 12 Families Species Agaonidae Platyscapa quadraticeps (Mayr, 1885) Battaglia et al. (1994), Herting (1972), Kozarazhevskaya and Mihajlovic (1983), Mendel et al. (1984) 725 Regime A parasitic/ predator Native range North America First Record Invaded countries in Europe 1921, IT AL, AT, BG, CH, CZ, DE, DK, FR, HU, IL, IT, MD, NL, PT, RO, RU, SI, SK, UA, 1953, ES DE, ES, IL, IT Habitat Aphelinus semiflavus Howard, 1908 A parasitic/ predator North America Aphytis abnormis (Howard, 1881) A parasitic/ predator North America 1953, FR ES, FR-COR, GR, HU G4 Aphytis acrenulatus DeBach & Rosen, 1976 A parasitic/ predator Africa 1994, IT IT I Aphytis chilensis Howard, 1900 A parasitic/ predator South America 1910, ES CY, DE, ES, FR, GR, IT-SIC I, G3, J100 Aphytis coheni DeBach, 1960 A parasitic/ predator Asia 1959, IL CY, GR, IL I Aphytis diaspidis (Howard, 1881) A parasitic/ predator North America 1952, F AT, CY, ES, FR, GR, IL, IT, NL, PL I, G3 Host References I2 Woolly apple aphid, Eriosoma lanigerum (Monophagous parasitoid) Del Guercio (1925) I, Aphids (Acyrtosiphon pisum, Macrosiphum, etc.) Diaspidids and coccids scale insects (Lepidosaphes, Coccus) Herting (1972), Janssen (1961), Thompson (1953) Herting (1972), Peck (1963), Stathas and Kontodimas (2001), Thompson (1953) Diaspidid scale insects Garonna (1994) (Aspidiella zingiberi and Rhizaspidiotus donacis )) Diaspidid scale Alexandrakis and insects (Aspidiotus, Neuenschwander (1979), Hemiberlesia etc.) Herting (1972), Liotta (1974), Mercet (1911), Thompson (1953), Viggiani (1994a) Chrysomphalus DeBach (1960), Rosen dictyospermi on Citrus and DeBach (1979), Wood (1962) Diaspidid scale insects Applebaum and Rosen (1964), Herting (1972), Rosen and DeBach (1979), Thompson (1953) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Status 726 Families Species Aphelinus mali (Haldeman, 1851) Status Native range Asia First Record Invaded countries in Europe 1959, IL BE, CY, CZ, DE, ES, FR, IL, NL A parasitic/ predator Aphytis lepidosaphes Compere, 1955 A parasitic/ predator Asia 1961, CY CY, ES, FR, FRCOR, GR, GRCRE, IL, IT I Aphytis lingnanensis Compere, 1955 A parasitic/ predator Asia 1966, IT AL, CY, ES, GR, IL, IT I Aphytis melinus DeBach, 1959 A parasitic/ predator Asia 1966, ITSIC I, J100 Aphytis mytilaspidis (Le Baron, 1870) A parasitic/ predator North America 1837, FR Aphytis yanonensis DeBach &Rosen, 1982 Cales noacki Howard, 1907 A parasitic/ predator parasitic/ predator Asia 1986, FR AL, BE, CY, CZ, DE, DK, ES, FR, GR, IL, IT-SIC, IT, PT BE, BG, CH, CY, CZ, DE, ES, FR, GB, GR, HR, HU, IT, ME, NL, PL, RO, RS, SE, SI, SK, UA, FR, GR C&S America 1970, IT A Regime Habitat I, J100 I, G3, J100 I, J100 ES, ES-CAN, FR, I, J100 GR, IL, IT, IT-SAR, IT-SIC, MT, PT Host References Diaspidid scale insects (Chrysomphalus ficus), Citrus, Ficus, Musa, Cucurbita Lepidosaphes beckii on Citrus DeBach (1960), Wood (1962) Scale parasitoidon citrus Aleurothrixus floccosus on Citrus Benassy and Pinet (1987) Argyriou (1974), Benassy et al. (1974), Rosen (1965), Rosen and DeBach (1979), Viggiani and Iannaconne (1972), Wood (1962) Aonidiella aurantii and Argov et al. (1995), Rosen other scales on Citrus and DeBach (1979), Viggiani (1994a) Aonidiella aurantii on Alexandrakis and Benassy Citrus (1981), Inserra (1971), Rosen and DeBach (1979), Viggiani (1994a) Diaspidid scale insects Rosen and DeBach (1979), Viggiani (1994a) Hymenoptera. Chapter 12 Families Species Aphytis holoxanthus DeBach, 1960 Carrero (1979), Del Bene and Gargani (1991), Onillon (1973), Spicciarelli et al. (1996) 727 First Record Invaded countries in Europe 1943, HU AT, DE, HU, RU, UA Coccobius fulvus (Compere & Annecke, 1961) Coccophagoides murtfeldtae (Howard, 1894) Coccophagoides utilis Doutt, 1966 Coccophagus bivittatus Compere, 1931 Coccophagus capensis Compere, 1931 Coccophagus ceroplastae (Howard, 1895) A parasitic/ predator North America 1986, FR FR I2, J100 A parasitic/ predator North America 1962, IT IT I A parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator North America Africa 1975, GR GR I 1960, IT IL, IT Africa Asia 1962, ITSIC 1975, FR Coccophagus cowperi Girault, 1917 A parasitic/ predator Africa Coccophagus flavoscutellum Ashmead, 1881 Coccophagus gossypariae Gahan, 1927 A parasitic/ predator A parasitic/ predator A A A Habitat I Host References Pupae of dipterous, chalcid and proctotrupids (hyperparasitoid) Diaspidid scales on ornemental plants and Citrus Pseudaulacaspis pentagona Erdös (1953), Herting (1978), Peck (1963), Thompson (1953) I Parlatoria oleae on olive tree Coccus hesperidum IL, IT-SIC I Saissetia oleae FR, IL I, J100 1963, IT GR, IL, IT I North America 1962, ITSIC IT-SIC I Saissetia oleae and Ceroplastes floridensis on Citrus Saissetia oleae and other coccids, (sometimes hyperparasitoid) Coccus oleae Argyriou and Kourmadas (1979) Herting (1972), Zinna (1961) Argov and Rössler (1988), Peck (1963) Argov and Rössler (1988), CIBC (1976) North America 1990, IT DE, IT I Gossyparia spuria (Eriococcidae) Benassy and Pinet (1987) Peck (1963) Ben-Dov (1978) Monastero (1962) Viggiani (1998), Viggiani (1999), Viggiani and Romagnoli (1995) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Native range North America 728 Families Status Regime Species Centrodora speciosissima A parasitic/ (Girault, 1911) predator Status Coccophagus silvestrii Compere, 1931 Coccophagus varius (Silvestri, 1915) Encarsia acaudaleyrodis Hayat, 1976 Encarsia aurantii (Howard, 1894) Encarsia azimi Hayat, 1986 Encarsia berlesei (Howard, 1906) A A Regime parasitic/ predator parasitic/ predator Native range Asia First Record Invaded countries in Europe 1973, IT IT Habitat I Pseudococcus fragilis Viggiani (1975a) Asia 1979, IT IT, I Coccus hesperidum Viggiani (1980) Saissetia oleae on Citrus Annecke and Mynhardt (1979b), Mazzone and Viggiani (1983) scales on Citrus, Vine, Carrero (1980), Faber and Populus and others Sengonca (1997), Montiel (polyphagous) and Santaella (1995), Oncuer (1974), Panis et al. (1977), Paraskakis et al. (1980) Various coccids on Viggiani and Mazzone Citrus (1979) Saissetia oleae Mazzone and Viggiani (1983) Aleyrodidae Hernández-Suárez et al. (2003) Diaspidid scale insects Howard (1895) (polyphagous) Aleyrodidae on various Gonzalez Zamora et al. cultivated plants (1996), Kirk et al. (1993) Pseudaulacaspis Ferrière (1961), Howard pentagona (1912), Silvestri (1908) parasitic/ predator Africa 1978, IL IL, IT I C parasitic/ predator Cryptogenic 1826, SE AL, BE, DE, ES, FR, IL, NL, PT, SE I, J100 A parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator Asia1972, FR Temperate Africa 1983, IT CZ, FR, I, J100 IL, IT I Asia ES-CAN J100 CH, DE, FR, HU, IT, PL ES, ES-CAN, IT, I, G3 AL, AT, BG, CH, DE, ES, FR, GR, HR, HU, IT, ITSAR, IT-SIC, ME, RU, SI, YU I A A A A A North America Asia Asia 1999, ESCAN 1941, IT 2001, IT 1906, IT I, J100 References 729 A Host Hymenoptera. Chapter 12 Families Species Coccophagus gurneyi Compere, 1929 Coccophagus matsuyamensis Ishihara, 1977 Coccophagus saissetiae (Annecke & Mynhardt,1979) Coccophagus scutellaris (Dalman, 1825) First Record Invaded countries Habitat in Europe 1915, NL BE, DE, ES, FR, NL J100 1962, IT IT I Cryptogenic 1917, IT CH, DE, ES, FR, IL, IT I Encarsia formosa (Gahan, 1924) A parasitic/ predator C&S America 1964, BU Encarsia guadeloupae Viggiani, 1987 A parasitic/ predator C&S America 2000, ESCAN AL, AT, BE, BG, I, J100 CH, CZ, DE, DK, EE, ES-CAN, FI, FR, GB, HU, IE, IL, IT, IT-SAR, IT-SIC, IT, LT, MT, NL, NO, PL, PT, RO, RS, SE, SK ES-CAN I Encarsia herndoni (Girault, 1935) A parasitic/ predator Asia 1987, FR Encarsia hispida De Santis, 1948 Encarsia inquirenda (Silvestri, 1930) A parasitic/ predator parasitic/ predator South 1992, IT America Asia 1979, ES Temperate A AL, ES, FR-COR, IT, IT-SIC I, J100 ES-BAL, ES-CAN , FR, IT, ES, IL, IT I, J100 I2 Host Scals on olive, Citrus, etc (polyphagous) Pseudaulacaspis pentagona Scales on Laurus, Citrus, Populus, Crataegus, Malus Whiteflies References Ghesquière (1933), Smits van Burgst (1915) Peck (1963) Gerson (1967), Herting (1972), Malenotti (1917), Neuffer (1962), Thompson (1953) Burnett (1962), Gerling (1966), Kowalska (1969), Lenteren et al. (1976), Scopes (1969), Stenseth (1976), Viggiani (1987) Nijhor, 2000 #587} Aleurodicus dispersus and Lecanoideus floccissimus Insulaspis gloverii, scale Benassy and Brun (1989), on Citrus Liotta et al. (2003), Maniglia et al. (1995), Viggiani (1987) Bemisia Nijhof et al. (2000) Lepidospahes glovenii on Citrus, against Viggiani (1987) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Native range Cryptogenic Asia 730 Families Status Regime Species Encarsia citrina (Craw, C parasitic/ 1891) predator Encarsia diaspidicola A parasitic/ (Silvestri, 1909) predator Encarsia fasciata C parasitic/ (Malenotti, 1917) predator Families Species Encarsia lahorensis (Howard, 1911) Status Regime Native range Asia First Record Invaded countries in Europe 1973, IT FR, GR, IL, IT, ITSAR, IT-SIC, RU, parasitic/ predator Encarsia lounsburyi (Berlese & Paoli, 1916) A parasitic/ predator Africa 1922, IT Encarsia meritoria Gahan, 1927 Encarsia pergandiella Howard, 1907 A parasitic/ predator parasitic/ predator North America Asia? Encarsia perniciosi (Tower, 1913) A parasitic/ predator Encarsia porteri (Mercet, 1928) Encarsia protransvena Viggiani, 1985 Encarsia sophia (Girault & Dodd,1915) A parasitic/ predator parasitic/ predator parasitic/ predator A A A Host I, J100 Citrus whitefly, Dialeurodes citri (specific parasitoid) I, J100 Insulaspis gloverii scale on Citrus 1990, IT AL, CH, CY, ES, ES-BAL, FR, FRCOR, FR, GR, IL, IT, NL, PT IT, IT-SIC I 1978, IT FR, IL, IT, IT-SIC I Bemisia tabaci on Gossypium Bemisia Asia 1946, IT AL, AT, BG, CH, CZ, DE, DK, YU, FR, GR, GL, IT, IT-SIC, RO, RS, SK, YU I San Jose scale South America North America Asia 1993, IT IT I 1998, ES ES, IT I 1992, IT ES, ES-CAN, IL, IT, I Aleyrodidae and various insect eggs Aleyrodidae and scale insects Bemisia and whiteflies References Pappas and Viggiani (1979), Viggiani (1981), Viggiani and Mazzone (1977a), Viggiani and Mazzone (1978) Viggiani (1987) Viggiani (1987) Buijs et al. (1981), Rivnay and Gerling (1987), Viggiani (1987) Bénassy et al. (1965), Bénassy et al. (1968), Gambaro (1965), Mathys and Guignard (1962), Neuffer (1962), Neuffer (1968) Viggiani and Gerling (1994b) Giorgini (2001), Polaszek et al. (1999) Gonzalez Zamora et al. (1996), Hernández-Suárez et al. (2003), Pedata and Viggiani (1993), Viggiani and Gerling (1994a) Hymenoptera. Chapter 12 A Habitat 731 First Record Invaded countries Habitat in Europe 1987, IL DE, ES, IL, IT, MT, I PL Eretmocerus corni Haldeman, 1850 Eretmocerus debachi Rose & Rosen, 1992 Eretmocerus eremicus Rose & Zolnerowich, 1997 A parasitic/ predator parasitic/ predator parasitic/ predator North America North America North America 1963, IT GR, IT I 1991, IT IL, IT, IT-SIC, I 1994, CZ BE, CH, CZ, DK, ES, FI, FR, DE, GR, HU, IT, LT, MT, NL, NO, PL, PT, SK I, J100 Eretmocerus haldemani Howard, 1908 A parasitic/ predator Asia 1968, FRCOR FR-COR, UA I Eretmocerus paulistus Hempel, 1904 Marietta carnesi (Howard, 1910) Pteroptrix chinensis (Howard, 1907) Pteroptrix orientalis (Silvestri, 1909) Pteroptrix smithi (Compere 1953) A parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator North America Asia 1970, ES AL, ES I 1987, ES IT, ES I Asia 1974, IT IT, RU I Asia 1909, IT IT I Asia 1968, IL IL, IT I A A A A A A Host Bemisia References Abd-Rabou (1999), Albert and Schneller (1994), Argov and Rössler (1988), Baraja et al. (1996), Bednarek and Goszczynski (2002), Mifsud (1997) Menteelos (1967) Siphoninus phillyreae (Aleyrodidae) Parabemisia myricae on Rose and Rosen (1992) citrus Bemisia, Trialeurodes Berndt et al. (2007), Gerling et al. (2001), Gonzalez et al. (2008), Lacordaire and Dussart (2008), Mary (2005), Rose and Zolnerowich (1997), Stansly et al. (2005) Aleyrodids (Bemisia, Chumak (2003), Onillon Trialeurodes) on Citrus, (1969) Solanum, .. Aleurothrixus floccosus DeBach and Rose (1976a), in Citrus groves DeBach and Rose (1976b) Hyperparasitoid Rosen (1962) Mytilococcus beckii on Citrus Chrysomphalus dictyospermi Chrysomphalus aonidum Liao et al. (1987), Viggiani (1975a) Viggiani and Garonna (1993) Flanders (1969), Viggiani (1975a) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Native range North America 732 Families Status Regime Species Eretmocerus californicus A parasitic/ Howard, 1895 predator Status Regime Native range First Record in Europe Invaded countries Habitat Host References C parasitic/ predator Cryptogenic Unknown, GB GB J Grain beetles (Cryptolestes) Finlayson (1950) C parasitic/ predator parasitic/ predator parasitic/ predator Cryptogenic North America North America Unknown, GB Unknown, SE Unknown GB J Fitton et al. (1978) SE J Tribolium confusum ( Larval parasitoid ) Anthrenus ES, FR, IL, IT J Various grain beetles (Cucujidae) Tussac and Blasco-Zumeta (1999) A parasitic/ predator Asia1965, CZ Temperate E, I1, I2, J100 Aphids in greenhouses Clausen (1978), Stary (1975), Stary and Remaudiere (1973), Stary et al. (1977){ Aphidius smithi Sharma & Subba Rao, 1959 A parasitic/ predator Asia1960, PL Temperate I Acyrthosiphon kondoi and A. pisum Pennacchio (1989) Cotesia hyphantriae (Riley, 1887) Cotesia marginiventris (Cresson,1865) A parasitic/ predator parasitic/ predator North America North America 1953, YU AL, AT, BE, CH, CZ, DE, DK, ES, FI, , FR, FR-COR, GB, GR, HU, IE, IT, LT, MT, NL, NO, PL, PT, PTMAD, SE, SK, AL, BG, CH, CY, CZ, DE, DK, ES, ES-CAN, FI, GR, HR, HU, IE, IL, IT, IT-SIC, LT, MD, NL, PL, PT, PTMAD, RU, SK, UA YU G4 Hyphantria cunea Glavendekic (2000) 1993, FR BE, DE, ES, FR, NL J100 grasslands (N)greenhouses (I) Clausen (1978) A A A Gordh and Moczar (1990) Hymenoptera. Chapter 12 Families Species Bethylidae Cephalonomia waterstoni Gahan, 1931 Holepyris sylvanidis (Brèthes, 1913) Laelius utilis Cockerell, 1920 Plastanoxus laevis (Ashmead, 1893) Braconidae Aphidius colemani Viereck, 1912 733 Regime Native range Africa First Record Invaded countries in Europe Unknown, IT IT A parasitic/ predator A parasitic/ predator Australasia 1932, ES A parasitic/ predator parasitic/ predator North America North America Unknown, GB 1933, FR C parasitic/ predator Cryptogenic Macrocentrus ancylivorus (Rohwer, 1923) Microgaster pantographae Muesebeck, 1922 Opius dimidiatus Ashmead, 1889 A parasitic/ predator A Pauesia cedrobii Starý & Leclant 1977 Habitat Host References I fruit-Infesting Tephritidae Clausen (1978) ES, ES-CAN, IL I fruit-Infesting Tephritidae Clausen (1978) GB I Cephus pygmeus Clausen (1978) FR, IT I Cydia molesta van Achterberg (1993) 1965, CZ AL, BG, CZ, DK, ES, FR FR-COR, IT, IT-SIC, PT E, I Aphids North America 1930, ITSAR FR-COR, IT-SAR, i Ancylis comptana Barbagallo et al. (1983), Costa and Stary (1988), Kavallieratos and Lykouressis (1999), Ortu and Prota (1983), Stary et al. (1985), Steenis (1992), Tremblay et al. (1978) Labeyrie (1957) parasitic/ predator North America Unknown, GB GB I Tortricid moths Fitton et al. (1978) A parasitic/ predator North America Unknown, NL NL I1 van der Linden (1986) A parasitic/ predator Africa 1987, FR FR, IL G1, I2 Liriomyza trifolii (Solitary endoparasitoid) Cedrodium on Cedrus A Fabre and Rabasse (1987), Remaudière and Stary (1993) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Status 734 Families Species Diachasmimorpha fullawayi (Silvestri, 1912) Diachasmimorpha tryoni (Cameron, 1911) Heterospilus cephi Rohwer, 1925 Hymenochaonia delicata (Cresson 1872) Lysiphlebus testaceipes (Cresson, 1880) Cynipidae Dryocosmus kuriphilus Yasumatsu, 1951 Dryinidae Neodryinus typhlocybae (Ashmead, 1893) Encyrtidae Adelencyrtus aulacaspidis (Brèthes, 1914) Aenasius flandersi Kerrich, 1967 Status A A A Regime parasitic/ predator parasitic/ predator parasitic/ predator Native range Asia First Record Invaded countries in Europe 1930, ES ES, IT Habitat North America Africa Unknown, DE 1914, IT DE I FR, GL, IT G4 G3 Host References Grey pine aphid, Schizolachnus pineti Phyllotreta leaf beetles (adults) Fruit-Infesting Tephritidae Quilis Pérez (1931) Haeselbarth (2008) Clausen (1978), Delanoue (1960) A parasitic/ predator North America Unknown AT, BE, CH, DK, FI, GR, HR, RS I Cecidomyidae Dessart (1994) A parasitic/ predator Africa 1912, IT GR, IL, IT I Fruits Greathead (1976), Podoler and Mazor (1981), Thompson (1953) A phytophagous Asia2002, IT Temperate CH, FR, HU, IT, SI G1, I2 Castanea Anonymous (2005), Breisch and Streito (2004), Csoka et al. (2009), Forster et al. (2009), Graziosi and Santi (2008) A parasitic/ predator North America 1994, IT CH, FR, IT, SI Metcalfa pruinosa Malausa (1999), Malausa et al. (2003) A parasitic/ predator South America 1930, FR Various Diaspididae Trjapitzin (1989) A parasitic/ predator South America 1999, ESCAN BG, CH, CZ, DE, G3, G4 ES, FR, GB, HR, HU, IT, RU, SI, UA ES-CAN I Phenacoccus manihoti Baez and Askew (1999) I Hymenoptera. Chapter 12 Families Species Pauesia unilachni (Gahan, 1927) Perilitus vittatae (Muesebeck, 1936) Psyttalia concolor (Szépligeti, 1910) Ceraphronidae Aphanogmus bicolor Ashmead, 1893 Chalcididae Dirhinus giffardii Silvestri,1913 735 Native range Asia? First Record Invaded countries in Europe 1966, ITFR, ES, ES-CAN, SIC GR, IL, IT, IT-SIC, PL A parasitic/ predator Aloencyrtus saissetiae (Compere,1939) Anagyrus agraensis Saraswat,1975 Anagyrus fusciventris (Girault, 1915) Anagyrus sawadai Ishii,1928 Anagyrus subflaviceps (Girault, 1915) Anicetus annulatus Timberlake, 1919 Anicetus ceroplastis Ishii,1928 Anthemus hilli Dodd, 1917 Avetianella longoi Siscaro, 1992 A parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator Africa 1987, IL Asia 1987, IL Bothriophryne fuscicornis Compere, 1939 Clausenia purpurea Ishii,1923 A A A A A A A A A A Regime Host References I, J100 Citrus leafminer, Phyllocnistis citrella, in Citrus orchards IL I IL I Saissetia oleae on citrus. Nipaecoccus viridis Argov and Rössler (1996), Michelakis (1997), Siscaro et al. (1997), Siscaro and Mazzeo (1997), Urbaneja et al. (2000) Argov and Rössler (1988) BE, DE, DK, ES, FR, DE, IT, NL IL J100 Australasia 1994, PT ES, IL, PT North America Asia 1977, HR Australasia 1983, IT Habitat Bar-Zakay et al. (1987) Viggiani and Battaglia (1983) Blumberg et al. (1999b) I pseudococcids on Cycas, coffee, Citrus Citrus mealybug, Pseudococcus cryptus Pseudococcids AL, HR I Scale insects on Citrus 1989, IL IL I Ceroplastes floridensis Hoffer (1970), Hoffer (1982) Blumberg (1977) Australasia 1954, ES ES I Chionaspis graminis Gerling et al. (1980) Australasia 1990, PT IT-SIC, IT, PT I, G1 parasitic/ predator Africa 1972, IL CZ, IL, SK I, G Phoracantha semipunctata (Oophagous) Various Coccidae Farrall et al. (1992), Longo et al. (1993), Siscaro (1992) Kfir and Rosen (1980) parasitic/ predator Asia 1974, IL IL, IT I Citriculus mealybug Pseudococcus cryptus Guerrieri and Pellizzari (2009), Rosen (1974) Asia 1996, IL I Simutnik et al. (2005) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Status 736 Families Species Ageniaspis citricola Logvinovskaya, 1983 Families Species Coccidencyrtus malloi Blanchard, 1964 Coccidoxenoides perminutus Girault, 1915 Status A A Regime parasitic/ predator parasitic/ predator Native range South America Asia First Record Invaded countries in Europe 1999, FR FR, IT Habitat Host J100 Diaspis boisduvalii 1956, IT CY, GB, IL, IT I, J100 Planococcus ficus and P. citri J References Panis and Pinet (1999a) A parasitic/ predator South America 1988, FR F, IT Comperiella bifasciata Howard, 1906 A parasitic/ predator Asia 1990, IT Comperiella lemniscata Compere & Annecke, 1961 Copidosoma floridanum (Ashmead, 1900) A parasitic/ predator Asia 1989, IT BE, CY, CZ, ES, I, J100 FR, GR, HU, IL, IT, IT-SIC, MD, NL, RU, UA ES, IL, IT I A parasitic/ predator North America 1920, GB Copidosoma koehleri Blanchard, 1940 Diversinervus cervantesi (Girault,1933) Diversinervus elegans Silvestri, 1915 A parasitic/ predator parasitic/ predator parasitic/ predator C&S America Asia 1994, IT BG, CZ, DE, ES, I ES-CAN, FR, DE, GB, GR-CRE, HU, IT, NL, PT, RU, RS, SE, SK AL, CY, GR, IT I 1982, IL IL I soft scale insects Guerrieri (1995), Guerrieri and Noyes (2005) Rosen and Alon (1983) Africa 1977, IT ES, FR, GR, IL, IT I Encyrtus fuscus (Howard, 1881) A parasitic/ predator North America 1901, IT IT I, G3 black scale, Saissetia oleae, on olive, Citrus (polyphagous) Lecanium scales Kfir and Rosen (1980), Panis (1983), Viggiani and Mazzone (1977b) Noyes and Hayat (1994) A A Phtorimea operculella 737 Comperia merceti (Compere, 1938) Hymenoptera. Chapter 12 Fry (1989), Noyes and Hayat (1994), Trjapitzin (1978), Viggiani (1975a), Zinna (1960) Supella longipalpa Goudey-Perrière et al. (1988), Goudey-Perrière et al. (1991) Aonidiella aurantii & Bénassy and Bianchi A. citrina on Citrus & (1974), Liotta and Salvia passionfruit (1991), Orphanides (1996) Chrysomphalus Battaglia (1988), Garonna dictyospermi and Viggiani (1989), Pina et al. (2001) Noctuid moths Guerrieri and Noyes (Polyembryonic) (2005), Noyes (1988) Status A parasitic/ predator Asia Metaphycus anneckei Guerrieri & Noyes, 2000 A parasitic/ predator Africa 1973 CY, ES, GR, IL, IT, PL, PT I2 Metaphycus flavus (Howard, 1881) A parasitic/ predator North America 1915, FR AL, CY, CZ, FR, ME, PT-MAD, PT, RU, ES-BAL I A parasitic/ predator parasitic/ predator Native range Africa Africa Host Coccids (Saissetia spp.) on Citrus, Ficus Mealybugs (Planococcus citri) on many host plants (polyphagous) Coccids on Nerium oleander, Asteraceae, Cupressus spp.., Leonotis leoneurus, Olea europaea, Leucadendron pubescens, Lycium tetrandrum Coccids on Nerium oleander, Asteraceae, Cupressus spp., Leonotis leoneurus, Olea europaea, Leucadendron pubescens, Lycium tetrandrum soft scales (Facultative gregarious parasitoid) References Embleton (1902) Krambias and Kotzionis (1980), Longo and Benfatto (1982), Luppino (1979), Mineo and Viggiani (1976), Viggiani (1975b) Trjapitzin (1989) Guerrieri and Noyes (2000) Monaco and D’Abbicco (1987), Noguera et al. (2003), Orphanides (1988), Tena-Barreda and Garcia-Mari (2006), Velimirovic (1994) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Metaphycus angustifrons Compere,1957 First Record Invaded countries Habitat in Europe 1901, GB BE, DE, DK, ES, I, J100 FR, GB, IL, NL I, J100 1959, IT AT, BA, BE, CY, CZ, DE, DK, ES, FI, FR, DE, GB, GR, IE, IL, IT, ITSAR, IT-SIC, NL, NO, PL, PT, SE, YU 1988, IL IL I2 A Regime 738 Families Species Encyrtus infelix (Embleton, 1902) Leptomastix dactylopii Howard, 1885 Status First Record Invaded countries in Europe 1993, ES ES I Africa 1978, IT AT, BE, CH, CY, DE, DK, ES, FR, FR-COR, GR, IL, IT, NL, SE J100 Metaphycus inviscus Compere,1940 A parasitic/ predator Africa 1987, ITSAR ES, ES-BAL, IL I2 Metaphycus lounsburyi (Howard, 1898) A parasitic/ predator Africa 1973, IT CY, DK, ES, FR, IL, I2, J100 IT, IT-SIC, NL, PL Black scale, Saissetia oleae, polyphagous on olive, citrus Metaphycus luteolus (Timberlake, 1916) A parasitic/ predator North America 1989, IT ES, IT, UA Fruit scales Metaphycus maculipennis (Timberlake, 1916) Metaphycus orientalis (Compere, 1924) A parasitic/ predator North America 1988, IT DE, ES, FR, GR, IT, RS A parasitic/ predator Asia 1989, BE BE A parasitic/ predator parasitic/ predator Habitat I2 Host References Protopulvinaria pyriformis on avocado Scale insects. Only in greenhouses Guerrieri and Noyes (2000) Argyriou and Katsoyannos (1976), Carrero (1980), Mazzone and Viggiani (1983), Montiel and Santaella (1995), Panis (1983), Panis et al. (1977), Stratopoulou and Kapatos (1984), Viggiani (1978) Argov and Rössler (1988), Guerrieri and Noyes (2000) Argyriou and Michelakis (1975), Canard and Laudeho (1977), Monaco (1976), Monaco and D’Abbicco (1987), Orphanides (1988), Panis (1977), Panis and Marro (1978), Tena-Barreda and Garcia-Mari (2006) Guerrieri and Noyes (2000), Viggiani and Guerrieri (1988) Guerrieri and Noyes (2000) Black scale, Saissetia Coccidae on Vitis I Coccidae on Citrus Guerrieri and Noyes (2000) 739 Native range Africa A Regime Hymenoptera. Chapter 12 Families Species Metaphycus galbus Annecke, 1964 Metaphycus helvolus (Compere, 1926) Status Regime Native range Africa First Record Invaded countries in Europe 1960, IT ES-CAN, ES, GR, IL, IT Habitat parasitic/ predator Metaphycus swirskii Annecke & Mynhardt, 1979 A parasitic/ predator Africa 1976, IT ES , FR, GR, GRCRE, IL, IT, NL I2 Microterys clauseni Compere,1926 Microterys nietneri (Motschulsky, 1859) Microterys speciosus Ishii,1923 Neodusmetia sangwani (Subba Rao,1957) Ooencyrtus kuwanae (Howard, 1910) A parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator Asia 1987, IL IL I Asia 1989, BG BG, PT-AZO I2 Asia 1987, IL IL I Asia 1974, IL IL E Asia 1932, PT Temperate AT, BA, BG, CH, CZ, DE, ES, FR, IT-SAR, MD, PL, PT, RO, RU, SK, UA, YU G1 Plagiomerus diaspidis Crawford, 1910 A parasitic/ predator North America 1994, ITSIC ES-CAN, FR, ITSIC, PT-MAD I Diaspididae on Opuntia Prochiloneurus pulchellus Silvestri, 1915 A parasitic/ predator Africa 1972, IL IL, IT I scale insects (polyphagous) A A A References fruit scales Argov and Rössler (1988), Blumberg et al. (1993), Guerrieri and Noyes (2000), Noyes and Hayat (1994), Trjapitzin (1989) scales on Ficus, Citrus, Annecke and Mynhardt Coffee, Solanum (1979a), Panis (1981), Viggiani and Mazzone (1977b) Ceroplastes floridensis Argov and Rössler (1988) on Citrus Coccus Simoes et al. (2006) Ceroplastes floridensis on Citrus Rhodesgrass scale, Antonina graminis Lymantria dispar Argov and Rössler (1988) Gerson et al. (1975) Bjegovic (1962), Keremidchiev et al. (1980), Mihalache et al. (1995), Milanovic et al. (1998), Roversi et al. (1991) Bue and Colazza (2005), Panis and Pinet (1999b), Russo and Siscaro (1994) Trjapitzin (1989) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) A A I2 Host 740 Families Species Metaphycus stanleyi Compere, 1940 Status First Record Invaded countries in Europe 1964, IL IL, RU I, J100 1998, IL IL, RU I, J100 1986, IL IL I2 Australasia 2006, FRCOR FR, FR-COR, GB, IE, IT I2 Rhopus nigroclavatus (Ashmead, 1902) Tachinaephagus zealandicus Ashmead, 1904 A parasitic/ predator parasitic/ predator North 1978, ES America Australasia 2002, PTMAD ES I DK, IT, PT-AZO, PT-MAD J Tetracnemoidea brevicornis (Girault, 1915) Tetranecmoidea peregrina (Compere, 1939) Tineophoctonus armatus (Ashmead, 1888) Zarhopalus sheldoni Ashmead, 1900 A parasitic/ predator Australasia 1987, IT FR, IT I, J100 A parasitic/ predator C&S America 1994, PT ES, FR, IL, IT, PT I, J100 A parasitic/ predator parasitic/ predator North America North America 1963, ES ES, IT J citrus mealybug, Pseudococcus calceolariae citrus mealybug, Pseudococcus calceolariae Anobiidae 1945, RU RU J100 Pseudococcus comstocki A A A A A A Regime parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator Native range Tropical, subtropical AsiaTemperate Africa Habitat Host References Pseudococcids (Vitis, Solanum) Pseudococcids on Citrus Nipaecoccus viridis on Citrus Ctenarytaina eucalypti on Eucalyptus Noyes and Hayat (1994), Walton and Pringle (2002) Blumberg et al. (1999a) scale insects on Poaceae Musca domestica in poultry houses Bar-Zakay et al. (1987) Bennett (2005), Chauzat et al. (2002), Costanzi et al. (2003a), Costanzi et al. (2003b), Malausa and Girardet (1997), Schnee et al. (2006) Trjapitzin (1989) Japoshvili and Noyes (2006), Koponen and Askew (2002), Turchetto et al. (2003) Laudonia and Viggiani (1986a) Hymenoptera. Chapter 12 Families Species Pseudaphycus angelicus (Howard, 1898) Pseudaphycus malinus Gahan,1946 Pseudectroma signatum (Prinsloo,1982) Psyllaephagus pilosus Noyes, 1988 Trjapitzin (1989) Trjapitzin (1989) Noyes and Hayat (1994) 741 Aprostocetus diplosidis Crawford, 1907 Aprostocetus microcosmus (Girault, 1917) Aprostocetus sicarius (Silvestri, 1915) Astichus trifasciatipennis (Girault, 1913) Ceranisus americensis (Girault, 1917) Ceranisus russelli (Crawford, 1911) Chaenotetrastichus semiflavus (Girault, 1917) Chouioia cunea Yang, 1989 Chrysocharis ainsliei Crawford, 1912 Chrysocharis oscinidis Ashmead, 1888 A A A A A A A A A A A A Regime parasitic/ predator parasitic/ predator Native range First Record in Europe Invaded countries Habitat Host References Australasia 1974, IT GB, IT I, J100 fruit flies, Anastrepha Graham (1991), Viggiani (1975a) Coccidae (Ceroplastes) Argyriou and Kourmadas on fruit trees (1980), Avidov et al. (1963), Domenichini et al. (1964) Contarinia sorghicola Priore and Viggiani (1965) Africa 1962, IL FR, GR, IL, IT I parasitic/ predator parasitic/ predator North America North America 1964, IT IT E 1977, ESCAN ES-CAN I Cecidomyiidae on Poaceae Graham (1987) parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator Africa 1962, IL IL, ME I Bactrocera oleae Australasia 1989, IT IT G5 North America North America North America 1994, NL NL I Gracillariidae on Robinia pseudoacacia Thrips Avidov et al. (1963), OILB (1971) Serini (1990) 1954, GB GB I Thrips Thompson (1955) 1995, DE DE G Pompilidae Vidal (1996) parasitic/ predator parasitic/ predator parasitic/ predator Asia 1990, IT IT G1 Hyphantria cunea Boriani (1991) North America North America 1984, IT DK, IT I 1984, NL FR, NL I Phytomyza on artichokes Liriomyza Hansson (1985), Ikeda (1996) Fry (1989), Woets and Linden (1985) Loomans et al. (1995) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Status 742 Families Species Eulophidae Aceratoneuromyia indica (Silvestri, 1910) Aprostocetus ceroplastae (Girault, 1916) Families Species Cirrospilus ingenuus Gahan,1932 Citrostichus phyllocnistoides (Narayanan, 1960) Elachertus cidariae (Ashmead, 1898) Euderus cavasolae (Silvestri, 1914) Galeopsomyia fausta LaSalle, 1997 Goetheana shakespearei Girault, 1920 Hyssopus thymus Girault, 1916 Leptocybe invasa Fisher & LaSalle, 2004 A A Regime parasitic/ predator parasitic/ predator Native range Asia Asia First Record Invaded countries Habitat in Europe 1994, IL CY, ES, IL, PTI MAD, PT 1995, IL ES-BAL, GR, IL, IT, I IT-SIC, IT, PT A parasitic/ predator North America 1971, IT IT G5 A parasitic/ predator parasitic/ predator North America C&S America 1988, CZ CZ, NO I 1985, IT IT, RU I1 parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator phytophagous North America Africa 1962, YU YU G1 1954, IT IT I C&S 1999, ES America Australasia 1992, ES ES I2 ES I North 1970, DE America Australasia 2003, PT DE G3, I2 ES, FR, FR-COR, IL, IT, PT G1 A A A A A A A Host References Phyllocnistis citrella in Citrus orchards Phyllocnistis citrella in Citrus orchards Argov and Rössler (1996), Vercher et al. (2000)P Argov and Rössler (1996), Barbagallo et al. (2000), Michelakis and Vacante (1997), Vercher et al. (2000) Parectopa robiniella on Vidano and Marletto Robinia (1972) Leafminer parasitoid Hagvar et al. (1994), Kalina (1989) Colorado potato beetle Laudonia and Viggiani (1986b), Yefremova (2002) fall webworm in Tadic MD (1964) deciduous trees Bactrocera oleae Thompson (1955) Phyllocnistis citrella on Citrus Thrips Rhyacionia buoliana pine stands gall-former on Eucalyptus Hymenoptera. Chapter 12 Closterocerus cinctipennis Ashmead, 1888 Diglyphus begini (Ashmead, 1904) Edovum puttleri Grissell, 1981 Status Vercher et al. (2000) 743 Viggiani and Nieves Aldrey (1993) Konig and Bogenschutz (1971) Anagnou-Veroniki et al. (2008), Kim et al. (2008), Mendel et al. (2004), Protasov et al. (2008) Tetrastichomyia clisiocampae (Ashmead, 1894) Thripobius javae (Girault, 1917) Eupelmidae Anastatus japonicus Ashmead, 1904 Anastatus tenuipes Bolivar & Pieltain, 1925 Eupelmus afer Silvestri, 1914 Regime A phytophagous A parasitic/ predator phytophagous parasitic/ predator A Native First Record Invaded countries range in Europe Australasia 2000, IT ES, FR, FR-COR, GR, IL, IT, PT North 1944, CZ America Australasia 1968, IL Habitat Host G1 gall-former on Eucalyptus camaldulensis (N), other Eucalyptus (I) CZ, DE, GB I ES, FR-COR, IL, IT, IT-SAR, PT CY, ES, ES-BAL, GR, IL, IT, IT-SIC, PT G1 I2 Phyllotreta zimmermanni gall-former on Eucalyptus Phyllocnistis citrella on Citrus References Branco et al. (2009), Protasov et al. (2007a), Protasov et al. (2007b), Rizzo et al. (2006), Sasso et al. (2008) Boucek (1965) A parasitic/ predator North America 1966, IT IT G1, I Lepidoptera Boucek (1977a), Rasplus (1992) Argov and Rössler (1996), Barbagallo et al. (2000), Michelakis and Vacante (1997), Siscaro et al. (1999) Domenichini (1967) A parasitic/ predator Asia 1995, IT BE, DE, DK, FR, IL, IT, IT-SIC, NL J100 Greenhouse thrips on Citrus, Viburnumn, Vitis and others Viggiani and Bernardo (1996), Wysoki et al. (2000) A parasitic/ predator parasitic/ predator Asia 1920, HU CZ, HU, SK, YU G1 Ruschka (1921) Africa 1999, IT IT J Lymantria and forest moths Supella longipalpa (Blattidae) parasitic/ predator Africa 1974, IT IT I Bactrocera oleae Viggiani (1975a) A A A Australasia 1995, IL Russo et al. (2000) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Pediobius phyllotretae (Riley, 1884) Quadrastichodella nova Girault, 1922 Semielacher petiolata (Girault, 1915) Status 744 Families Species Ophelimus maskelli (Ashmead 1900) Status Native First Record Invaded countries range in Europe Australasia 1964, IT IT, SK, UA, YU A parasitic/ predator I, I1, F5 Eupelmus longicorpus Girault, 1915 Eurytomidae Bruchophagus sophorae Crosby & Crosby, 1929 Eurytoma aloineae (Burks, 1958) Eurytoma orchidearum (Westwood, 1869) Prodecatoma cooki (Howard, 1896) Tetramesa albomaculatum (Ashmead, 1894) A parasitic/ predator Australasia 1987, ES ES A phytophagous Asia 1960, RO A phytophagous phytophagous phytophagous phytophagous Africa North America North America North America A A A Regime Habitat Host References I sorghum midge (Cecidomyiidae) and other midge on Poaceae midge on Poaceae Boucek (1977b), Kalina (1989), Priore and Viggiani (1965), Trjapitzin (1978) Bouček (1988) BG, HU, RO, RS, RU, SK, UA, YU I2 Sophora seeds Grubik (1992), Mihajlovic (1983), 3871996477 1957, DE DE J100 Aloe Burks (1958) 1962, FR DK, FR, NL J100 1886, AT AT I Cattleya and other orchids Grape wasp, Vitis Gijswijt (2003), Peck (1963)P Howard (1896) 1977, GB BG, DE, GB, SE I1 Wheat and Poaceae Boucek and Graham (1978), Hedqvist (2003), Stojanova (2004), Vidal (2001) Popescu (2004), Porchinsky (1881), Walker (1871) Zerova (1978) A phytophagous North America 1870, IT ES, HU, IL, IT, RO, I1 RU, UA wheat and Poaceae Tetramesa swezeyi (Phillips & Poos, 1922) Figitidae Aganaspis daci (Weld, 1951) A phytophagous Unknown 1977, RU RU, UA I1 wheat and Poaceae A parasitic/ predator Africa 1970, FR FR, GR_NEG I Bactrocera oleae Nunez-Bueno (1982), Papadopoulos and Katsoyannos (2003) 745 Tetramesa maderae (Walker, 1849) Hymenoptera. Chapter 12 Families Species Eupelmus australiensis (Girault, 1913) Cardiocondyla wroughtoni (Forel, 1890) Crematogaster brevispinosa Mayr, 1870 Hypoponera ergatandria (Forel, 1893) Hypoponera punctatissima (Roger, 1859) Regime Native range parasitic/ predator parasitic/ predator C&S America Africa 1874, CH CH, DE, FR, UA J100 Greenhouses Forel (1874) 1894, PT ES-CAN, PT-MAD G, I2, J1, X24 Natural sites and gardens, arid sites A parasitic/ predator Africa 1981, ESCAN I2, X24, J1 Gardens, houses, buildings A parasitic/ predator Africa 1930, IL CY, ES, ES-CAN, IL, IT, IT-SAR, ITSIC PT-MAD ES-CAN, IL Heinze and Trenkle (1997), Kluger (1983), Reyes-Lopez et al. (2008), Wetterer et al. (2007) Finzi (1936), Mei (1995), Wetterer et al. (2007) I2 A parasitic/ predator Asia 1982, IL IL H5, J Miscelleanous habitats, Seifert (2003) disturbed areas, beaches Miscelleanous habitats, Kluger (1983) disturbed areas A parasitic/ predator C&S America 1935, CZ CZ J100 Greenhouses Šefrová and Laštůvka (2005) A parasitic/ predator parasitic/ predator C&S 1952, DE America Tropical, 1847, PT subtropical DE, FR J Sparse or no vegetation, buildings Antropophilic, in greenhouses or other heated biuldings, gardens in Madeira Geiter et al. (2002) A A A First Record in Europe Invaded countries Habitat AT, BE, BG, CH, J, J100, CZ, DE, DK, ES, I2, X24 ES-CAN, FR, FRCOR, GB, GR, HU, IE, IS, IT, LU, MT, NL, NO, PT, PT-AZO, PT-MAD, RO, RS, RU, SE, SK, UA, YU Host References Blacker (2007), Boer et al. (2003), Boer et al. (2006), Carniel and Governatori (1994), Czechowska and Czechowski (1999b), Dessart and Cammaerts (1995), Jones (1997), Seifert (1982), Wetterer et al. (2007) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Cardiocondyla mauritanica Forel, 1890 Cardiocondyla obscurior (Wheeler, 1929) Status 746 Families Species Formicidae Brachymyrmex heeri Forel, 1874 Cardiocondyla emeryi Forel, 1881 Families Species Lasius neglectus Van Loon, Boomsma & Andrasfalvy, 1990 Status Regime Native First Record Invaded countries range in Europe Asia1973, HU BE, BG, CZ, DE, Temperate ES, FR, GL, HU, PL, PT Habitat parasitic/ predator Lasius turcicus Sanctchi, 1921 A parasitic/ predator Asia1970, HU Temperate Linepithema humile (Mayer, 1868) A parasitic/ predator C&S America 1847, PT Linepithema leucomelas Emery, 1894 Monomorium andrei Saunders, 1890 Monomorium destructor (Jerdon, 1851) A parasitic/ predator parasitic/ predator parasitic/ predator C&S America Africa 1955, AT AL, BE, BG, CZ, I2, X24 DE, DK, EE, ES, ES-CAN, FR, GR, HU, IT, PL, RO BE, BG, CH, CZ, J, G, I2 DE, ES, ES-CAN, FR, FR-COR, GB, IT, IT-SAR, IT-SIC, PL, PT, PT-AZO, PT-MAD AT J100 1924, ES ES, ES-BAL Asia 1892, ESBAL ES-BAL, PL, PT A A I2, X24 Polygynous species, parks and gardens Gardens References Boomsma et al. (1990), Czechowska and Czechowski (1999a), Czechowska and Czechowski (2003), Dekoninck et al. (2002), Espadaler (1999), Markó (1988), Neumeyer (2008), Schultz and Busch (2009), Seifert (1992), Seifert (2000), Van Loon et al. (1990) Seifert (1996) Giraud et al. (2002), Suarez et al. (2001), Wild (2004), Wild (2009) Gardens, greenhouses Wild (2007) J Urban environment J1 Urban environment Reyes Lopez and Luqque Garcia (2003) Boer and Vierbergen (2008), Salgueiro (2003), Šefrová and Laštůvka (2005), Wetterer (2009a), Yarrow (1967) 747 Various habitats indoors and outdoors Hymenoptera. Chapter 12 A Host First Record Invaded countries in Europe 1982, DE DE Monomorium salomonis (Linnaeus, 1758) A parasitic/ predator tropical 1881, FRL Pachycondyla darwinii Forel, 1893 Paratrechina bourbonica (Forel, 1886) Paratrechina flavipes (Smith, 1874) Paratrechina jaegerskioeldi (Mayr, 1904) Paratrechina longicornis (Latreille, 1802) A parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator Unknown A parasitic/ predator Africa 1847, ESMAD CH, CZ, DE, ES, H, I2, J1, Houses, buildings, ES-CAN, FI, FR, J100 plant cavities, trees, GB, IL, IT, MT, PTdebris, rotten wood AZO, PT-MAD C parasitic/ predator Cryptogenic 1881, FI CY, CZ, DE, FI, FR, GB, GR, NL, RU, SE, UA Paratrechina vividula (Nylander, 1846) A A A 1892, ES Unknown, MT Tropical, Unknown, subtropical GB Asia1952, DE Tropical Africa 1989, ESMAD Habitat Host References J100 Greenhouses Sellenschlo (1991) AT, BG, CH, CZ, DE, DK, EE, ES-CAN, FR, FRCOR, GB, HU, IL, IT, IT-SAR, IT-SIC, LT, ME, NL, NO, PT-MAD, PT, RS ES, ES-BAL, FR, GB, IT, IT-SAR, IT-SIC, MT MT J1, J100, X25, I2 Stored products antropophilic, mainly indoors, gardens in Madeira Markó et al. (2006), Salgueiro (2003) F6, J100 Garrigue Salgueiro (2003) U Forested areas GB U Cosmopolitan, tropics Fitton et al. (1978) DE, ES J1 Buildings ES, ES-CAN, GRCRE, PT-MAD J2, I2, X24 Low constructed buildings, gardens J, J100 Constructed areas, greenhouses Espadaler and Colllingwood (2000) Collingwood (1993), Espadaler and Bernal (2003), Kluger (1988) Collingwood et al. (1997), Espadaler and Bernal (2003) , Freitag et al. (2000), Heinze (1986), Tinaut and Año (2000) Collingwood and Hughes (1987) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Native range AsiaTropical tropical 748 Families Status Regime Species Monomorium floricola A parasitic/ (Jerdon, 1851) predator Monomorium pharaonis A parasitic/ (Linnaeus, 1758) predator Native range C&S America Tropical, subtropical Africa Pheidole noda (Smith, 1874) Pheidole teneriffana Forel, 1893 A parasitic/ predator parasitic/ predator Asia 2003, IT Africa 1893, ESBAL Plagiolepis alluaudi (Emery, 1894) Plagiolepis exigua Forel, 1894 Plagiolepis obscuriscapa Santschi, 1923 Pyramica membranifera (Emery, 1869) A parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator AsiaTemperate Tropical, subtropical C&S America Africa Strumigenys lewisi Cameron, 1886 Strumigenys rogeri Emery, 1890 A parasitic/ predator parasitic/ predator A A A A A First Record Invaded countries in Europe 1952, DE CH, DE, DK, FR, GB 1883, FR FR, DE 1847, PTMAD Habitat Host J100 Greenhouse J100 Sparsely wooded area (N), greenhouse(I) Gardens, urban I2, J1, DE, ES, ES-CAN, FR, GB, GR, GRJ100 CRE, IT, ME, PT, PT-AZO, PT-MAD, RO, YU IT I2 Nursery References Longino and Cox (2009) Bernard (1968), Limonta and Colombo (2003) I2, X24 Disturbed areas 1915, IE ES, ES-BAL, ESCAN, GR, GRCRE, GR_SEG, GR, IT-SIC CH, DE, FR, IE Limonta and Colombo (2003) De Haro et al. (1986), Gomez and Espadaler (2006) J100 Greenhouses Geiter et al. (2002) 1952, DE DE J100 Greenhouses Geiter et al. (2002) Unknown IT, RO U Unknown Moscaliuc (2009) 1989, PTMAD PT-MAD I2, X24 Gardens Asia 1996, MT MT J100 Greenhouses Espadaler (1979), Espadaler and Lopez Soria (1991) Schembri and Collingwood (1995) Africa Unknown DE, GB J100 Greenhouses Hymenoptera. Chapter 12 Families Status Regime Species Pheidole bilimeki Mayr A parasitic/ 1870 predator Pheidole guineensis A parasitic/ (Fabricius, 1793) predator Pheidole megacephala A parasitic/ (Fabricius, 1793) predator 749 Technomyrmex albipes (Smith, 1861) Technomyrmex detorquens (Walker, 1859) Temnothorax longispinosus Roger, 1863 Tetramorium bicarinatum (Nylander, 1846) Tetramorium insolens (Smith, 1861) A Tetramorium lanuginosum Mayr, 1870 Tetramorium simillimum (Smith, 1851) A A Regime parasitic/ predator parasitic/ predator Native range North America Tropical, subtropical First Record Invaded countries in Europe 1989, PTPT-MAD MAD 1984, DE AT, CH, DE, FI, GB, RU detrivorous AsiaTropical parasitic/ Asia predator 1989, PTMAD 1937, CZ AT, NL, PT-MAD A parasitic/ predator North America A parasitic/ predator AsiaTropical A parasitic/ predator A A A Habitat I2, X24 J1, J100 Host Gardens; predator on collembola stored products, antropophilic, indoors only Gardens, houses AT, CZ, DE I2, X24, J1 J100 Unknown, ES ES D6 Oak and mixed woodland 2003, IT DE, IT, PT-AZO, SE J100 Nurseries Asia, Unknown ATstralasia AT, FR, NL, PL J100 Greenhouses parasitic/ predator Asia IL, MT J100 Greenhouses s parasitic/ predator Tropical, Unknown subtropical DE, EE, FR, GB, IL, PL, PT-AZO, PT-MAD, GB J100 Greenhouses Unknown Greenhouses, houses References Geiter et al. (2002) Boer and Vierbergen (2008), Espadaler and Espejo (2002), Hogmo (2003b), Jucker et al. (2008), Scheurer and Liebig (1998), Sorvari (2002), Vipin et al. (1999), Wetterer (2009b) Boer and Vierbergen (2008) Šefrová and Laštůvka (2005) Högmo (2003a), Reyes and Espadaler (2005), Wetterer et al. (2004) de Jonge (1985), Radchenko et al. (1998), Radchenko et al. (1999) Reyes and Espadaler (2005), Schembri and Collingwood (1995) Bernard (1968), Wetterer et al. (2006) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Status 750 Families Species Strumigenys silvestrii Emery, 1906 Tapinoma melanocephalum (Fabricius, 1793) Status Polynema striaticorne Girault, 1911 Pamphiliidae Cephalcia alashanica (Gussakovskij, 1935) Regime Native range First Record in Europe Invaded countries Habitat Host References A parasitic/ predator AsiaUnknown Temperate FR, R I Stem borers (Pyralidae) A parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator North America North America North America North America North America Unknown AT, FR, RU G3 Sawflies on Tsuga Unknown FR, NO G3 Sawflies Unknown AT, DE, FI, GB, NL, NO, PL, RU AT, PL, RU, SE G3 G3 Sawflies (Diprionidae Hedstrom (1987), Jussila and others) (1989), Phillips (1997) Xylophagous beetles Hedstrom (1987) Unknown, DE DE I Apple tortricid A phytophagous Asia1970, DK Temperate DK I, E Pollinator of fruit trees Kristjansson and Rasmussen (1990) A parasitic/ predator Australasia 1977, IT ES, FR, IT, PT I2 A parasitic/ predator North America IT I2 Eucalyptus snout-beetle Arzone and Vidano Gonipterus scutellatus (1978), Cadahia (1986), (egg Parasitoid) Rivera et al. (1999), Vaz et al. (2000) Ceresa bubalus Vidano (1968) A phytophagous 1986, NL AsiaTemperate NL G3 Picea A A A A Unknown 1966, IT Gokhman (1996) Biermann (1973) 751 Battisti and Sun (1996), Gossner et al. (2007), Holusa et al. (2007), Jachym (2007), Shinohara and Zombori (2003) Hymenoptera. Chapter 12 Families Species Ichneumonidae Auberteterus alternecoloratus (Cushman, 1929) Cryptus luctuosus Cresson, 1864 Cteniscus dorsalis Cresson, 1864 Delomerista novita (Cresson, 1870) Ephialtes spatulatus (Townes, 1960) Itoplectis conquisitor (Say,1835) Megachilidae Osmia cornifrons (Radoszkowski, 1887) Mymaridae Anaphes nitens (Girault, 1928) A Mesopolobus spermotrophus Husey, 1960 A A A First Record in Europe Invaded countries Habitat Host References North America 1876, AT AT G Ptilinus (Anobiidae) Giraud and Laboulbène (1878) North America 1980, IT IT J100 Trialeurodes vaporariorum Tropical, 1971, FR subtropical ES, FR, IT, IT-SIC J100 Aleurothrixus floccosus Manzano et al. (2002), Viggiani (1997), Vis and Lenteren (2008) DeBach and Rose (1976a), Liotta et al. (2003) Cryptogenic 1911, AT AT, BE, CH, CZ, J DE, FR, GB, GR, HU, IL, IT, PT, RO, RU, RS, SE, SK Stored products beetles parasitic/ predator parasitic/ predator parasitic/ predator Africa 1957, IT IT I Bactrocera oleae Beratlief (1967), Boucek (1977b), Boucek and Graham (1978), Frilli (1965), Garrido-Torres and Nieves-Aldrey (1990), Hedqvist (2003), Kalina (1989), Mitroiu (2001), Ruschka (1912) Thompson (1958) Africa 1974, IT IT I Bactrocera oleae Viggiani (1975a) North America 1953, GB BE, DK, FR, GB, NL, PL, SE G3 Megastigmus seed chalcid in Abies seeds parasitic/ predator North America 1952, GB BE, CZ, DE, FR, GB, IT, LU, NL, PL, SE G3 Megastigmus seed chalcid in Douglas-fir seeds Bak (1999), Pettersen (1976), Skrzypczynska (1989), Wisniowski (1987) Graham (1969) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Halticoptera daci Silvestri, 1914 Mesopolobus modestus (Silvestri, 1914) Mesopolobus pinus Hussey, 1960 Native range 752 Families Status Regime Species Perilampidae Steffanolampus A parasitic/ salicetum (Steffan, predator 1952) Platygastridae Amitus fuscipennis A parasitic/ MacGown & Nebeker, predator 1978 Amitus spiniferus A parasitic/ (Brèthes, 1914) predator Pteromalidae Anisopteromalus C parasitic/ calandrae (Howard, predator 1881) Status Muscidifurax raptor Girault & Sanders, 1910 Paracarotomus cephalotes Ashmead, 1894 Spalangia cameroni, Perkins 1910 A parasitic/ predator Native First Record Invaded countries range in Europe C&S 1980, IL IL America Australasia 1973, IT ES, ES-CAN, FR, GR, IL, IT, IT-SIC, IT North 1954, CZ CZ, DE, DK, ES, America IT, RO A parasitic/ predator North America 1976, FR FR, IT, RU, A parasitic/ predator North America 1969, DK Theocolax elegans (Westwood, 1874) C parasitic/ predator Cryptogenic Urolepis rufipes (Ashmead, 1896) A parasitic/ predator North America A parasitic/ predator parasitic/ predator parasitic/ predator Scelionidae Duta tenuicornis (Dodd, 1920) Gryon leptocorisae (Howard, 1885) Telenomus busseolae Gahan, 1922 A A A A Regime parasitic/ predator parasitic/ predator Habitat Host References U Seed-beetles Boucek (1991) G, I2, F Scales, Quercus, Citrus, Fagus, Olea (Highly polyphagous) Musca domestica and stable files Raspi (1988), Simoes et al. (2006), Stratopoulou et al. (1981) Fabritius (1978), Fabritius (1981), Rutz and Axtell (1979) Boucek (1976), Dzhanokmen (1984) CY, CZ, DE, DK, J ES, IT, MD, RO, SE Musca domestica and stable files 1957, DE BE, DE, GR, J Stored products beetles 1989, DE DE, DK, SE J house flies (pupae) Falco et al. (2006), Gibson (2009), Maini and Bellini (1991), Tormos et al. (2009) Eliopoulos et al. (2002), Mitroiu (2001), Thompson (1958) Gibson (2000), Hedqvist (2003), Skovgard and Jespersen (1999) Australasia 1989, HU HU, MD I North America Africa Unknown DK, FR, IT I Unknown, IT IT I Crickets (Egg parasitoid) Stenocoris (Egg parasitoid) Stem borers (Egg parasitoid) J Hymenoptera. Chapter 12 Families Species Monoksa dorsiplana Boucek, 1991 Moranila californica (Howard, 1881) Popovici (2005) Mineo (1981) 753 Conti and Bin (2000), Gullu and Simsek (1995), Laudonia et al. (1991) Regime Native range A parasitic/ predator North America Unknown ES, FR, IT U Scale insects (Hyperparasitoid via Encyrtids) A phytophagous phytophagous North America North America 1995, GB GB, IT G3 Conifers 1885, FR BE, CH, DE, DK, FR, GB, GR, HU, IE, IL, IT, LU, NL, PT, SE, SK G3, I2 phytophagous phytophagous phytophagous North America North America North America 1957, GB GB G, I2 1991, GB GB, IS, NL, PL G3 Viitasaari and Midtgaard (1989) Conifer trunks (mainly Hayes (1982), Hellrigl Abies) (1984), Kirk (1974), Midtgaard (1986), Schwarz (1994), Viitasaari and Midtgaard (1989) Fagus, Quercus, Acer, Winter (1988) Betula, etc Conifers Witmond (2001) 1944, GB GB G3 Conifers Fitton et al. (1978) AT, CH, DE, ES, FR, FR-COR, HR, IT, SI AT, BE, DE, ESCAN, FR, FRCOR, HR, IT, LU, PT-MAD, PT, UA E, X25 Crickets in grasslands (predatory) C3, X25 Adults nectar at flowers and mud nests are built in sheltered locations such as garages and underneath bridges Pagliano et al. (2000), Scaramozzino and Pagliano (1987) Bitsch et al. (1997), Pagliano et al. (2000) A First Record in Europe Tremex columba (Linnaeus, 1763) Urocerus albicornis (Fabricius, 1781) Urocerus californicus Norton, 1869 Sphecidae Isodontia mexicana (Saussure, 1867) A A parasitic/ predator North America 1960, FR Sceliphron cementarium (Drury, 1773) A parasitic/ predator North America 1945, FR A A Invaded countries Habitat Host References Woolley (1988) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Siricidae Sirex areolatus (Cresson, 1867) Sirex cyaneus cyaneus Fabricius, 1781 Status 754 Families Species Signiphoridae Chartocerus niger (Ashmead, 1900) Families Species Sceliphron curvatum (Smith, 1870) Sceliphron deforme (Smith, 1856) Pachynematus (Larinematus) itoi Okutani, 1955 Torymidae Eridontomerus isosomatis (Riley, 1882) Megastigmus aculeatus nigroflavus Hoffmeyer, 1929 Regime Native First Record Invaded countries range in Europe Asia1979, AT AT, BG, CH, ,CZ, Temperate DE, FR, FR-COR, HR, HU, IT, ITSAR, IT-SIC,RS, SI, UA, Host References Adults nectar at flowers and mud nests are built in Sheltered locations such as garages and underneath bridges, predatory Adults nectar at flowers and mud nests are built in sheltered locations such as garages and underneath bridges, predatory Bitsch and Barbier (2006), Bogusch et al. (2005), Castro (2007), Cetkovic et al. (2004), Ebmer (1995), Gonseth et al. (2001), Rahola (2005), van der Vecht (1984) Cetkovic et al. (2004) AT, BE, BG, CH, G, I2 CZ, DE, ES, FI, FR, GB, GR, HR, HU, IT, LT, MD, NL, PL, RO, SK, UA AT G3, G5 Robinia Ermolenko and Sem’yanov (1981), Markó et al. (2006) Larix Pschorn-Walcher and Zinnert (1971) 1912, HU CZ, HU, SK, UA I Tetramesa on Poaceae 1966, DE BG, DE, FR, RU F, I2, E5 Rosa Boucek (1968), Erdös (1954), Grissell (1995) Roques and Skrzypczynska (2003) A parasitic/ predator A parasitic/ predator Asia1998, ME Temperate FR, ME A phytophagous North America A phytophagous Asia1971, AT Temperate A parasitic/ predator phytophagous North America North America A Habitat 1825, DE C3, X25 C3, X25 Hymenoptera. Chapter 12 Tenthredinidae Nematus (Pteronidea) tibialis Newman, 1837 Status 755 A phytophagous Native range North America Megastigmus borriesi Crosby, 1913 A phytophagous Asia1969, FINTemperate ALA DK, FI-ALA, RU Megastigmus lasiocarpae Crosby, 1913 Megastigmus milleri Milliron, 1949 A phytophagous phytophagous North America North America 1969, FINALA 1952, GB FIN-ALA DK, FR, NL, GB G3, G4, X11 Abies Megastigmus nigrovariegatus Ashmead, 1890 Megastigmus pinsapinis Hoffmeyer, 1931 A phytophagous North America 1987, FR FR E5 Rosa A phytophagous Africa 1858, FR ES, FR, IT G3, G4, X11 Cedrus Megastigmus pinus Parfitt, 1857 A phytophagous North America 1931, GB BE, CZ, DE, DK, G3, G4, FR, GB, IE, NL, SE X11 Abies Megastigmus rafni Hoffmeyer, 1929 A phytophagous North America 1930, GB BE, DE, DK, FR, GB, NL G3, G4, X11 Abies Megastigmus schimitscheki Novitzky, 1954 Megastigmus specularis Walley, 1932 A phytophagous Asia1990, FR Temperate FR G3, G4 Cedrus A phytophagous North America DK, FI, FR, RU, SE G3, G4, X11 A Regime First Record Invaded countries in Europe 1954, DE CZ, DE, DK, FR, GB, PL, RU 1920, FINALA Habitat Host G3, G4, X11 Picea, Pinus strobus X11 Abies Abies Abies References Jensen and Ochsner (1999), Roques and Skrzypczynska (2003) Annila (1970), Jensen and Ochsner (1999), Ochsner (1998) Annila (1970) Jensen and Ochsner (1999), Roques and Skrzypczynska (2003) Roques and Skrzypczynska (2003) Pintureau et al. (1991), Roques and Skrzypczynska (2003), Skrzypczynska and Mazurkiewicz (2002) Jensen and Ochsner (1999), Roques and Skrzypczynska (2003) Jensen and Ochsner (1999), Roques and Skrzypczynska (2003) Roques and Skrzypczynska (2003) Jensen and Ochsner (1999), Roques and Skrzypczynska (2003) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Status 756 Families Species Megastigmus atedius Walker, 1851 Native range North America Megastigmus transvaalensis (Hussey, 1956) Trichogrammatidae Megaphragma mymaripenne Timberlake, 1924 Oligosita distincta (Silvestri, 1915) Oligosita sanguinea (Girault, 1911) Trichogramma achaeae Nagaraja & Nagarkatti, 1970 Trichogramma chilonis Ishii, 1941 A phytophagous Africa A parasitic/ predator AsiaTropical 1995, IT IT-SIC, IT I A parasitic/ predator parasitic/ predator parasitic/ predator Africa 1939, FR FR, SE I North America Asia 1949, HU HU I 1987, FR FR I A parasitic/ predator Asia 1985, DE DE, RO I1 Trichogramma dendrolimi Matsumura, 1926 A parasitic/ predator Asia 1978, BG AT, BE, BY, BG, DE, FR, GR, HU, IT, LT, LV, MD, RO, RU, UA I, G A A First Record Invaded countries Habitat in Europe G3, G4, 1896, GB AT, BE, CH, CZ, X11 DE, DK, EE, ES, FI, FR, GB, HU, IE, IT, ME, NL, NO, PL, PT, RO, RS, RU, SE, SK, UA 1962, ESES, ES-CAN, FR, I2, G5 CAN PT Host References Pseudotsuga Mailleux et al. (2008), Roques and Skrzypczynska (2003) Schinus Grissell and Prinsloo (2001), Scheffer and Grissell (2003) Thrips (Egg parasitoid) Sinacori et al. (1999), Viggiani and Bernardo (1996) Leafhoppers (Egg Hedqvist (2003), Nowicki parasitoid) (1940) Cicadellid in wheat Erdös (1956) (Egg parasitoid) Stem-borer (Egg Voegelé et al. (1988) parasitoid) Hymenoptera. Chapter 12 Families Status Regime Species Megastigmus A phytospermotrophus Wachtl, phagous 1893 Cabbage moths, Glas and Hassan (1985) cotton bollworm, maize pyralid, armyworm Lepidoptera, e.g. Babi et al. (1984), Wetzel Epichoristodes acerbella Dickler (1994) 757 C parasitic/ predator Trichogramma perkinsi Girault, 1912 Trichogramma pretiosum Riley, 1879 Uscana johnstoni (Waterston, 1926) Uscana semifumipennis Girault, 1911 Vespidae Vespa velutina nigrithorax du Buysson, 1905 A C A A A Regime Native range Cryptogenic First Record Invaded countries in Europe 1957, CZ CZ, DE, ES, FR, GB, GR, IT Habitat parasitic/ predator parasitic/ predator parasitic/ predator parasitic/ predator Asia 1984, FR FR I1 Cryptogenic Africa 1975, GR ES, GR, YU I1 1970, RO RO J Maize borer and forest CIBC (1976), Herting moths (1975), Thompson (1958), Viggiani and Laudonia (1989) Lepidopteran pests Voegelé et al. (1988) (highly polyphagous) Cotton leafworm Danon (1989), Stavraki (1976) Bruchinae Botoc (1971) North America 1963, HU HU J Bruchinae Reichart (1964) parasitic/ predator Asia2004, FR Temperate FR G Woodland Haxaire et al. (2006), Villemant et al. (2006) I1, G Host References Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Status 758 Families Species Trichogramma minutum Riley, 1871 Table 12.2. Hymenoptera species alien in Europe. List and characteristics. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Last update 01/03/2010. Families Species Aphelinidae Eretmocerus mundus Mercet, 1931 First Record Invaded countries Habitat Host References E parasitic/ predator Mediterranean region Unknown DE, NL J100 Cotton whitefly, Bemisia, Trialeurodes Drost et al. (1996) E phytophagous phytophagous phytophagous phytophagous phytophagous Europe 2001, DK DK, PT I Pedersen (1996) Europe 1987, DK DK, PT I Europe 2005, AL AL, GL I Europe 1959, IS IS I Europe 1979, IS IS I Pollinator of various cultivated plants Pollinator of various cultivated plants Pollinator of various cultivated plants Pollinator of various cultivated plants Pollinator of various cultivated plants Prys-Jones et al. (1981) E E E E Pedersen (1996) Prys-Jones et al. (1981) E phytophagous Europe 2000, GB GB I2 Berberis Fitton et al. (1978) E parasitic/ predator Europe 2005 PTAZO PT-AZO, GB J Insects in wood furnitures; cause dermatitis in human by stings Fitton et al. (1978) E phytophagous Europe 1905, GB I2, D2 Athyrium ferns (Leaf miner) Schedl (1974) 759 Blasticotomidae Blasticotoma filiceti Klug 1834 Native range Hymenoptera. Chapter 12 Apidae Apis mellifera carnica (Pollmann, 1879) Apis mellifera ligustica (Spinola, 1806) Apis mellifera mellifera Linnaeus, 1758 Bombus hortorum (Linnaeus, 1761) Bombus lucorum (Linnaeus, 1761) Argidae Arge berberidis Schrank, 1802 Bethylidae Sclerodermus domesticus Klug, 1809 Status Regime Status Regime Native range First Record Invaded countries Habitat Host References parasitic/ predator Asia1915, HU Temperate AT, HR, HU, IT F6 Bees Pagliano et al. (2000) E phytophagous phytophagous phytophagous phytophagous phytophagous phytophagous Europe 1735, GB GB, IE G Quercus Fitton et al. (1978) Europe G,I2 Quercus Fitton et al. (1978) Europe Unknown, GB GB 1735, GB GB G Quercus Fitton et al. (1978) Europe 1735, GB GB I2 Quercus Fitton et al. (1978) Europe Unknown GB, IE I2 Quercus Fitton et al. (1978) Europe 1993, GB GB G Quercus Fitton et al. (1978) Europe Unknown, IE Unknown, GB Unknown, GB Unknown, GB Unknown IE G3 Pinus Fitton et al. (1978) GB G3 Pinus Fitton et al. (1978) GB G3 Picea Fitton et al. (1978) GB G3 Pinus Fitton et al. (1978) IE, GB G3 Pinus Fitton et al. (1978) E E E E E E E E E E phytophagous phytophagous phytophagous phytophagous phytophagous Europe Europe Europe Europe Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) E 760 Families Species Chrysididae Chrysis marginata Mocsary, 1889 Cynipidae Andricus corruptrix (Schlechtendal, 1870) Andricus grossulariae Giraud,1859 Andricus kollari (Hartig 1843) Andricus lignicola (Hartig,1840) Andricus quercuscalicis (Burgesdorff 1783) Aphelonyx cerricola (Giraud 1859) Diprionidae Diprion pini (Linnaeus, 1758) Diprion similis (Hartig, 1836) Gilpinia hercyniae (Hartig, 1837) Gilpinia virens (Klug, 1812) Neodiprion sertifer (Geoffroy, 1785) Families Species Encyrtidae Ageniaspis fuscicollis (Dalman, 1920) Status Regime First Record Invaded countries Habitat Host References AU, BE, BY, CH, CZ, DE, DK, EE, ES-CAN, FI, GB, HU, IS, IE, LT, LV, LU, MD, NL, NO, NO-SVA, PL, PT-AZO, PT-MAD, RO, RU, SE, SK, UA CZ, ES-CAN, FR, HR, IL, MD, ME, NL, PT, RU, SE, YU I Prays oleae on Citrus and yponomeutids Koscielska (1963), Kuhlmann (1994), Nénon (1978) J100 Pseudococcids on Citrus and many crops Tingle and Copland (1988) E parasitic/ predator Mediterranean region 1735, GB E parasitic/ predator Mediterranean region 1994, PT Eulophidae Thripastichus gentilei (Del Guercio, 1931) E parasitic/ predator Europe 1930, IT DE, FR, IT, YU I Thrips Del Guercio (1931), Domenichini et al. (1964), Herting (1971) Eurytomidae Bruchophagus robiniae Zerova, 1970 E parasitic/ predator Europe 1969, UA BG, UA, G5 Seed feeder on Robinia pseudoacacia Stojanova (1997), Zerova (1970) Formicidae Aphaenogaster senilis Mayr, 1853 E parasitic/ predator 2005, PTAZO PT-AZO, U Natural habitat, garrigue Wetterer et al. (2004) E parasitic/ predator parasitic/ predator parasitic/ predator Mediterranean region Europe Unknown DE, GB J Trees Bernard (1968) E1, H5 Warm, dry, stony environnements Meadows, dry grasslands, Forest borders Collingwood (1958) Anagyrus pseudococci (Girault, 1915) E E Europe Europe Unknown, IE IE Unknown, IE IE E1, E5 Collingwood (1958) 761 Crematogaster scutellaris (Olivier, 1792) Lasius alienus (Foerster, 1850) Lasius flavus (Fabricius, 1781) Hymenoptera. Chapter 12 Native range E E parasitic/ predator parasitic/ predator E parasitic/ predator A phytophagous E parasitic/ predator phytophagous phytophagous phytophagous phytophagous phytophagous E E E E E Native range Europe Mediterranean region Europe First Invaded countries Record Unknown, IE IE Unknown BE, BG, DE, GB, HU, PL, RU Habitat Host References E5 Trunks and stumps, forest borders Dry and warm areas Edwards (1997) G Geiter et al. (2002) 1847, PTMAD ES-CAN, GB, PT-AZO, G, J1, I2 Gardens, urban, arid PT-MAD sites Wetterer et al. (2004) Europe Unknown RU I Pollinator of alfalfa Pesenko and Astafurova (2003) Europe Unknown GB G3 Pinus Fitton et al. (1978) Europe 1986, NL BE, NL G3 Larix Magis (1988) Europe 1986, NL NL G3 Picea Europe 1988, BE BE, LU G3 Picea van Achterberg and van Aartsen (1986) Magis (1988) Europe 1986, NL BE, NL G3 Picea Magis (1988) Europe 1941, NL BE, DK, GB, LT, NL, SE, UA G3 Larix Billany and Brown (1980) PT-AZO J Stored products weevils, Sitophilus, in grain G3 Conifers E parasitic/ predator Europe 2005, PTAZO E phytophagous Europe Unknown, GB GB Fitton et al. (1978) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Tetramorium caldarium (Roger, 1857) Megachilidae Megachile rotundata (Fabricius, 1787) Pamphiliidae Acantholyda erythrocephala L. 1758 Acantholyda (Itycorsia) laricis (Giraud, 1861) Cephalcia abietis (Linnaeus, 1758) Cephalcia alpina (Klug, 1808) Cephalcia erythrogaster (Hartig, 1837) Cephalcia lariciphila (Wachtl, 1898) Pteromalidae Lariophagus distinguendus (Förster, 1841) Siricidae Sirex juvencus (Linnaeus, 1758) Status Regime 762 Families Species Lasius fuliginosus (Latreille, 1798) Ponera coarctata (Latreille, 1802) Athalia rosae (Linnaeus, 1758) Hoplocampa brevis (Klug, 1816) Nematus (Pteronidea) spiraeae Zaddach, 1883 Pachynematus (Epicenematus) montanus (Zaddach, 1883) Pachynematus (Larinematus) imperfectus (Zaddach, 1876) Pachynematus (Pikonema) scutellatus (Hartig, 1837) Status Regime E E E phytophagous phytophagous phytophagous Native range Europe First Record Unknown Europe Europe Invaded countries Habitat Host References GB G3 Pinus, Abies, Larix Fitton et al. (1978) Unknown, GB GB 1951, GB GB G3 Conifers Fitton et al. (1978) G3 Conifers Fitton et al. (1978) E phytophagous Europe Unknown, GB GB I2 Viola Fitton et al. (1978) E phytophagous Europe 1953, GB G3, I2 Larix E phytophagous phytophagous phytophagous phytophagous Europe I,J Brassica, Sinapis Europe Unknown, GB GB 1935, GB GB Leston (1988), Piekarczyk and Wright (1988), Speight (1979) Fitton et al. (1978) I2, G5 Pyrus Fitton et al. (1978) Europe 1824, GB I2 Spiraea, Aruncus Fitton et al. (1978) Europe Unknown, GB GB G3 Picea Fitton et al. (1978) E E E DK, EE, GB, HU, IE, SE GB E phytophagous Europe 1915, DK BE, DK, GB, HU, LV, NL G3, G5 Larix Fitton et al. (1978) E phytophagous Europe Unknown GB, IE G3 Picea Fitton et al. (1978) Hymenoptera. Chapter 12 Families Species Sirex noctilio Fabricius, 1773 Urocerus gigas (Linné, 1758) Xeris spectrum (Linnaeus, 1758) Tenthredinidae Ametastegia (Protemphytus) pallipes (Spinola, 1808) Anoplonyx destructor Benson, 1952 763 Native range Europe Europe First Invaded countries Record Unknown, GB GB 1846, GB GB Habitat Host References I,J Digitalis, Plantago Fitton et al. (1978) I2, G1 Polygonatum Fitton et al. (1978) Unknown, IE IE G3 Picea Europe Unknown, GB GB G3 Picea Fitton et al. (1978) Europe 1906, GB DK, EE, ES, GB, IE, LV, G3, I2, NL, NO, SE FB Larix Fitton et al. (1978) Europe 1954, GB GB G3 Larix Fitton et al. (1978) Europe Unknown, GB GB G3 Picea Fitton et al. (1978) Europe 1949, GB GB G3 Picea Fitton et al. (1978) Europe 1915, DK BE, BY, DK, EE, GB, NL, SE, GB G3, I2, FB Larix Fitton et al. (1978) Europe 1915, DK BE, DK, EE, ES, GB, HU, IE, ME, NL, RS, SE, UA G3, FB, I2 Larix Fitton et al. (1978) Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Europe 764 Families Status Regime Species Pachyprotasis variegata E phyto(Fallen, 1808) phagous Phymatocera aterrima E phyto(Klug, 1816) phagous Pristiphora E phyto(Lygaeonematus) abietina phagous (Christ, 1791) Pristiphora E phyto(Lygaeonematus) phagous compressa (Hartig, 1837) Pristiphora E phyto(Lygaeonematus) phagous erichsonii (Hartig, 1837) Pristiphora E phyto(Lygaeonematus) glauca phagous Benson, 1954 Pristiphora E phyto(Lygaeonematus) saxesenii phagous (Hartig, 1837) Pristiphora E phyto(Lygaeonematus) phagous subarctica (Forsslund, 1936) Pristiphora E phyto(Lygaeonematus) wesmaeli phagous (Tischbein, 1853) Pristiphora E phyto(Oligonematus) laricis phagous (Hartig, 1837) Trichogrammatidae Trichogramma brassicae Bezdenko, 1968 First Record 1995, FI Invaded countries Habitat Host References EE, FI FA, I2 Lindqvist (1974) Europe 2004, GB GB G3, G4 Spiraea chamaedryfolia Tilia Fitton et al. (1978) Europe 1946, GB GB I2 Thalictrum Fitton et al. (1978) Europe Unknown, GB GB G3 Picea Fitton et al. (1978) Europe Unknown, GB GB G3 Picea Fitton et al. (1978) Europe 1879, GB IE, GB Larix Europe 1943, IE IE, GB G3, G4,X11 G3, G4,X11 G5, I2, X15 Roques and Skrzypczynska (2003) Roques and Skrzypczynska (2003) Rasplus et al. (2000), Roques and Skrzypczynska (2003) Asia1915, SI Temperate AL, BA, BG, ES, FRCOR, FR, GR, HR, IL, IT, ME, MT, PT, RO, RS, SI AT, BG, CH, DE, ES, FR, NL, RO E parasitic/ predator Europe 1996, DE E parasitic/ predator parasitic/ predator Europe Unknown, IS IS Unknown FÖ, IS E Eurasia Abies Cupressus I1 Ostrinia corn borer but highly polyphagous Pintureau (2008) G3, G4 Woodland Olafsson (1979) H, X25 Woodland Olafsson (1979) 765 Vespidae Vespula germanica (Fabricius, 1793) Vespula vulgaris (Linné, 1758) Native range Europe Hymenoptera. Chapter 12 Families Status Regime Species Pristiphora (Pristiphora) E phytoangulata Lindqvist, 1974 phagous Pristiphora (Pristiphora) E phytoleucopus (Hellén, 1948) phagous Pristiphora (Pristiphora) E phytothalictri (Kriechbaumer, phagous 1884) Pristiphora (Sharliphora) E phytoamphibola (Förster, phagous 1854) Pristiphora (Sharliphora) E phytonigella Förster, 1854) phagous Torymidae Megastigmus pictus E phyto(Förster, 1841) phagous Megastigmus suspectus E phytoBorries, 1895 phagous Megastigmus wachtli E phytoSeitner, 1916 phagous 766 Jean-Yves Rasplus et al. / BioRisk 4(2): 669–776 (2010) Table 12.3. Number of alien Hymenoptera per European countries. Countries Italy mainland France mainland Spain mainland Israel Germany mainland Greece mainland Great Britain Czech Republic Netherlands Denmark Italy Sicily Portugal mainland Russia Belgium Austria Hungary Spain Canary islands Switzerland Poland Sweden Cyprus Bulgaria Ukraine France Corsica Romania Portugal Madeira Slovakia Albania Former Yougoslavia Serbia N 144 111 90 82 80 50 45 41 40 36 36 35 33 32 31 30 30 30 26 23 23 22 22 19 18 18 18 17 14 14 Countries Finland mainland Italy Sardinia Montenegro Spain Balearic islands Croatia Norway mainland Ireland Malta Moldova Slovenia Lithuania Portugal Azores Greece Crete Estonia Luxemburg Greenland Iceland Belarus FinlandAland Greece South Aegean Isl Latvia Bosnia Feroe Islands Greece North Aegean Isl Norway Svalbard Andorra FYRM Macedonia Greece Ionian islands Lichtenstein N 13 13 11 11 10 10 10 8 8 8 7 7 6 5 4 3 2 2 2 2 1 1 1 1 1 0 0 0 0 A peer reviewed open access journal BioRisk 4(2): 767–791 (2010) RESEARCH ARTICLE doi: 10.3897/biorisk.4.59 BioRisk www.pensoftonline.net/biorisk Thrips (Thysanoptera) Chapter 13.1 Philippe Reynaud Laboratoire national de la protection des végétaux, Station d’Angers, 7 rue Jean Dixméras, 49044 Angers Cedex 01, France Corresponding author: Philippe Reynaud (philippe.reynaud@agriculture.gouv.fr) Academic editor: David Roy | Received 27 January 2010 | Accepted 25 May 2010 | Published 6 July 2010 Citation: Reynaud P (2010) Thrips (Thysanoptera). Chapter 13.1. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(2): 767–791. doi: 10.3897/biorisk.4.59 Abstract Thrips (Order Thysanoptera) are found worldwide and include almost 6000 species. Several of them are notorious for causing extensive crop damage (by feeding on leaf tissue or by vectoring viral disease). Their small size (usually less than 2 millimeters) and cryptic habits have facilited invasions and establishment in Europe in the wild or in greenhouses. Fifty-two alien species, belonging to four families have been recorded within Europe. Species introduced before 1950 mostly originate from America, tropical and subtropical areas and subsequent arrivals generally originate from Asia (and from America to some extent). Five countries host more than 30% of the European alien thrips fauna and two alien thrips occur in more than 50% of the countries and islands of Europe. Keywords Thysanoptera, thrips, alien, Europe 13.1.1. Introduction Thrips (Order Thysanoptera) are ubiquitous, small to minute (a few millimeters long) and slender-bodied insects with fringed wings. The morphology is reduced: thrips have only one functional mandibular stylet, the second being greatly reduced, thus forming asymmetrical suctorial mouthparts compacted within a short cone-shaped rostrum. About 50% of the known species of Thysanoptera feed on fungi, approximately 40% feed on living tissues of dicotyledonous plants or grasses, and the remainder exploit Copyright Philippe Reynaud. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 768 Philippe Reynaud / BioRisk 4(2): 767–791 (2010) mosses, ferns, gymnosperms, cycads, or are predatory (Morse and Hoddle 2006). Less than 1% of described thrips species are serious pests and most economic literature deals with just four species (Mound and Teulon 1995). The almost 6000 known species of thrips are at present arranged into two suborders (Terebrantia and Tubulifera) and nine families, but disagreement exists concerning the family classification system (Mound 2007). Phlaeothripidae is the largest family and the sole family in the suborder Tubulifera with about 3500 described species (Mound and Morris 2007). The other eight families are all included in the suborder Terebrantia (2400 species). Members of the Merothripidae (15 species) and Uzelothripidae (1 species) are all very small thrips associated with fungal hyphae in warm countries. In contrast, members of the Melanthripidae (65 species) are usually large and robust, and they all breed in flowers, and occur in temperate areas. The Aeolothripidae (190 species) is a rather larger family of mainly phytophagous species feeding on flowers, or non-obligate predators of other arthropods. The species of the next three families are poorly known, Fauriellidae (5 species) from California, southern Europe and South Africa. Adiheterothripidae (6 species) are known only from the flowers of date palms, Phoenix dactylifera and Heterothripidae (71 species), are found only in the New World and, with one exception, all species live within flowers. The eighth family, with nearly 2100 known species is by far the largest within Terebrantia : Thripidae are found worldwide and include almost all of the pest species of thrips, many of them feed and breed on both leaves and in flowers. 13.1.2 Taxonomy of the Thysanoptera species alien to Europe The 52 species of Thysanoptera alien to Europe belong to four different families (Table 13-1) but two of them (Phlaeothripidae and Thripidae) include more than 99% of the alien species. Suborder Tubulifera Phlaeothripidae: The traditional classification of Tubulifera comprises a single family with two subfamilies. All members of the smaller subfamily, the Idolothripinae, feed on fungal spores and live on dead twigs, in leaf litter or within the bases of grass and sedge tussocks. The spore-feeding Nesothrips propinquus is the unique alien species among less than 30 european species and is widely distributed in countries occuring along the sailing route from New Zealand to Europe, presumably in hay and straw (Mound 2006). It can be found on citrus fruits in its native habitat but there is no evidence of producing any damage (Blank and Gill 1997). Phlaeothripinae is the main subfamily of Phlaeothripidae, with 2800 species (Mound and Morris 2007). They exhibit a wide range of biologies: a few are predatory, some are flower feeders but in most cases, they are leaf feeding or associated with fungi in leaf litter or on dead wood. Fourteen species Thrips (Thysanoptera). Chapter 13.1 769 belonging to ten genera are here considered to be alien species in Europe (from a total of around 180 native species). Among them, five species prey upon small arthropods (including scale insects), five species are detrivorous and four species are known to be phytophagous, including Gynaikothrips ficorum which is recognized as a pest on Ficus (preferred host) and other hosts. Suborder Terebrantia Merothripidae: This family of three genera, with 15 fungus-feeding species that live on dead twigs and in leaf-duff, is found mainly in the Neotropics (Hoddle et al. 2004). Merothrips floridensis is the unique representant of this family in Europe. This is an interesting example of a small and usually wingless species with a scattered distribution, probably associated with trading routes and commercial traffic of hay, dead wood and living plants (Mound 1983). Aeolothripidae: Until recently, Melanthripidae was included in this family. However, a morphology-based distinction with the Aeolothripidae is now well supported (Mound and Morris 2007). Typical Aeolothripidae are generally regarded as facultative predators on other small arthropods but with a few exceptions. They are mainly distributed in the temperate parts of the world, although members of several genera are restricted to the tropics. This is the case of the two alien species of ant-mimicking thrips (Franklinothrips vespiformis and Franklinothrips megalops) recorded in Europe, that have been marketed or tested as biocontrol agents in glasshouses (Mound and Reynaud 2005). Thripidae: Four sub-families are currently recognized worldwide. Each of these is represented by alien species in Europe. Dendrothripinae are small in size and live on young leaves. They have been defined by the presence of a remarkably elongate metasternal endofurca associated with a jumping habit. There are two alien species, Leucothrips nigripennis and Pseudodendrothrips mori, compared to eight native species. Panchaetothripinae are strongly reticulate thrips and are regarded as leaf feeders with a tropical or subtropical distribution. They are well represented amongst alien species (eight species) because they are able to breed on ornemental plants in European greenhouses. There are no native species in Europe with one exception in the canary Islands and Madeira. Sericothripinae are a small sub-family in Europe with only two genera and eight species, including one recently described alien (Neohydatothrips samayunkur). The species are all phytophagous in flowers and on leaves. The subfamily Thripinae is the main sub-family in Europe with 59 genus and more than 240 native species and the main group of aliens in Thysanoptera with 18 genera and 24 species. Thripinae feed and breed both on leaves and in flowers and a few are specialized predators. Some thrips species transmit plant viruses. They are all included in this subfamily. Thrips-transmitted viruses can cause signiﬁcant diseases of many crop plants and their impact worldwide is immense. In Europe, seven thrips species are known vectors of 770 Philippe Reynaud / BioRisk 4(2): 767–791 (2010) Figure 13.1.1. Relative importance of the families of Thysanoptera in the alien and native entomofauna in Europe. Families are presented in a decreasing order based on the number of alien species. Species alien to Europe include cryptogenic species. The number over each bar indicates the number of species observed per family. virus including five alien species: three species of Frankliniella, one species of Thrips and Microcephalothrips abdominalis (Jones 2005). Western ﬂower thrips, Frankliniella occidentalis is one of the most important pests of greenhouse crops, especially in ornamental species. 13.1.3 Temporal trends of introduction in Europe of alien thrips Because of their small size, ability to reach high numbers, cryptic behavior, egg deposition inside plant tissue (e.g., all Terebrantia), and a propensity to secrete themselves in tight spaces (Morse and Hoddle 2006), thrips remain inconspicuous insects. The accurate recognition of alien Thysanoptera species is also a major challenge because of the difficulty of a morphometric identification (close morphological similarity) for non-specialists. There is also a lack of taxon specialists that are needed to study newly recorded species, confounded by the lack of identification keys in local monographs. Thrips identification requires significant experience, encyclopaedic knowledge, a good reference collection and relevant literature. Molecular and visual online-identification tools of the main pest thrips are now available but are not yet widely used. For the reason above, it is likely that the real number of of alien thrips species present in Europe is greatly underestimated. The date of the first record in Europe is also unknown for seven species (13.5%). The first alien thrips species (Heliothrips haemorrhoidalis, called the greenhouse thrips) was discovered and originally described by Bouché in Germany in the first half of the 19th century from specimens taken from a greenhouse. This species was probably introduced into Europe on ornamental plants from tropical America. H. haemorrhoidalis is now widespread in Europe indoors and Thrips (Thysanoptera). Chapter 13.1 771 can be found outdoors in the southern countries. Before the First World War, seven different tropical thrips were recorded as minor pests or useful predators, always collected under protected conditions. The first outdoor alien species collected in Europe was the Thripinae Stenchaetothrips biformis, a major pest of rice in Asia, described in England and collected later in several european countries. S. biformis sensu stricto is common in vegetative shoots of Phragmites australis in temperate Europe, even though S. biformis ‘rice form‘ is common on Oryza sativa in Asia and South America (Vierbergen 2004). From 1950, a clear acceleration of thrips introductions is evident (Figure 13.1.2), with a new alien species every two years on average and as many as one new alien species per year during the period 1975 - 1999. The main event during this period was the occurrence of the western flower thrips Frankliniella occidentalis in the Netherlands in 1983, originating from western North America. By 1986, it was reported in Sweden and Denmark and, by 1987, it had reached France and Spain. Since then, it has been reported from most European countries and has become a major pest of agricultural and horticultural crops throughout. Since 2000, three non-native Thysanoptera are recorded, with a somewhat smaller rate of discovery compared with the previous period. 13.1.4. Biogeographic patterns of the thrips species alien to Europe 13.1.4.1 Origin of alien species Exact knowledge of the geographical origin of alien thrips species is a vital step in enforcement of scientifically based plant quarantine and free trade protocols. Unfortunately, the area of origin of alien thrips remains unclear in 13.5% of cases. Many alien species were first described in Europe, but were undoubtedly native from other continents. Kelly’s citrus thrips (KCT) was thus ﬁrst collected in October 1914 in Queensland (Australia), described as Physothrips kellyanus by Bagnall in 1936 and known only from Australia in the last 36 years. After taxonomic studies, KCT was transferred to Pezothrips, a new genus including nine Palaearctic species. The morphological similarity of KCT to the eight Pezothrips species from the southern Palaearctic suggests that P. kellyanus itself originated in that part of the world. But KCT is not known to breed on any endemic plant in Mediterranean countries even when KCT larvae and adults have been found on australian endemic plants such as Myoporum insulare (Myoporaceae) (Webster et al. 2006). KCT is a good example of a thrips species with an unclear origin. The spread may have had more than one origin and the source of reintroductions of many plant pests and pathogens has changed over time. For example, Frankliniella occidentalis originally from the USA, was introduced to the UK from the Netherlands, and is reintroduced from several tertiary sources, such as Kenya (Perrings et al. 2005). Alien thrips come mainly (65.4%) from Asia, Central and South America and North America (Figure 13.1.3). Temporal analysis shows that Central and South America and Africa were the main source of introductions before 1900, followed by species 772 Philippe Reynaud / BioRisk 4(2): 767–791 (2010) Figure 13.1.2. Temporal changes in the mean number of records per year of Thysanoptera species alien to Europe from 1492 to 2007. The number over each bar indicates the absolute number of species newly recorded per time period. of mainly tropical, subtropical and Australasian origins between 1900 and 1950. After that date, non-indigenous thrips mostly originate from Asian and secondarily from North America. 13.1.4.2 Distribution of alien species in Europe Figure 13.1.4 presents the colonization of European countries and main islands by alien thrips. Countries can be divided into the following categories: – 13 countries with no known alien species. They include particulary small countries, some small southern islands, northern islands and a large northern country, Belarus. – 21 countries which host less than 10% of the known invasive thrips in Europe. This category comprises large countries, probably poorly sampled by entomologists (Greece) or northern countries (Poland, Ukraine, Austria) and large islands which have been poorly surveyed. – 17 countries with 10% to 30% of the known invasive thrips. This group generally consist of large countries (Germany, Spain, Sweden, Norway, Finland) but also includes small southern islands (Azores, Madeira, Canary islands) well sampled by entomologists and with a favourable climate for exotic thrips. – 5 countries with more than 30% of the known European alien thrips fauna. Three large countries are involved, two with varied but favourable climate (Italy and France) and two with a long tradition of thysanopterologists (Great Britain and Thrips (Thysanoptera). Chapter 13.1 773 Figure 13.1.3. Origin of the 52 alien species of Thysanoptera established in Europe. Numbers indicate the relative proportion of alien species originating from a given region. Germany). Lastly, Netherlands, owing to its open economy and international trade, records 20 alien thrips species. Surprisingly, there is no significant relationship between country surface area and number of alien species (Figure 13.1.5, r2 = 0.2522). For instance, Netherlands and Italy harbour the same number of non-native thrips, but Netherland surface is only 14% of of the area of Italy. Only two alien thrips (Frankliniella occidentalis and Heliothrips haemorrhoidalis) occur in more than 50% of the countries and islands of Europe and a quarter of the species are known from a single country. There is no clear relationship between the date of first record and the number of contaminated countries. 13.1.5. Pathways of introduction in Europe of alien thrips species Adults and larvae of Thysanoptera are very small, highly thigmotactic, and often lay minute eggs within plant material (e.g. petioles, stems, leaves and fruit) making rapid visual detection impossible. As a consequence, accidental introduction in Europe is the rule for non-native Thysanoptera (94%) and intentional introduction is confirmed for only three species (Franklinothrips vespiformis, Franklinothrips megalops and Karnyothrips melaleucus). The global trade in ornamental greenhouse plants is clearly the main pathway for non-native thrips: all widespread alien species in Europe are greenhouse pests or predators. It also means that after introduction, domestic trade of ornamental plants inside Europe is a major pathway for the transport of thrips. Greenhouse environments eliminate climatic barriers to establishment (e.g., H. haemorrhoidalis) and may also provide important overwintering sites from which outdoor populations establish in spring to attack vegetable crops (e.g., F. occidentalis in northern Europe) (Morse and Hoddle 2006). 774 Philippe Reynaud / BioRisk 4(2): 767–791 (2010) Figure 13.1.4. Comparative colonization of continental European countries and islands by the thrips species alien to Europe. Archipelago: 1 Azores 2 Madeira 3 Canary islands. 13.1.6. Ecosystems and habitats invaded in Europe by alien thrips species Although thrips are known as inhabitants of flowers, they are also abundant and diverse in other microhabitats. They are phytophagous insects, sap suckers (some of which feed on aquatic plants), but can also work as decomposers, fungivores, pollinators, predators on insects and mites, whilst one species was recently discovered as an ectoparasite under the wings of a bug. Alien thrips are mostly phytophagous (75%) and seldom predators (13.5%) or detritivores (11.5%). Cultivated habitats are preferentially (94.2%) invaded by exotic thrips, including greenhouses that provide suitable habitat for 55.8% of the invasive species in Europe (Figure 13.1.5). Nevertheless, we can assume that thrips species such as spore and fungal feeders are underestimated in faunal studies, because these ecosystems are usually less investigated by thysanopterologists. Similarly, the wild flora that surrounds areas of crops is rarely Thrips (Thysanoptera). Chapter 13.1 775 Figure 13.1.5. Relationships bewteen the size of the European countries and the number of alien Thysanoptera observed in the country. best fit: Y= 2E-05x + 3.5957; r= 0.2522) sampled. It may also be important in facilitating the spread and colonization of new ecosystems. The remaining habitats (13.5%) include deciduous wooded habitats, dry grasslands or unknown habitats. 13.1.7. Ecological and economic impact of alien thrips species Three major food sources are used by thrips: fungal hyphae and spores, green leaves, and flowers with or without leaves as well. A few species are also predators, and a very few feed only on mosses (Mound and Marullo 1996). More than 95% of Terebrantia are associated with vascular plants, whereas about 60% of Tubulifera species are fungivores (Mound 2002). But of an estimated 8000 extant species of thrips (Lewis 1997) and more than 5500 species that are described, scarcely 1% are recorded as serious pests, mainly in the Thripidae family. Thrips can affect plants by direct feeding, which may leave visible signs of damage, such as leaf silvering. Many tubuliferans also cause galls1. A few thrips transmit plant viruses and can cause signiﬁcant diseases of many crop plants and their impact worldwide has been judged to be substantial (Jones 2005). Thrips can also be considered as pests through their habit of crawling into small spaces, a behavior known as thigmotaxis. This behaviour can trigger smoke detectors and fire alarms and thus cause considerable inconvenience. Similarly, thrips can invade computers, watches, paintings, polystyrene building insulation, hypodermic needles in manufacture, and many other unlikely places (Hoddle et al. 2008). Thrips may also become a nuisance when they swarm and land on exposed areas of skin but humans 1 Not all plant feeding by thrips is disadvantageous: attempts have been made in USA to control alligator weed (Alternanthera philoxeroides) by Amynothrips andersoni imported from Argentina. 776 Philippe Reynaud / BioRisk 4(2): 767–791 (2010) Figure 13.1.6. Main European habitats colonized by the established alien species of Thysanoptera. The number over each bar indicates the absolute number of alien thrips recorded per habitat. Note that a species may have colonized several habitats. are usually unintended, occasional, short-term hosts without medical consequences (Faulde et al. 2007). Throughout the world, only six of the 210 described species of Frankliniella are known to be vectors of viruses, only four of the 290 species of the genus Thrips, and just one of the 100 species of Scirtothrips. In addition, one species of Ceratothripoides and Microcephalothrips abdominalis are known to transmit virus. Thrips transmit plant viruses in the Tospovirus, Ilarvirus, Carmovirus, Sobemovirus and Machlomovirus genera (Jones 2005). Of over 52 species of alien thrips, less than 10 can be considered as having an impact on human activities. The ecology and biology of other species is generally poorly known and ecological and economic impact cannot be evaluated. Various members of the genus Franklinothrips are of economic importance (Mound and Reynaud 2005). F. vespiformis is recently marketed in continental Europe and Israel as a biocontrol agents in greenhouses for the control of thrips and mite pests; its prey also includes whiteﬂies and leafminers (Larentzaki et al. 2007). Frankliniella occidentalis (the Western flower thrips) is a major worldwide crop pest with a huge economic impact and has become a key pest in a large range of agricultural and floricultural production areas in the world (see factsheet 14.78). It has a very extensive host range including field crops, orchards, greenhouse crops and weeds. The Western ﬂower thrips is considered as the most important thrips vector of diseases. It transmits Chrysanthemum stem necrosis virus (CSNV), Groundnut ringspot virus (GRSV), Impatiens necrotic spot virus (INSV), Tomato chlorotic spot virus (TCSV) and Tomato Thrips (Thysanoptera). Chapter 13.1 a 777 b c Figure 13.1.7. Adults of some Thysanoptera alien to Europe. a Echinothrips americanus b Gynaikothrips ficorum c Pezothrips kellyanus (credit: Philippe Reynaud, LNPV). spotted wilt virus (TSWV). There is also an indirect economic effect when introduced into a new area. For example, western flower thrips is a major economic driving force of greenhouse and field crop IPM research. F. occidentalis is restricted to glasshouses in northern Europe, but has established outdoors in areas with milder winters. The international spread of the western flower thrips occurred predominantly by the movement of horticultural material, such as cuttings, seedlings and potted plants. Within Europe, an outward spread from the original outbreak in the Netherlands (1983) is discernible. The speed of spread was 229 +/- 20 km/year (Kirk and Terry 2003). Chemical control is difficult, because F. occidentalis is resistant to most pesticides, but some predatory mites and minute Pirate bugs provide effective biological control under glasshouses. Two other North American Frankliniella species are known in Europe, but with a very limited distribution and without economic impact. The potential introduction of the Melon thrips (Thrips palmi) represents a continuous threat to glasshouse ornamental and vegetable crops in Europe (see factsheet 14.80). Numerous interceptions have been reported on cut ﬂowers and fruit vegetables and several outbreaks were found in glasshouses in the Netherlands and UK since 1988. The potential of adults and larvae to survive an entire winter oudoors in the UK is very limited however (McDonald et al. 2000), which has favoured successful control and eradication of all these outbreaks. T. palmi is considered to be absent in Europe, although it was detected outdoors within ﬂowers of kiwi fruit (Actinidia deliciosa) in Portugal in 2004, but in later surveys the 778 Philippe Reynaud / BioRisk 4(2): 767–791 (2010) pest was no longer found. The palm thrips is essentially a tropical species, and therefore most parts of Europe are not suitable for its establishment. We can assume, however, that most of southern Europe could harbour this species outdoors and the species could establish indoors in other places. High developmental and reproductive rates at glasshouse temperatures allows rapid build-up of populations, even from small numbers of females (Cannon et al. 2007). Vector of alien topospovirus, the Melon thrips has been implicated in the transmission of at least six plant viruses. T. palmi is a quarantine organism for the EU and as such requires eradication wherever it is found. Several other alien thrips species occur indoor in Europe with a low economic impact, including Hercinothrips femoralis, Heliothrips haemorrhoidalis and Echinothrips americanus. These species are found in the wild in tropical and subtropical regions, but are restricted to glasshouses in western Europe, with the exception of H. haemorrhoidalis (also called the greenhouse thrips). The greenhouse thrips can also live in the wild in southern Europe. It has many hosts, including ornamental shrubs and field crops (citrus, avocado and tea) but preferred hosts in Southern Europe are Myrtus communis and Viburnum tinus. E. americanus was recently introduced from the USA, where it is seldom a pest, into Europe (Netherlands). However, in Europe it has more than 50 known food plants, including ornamental and woody plants and vegetables. The species is often found in sizable numbers without showing obvious damage symptoms to the plant (Vierbergen et al. 2006) and seems to be highly susceptible to insecticides (Karadjova and Krumov 2003). H. femoralis (the sugar beet thrips) is a minor polyphagous pest under glasshouses that feeds on more than 50 hostplants but is also an important pest almost everywhere where bananas are grown (Trdan et al. 2007). The genus Gynaikothrips includes about 40 species, with two related pest species (G. ficorum and G. uzeli). The same common name (Cuban Laurel Thrips) is used for these two leaf-galling thrips species on decorative Ficus trees distributed worldwide by the horticultural trade. But only Gynaikothrips ficorum is at the present time known as an alien species in Europe. These two species can only be differenciated by a microscopic examination of the pronotal posteroangular pair of setae. According to Mound et al. (Mound et al. 1995), G. ficorum is the primary gall maker on Ficus microcarpa while G. uzeli is the primary gall maker on F. benjamina. G. ficorum was first described from Algeria, but is native of Southeast Asia. Adults vary from about 2.6 mm to 3.6 mm in length and are dark yellowish-brown to black. Infested, curled leaves become hard and tough, then gradually yellower and browner and eventually drop from the plant prematurely. Finally, the ornamental value of the plant is reduced. The Cuban Laurel Thrips is a minor pest in Europe and only under glasshouses, but adults can be a nuisance in North Africa on Ficus microcarpa planted in cities, by flying into people’s eyes or irritating their skin (Mumcuoglu and Volman 1988). The Composite thrips Microcephalothrips abdominalis, the only species in the genus, is a light-brown species characterized by an unusual small head in relation to the pronotum. It lives on Compositae flowers throughout its life, where it is considered as an important pollinating agent. M. abdominalis is known to transmit TSV (Greber et al. 1991), a serious disease of peanut and sunﬂower in India (Jones 2005) but this virus Thrips (Thysanoptera). Chapter 13.1 779 is not a quarantine pest for EU. It has been suggested that this pantropical species is native to the New World and has been transported elsewhere by man (Stannard 1968). This species has been known from Italy since 1994 but has subsequently shown a slow rate of spead in Europe. The Composite thrips is considered as a minor pest but is not reported yet as a pest in Europe. References Aitkenhead P (1951) The Gladiolus Thrips - a Pest new to Britain. Agriculture 57, 11: 517–523. Anonymous (2004) First report of Thrips palmi in Portugal. EPPO Reporting Service 144: 2. Bagnall RS (1909) On the Thysanoptera of the Botanical Gardens, Brussels. Annales de la Société entomologique de Belgique 53: 171–176. 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Reuter OM (1891) Thysanoptera funna i finska orangerier. Meddelanden af Societatis pro Fauna et Flora Fennica 17: 161–167. Reuter OM (1904) Ein neues Warmhaus-Thysanopteron. Meddelanden af Societatis pro Fauna et Flora Fennica 30: 106–109. Reynaud P (1998) Echinothrips americanus. Un nouveau thrips des serres importé en France. Phytoma 507: 36–38. Reynaud P, Bertaux F, Martinez M (2001) Premier signalement en Europe de Neohydatothrips samayunkur (Kudo) (Thysanoptera, Thripidae). Nouvelle Revue d’Entomologie 18: 91–93. Thrips (Thysanoptera). Chapter 13.1 783 Sakimura K (1967) Redescription of Anaphothrips orchidaceus and A. orchidearum (Thysanoptera: Thripidae). The Florida Entomologist 50: 89–97. Stannard LJ (1968) The Thrips, or Thysanoptera, of Illinois. Bulletin of the Illinois Natural History Survey 29: 213–552. Strapazzon A (1999) Italian faunal records 368. Bollettino Della Societa Entomologica Italiana 131: 259. Streito JC, Martinez M (2005) Nouveaux ravageurs, 41 espèces depuis 2000. Phytoma - La défense des Végétaux 586: 16–20. Trdan S, Jovic M, Andjus L (2005) Palm thrips, Parthenothrips dracaenae (Heeger) (Thysanoptera: Thripidae), in Slovenia: still a pest of minor importance? Acta agriculturae Slovenica 85: 211–217. Trdan S, Kuznik L, Vidrih M (2007) First results concerning the efficacy of entomopathogenic nematodes against Hercinothrips femoralis (Reuter). Acta Agriculturae Slovenica 89: 5–13. Varga L (2008) Hercinothrips femoralis (Reuter, 1891) – a new pest thrips (Thysanoptera: Panchaetothripinae) in Slovakia. Plant Protection Science 44: 114–118. Vierbergen G, Mantel WP (1991) Contribution to the knowledge of Frankliniella schultzei (Thysanoptera: Thripidae). Entomologische Berichten 51: 7–12. Vierbergen G (1996) Annual Report 1996. Wageningen, Netherlands: Plant Protection Service, Diagnostic Centre. 114 pp. Vierbergen G (1998) Echinothrips americanus Morgan, a new thrips in Dutch greenhouses (Thysanoptera: Thripidae). Proceedings of the section Experimental and Applied Entomology of the Netherlands Entomological Society (N.E.V.) 9: 155–160. Vierbergen G (2004) Eight species of thrips new for the Netherlands and some taxonomical changes in Stenchaetothrips, Thrips and Hoplothrips (Thysanoptera). Acta Phytopathologica et Entomologica Hungarica 39: 199–209. Vierbergen G, Cean M, Szeller IH, Jenser G, Masten T, Simala M (2006) Spread of two thrips pests in Europe: Echinothrips americanus and Microcephalothrips abdominalis (Thysanoptera: Thripidae). Acta Phytopathologica et Entomologica Hungarica 41: 287–296. Webster KW, Cooper P, Mound LA (2006) Studies on Kelly’s citrus thrips, Pezothrips kellyanus (Bagnall) (Thysanoptera: Thripidae): sex attractants, host associations and country of origin. Australian Journal of Entomology 45, 1: 67–74. Wilson TH (1975) A monograph of the subfamily Panchaetothripinae (Thysanoptera: Thripidae). Memoirs of the American Entomological Institute 23: 1–354. Zur-Strassen R (1965) Einige neue terebrante Thysanopteren-Arten von den Kanarischen Inseln (Ins., Thysanoptera). Commmentationes biologicae Societas scientiarus fennica 28: 3–41. Zur-Strassen R (1973a) Ergebnisse der Forschungsreise auf die Azoren 1969. III Zur Faunistik und Zoogeographie der Thysanopterenfauna der Azoren im Mittel-Atlantik. Boletin do Museo Municipal do Funchal 27, 117: 26–50. Zur-Strassen R (1973b) Uber einige zumeist floricole Fransenfluger ans dem sudlichen Andalusien (Spaniena) (Ins Thysanoptera). Senckenbergiana biologia 54: 327–338. Zur-Strassen R (1982) Thysanopterologische Notizen (6) (Insecta: Thysanoptera). Senckenbergiana Biologica 63: 191–209. 784 Philippe Reynaud / BioRisk 4(2): 767–791 (2010) Zur-Strassen R (1986a) Frankliniella occidentalis (Pergande 1985), ein nordamerikanischer Fransenflügler (Thysanoptera) als neuer Bewohner europäischer Gewächshäuser. Nachrichtenblatt Deutschen Pflanzenschutzdienstes 38: 86–88. Zur-Strassen R (1986b). Thysanopteran auf Inseln der Nordlichen Sporaden in der Agais (Griechenland). Senckenbergiana Biologica 67: 85–129. Zur-Strassen R (1995) Dorcadothrips billeni n. sp. (Insecta: Thysanoptera), ein neuer terebranter Fransenflugler von Wasserfarn. Mitteilungen der Entomologischen Gesellschaft Basel 45: 148–153. Zur-Strassen R (1996) New data on systematics and distribution of some West Palaearctic Terebrantia species (Thysanoptera). Entomologische Nachrichten Und Berichte 40: 111–118. Zur-Strassen R (2003) Die terebranten Thysanopteren Europas und des Mittelmeer-Gebietes. Die Tierwelt Deutschlands 74. Keltern, Germany: Verlag Goecke and Evers. 277 pp. Zur-Strassen R, Borges PAV (2005) Thysanoptera. In: Borges PAV, Cunha R, Gabriel R, Martins AMF, Silva L, Vieira V (Eds) A list of the terrestrial fauna (Mollusca and Arthropoda) and flora (Bryophyta, Pteridophyta and Spermatophyta) from the Azores. Horta, Angra do Heroísmo and Ponta Delgada: Direcção Regional de Ambiente e do Mar dos Açores and Universidade dos Açores, 189–190. Table 13.1.1. List and main characteristics of the Thysanoptera species alien to Europe. Status: A: Alien to Europe; C: cryptogenic species. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Only selected references are given. Last update 03/02/2010. Family Status Regime Native Species range Aeolothripidae Franklinothrips megalops A predator Africa (Trybom, 1912) Merothripidae Merothrips floridensis Watson, 1927 Phlaeothripidae Aleurodothrips fasciapennis (Franklin, 1908) Bagnalliella yuccae (Hinds, 1902) Eurythrips tristis Hood, 1941 Gynaikothrips ficorum (Marchal, 1908) Haplothrips gowdeyi (Franklin, 1908) Invaded countries Habitat Hosts Unknown BG, ES, NL J100 Greenhouses thrips and black vine thrips References Zur-Strassen (2003), Mound and Reynaud (2005) Zur-Strassen (2003) A predator C&S America Unknown BE, CH, DE, DK, FR, IL, NL, PT-MAD, SE J100 Frankliniella occidentalis and two- spotted spider mite, Tetranychus urticae Koch (Acari: Tetranychidae). A detritovorous C&S America 1955, FR ES, FR, PT-AZO I Citrus (fungivorous) Bournier (1960), ZurStrassen and Borges (2005) C predator Cryptogenic 1908, BE BE, DE J100 Aonidella, Crysomphalus and other scales Bagnall (1909), Geiter et al. (2002) A phytoNorth phagous America unknown North America phytoAsiaphagous Tropical 1957, FR FR, HU, IT, RO, UA I2 Yucca Jenser (1989) U Sporophagous Zur-Strassen and Borges (2005) Bournier (1983), Pelikán (1991), Laudonia and Viggiani (2005) A A A phytophagous Africa 2005, PT- PT- AZO AZO 1983, FR- CZ, DE, FR-COR, GRCOR CRE, IL, IT, IT- SAR, IT- SIC, NL, PT, PTMAD 1978, GR CY, ES, ES- CAN, GR, PT-AZO, PT-MAD I2, J100 Ficus I Solenaceae, Apiaceae 785 Zur-Strassen (1986b), Zur-Strassen and Borges (2005) Thrips (Thysanoptera). Chapter 13.1 Franklinothrips vespiformis (Crawford, 1909) 1st record in Europe Status Regime A C C phytophagous detritovorous detritovorous 1st record Invaded countries in Europe 2001, ES ES Habitat Hosts References I2 Crataegus oxyacantha Berzosa et al. (2001) Cryptogenic Cryptogenic 1954, RO CZ, RO G Prunus armeniacum Pelikán (1990) 1939, GB CZ, GB, NO, SE X16 Kobro and Rafoss (2006), Mound et al. (1976) North America North America C&S America 1974, ES ES X13 1919, AL AL, CY, ES, IT- SAR, PT I2 1911, DK DK, ES- CAN, IT, PTAZO, PT-MAD, Polystictus abietinus fungus on dead pine branches Predator (sparsely wooded land) Fiorinia fioriniae (scale) on many ornamentals Coccidae, Diaspididae scales (Howardia biclavis) Karnyothrips americanus (Hood, 1912) Karnyothrips flavipes (Jones, 1912) Karnyothrips melaleucus (Bagnall, 1911) A predator A predator A predator Nesothrips propinquus (Bagnall, 1916) A detritovorous Australasia 1974, PTAZO ES- CAN, NL, PT-AZO, I PT-MAD Podothrips semiflavus Hood, 1913 Suocerathrips linguis Mound & Marullo, 1994 Thripidae Anaphothrips sudanensis Trybom, 1911 A parasitic/ predator detritovorous North America Cryptogenic 1964, CY CY I 1994, GB BE, GB J100 A phytophagous Unknown ES, CY E1, F6 Grasses, cereals Zur-Strassen (2003) A phytophagous Tropical, subtropical C&S America 1969, P-AZO IT, PT-AZO, PT-MAD I Cyathula prostrata (folivorous) and young coconut fruits Zur-Strassen (1973a), Zur-Strassen and Borges (2005) Anisopilothrips venustulus (Priesner, 1923) C J100 Berzosa (1988) Priesner (1919), Canale et al. (2003) Bagnall (1911), Mound and Marullo (1994), Zur-Strassen and Borges (2005) Sporophagous Mound (1974), ZurStrassen and Borges (2005) Aspidiella sacchari (coccid Priesner (1964b) scale) Penicilium species living Mound and Marullo on Sansevieria surface (1994) Philippe Reynaud / BioRisk 4(2): 767–791 (2010) Native range Asia 786 Family Species Haplothrips rivnayi Priesner, 1936 Hoplothrips lichenis Knechel, 1954 Hoplothrips unicolor (Vuillet, 1914) Native range C&S America 1st record Invaded countries in Europe 1907, GB BE, DE, DK, FR, GB, NO, SE C&S America North America C&S America 1998, I AsiaTropical AsiaTropical Unknown, NL NL 1975, NL NL J100 Orchidaceae (Vanda) J100 Orchidaceae AsiaTropical North America 1994, DE DE J100 1996, FR AT, BE, BG, DE, DK, FR, FR-COR, GB, IT, NL, NO, SE, SI J100 Microsorum pteropus (Oriental water fern) Hibiscus (but polyphagous on ornemental crops) Frankliniella schultzei (Trybom, 1910) C phytophagous Cryptogenic 1988, NL NL J100 Frankliniella fusca (Hinds, 1902) A phytophagous North America 1964, NL NL J100 AsiaTropical IT, SE Habitat Hosts References J100 Orchidaceae Bagnall and John (1935), Sakimura (1967) J100 Spathiphyllum Colombo et al. (1999) Navel oranges exports (contaminant) Anthurium, banana, Citrus, orchids Zur-Strassen (2003) Unknown, GB J100 GB 1935, F BE, CZ, DE, DK, FI, J100 FR, GB, IL, IT, NO, NL, PT-MAD, SE 1996, NL IT, NL J100 Araceae, Piper Bagnall and John (1935), Del Bene and Gargani (2001) Vierbergen (1996) Mantel and van de Vrie (1988) Mound (1976) Zur-Strassen (1995) 787 Reynaud (1998), Vierbergen (1998), Vierbergen et al. (2006), Zur-Strassen (2003) Polyphagous, recorded as Vierbergen and Mantel a pest of vegetables and (1991) ornemental crops Polyphagous, reported to Mantel and van de Vrie cause direct damage to (1988) peanuts and cotton Thrips (Thysanoptera). Chapter 13.1 Family Status Regime Species Aurantothrips A phytoorchidaceus (Bagnall, phagous 1909) Bradinothrips musae A phytoHood, 1956 phagous Caliothrips fasciatus A phyto(Pergande, 1895) phagous Chaetanaphothrips A phytoorchidii (Moulton, phagous 1908) Copidothrips A phytooctarticulatus (Schmutz, phagous 1913) Dichromothrips corbetti A phyto(Priesner, 1936) phagous Dichromothrips A phytophalaenopsidis phagous Sakimura, 1955 Dorcadothrips billeni A phytoZur-Strassen, 1995 phagous Echinothrips americanus A phytoMorgan, 1913 phagous Habitat Hosts References Zur-Strassen (1986a), Kirk I2, J100 Polyphagous (Plants, and Terry (2003) trees- Populus); flowers and leaves; vector tobacco streak ilarvirus (TSV) and tomato spotted wilt virus (TSWV) Heliothrips haemorrhoidalis (Bouché, 1833) A phytophagous C&S America Hercinothrips bicinctus (Bagnall, 1919) A phytophagous Tropical, subtropical Hercinothrips femoralis (Reuter, 1891) A phytophagous C&S America 1891, FI BE, CZ, DE, DK, ES, J100 ES-CAN, FI, FR, GB, HU, IL, IT, LV, MD, NL, RO, SE, SK, SI, UA Leucothrips nigripennis Reuter, 1904 A phytophagous C&S America 1904, FI AL, BE, CZ, DE, DK, FI, FR, GB, NL I2, J100 Polyphagous (Citrus, avocados, ornamental plants) in urban , agricultural and modified habitats, rarely forests, mainly greenhouses Bouché (1833), Mound et al. (1976), Zur-Strassen (2003), Zur-Strassen and Borges (2005) J100 Bagnall (1919), Mound et al. (1976), Wilson (1975), Zur-Strassen and Borges (2005) Reuter (1891), Mound et al. (1976), Varga (2008) J100 Musa spp., passionfruit (folivorous) Polyphagous (banana, beet, celery, Commelina diffusa, Crinum, Chrysanthemum, dwarf milo maize, eggplant, Emilia sonchifolia, Erechtites hieracifolia, grass, orchids, pineapple, Plantago major) Ferns Reuter (1904), Mound (1999) Philippe Reynaud / BioRisk 4(2): 767–791 (2010) 1st record Invaded countries in Europe 1983, NL AL, AT, BE, BG, CH, CZ, DE, DK, EE, ES, FI, FR, GB, GR, HR, HU, IE, IL, IT, IT-SAR, IT-SIC, LT, LV, NL, NO, PT, RO, RS, SE, SK, SI, UA 1833, DE AL, AT, BE, BG, CH, CZ, DE, DK, ES, FI, FR, FR- COR, GB, GR, HU, IL, IT, IT-SAR, IT-SIC, LT, LV, MD, MT, NL, NO, PT, PT-AZO, PTMAD, RO, SE, SI, SK, UA 1907, BE BE, DE, DK, ES, ESCAN, FR, GB, HU, IT, NL, PT-AZO, PT-MAD 788 Family Status Regime Native Species range Frankliniella occidentalis A phytoNorth (Pergande, 1895) phagous America 1st record Invaded countries in Europe 1999, IT ES-CAN, HU, IT, SI I2 Neohydatothrips samayunkur (Kudo, 1995) Organothrips indicus Bhatti, 1974 A phytophagous A phytophagous Palmiothrips palmae (Ramakrishna, 1934) A Parthenothrips dracaenae (Heeger, 1854) Habitat Hosts Asteraceae (Bidens formosa -cosmos, Chrysanthemum, Helianthus, Pyrethrum, Tagetes, Zinnia) Marigold (Tagetes sp.) 2000, FR FR I 1985, DE DE J100 phytophagous AsiaTropical 1965, ESCAN ES-CAN, IL I2 A phytophagous Africa 1852, AT J100 Pezothrips kellyanus (Bagnall, 1916) Phibalothrips peringueyi (Faure, 1925) C phytophagous phytophagous 1981, GR I2 Citrus E Grasses Plesiothrips perplexus (Beach, 1896) A phytophagous Cryptogenic Tropical, subtropical C&S America AT, BE, BG, CH, CZ, DE, DK, ES, FI, FR, GB, GR, HU, IS, IT, LV, MD, NL, NO, RO, SE, SI ES, FR, GR, IT-SIC, IL, NL IT, IT-SIC Water hyacinth (Eichhornia crassipes) in warmed aquarium (aquatic species) Phoenix flowers, including date palm, Phoenix dactilifera Dracena, Ficus 1975, PTMAD IT, PT-AZO, PT-MAD E Poaceae Pseudodendrothrips mori (Niwa, 1908) A phytophagous AsiaTropical 1974, IT ES, FR, IT, SI I2 Morus A 1985, ITSIC Strapazzon (1999), Vierbergen et al. (2006) Reynaud et al. (2001) Mound (2000) Zur-Strassen (1965) Heeger (1854), Trdan et al. (2005) Zur-Strassen (1986b), Zur-Strassen (2003) Zur-Strassen (1996), ZurStrassen (2003) Zur-Strassen (1982), Zur-Strassen and Borges (2005) Cappellozza and Miotto (1975), Vierbergen et al. (2006) 789 Tropical, subtropical Asia References Thrips (Thysanoptera). Chapter 13.1 Family Status Regime Native Species range Microcephalothrips A phytoTropical, abdominalis (Crawford, phagous sub1910) tropical Stenchaetothrips biformis (Bagnall, 1913) A phytophagous AsiaTropical Stenchaetothrips spinalis Reyes, 1994 Thrips australis (Bagnall, 1915) A phytophagous phytophagous Asia1999, FR Temperate Australasia 1930, CY A 1st record Invaded countries in Europe 1982, GB GB J100 Philodendron Palmer and Mound (1985) 1995, DE DE, SE J100 1909, BE BE, CZ, DE, DK, FI, J100 FR, IT, LV, NO, NL, PTMAD, SE CZ, GB, IT, NL, PL, RO J100 Microsorum pteropus (Oriental water fern) Avocado, onions, … Billen and Zur-Strassen (1995) Bagnall (1909), Hoddle and Mound (2003) Growing tips of seedling rice, Oryza sativa (larva, adult); secondary hosts: maize, Zea mays, wild sugarcane, Saccharum spontaneum, wild grasses (Agropyron- wheatgrass, Festuca-fescues, Pennisetta) Bambusoideae Bagnall (1913), Kucharczyk and Zawirska (2001), Vierbergen (2004) 1913, GB FR CY, ES, ES-CAN, FR, GR, IT, IT-SIC, PT, PTAZO, PT-MAD Thrips palmi Karny, 1925 A phytophagous AsiaTropical 1995, PT CZ, NO, PT Thrips simplex Morrison, 1930 A phytophagous Africa 1946, FR AT, BG, CH, CZ, DE, ES, ES-CAN, FR, GB, HU, IL, IT, NO, NL, PT, PT-AZO, RO, SE, SI, UA Habitat I2 Hosts References Streito and Martinez (2005) I2, F6 Eucalyptus, Melaleuca Priesner (1964a), Priesner (1964b), Zur-Strassen (1973b), Zur-Strassen and Borges (2005) I, J Quarantine pest, Anonymous (2004), polyphagous but a threat Cannon et al. (2007) to glasshouse ornamental and vegetable crops in Europe I2, J100 Gladiolus, polyphagous in Aitkenhead (1951), greenhouses Bournier (1954), ZurStrassen and Borges (2005), Milevoj et al. (2008) Philippe Reynaud / BioRisk 4(2): 767–791 (2010) Native range C&S America AsiaTropical Cryptogenic 790 Family Status Regime Species Psydrothrips kewi A phytoPalmer & Mound, 1985 phagous Pteridothrips pteridicola A phyto(Karny, 1914) phagous Scirtothrips longipennis C phyto(Bagnall, 1909) phagous Table 13.1.2. List and main characteristics of some Thysanoptera species alien in Europe. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Only selected references are given. Last update 03/02/2010 Family Species Aeolothripidae Aeolothrips fasciatus (L., 1758) Regime Native range Invaded countries Habitat Hosts References G3 Pinus U Dead wood or leaf-litter Europe ES- CAN, PT- AZO PT- AZO G Liothrips vaneeckei Priesner, 1920 phytophagous Europe GB J100 Dead wood of broadleaved trees, feeding on fungi (possibly Peniophora) Lilly bulbs Pelikán and Schliephake (1994) Zur-Strassen and Borges (2005) Zur-Strassen and Borges (2005) Bagnall (1933), Mound et al. (1976) Thripidae Aptinothrips rufus Haliday, 1836 phytophagous Europe PT- AZO I Grasses, cereals phytophagous Europe PT- AZO I mycophagous Europe AT U Alopecurus pratensis, Lilium, clover, peach, pear, apple, grasses, wheat Fungi infecting weeds phytophagous Europe PT- AZO E, I, J Poaceae phytophagous Europe GB G3, G4 Melilotus phytophagous Europe GB I1, I2, FA, E2, E5 Polyphagous (weeds, flowers, trees and crops) Chirothrips manicatus Haliday, 1836 Euphysothrips minozzii Bagnall, 1926 Limothrips cerealium Haliday, 1836 Odontothrips meliloti Priesner, 1951 Thrips tabaci Lindeman, 1889 PT- AZO E, I phytophagous Europe GB phytophagous Europe mycophagous Europe mycophagous Mound et al. (1976) Zur-Strassen and Borges (2005) Zur-Strassen and Borges (2005) Zur-Strassen (2003) Zur-Strassen and Borges (2005) Pitkin (1972), Mound et al. (1976) Bagnall (1923) 791 DE, CZ Rhipidothrips gratiosus Uzel, 1895 Phlaeothripidae Apterygothrips pinicolus Pelikan & Schliephake, 1994 Hoplandrothrips consobrinus (Knechtel, 1951) Hoplothrips ulmi (F., 1781) Europe Thrips (Thysanoptera). Chapter 13.1 Zur-Strassen and Borges (2005) I,J Both a pollen feeder and a predator of onion thrips; Taraxacum officinale, Trifolium repens, Epilobium angustifolium, Grasses Grasses, wild oats predator/ phytophagous A peer reviewed open access journal BioRisk 4(2): 793–805 (2010) doi: 10.3897/biorisk.4.46 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Psocids (Psocoptera) Chapter 13.2 Nico Schneider 79, rue Tony Dutreux, L-1429 Luxembourg-Bonnevoie, Luxemburg Corresponding author: Nico Schneider (nico.schneider@education.lu) Academic editor: David Roy | Received 1 January 2010 | Accepted 25 May 2010 | Published 6 July 2010 Citation: Schneider N (2010) Psocids (Psocoptera). Chapter 13.2. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(2): 793–805. doi: 10.3897/biorisk.4.46 Abstract Among the 231 species of Psocoptera present in Europe, 49 (21.2%) are considered to be of alien origin. They include 29 exotic introduced species and 20 cryptogenic species. Most of the exotic species originated from tropical and subtropical areas, essentially from Africa. Many of them are food pests, moving along with stored products. Thirty-nine of these species occur in buildings in Europe. Keywords Psocoptera, psocids, domestic, stored products, alien, Europe 13.2.1 Introduction Psocoptera (commonly called psocids) are one of the smaller orders of paraneopteran insects. Many species are arboreal, but a few are more usually found on low vegetation or in litter. All feed on microflora and organic debris. Some are found in nests of birds and mammals, within aggregations of other insects or associated with human habitations. The head of these usually soft bodied pterygote insects (with a body length of 0.67 mm to 8 mm) is globulous with an usually prominent clypeus and projecting eyes, long and filiform antennae and biting mouthparts, the laciniae being characteristic for the order. Adults have usually four wings with simple venation. However, many species are brachypterous, micropterous or apterous (Lienhard 1998, Lienhard and Smithers 2002, Mockford 1993, New 2005). Copyright Nico Schneider. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 794 Nico Schneider / BioRisk 4(2): 793–805 (2010) 13.2.2 Taxonomy of Psocoptera species alien to Europe According to Lienhard (1998) (Lienhard 1998) a total of 231 species in 25 families of psocopterans are present in Europe. Forty-nine of these are not native, 29 as aliens to Europe and 20 as cryptogenic, globally accounting for 21,2% of the total psocopteran fauna in Europe (Figure 13.2.1). They are included in 12 families: Lepidopsocidae (5), Trogiidae (4), Psoquillidae (3) and Psyllipsocidae (5) belonging to the suborder Trogiomorpha; Liposcelididae (14) and Pachytroctidae (2) belonging to the suborder Troctomorpha; and Caeciliusidae (1), Ectopsocidae (10), Elipsocidae (1), Lachesillidae (2), Peripsocidae (1) and Psocidae (1) belonging to the suborder Psocomorpha. Details for each family are as follows. Lepidopsocidae: Adult wings of lepidopsocids, which belong to the family group Atropetae, are generally pointed apically when fully developed. Body and forewings are generally covered with scales, but occasionally with dense setae Mockford 1993. The five species found in Europe are all alien. Trogiidae: Species in this family, which belongs to the family group Atropetae, are apterous, micropterous or brachelytropterous. Body and forewings lack scales and dense setae. Four of the 19 species found in Europe are cryptogenic (21%). These four species are brachelytropterous, presenting short, leathery winglets similar to short elytra. Their habitats are associated with humans, e.g. within buildings and glasshouses (Lienhard 1998, Mockford 1993). Psoquillidae: Body and forewings of species in this family, which also belongs to the family group Atropetae, do not bear scales. The three species currently found in Europe are not believed to be native, two of them being with certainty of alien origin and the third cryptogenic. All occur within buildings in Europe. Psyllipsocidae: This family belongs to the family group Psocatropetae. The five species found in Europe (100%) are either alien (Baz 1990) or cryptogenic species (Baz 1988). All are usually found in buildings and glasshouses in Europe. Liposcelididae: These psocids belong to the family group Nanopsocetae. They are characterised by a flattened body and antennae with secondary annulations on their flagellum. Fourteen species out of the 39 (36%) found in Europe are either aliens (Broadhead 1950) or cryptogenic species (Broadhead 1954b). They include exclusively apterous species, all of which being occur in buildings. Pachytroctidae: The body shape of the species of this family, which belongs to the family group Nanopsocetae, is not flattened dorsoventrally and the basal flagellar segments are not secondarily annulated (New 2005). Only three species live in Europe, and two of them are not natives (67%). Caeciliusidae: This family belongs to the family group Caeciliusetae and to the superfamily Caecilioidea. The species are characterised by the presence of at least one, or more commonly two or three, ventral abdominal vesicles (Mockford 1993). The family was once named Caeciliidae, but the latter name was changed because of homonymy with a family of amphibians (Lienhard 1998). Only one species out of the 15 (7%) found in Europe is cryptogenic. Lacroixiella martini, is only known by the two syntypes collected by Lacroix in 1918 in a french military hospital (Lacroix 1919). Psocids (Psocoptera). Chapter 13.2 795 Figure 13.2.1. Taxonomic overview of the Psocoptera species alien to Europe compared to the native fauna. Species alien to Europe include cryptogenic species. Families are listed in a decreasing order based on the number of alien species. The number over each bar indicates the number of species observed per family. Lachesillidae: These psocids belong to the family groupe Homilopsocidea. The species have an areola postica* in their forewings characterized by a very sparse and short ciliation on its veins and margin. The lacinial* tip of the Lachesillidae is slender and bicuspid *(Lienhard 1998, Mockford 1993). Two species out of the 12 found in Europe are aliens (17%). Ectopsocidae: Members of this family, which belongs to the family group Homilopsocidea, are characterised by the absence of an areola postica in their wings such as in the family Peripsocidae. Ten out of the 14 species found in Europe (71%) are either alien (Broadhead 1950) or cryptogenic species (Baz 1988). Six of these species are found occurring within buildings, either regularly (Baz 1990) or sometimes (Baz 1990), compared to only one of the 4 native species. Peripsocidae: Species of this family, which also belongs to the family group Homilopsocidea, are also characterised by their absence of an areola postica in their wings. Peripsocus bivari is the only alien among nine species found in Europe (11%). 796 Nico Schneider / BioRisk 4(2): 793–805 (2010) Figure 13.2.2. Geographic origin of the Psocoptera species alien to Europe. Numbers indicate the relative proportion of alien species originating from a given region. Elipsocidae: Veins and wing margins of species in this family, which also belongs to the family group Homilopsocidea, bear setae but the hindwing marginal setae are limited to the radial fork (New 2005). Propsocus pulchripennis, occurring out-of-doors in France, Great Britain and Madeira, is the only alien among 24 species found in Europe (4%). The native range of this widespread species is the coastal regions of subtropical countries (Lienhard 1998, Lienhard and Smithers 2002). Psocidae: This family belongs to the family group Psocetae. Its members are easily recognised by their wing venation, where the areola postica is fused to the M-vein. The Psocidae fauna of Europe includes 34 species but Trichadenotecnum innuptum is the only alien among them (3%). 13.2.3 Temporal trends of introduction in Europe of alien psocids The cryptogenic species Trogium pulsatorium was already known as Termes pulsatorium in the 10th edition of Systema Naturae by Linnaeus in 1758 (Linnaeus 1958). From 1850 to 1874, three other cryptogenic species and the alien Psoquilla marginepunctata were recorded from Europe. One alien and one cryptogenic species followed in 1899. In the 20th century, seven species on our list were recorded for the first time in Europe from 1900 to 1924, 18 from 1925 to 1949, seven from 1950 to 1974 and finally ten from 1975 to 1999. In the 21st century, no new alien has arrived in Europe to date. 13.2.4 Biogeographic patterns of the Psocoptera species alien to Europe The distribution status is only known for 29 species out of 49, 40.8% being thus considered as cryptogenic. Figure 13.2.2 details the probable regions of origin. Most Psocids (Psocoptera). Chapter 13.2 797 Figure 13.2.3. Colonization of continental European countries and main European islands by the Psocoptera species alien to Europe. Archipelago: 1 Azores 2 Madeira 3 Canary islands. species appear to originate from Africa (9 species, 18.4 %), far beyond the other continents but for seven species (14.3%) we only know that they arrived from tropical and subtropical regions. Central and Western Europe appear to be more colonized by alien psocids (Figure 13.2.3). 13.2.5 Pathways of introduction in Europe of alien psocids, invaded habitats and known impacts The main pathway of introduction is trade. Lachesilla pacifica is probably dispersed by wind. Most aliens and cryptogenic species are found in warehouses in stored products. Many of them are food pests. Forty species (88.9 %) are associated with buildings in Europe. 798 a Nico Schneider / BioRisk 4(2): 793–805 (2010) b Figure 13.2.4. Alien psocids. a Ectopsocus briggsi McLachlan, 1899 (Credit: Tom Murray 2008) b Liposcelis bostrychophila Badonnel, 1931 (Credit: Joyce Gross 2006). Acknowledgments I thank Dr. Charles Lienhard (Muséum d’histoire naturelle de Genève, Switzerland), and Drs. Alain Roques and David Lees (INRA, Centre de Recherche d’Orléans, France). Ms. Joyce Gross (BSCIT, University of California, Berkeley, USA) and Mr. Tom Murray (Groton, Massachusetts, USA). References Badonnel A (1943) Psocoptères. Faune de France 42. Paris: Lechevallier et fils. 164 pp. Baz A (1988) Psocopteros de Azores: nuevas citas, descripciones y sinonimias; Boletim da Sociedade Portuguesa de Entomologia 93: 1–15. Baz A (1990) Nanopsocus oceanicus Pearman, 1928, nueva especie para la fauna de Europa (Psocoptera, Pachytroctidae). Nouvelle Revue d’Entomologie 7: 251–254. Bigot L (1982) Structure et dynamique des populations frondicoles d’insectes Coléoptères et Psocoptères dans la Forêt de la Sainte-Baume. Bulletin du Muséum d’Histoire naturelle, Marseille 42: 7–18. Broadhead E (1947) New species of Liposcelis Motschulsky (Corrodentia, Liposcelidae) in England .Transactions of the Royal Entomological Society of London 98: 41–58. Broadhead E (1950) A revision of the genus Liposcelis Motschulsky with notes on the position of this genus in the order Corrodentia and on the variability of ten Liposcelis species. Transactions of the Royal Entomological Society of London 98: 41–58. Broadhead E (1954a) a A new parthenogenetic psocid from stored products, with observations on parthenogenesis in other psocids. Entomologist’s Monthly Magazine 90: 10–16. Broadhead E (1954b) The infestation of warehouses and ships’ holds by psocids in Britain. Entomologist’s Monthly Magazine 90: 103–105. Psocids (Psocoptera). Chapter 13.2 799 Broadhead E (1955) Two new psocid species from stored products in Britain. Proceedings of the Royal Entomological Society of London (B) 24(1/2): 7–12. Danks L (1955) Psocoptera of the Batumi and Sochi Botanic Gardens. Entomologicheskoe Obozrenie 34: 180–184. Eichler W (1938) Thylacopsis madagascariensis, Brachymyrmex heeri und Ptilodactyla luteipes in einem Dahlemer Gewächshaus (Studien zur deutschen Gewächshausfauna II). Zoologischer Anzeiger 122: 330–333. Enderlein G (1905) Morphologie, Systematik und Biologie der Atropiden und Troctiden, sowie Zusammenstellung aller bisher bekannten recenten und fossilen formen. Results of the Swedish Zoological Expedition to Egypt and the White Nile, 1901 18, 1–58. Enderlein G (1906) Zehn neue aussereuropäische Copeognathen. Stettiner Entomologische Zeitung 67: 306–316. Günther KK (1974) Staubläuse, Psocoptera. Die Tierwelt Deutschlands 61. Jena: G. Fischer. 314 pp. Hagen H (1865) Synopsis of the Psocina without ocelli. Entomologists’ Monthly Magazine 2: 121–124. Harrison JWH (1916) A November week at Grange-over-sands V Psocoptera collected by Richard S Bagnall, FLS. Lancashire Naturalist 9: 197–109. Heyden CHG von (1850) Zwei neue deutsche Neuropteren-Gattungen. Stettiner Entomologische Zeitung 11: 83–85. Heymons R (1909) Ein neuer Troctes als Schädling in Buchweizengrütze (Corrod). Deutsche Entomologische Zeitschrift 1909: 452–455. Jentsch S (1939) Beiträge zur Kenntnis der Überordnung Psocoidea 8 Die Gattung Ectopsocus (Psocoptera). Zoologische Jahrbücher (Abteilung Sytematik) 73: 111–128. Lacroix J (1919) Description d’un Psocide nouveau de France. Bulletin de la Société entomologique de France 1919: 80–81. Lienhard C (1977) Die Psocopteren des schweizerischen Nationalparks und seiner Umgebung (Insecta: Psocoptera). Ergebnisse der wissenschaftlichen Untersuchungen im Schweizerischen Nationalpark 14: 417–551. Lienhard C (1985) Sur quelques espèces intéressantes de Psocoptères du Bassin Lémanique et du Valais. Bulletin Romand d’Entomologie 3: 73–79. Lienhard C, (1986) Beitrag zur Kenntnis der Psocopteren-Fauna Ungarns (Insecta). Annales Historico-naturales Musei nationalis Hungarici 78: 73–78. Lienhard C (1994) Staubläuse (Psocoptera) – ungebetene Gäste in Haus und Vorrat. Mitteilungen der Entomologischen Gesellschaft Basel, Neue Folge 44: 122–160. Lienhard C (1996) Psocoptères nouveaux ou peu connus de quelques îles atlantiques (Canaries, Madère, Açores, Ascension) et de l’Afrique du Nord (Insecta: Psocoptera). Boletim do Museu Municipal do Funchal (Historia Natural) 48: 87–151. Lienhard C (1998) Psocoptères euro-méditerranéens Faune de France 83. Paris : Fédération Française des Sociétés de Sciences Naturelles. 517 pp. Lienhard C, Schneider, N (1993) Dorypteryx longipennis Smithers, un psoque domicole nouveau pour l’Europe (Psocoptera: Psyllipsocidae). Bulletin et Annales de la Société Royale Belge d’Entomologie 129: 129–137. 800 Nico Schneider / BioRisk 4(2): 793–805 (2010) Lienhard C, Smithers CN (2002) Psocoptera (Insecta): World Catalogue and Bibliography. Instrumenta Biodiversitatis 5. Genève: Muséum d’Histoire Naturelle. 745 pp. Lienhard C (2002–2009). Additions and Corrections to Lienhard & Smithers, 2002 Psocoptera (Insecta) : World Catalogue and bibliography. Psocid News 4 (2002), 6 (2004), 7 (2005), 8 (2006), 9 (2007), 10 (2008) and 11 (2009). http://www.psocodea.org/psocid_news. Linnaeus C von (1758) Systema Naturae per Regna tria Naturae, secundem Classes, Ordines, Genera, Species, cum Characteribus, Differentis, Synonymis, Locis. Tom.1, Editio decima, reformata. Holmiae: Laurentii Salvii. 824 pp. McLachlan R (1899) Ectopsocus briggsi, a new genus and species of Psocidae found in England. Entomologists’ Monthly Magazine 35: 277–278. Mockford E L (1993) North American Psocoptera (Insecta). Flora and Fauna Handbook 10. Gainesville, Florida: Sandhill Crane Press. 455 pp. Motschulsky V von (1852) Etudes entomologiques. (Psocids: Excursions entomologiques de 1852, jusqu’au 1 Juillet). Helsingfors: 19–20. New TR (2005) Psocids, Psocoptera (Booklice and barklice). Handbooks for the Identification of British Insects Vol 1, Part 7. London: Royal Entomological Society. 146 pp. Pearman JV (1925) Additions to the British psocid fauna. Entomologist’s Monthly Magazine 61: 124–129. Pearman JV (1929) New species of Psocoptera from warehouses. Entomologist’s Monthly Magazine 65: 104–109. Pearman JV (1931a) A new species of Lepinotus (Psocoptera). Entomologist’s Monthly Magazine 67: 47–50. Pearman JV (1931b) More Psocoptera from warehouses. Entomologist’s Monthly Magazine 67: 95–98. Pearman JV (1932) A new species of Tapinella (Psocoptera). Stylops 1: 240–242. Pearman JV (1942) Third note on Psocoptera from warehouses. Entomologist’s Monthly Magazine 78: 289–292. Pearman JV (1946) A specific characterization of Liposcelis divinatorius (Müller) and mendax sp n (Psocoptera). Entomologist 79: 235–244. Ribaga C (1899) Descrizione di nuovo genere e di nuova specie di Psocidi trovato in Italia. Rivista di Patologia Vegetale 8: 156–159. Ribaga C (1904) Sul genere Ectopsocus MacLachl e descrizione di una nuova varietà dell’ Ectopsocus briggsi MacLachl. Redia 1: 294–298. Ribaga C (1907) Copeognati nuovi. Redia 4: 181–189. Selys-Longchamps E de (1872) Notes on two new genera of Psocidae. Entomologist’s Monthly Magazine 9: 145–146. Titschack E (1930) Die Copeognatha, Megaloptera, Neuroptera und Mecoptera der näheren und weiteren Umgebung Hamburgs. Verhandlungen des Vereins für Naturwissenschaftliche Heimatforschung 21: 104–127. Table 13.2.1. List and main characteristics of the Psocoptera species alien to Europe. Status: A: Alien to Europe; C: cryptogenic species. Country codes abbreviations refer to ISO 3166 (see Appendix I). Habitat abbreviations refer to EUNIS (see Appendix II). Last update 31/ 12/ 2009 Ectopsocus maindroni Badonnel, 1935 Ectopsocus meridionalis Ribaga, 1904 Native range 1st record in Europe Invaded countries Habitat U References C Unknown 1918, FR FR A Asia 1955, RU A ?Australia 1991, IE AT, CH, DE, HR, HU, IL, IT, RU, YU G, I, J, X Danks (1955), Lienhard (1998), Lienhard and Smithers (2002), Lienhard (2002)–(2009) GB, IE G Lienhard (1998), Lienhard and Smithers (2002) C Unknown 1899. GB A Tropical, subtropical 1954, GB C Unknown 1904, IT A Africa, Asia A Africa, Asia 1984, PTAZO 1929, GB A Australia A ?Africa A C. & S. America A A AT, BE, CH, CY, CZ, DE, EE, ES, ES- G, I, X CAN, FI, FR, GB, GR, HR, HU, IE, IL, IT, LU, ME, MK, NL, NO, PT, PTAZO, PT-MAD, PL, RS, RU, SE, YU GB, IT J AT, CH, CY, CZ, DE, ES, ES-CAN, J, X FR, GR, HR, HU, IE, IL, IT, LU, ME, MK, MT, RO, RS, YU CH, PT-AZO J Lacroix (1919), Lienhard (1998), Lienhard and Smithers (2002) Lienhard (1998), Lienhard and Smithers (2002), Lienhard (2002)–(2009), McLachlan (1899) Broadhead (1954b), Lienhard (1998), Lienhard and Smithers (2002) Lienhard (1998), Lienhard and Smithers (2002), Lienhard (2002)–(2009), Ribaga (1904) Lienhard (1994), Lienhard (1998), Lienhard and Smithers (2002), Mockford (1993) Lienhard and Smithers (2002), Pearman (1929) CH, GB, PT-AZO J PT-MAD G ES, ES-CAN, IT, PT-AZO, PT-MAD J DE, ES G, J Tropical, subtropical 1981, PTMAD FR, GB, PT-MAD X Baz (1990), Bigot (1982), Lienhard (1998), Lienhard and Smithers (2002), Lienhard (2002)–(2009) North America CH, FR G Lienhard (1998), Lienhard and Smithers (2002) 1992, PTMAD 1906, ESCAN 1928, DE 1986, CH Lienhard (1996, 1998), Lienhard and Smithers (2002) Enderlein (1906), Lienhard (1998), Lienhard and Smithers (2002) Jentsch (1939), Lienhard (1998), Lienhard and Smithers (2002) 801 Ectopsocus pumilis (Banks, 1920) Ectopsocus richardsi (Pearman, 1929) Ectopsocus rileyae Schmidt & Thornton, 1993 Ectopsocus strauchi Enderlein, 1906 Ectopsocus titschacki Jentsch, 1939 Elipsocidae Propsocus pulchripennis (Perkins, 1899) Lachesillidae Lachesilla pacifica Chapman, 1930 Status Psocids (Psocoptera). Chapter 13.2 Family Species Caeciliusidae Lacroixiella martini (Lacroix, 1919) Ectopsocidae Ectopsocopsis cryptomeriae (Enderlein, 1907) Ectopsocus axillaris (Smithers, 1969) Ectopsocus briggsi McLachlan, 1899 C Unknown 1852, RU Liposcelis corrodens (Heymons, 1909) C Unknown 1909, DE Habitat References ES-CAN, PT-MAD G, I, X Lienhard (1998), Lienhard and Smithers (2002) DE J Eichler (1938), Lienhard (1998), Lienhard and Smithers (2002) GB J GB J FR, GB, IE, PT-MAD J, X Broadhead (1955), Lienhard (1998), Lienhard and Smithers (2002) Broadhead (1954b), Lienhard (1998), Lienhard and Smithers (2002) Harrison (1916), Lienhard (1998), Mockford (1993) DE, GB J Lienhard (1998), Lienhard and Smithers (2002), Selys-Longchamps (1872) ES-CAN J Lienhard (1996) GB J GB J Lienhard (1998), Lienhard and Smithers (2002), Pearman (1931b) Broadhead (1955), Lienhard (1998), Lienhard and Smithers (2002) Badonnel (1943), Lienhard (1998), Lienhard and Smithers (2002) AT, BE, CH, CY, CZ, DE, ES, ESJ CAN, FI, FR, GB, GR, HR, HU, IE, IL, IT, LU, MK, MT, NL, NO, PT, PTAZO, PT-MAD, PL, RO, RS, SE, YU AT, BE, CH, CY, CZ, DE, ES, ESJ CAN, FI, FR, GB, GR, HR, IT, LU, MK, NO, PL, PT, RO, RS, SE, RU, YU AT, BE, CH, CY, CZ, DE, ES, FI, FR, G, J GB, GR, HR, HU, IE, IT, LU, MK, MT, NL, NO, PL, PT, PT-AZO, PTMAD, RO, RS, SE, YU Broadhead (1950), Lienhard (1998), Lienhard and Smithers (2002), Lienhard (2002)–(2009), Motschulsky (1852) Heymons (1909), Lienhard (1998), Lienhard and Smithers (2002) Nico Schneider / BioRisk 4(2): 793–805 (2010) Liposcelis brunnea Motschulsky, 1852 Invaded countries 802 Family Status Native range 1st record Species in Europe Lachesilla tectorum A Tropical, subtropical 1992, PTBadonnel, 1931 MAD Lepidopsocidae Echmepteryx A Tropical, subtropical 1938, DE madagascariensis (Kolbe, 1885) Lepolepis bicolor Broadhead, A Africa, Asia 1945, GB 1955 Nepticulomima sakuntala A Asia, tropical 1954, GB Enderlein, 1906 Pteroxanium kelloggi A North America 1916, GB (Ribaga, 1905) Soa flaviterminata A Tropical, subtropical 1930, DE Enderlein, 1906 Liposcelididae Belaphotroctes ghesquierei A ?Africa 1993, ESBadonnel, 1949 CAN Embidopsocus minor A Africa 1931, GB (Pearman, 1931) Liposcelis albothoracica A Africa 1955, GB Broadhead, 1955 Liposcelis bostrychophila C Unknown 1943, FR Badonnel, 1931 Family Species Liposcelis decolor (Pearman, 1925) Status Native range Unknown Liposcelis entomophila (Enderlein, 1907) C Unknown Liposcelis mendax Pearman, 1946 A Africa Liposcelis obscura Broadhead, 1954 Liposcelis paeta Pearman, 1942 A ?Africa C Unknown C Unknown A ?Asia C Unknown A Tropical, subtropical 1988, ES CY, ES, ES-CAN J C Unknown 1932, GB ES-CAN, GB J A ?Africa 1979, PTAZO ES-CAN, FR, PT-AZO,PT-MAD G, X Baz (1988), Lienhard (1996, 1998), Lienhard and Smithers (2002) A North America 1965, HU CH, HU, IT G, X Lienhard (1986, 1998), Lienhard and Smithers (2002) A C. & S. America 1865, ?DE BE, CZ, ?DE, GB, IT, PT-AZO G, J Günther (1974), Hagen (1865), Lienhard and Smithers (2002) Liposcelis paetula Broadhead, 1950 Liposcelis pearmani Lienhard, 1990 Liposcelis pubescens Broadhead, 1947 Pachytroctidae Nanopsocus oceanicus Pearman, 1928 Tapinella castanea Pearman, 1932 Peripsocidae Peripsocus bivari Baz 1988 Psocidae Trichadenotecnum innuptum Betz, 1983 Psoquillidae Psoquilla marginepunctata Hagen, 1865 Baz (1990), Lienhard (1998), Lienhard and Smithers (2002) Lienhard (1998), Lienhard and Smithers (2002), Lienhard (2002)–(2009), Pearman (1932) 803 C Psocids (Psocoptera). Chapter 13.2 1st record Invaded countries Habitat References in Europe Broadhead (1950), Lienhard (1998), Lienhard and 1925, GB AT, BE, CH, CY, CZ, DE, EE, ES, ES- J Smithers (2002), Pearman (1925) CAN, FI, FR, GB, GR, HR, HU, IL, IT, LU, LV, MK, MT, NL, NO, PL, PT, PT-MAD, RO, SE, YU 1929, GB CH, CY, CZ, DE, ES, FI, GB, HR, IL, J Broadhead (1950), Lienhard (1998), Lienhard and IT, PT, PT-AZO, YU Smithers (2002), Lienhard (2002)–(2009), Pearman (1929) 1946, FR, CH, ES, ES-CAN, FR, GB, HR, IT, J Broadhead (1950), Lienhard (1998), Lienhard and GB YU Smithers (2002), Lienhard (2002)–(2009), Pearman (1946) 1954, GB GB J Broadhead (1954a), Lienhard (1998), Lienhard and Smithers (2002) 1940, GB BE, CZ, ES, GB, HR, IT, YU J Broadhead (1950), Lienhard (1998), Lienhard and Smithers (2002), Lienhard (2002)–(2009), Pearman (1942) 1945, GB ES-CAN, GB, IT, PT-MAD G, J Broadhead (1950), Lienhard (1998), Lienhard and Smithers (2002) 1945, GB AT, CH, CZ, DE, ES, FI, FR, GB, HR, J Broadhead (1950), Lienhard (1998), Lienhard (2002– HU, IL, IT, LU, NL, YU 2010), Lienhard and Smithers (2002) 1943, GB BE, CH, CZ, DE, GB, IT, LU, PTJ Broadhead (1947), Broadhead (1950), Lienhard AZO, YU (1998), Lienhard and Smithers (2002) Trogiidae Lepinotus inquilinus von Heyden, 1850 Lepinotus patruelis Pearman, 1931 Lepinotus reticulatus Enderlein, 1905 Trogium pulsatorium (Linnaeus, 1758) Native range A Africa 1st record in Europe 1931, GB GB Invaded countries Habitat References C Unknown 1929, GB GB A Africa 1973, CH C Unknown 1988, LU AT, BA, BE, CH, CZ, DE, DK, ES, J ES-CAN, FI, FR, GB, HR, HU, IE, IL, IT, LU, NO, PL, SE, SK, YU BE, CH, ES, IE, IT, LU, NL J C Unknown 1907, IT AT, BE, CH, CZ, DE, ES, FR, IT J A Tropical, subtropical 1899, IT ES-CAN, IL, IT, PT-MAD H, J C Unknown 1872, FR AT, BE, CH, CZ, DE, ES, ES-CAN, FI, FR, GB, GR, HR, HU, IE, IL, IT, LU, NL, NO, PL, PT, PT-AZO, PTMAD, RO, RU, SE, YU H, J C Unknown 1850, DE J Heyden (1850), Lienhard (1998), Lienhard and Smithers (2002) C Unknown 1930, GB J C Unknown 1905, DE Lienhard (1998), Lienhard and Smithers (2002), Pearman (1931a) Enderlein (1905), Lienhard (1998), Lienhard and Smithers (2002) C Unknown 1758, Europe AT, BE, CH, CZ, DE, DK, ES, ESBAL, ES-CAN, FI, FR, GB, GR, HR, HU, IS, IT, LU, NL, NO, PL, PTAZO, PT-MAD, RO, RU, SE, YU AT, BE, CH, CZ, DE, FI, FR, GB, IE, IT,LU, NO, PL, PT-AZO, SE AT, BE, CH, CY, CZ, DE, DK, ES, ES-CAN, FI, FR, GB, GR, HR, HU, IL, IS, IT, LU, MK, NL, PT, PT-AZO, PL, RO, RU, SE, YU AL, AT, BE, BG, CH, CY, CZ, DE, DK, EE, ES, ES-CAN, FI, FR, GB, GR, HR, HU, IE, IL, IS, IT, LT, LU, NL, NO, PL, PT-AZO, PT-MAD, RO, RU, SE, YU J Lienhard (1998), Lienhard and Smithers (2002), 38 J Lienhard (1998), Lienhard and Smithers (2002), Peramn (1929) J J Lienhard (1977, 1998, 2002- 2010), Lienhard and Smithers (2002) Lienhard (2002- 2010), Lienhard and Schneider (1993), Lienhard and Smithers (2002) Lienhard (1998), Lienhard and Smithers (2002), Mockford (1993), Titschak (1930) Lienhard (1998), Lienhard and Smithers (2002), Ribaga (1904) Lienhard (1998, 2002- 2010), Lienhard and Smithers (2002), Sélys- Longchamps (1872) Lienhard (1998), Lienhard and Smithers (2002) Nico Schneider / BioRisk 4(2): 793–805 (2010) Dorypteryx longipennis Smithers, 1991 Dorypteryx pallida Aaron,1883 Psocathropos lachlani Ribaga, 1899 Psyllipsocus ramburii SélysLongchamps, 1872 Status 804 Family Species Rhyopsocus disparilis (Pearman, 1931) Rhyopsocus peregrinus (Pearman, 1929) Psyllipsocidae Dorypteryx domestica (Smithers, 1958) Psocids (Psocoptera). Chapter 13.2 805 Table 13.2.2. List and characteristics of the Psocoptera species alien in Europe. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Last update 31/ 12/ 200 Family Species Caeciliusidae Enderleinella obsoleta (Stephens, 1836) Native range Invaded countries in Europe Habitat References Central Europe BE, BG, DK, FI, G FR, GB, IE, IT, LU, ME, MK, NL, NO, RO, RU, SE Lienhard (1998), Lienhard and Smithers (2002) Mediterranean region CH, GB Lienhard (1998), Lienhard and Smithers (2002) Central Europe BE, ES, FR, GB, IT, G, H, J PT, PT-AZO, PTMAD, RO Lienhard (1998), Lienhard and Smithers (2002), Lienhard (2002)–(2009) Liposcelididae Liposcelis rufa Broadhead, 1950 Mediterranean region CH, GB, PL G, J Lienhard (1998), Lienhard and Smithers (2002) Peripsocidae Peripsocus milleri (Tillyard, 1923) Atlantic coast of Europe IT, YU G, J Central Europe BE, ES, FI, FR, GB, GR, HR, IL, LU, NL, RO, RU, SE, YU G Lienhard (1998), Lienhard and Smithers (2002) Lienhard (1998), Lienhard and Smithers (2002) Trichopsocidae Trichopsocus clarus (Banks, 1908) Mediterranean region J, X Lienhard (1998), Lienhard and Smithers (2002) Trichopsocus dalii (McLachlan, 1867) Mediterranean region CH, CZ, DE, FI, GB, HU, IE, LT, NL, PL, RU, SE AT, BE, CH, CZ, DE, GB, HU, LU, PL, RU G Lienhard (1998), Lienhard and Smithers (2002) Trogiidae Cerobasis annulata (Hagen, 1865) Mediterranean region AT, BE, CH, CZ, DE, GB, LU, NL, NO, PL, RU G, J, X Lienhard (1998), Lienhard and Smithers (2002) Ectopsocidae Ectopsocus vachoni Badonnel, 1945 Lachesillidae Lachesilla greeni (Pearman, 1933) Peripsocus parvulus Kolbe, 1880 G, J A peer reviewed open access journal BioRisk 4(2): 807–831 (2010) doi: 10.3897/biorisk.4.68 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Dictyoptera (Blattodea, Isoptera), Orthoptera, Phasmatodea and Dermaptera Chapter 13.3 Jean-Yves Rasplus1, Alain Roques2 1 UMR Centre de Biologie et de Gestion des Populations, CBGP, (INRA/IRD/CIRAD/Montpellier SupAgro), Campus international de Baillarguet, CS 30016, 34988 Montferrier-sur Lez, France 2 NRA UR633, Zoologie Forestière. Centre de recherche d’Orléans, 2163 Avenue de la Pomme de Pin, CS 40001 Ardon, 45075 Orléans Cedex 2, France Corresponding authors: Jean-Yves Rasplus (rasplus@supagro.inra.fr), Alain Roques (alain.roques@orleans.inra.fr) Academic editor: David Lees | Received 26 March 2010 | Accepted 25 May 2010 | Published 6 July 2010 Citation: Rasplus J-Y, Roques A (2010) Dictyoptera (Blattodea, Isoptera), Orthoptera, Phasmatodea and Dermaptera. Chapter 13.3. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(2): 807–831. doi: 10.3897/ biorisk.4.68 Abstract For convenience, we treat all “polyneopteran” orders together. Five orders of hemimetabolous “Polyneoptera” include species alien to Europe, namely Blattodea, Isoptera, Orthoptera, Phasmatodea and Dermaptera. A total of 37 species alien to Europe have been recorded. These belong to 14 different families. Most of these species show a detritivorous feeding regime (22 spp.), whereas 12 species are phytophagous and two are predators. The majority of species were first observed between 1900 and 1975. Unlike other arthropod groups, the mean number of polyneopteran species newly recorded per year showed no acceleration since 1975. The alien “Polyneoptera” mostly originated from Central/ South America and Asia (10 species each, 27.0%), followed by Africa (7, 18.9%). Germany hosts the largest number of alien Polyneoptera (15 spp.), followed by Denmark (14), Spain (11) and France (10). All but one alien species represent unintentional introductions. More than 75% of the species are associated with artificial habitats (houses, buildings and greenhouses) and cultivated areas. Blattodea and Isoptera have huge economic and/ or medical importance. The cost of treatments and sanitary measures against termites and cockroaches, in particular, is significant in Europe. Keywords Alien, Orthoptera, grasshoppers, Blattodea, coackroaches, Isoptera, termites, Phasmatodea, walking sticks, Dermaptera, earwigs Copyright J.-Y. Rasplus, A. Roques. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 808 Jean-Yves Rasplus & Alain Roques / BioRisk 4(2): 807–831 (2010) 13.3.1 Introduction For convenience, we group all the orders belonging to the “Polyneoptera” assemblage into the same chapter. This non-monophyletic assemblage of eleven “orthopteroid” orders includes five orders which contain species alien to Europe. Some of these orders are very familiar such as grasshoppers (Orthoptera), cockroaches (Blattodea), termites (Isoptera), earwigs (Dermaptera) and walking sticks (Phasmatodea). Lesser known groups include web-spinners (Embioptera), angel insects (Zoraptera) and ice-crawlers (Grylloblattodea). We describe here the characteristics of the species alien to Europe. Blattodea contains over 4500 species worldwide, with about 150 species in Europe. They are among the most ancient winged insects, the earliest fossils dating back to the Carboniferous. The group is well defined by a combination of characters: eggs usually contained in oothecae (egg cases), leathery forewings, male genitalia asymmetrical and cerci* with one or more segments. Most cockroaches are tropical and found in a wide variety of habitats such as dead or decaying leaves or trees, caves, under stones, in nests of social insects etc. Cockroaches are mostly scavengers eating organic material. Less than 1% (30 species) are associated with humans, but these species contribute to the unpopular reputation of these insects. Cockroaches exhibit diverse reproductive biology. Most species have sexual reproduction, but some populations of Pycnoscelus surinamensis are parthenogenetic. These hemimetabolous insects produce hardened oothecae deposited on a substrate or membraneous oothecae that are incubated in a brood sac within the female’s body. Some species exhibit a high level of parental care. Isoptera consists of over 2600 species (mostly tropical). Termites are the oldest social insect group with complex societies dating back at least to the early Cretaceous (140 Mya). Only 12 species occur in Europe. Recent studies have shown that Isoptera are basically social cockroaches forming a monophyletic clade within the Blattodea, most likely the sister group of the Cryptocercidae (woodroaches) (Inward et al. 2007). Termites are the only hemimetabolous insects that exhibit true social behavior. They build large nests housing an entire colony. These colonies contain adult reproductives (one queen and one king) plus hundreds or thousands of immatures that serve as workers and soldiers. Termites are important decomposer animals in lowland tropical ecosystems. They mostly feed on dead plant material and are able to digest cellulose with the help of symbiotic gut symbionts. Orthoptera comprises more than 20000 species worldwide and 1044 species in Europe belonging to two suborders, Caelifera (grasshoppers) and Ensifera (ladykids). This group of median-sized insects is well characterized by (1) long hind legs modified for jumping; (2) hardened, leathery forewings (tegmina) which are spread in flight and covering membranous hindwings at rest; (3) unsegmented cerci; and (4), a pronotum usually with large descending lateral lobes. Orthopterans are common in most terrestrial habitats, but are more diverse in the tropics. They are mostly phytophagous and include some outstanding agricultural pests (locusts and certain katydids). Dictyoptera (Blattodea, Isoptera), Orthoptera, Phasmatodea and Dermaptera. Chapter 13.3 809 Phasmatodea (also known as Phasmida) comprises 3000 species worldwide with only 15 species known in Europe. Stick-insects are found in nearly all temperate and tropical ecosystems. Species are mostly nocturnal and phytophagous. Phasmatodea bears several common morphological characters that clearly define the order: an emarginated labrum, a pair of exocrine glands located inside the prothorax, and a thorax fused with the first abdominal sternum. Phasmids undergo an incomplete metamorphosis (four to eight instars), with the young nymphs resembling miniature, albeit wingless, adults. Dermaptera comprises about 1800 species and about 80 species in Europe. These small- to median-sized insects have the head prognathous* and are clearly characterized by two or more apomorphies: long unsegmented (not always forceps-like) cerci, and details of hindwing structure. The biology of Dermaptera is poorly known. Most species appear to be omnivorous but some are phytophagous and a few are predators. The development is hemimetabolous. Earwigs have larvae (four to five instars) that resemble the adult, except that the wings are only buds. Several characteristics group species in these orders together. The polyneopteran group treated here comprises mostly phytophagous species (consuming fresh plants, dead wood or leaves), but some species are detrivorous. None of the species alien to Europe is parasitic and very few are predators. These species are rarely transported with cultivated plants, even if eggs of stick-insects are introduced with soil. Consequently, polyneopterans are rarely introduced into Europe through the plant trade. Most species are relatively large and conspicuous, the smallest insects belonging to Isoptera and Dermaptera. All of them are hemimetabolous and consequently their larvae are biologically similar to adults. The diversity of these groups in the Holarctic region is relatively limited and most species are tropical. These characteristics may partly explain the relatively low number of species in the alien fauna that has colonized Europe, compared to worldwide Polyneopteran diversity. 13.3.2.Taxonomy of alien species A total of 37 species alien to Europe have been recorded. These species belong to five different orders and 14 different families (Table 13.3.1; Figure 13.3.1). Blattodea account for 18 species and is the order with by far the greatest number of aliens to Europe. Eleven species belong to Orthoptera, four to Phasmatodea, while Dermaptera and Isoptera include two alien species each. Within Orthoptera, Ensifera are well representated with seven species (63% of Orthoptera). Among these alien species, 22 are detritivorous, 12 phytophagous and two are predators, the biology of one species being unknown. This results show that within invasive Polyneoptera, a majority of species are detritivorous or phytophagous (94%). Table 13.3.2 presents some species of the same orders considered as alien in Europe (native to a European region but introduced in another through human activity). 810 Jean-Yves Rasplus & Alain Roques / BioRisk 4(2): 807–831 (2010) Figure 13.3.1. Relative importance of the families of Blattodea, Isoptera, Orthoptera, Phasmatodea, and Dermaptera in the alien and native entomofauna in Europe. Families are presented per order in a decreasing ranking based on the number of alien species. Species alien to Europe include cryptogenic species. The number over each bar indicates the number of species observed per family. Blattodea Blaberidae. This small family contains ten species in Europe, all of them introduced from tropical countries. These cockroaches are ovoviviparous, some species being parthenogenetic. Several Blaberidae species have been introduced into urban areas of Europe. Among them, Blaberus atropos is a native to South America that exhibits a death’s-head markings on the mesonotum and metanotum. Nauphoeta cinerea lives mostly around the outside of buildings but also occurs in houses. Panchlora nivea is commonly associated with bananas and palm trees. This species was introduced in Northern Europe with shipments of bananas. Pycnoscelus surinamensis, a Malaysian cockroach, as been introduced several times to Europe. It occurs in greenhouses and cannot live outdoors. Its European populations appear to be parthenogenetic. This trait has been wrongly identified to explain the strong invasive ability of this cockroach (Grandcolas et al. 1996). Rhyparobia maderae, an afrotropical cockroach, was probably transported to southern Europe with banana shipments and occurs indoors. Blattellidae. Among the ca. 135 species of Blatellidae occurring in Europe, only two species, Nyctibora laevigata and Supella longiplapa, have an alien origin, both having been introduced from tropical regions. The last one is an afrotropical species with synanthropic habits, occurring in houses and greenhouses in Europe. These long- legged cockroaches carry the eggcase externally. Dictyoptera (Blattodea, Isoptera), Orthoptera, Phasmatodea and Dermaptera. Chapter 13.3 811 Blattidae. Only six species are known in Europe, all of them having been introduced from tropical or subtropical regions. Blatta orientalis, Periplaneta spp. and Neostylopyga rhombifolia are synanthropic species that have long been introduced to Europe. A more recent arrival is that of Shelfordella lateralis, the Turkestan cockroach, which has been discovered in 2007 in Cagliari, Sardinia. This species has previously been introduced in the 1970s in the Southern United States (California, Texas, Arizona) probably with military people coming back from the Middle East (Fois et al. 2009). These blattid species mostly develop indoors, in heated buildings but can also develop in greenhouses and in the city streets. Isoptera Kalotermitidae. This family comprises only four species in Europe, of which only Cryptotermes brevis is alien to Europe. This species infests dry wood and can damage woodwork, furniture and floors. C. brevis has been found both in Northern and Southern Europe but it has been more widely introduced to tropical countries. Recent studies showed that the early European shipment of exports from coastal Peru and Chile caused the release and initial dispersal of C. brevis from its natural range (Scheffrahn et al. 2009). Rhinotermitidae. This family comprises seven species in Europe, including one alien species originating from North America, Reticulitermes flavipes (= R. santonensis (Feytaud); see Austin et al. 2005), where it is considered to be a significant pest. Subterannean termites in the genus Reticulitermes Holmgren (Isoptera: Rhinotermitidae) are the major termite pests infesting wooden structures in Europe and the near East. Dermaptera Anisolabididae. This family comprises 12 species in Europe. Euborellia stali, of Asian origin, preys on stem borers associated with rice entering the borer tunnel. This widespread species has recently been introduced in Italy. Labiduridae. Only two species of Labiduridae are known from Europe, including a species originating from tropical/subtropical regions, Nala lividipes. This species is considered as a pest with local economic importance, but it is rare in Europe. Orthoptera Acrididae (Caelifera). This diverse family (about 350 species in Europe) only contains four species alien to Europe. Furthermore, the status of two of them, Notaustorus albicornis and Dociostaurus tartarus, is unclear, and these species could be native to Southeastern Europe. Bradyporidae (Ensifera). A total of 84 species occur in Europe, one of them being possibly alien to Europe, Ephippigerida nigromarginata, originating from Africa. 812 Jean-Yves Rasplus & Alain Roques / BioRisk 4(2): 807–831 (2010) Gryllidae (Ensifera). A total of 83 species of gryllids occur in Europe, but only one is an alien species. Gryllodes sigillatus is probably native to southwestern Asia and has been spread by commerce to different part of the world. This species is found indoors. Myrmecophilidae (Ensifera). This small family of crickets contains 11 European species, one having been possibly introduced to Europe, the cryptogenic Myrmecophilus americanus. Myrmecophilus ant crickets are symbionts associated with ant nests. They are kleptoparasitic and feed on food resources in ant nests and induce ants to regurgitate liquid food. M. americanus is associated with an invasive ant species Paratrechina longicornis. Phaneropteridae (Ensifera). Only one alien species, Topana cincticornis, has been recorded to be compared with the 149 species of this family native to Europe. This species, of South American origin, has only been observed in France (Morin 2001). Rhaphidophoridae (Ensifera). This family contains 53 species in Europe. Only one of them is alien to Europe, Tachycines asynamorus. This oriental species mostly develops indoors (houses, greenhouses) in Northern Europe but also outdoors during the summer in Southern Europe. Tettigoniidae (Ensifera). This family contains 221 species in Europe, two of them (namely Copiphora brevirostris and Phlugiola dahlemica) having been introduced from Central and South America. The latter species was described inhabiting greenhouses in the Botanical Gardens of Berlin (Weidner 1938). Phasmatodea Phasmatidae. The family contains only four species in Europe, all of them introduced and occurring in Southern Great Britain. Three of these species (The Prickly Stick Insect, Acanthoxyla geisovii, The Unarmed Stick Insect, Acanthoxyla inermis, and the Smooth Stick Insect, Clitarchus hookeri) arrived from New Zealand with plants, most likely as eggs in the soil (Lee 1993). The last species Carausius morosus is native of the Oriental region but was also introduced in Germany (Weidner 1981). Some stick insects used as pets may also have escaped from captivity but we have no data about that. In conclusion, the only group of polyneopterans with a significant number of introduced species compared to the native European fauna is that of cockroaches (Figure 13.3.1). Blaberidae and Blattidae are represented in Europe only by exotic species nonintentionally introduced by humans. 13.3.3 Temporal trends The dates of introduction of most alien cockroaches are largely unknown although it is likely that most of these synanthropic species were introduced to Europe long ago, Dictyoptera (Blattodea, Isoptera), Orthoptera, Phasmatodea and Dermaptera. Chapter 13.3 813 following human movements and trade. For instance, the first record for Blatta orientalis dates back to 1500 in a region corresponding at present to the Czech Republic. Finally, first records in Europe of alien Polyneoptera, excluding four species considered as cryptogenic, were obtained for 21 out of the 33 remaining alien species (64 %). Most of these 21 species were first observed between 1900 and 1975. Interestingly, the mean number of new records per year has not accelerated during the last 200 years, unlike most other groups of arthropods (Figure 13.3.2). On the average, less than one species was newly observed every five years during the period 1900 to 2006. 13.3.4. Biogeographic patterns Origin of alien species A region of origin could be traced for 35 (95%) of the alien Polyneoptera introduced to Europe. Central/South America and Asia, with 10 species each (27.0 %), provided equally the greatest part of these alien species followed by Africa (7 spp.; 18.9 %) (Figure 13.3.3). This pattern largely differs from the one observed in most other groups of insects where South America contributes much less to the alien fauna. Indeed, most Blattodea are of tropical origin and generally became sub-cosmopolitan species that occur in buildings and exceptionally outdoors in Europe. Within Orthoptera, most Ensifera also have a tropical origin and several species can presently survive only within greenhouses in Europe. To the contrary, Caelifera are mostly Palaearctic species that naturally occur in areas adjacent to Europe. Alien Isoptera originate from North and South America. Most alien Phasmatodea originate from Australasia and were introduced into England with plants. Distribution of alien species in Europe Alien polyneopteran species and families are not evenly distributed throughout Europe and large differences exist between countries (Figure 13.3.4; Table 13.3.3). The number of taxonomists and the intensity of studies and sampling may also have influenced these differences. Little information is available for some central and northeastern European countries, and consequently these areas appear to host comparatively less alien species. Germany hosts the largest number of alien Polyneoptera (15 spp.), followed by Denmark (14), Spain (11) and France (10). Most European countries host a low number of introduced species (five or less). No correlation with the country surface area has been found. However, it appears that northern countries in Europe host globally more alien species. 814 Jean-Yves Rasplus & Alain Roques / BioRisk 4(2): 807–831 (2010) Figure 13.3.2. Temporal changes in the mean number of new records per year of ‘Polyneoptera’ alien to Europe from 1492 to 2006. Cryptogenic species excluded. The number above the bar indicates the number of species introduced. 13.3.5. Main pathways to Europe The main pathway for introduction of most polyneopteran species alien to Europe is unknown. Where known, most introductions were unintentional. Whilst Blattodea species have followed humans and have long been introduced in Europe probably as stowaways as more recently observed for Blaberus atropos, Panchlora nivea, and Rhyparobia maderae found within banana shipments (Sein 1923). Some recent invaders also seem to have been introduced through wood transport (Isoptera) or introduction of plant material (Phasmatodea and Ensifera). Nauphoeta cinerea has been introduced intentionally and only one species (Euborellia stali) have been introduced for biological control purposes. 13.3.6. Most invaded ecosystems and habitats A large proportion of polyneopteran species alien to Europe (>75%) are associated with artificial habitats (houses, buildings and greenhouses) and cultivated areas (Figure 13.3.5). The proportion is somewhat lower (>55%) for the species alien to countries within Europe. These results are mostly linked to the strong associations of some Blattodea, Isoptera and Ensifera with humans. Only few species (10 spp.) have yet colonized natural and semi- natural habitats (grasslands, heathland or coastal habitats). Dictyoptera (Blattodea, Isoptera), Orthoptera, Phasmatodea and Dermaptera. Chapter 13.3 815 Figure 13.3.3. Origin of the species of Polyneoptera alien to Europe. 13.3.7. Ecological and economic impact While most ‘Polyneoptera’ species introduced to Europe have only limited ecological or economic impact, two orders are considered as important pests: Blattodea and Isoptera. Blattodea have great medical significance (Baumholtz et al. 1997) and several species of cockroaches represent a potential threat to human health and well-being. These species are the most common household insect pests and there are two areas of concern regarding their potential for causing disease in humans. First, cockroaches are recognized as being an important source of indoor allergens. These allergens are found in their body, saliva and faecal matter. They cause asthmatic reactions in humans and are also implied in skin reactions. In recent studies, a strong association has been found between the presence of cockroaches and increase in the severity of asthma symptoms in individuals who are sensitive to cockroach allergens. Finally, oedema of the eyelids and dermatitis has been attributed to cockroaches. Second, because of high humidity, high temperature and presence of food, cockroaches normally breed well in houses, grocery stores, restaurants and hospitals. They feed on a variety of foodstuffs (meat, grease, candies, chocolate, cheese, bread and other unprotected materials), regurgitate fluid from their mouth, and deposit faeces on foodstuffs. Because of their movement between waste and food materials, cockroaches can acquire, carry, and directly transfer to food and eating utensils the bacterial pathogens that cause food poisoning, diarrhea (Burgess and Chetwyn 1981), or typhoid. About 40 species of bacteria pathogenic to humans have been naturally found in or on cockroaches. Among them are found, several agents of dangerous infections such as bubonic plague (Yersinia pestis (Lehmann and Neumann) van Loghem), dysentery (Shigella alkalescens (Andrewes)), diarrhea (Shigella paradysenteriae Duval-Sonne), uri- 816 Jean-Yves Rasplus & Alain Roques / BioRisk 4(2): 807–831 (2010) Figure 13.3.4. Comparative colonization of continental European countries and islands by the ‘Polyneoptera’ species alien to Europe. Archipelago: 1 Azores 2 Madeira 3 Canary islands. nary tract infection (Pseudomonas aeruginosa (Schroeter) Migula), abscesses (Staphylococcus aureus Rosenbach), food poisoning (Clostridium perfringens (Veillon and Zuber) Hauduroy et al, Escherichia coli (Migula) Castellani and Chalmers, Enterococcus faecalis (Andrewes and Horder) Schleifer and Kilpper-Bälz, P. aeruginosa), gastroenteritis (Salmonella spp.), typhoid fever (Salmonella typhi (Schroeter) Warren and Scott), leprosy (Mycobacterium leprae (Hansen) Lehmann and Neumann), and nocardiosis (Actinomyces spp). Several species of helminths are also transmitted by cockroaches, among them Schistosoma haematobium, Taenia saginata Goeze, Ascaris lumbricoides L., Ancylostoma duodenale (Dubini), and Necator americanus (Stiles) (Goddeeris 1980). Helminth eggs have been found naturally occurring in cockroaches, or appear in the faeces (Cochran 1999). Furthermore several virus, protozoa and fungi have been reported as occurring naturally in cockroaches and could also be transmited by these insects. However, proving unequivocally that cockroaches transmit disease to humans remains difficult Dictyoptera (Blattodea, Isoptera), Orthoptera, Phasmatodea and Dermaptera. Chapter 13.3 817 Figure 13.3.5. Main European habitats colonized by the ‘Polyneoptera’ species alien to Europe and alien in Europe. The number over each bar indicates the absolute number of alien species recorded per habitat. Note that a species may have colonized several habitats. (Baumholtz et al. 1997). However, costs associated with cockroaches are also linked to their control, either directly or indirectly through the use of pesticides that may facilitate emergence of pathogen resistance to some chemicals. Cockroaches are suspected to be important agents in the transmission of antibiotic resistant microbes in livestock production systems. Livestock production uses antibiotics therapeutically but this facilitates the emergence of resistant bacteria that may subsequently affect the human population. Finally, cockroaches can also damage household items, by eating glue in wallpaper, books, and furniture. The second group of ‘Polyneoptera’ with huge economic impact is termites. Termites play a critical ecological and agricultural role and some of them are pests. Some species (e.g. Cryptotermes brevis) has been introduced by human activity to almost every part of the world and cause severe damage to wooden structures. Reticulitermes Holmgren (Isoptera: Rhinotermitidae) are the major termite pests infesting structures and trees in Europe and the near East (Lohou et al. 1997). This genus contains the most significant termite pests of North America (the R. flavipes (Kollar) complex) and Europe (the R. lucifugus (Rossi) complex), and significant pest species in Asia (R. speratus (Kolbe)). Consequently, some of these species are susceptible to become major pests if they are introduced to Europe in the future. In Germany, R. flavipes appears to have been introduced on multiple occasions from USA with pine (Pinus spp.) logs (Harris 1962; UNEP 2000; Weidner 1978). This species had caused significant damage and costs for repair and control. The overall cost of treatments against termites in Europe may account for 1 billion euros by 2005 (UNEP 2000) whilst the estimated cost of termite damage could reach $20 billion annually (Su 2002). 818 Jean-Yves Rasplus & Alain Roques / BioRisk 4(2): 807–831 (2010) a b c e d Figure 13.3.6. Some Polyneoptera alien to Europe. a Pycnoscelus surinamensis (Blattodea) (Credit : Tom Murray) b Nala lividipes (Credit : MNHN Paris) c Cryptotermes brevis (Isoptera) (Credit : RH Scheffrahn) d Gryllodes sigillatus (Orthoptera) (Credit : JJ Argoud) e late instar nymph of Ancanthoxyla geisovii (Phasmatodea). (Credit: R. Hoare). 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UNEP, FAO (2000) United Nations Environment Programme and Food and Agriculture Organization. Termite biology and management workshop (Geneva: UNEP). 822 Jean-Yves Rasplus & Alain Roques / BioRisk 4(2): 807–831 (2010) Uvarov BP (1944) A New Zealand Phasmid (Orthoptera) established in the British Isles. Proceedings of the Royal Entomological Society of London, B 13: 94–96. Vigna Taglianti A (2005) Insecta Dermaptera. In: Ruffo S, Stoch F (Eds) Checklist e distribuzione della fauna italiana. Memorie del Museo Civico di Storia naturale di Verona, 2 .serie, Sezione Scienze della Vita 16: 141–142. Weidner H (1937) Termiten in Hamburg. Zeitschrift fiir Pflanzenkrankhr 47: 593. Weidner H (1938) Die Geradflügler (Orthopteroidea und Blattoidea) der Nordmark und Nordwest-Deutschlands. Verhandlungen des Vereins für naturwissenschaftliche Heimatforschung zu Hamburg 26: 25–68. Weidner H (1978) Die Gelbfussige Bodentermite Reticulitermes flavipes (Kollar 1837) in Hamburg (Isoptera). Eine Dokumenation zur Geschichte der angewandten Entomologie in Hamburg. Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 6: 49–100. Weidner H (1981) Einschleppung von Heuschrecken (Saltatoria und Phasmida) nach Hamburg. Anzeiger für Schaedlingskunde Pflanzenschutz Umweltschutz 54: 65–67. Wetterer JK, Hugel S (2008) Worldwide spread of the ant cricket Myrmecophilus americanus, a symbiont of the Longhorn Crazy ant, Paratrechina longicornis. Sociobiology 52: 157–165. Table 13.3.1. Blattodea, Isoptera, Orthoptera, Phasmatodea and Dermaptera species alien to Europe. List and characteristics. Status: A Alien to Europe C cryptogenic species. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Last update 01/03/2010 BLATTODEA Blaberidae Blaberus atropos (Stoll, 1813) Status Regime Native range First Record in Europe Invaded countries Habitat Host A detritovorous Tropical, subtropical Unknown DE, DK J Blaberus parabolicus (Walker,1868) Henschoutedenia flexivitta (Walker, 1868) Nauphoeta cinerea (Olivier, 1789) A detritovorous detritovorous detritovorous C&S America Africa Unknown DK J Unknown DE, DK J Reared C&S America Unknown CZ, DE, DK, GB J Reared for reptile pet food Panchlora fraterna Saussure & Zehntner, 1893 Panchlora peruana Saussure, 1864 Phoetalia circumvagans (Burmeister, 1838) Phoetalia pallida (Brunner, 1865) Pycnoscelus surinamensis (Linnaeus, 1767) A detritovorous C&S America Unknown DK J A detritovorous detritovorous detritovorous detritovorous C&S America Tropical, subtropical Tropical, subtropical AsiaTropical 1912, DK DK J Unknown ES-CAN J Unknown DK, ES-CAN J 1950, CZ CH, CZ, ES-CAN, FR, J1 GB, IE, IL, IS, PL, PTAZO, PT-MAD A A A A A With banana in or near human habitations, In or near human habitations, Tropical and subtropical moist places References Cornwell (1968), Holst (1986), Princis (1947) Holst (1986), Princis (1947) Holst (1986), Princis (1947) Cornwell (1968), Šefrová and Laštůvka (2005) Holst (1986), Princis (1947) Holst (1986), Princis (1947) Bland et al. (1996) Princis (1947) Asshoff and Coray (2003), Chopard (1922), Cornwell (1968), Šefrová and Laštůvka (2005) Dictyoptera (Blattodea, Isoptera), Orthoptera, Phasmatodea and Dermaptera. Chapter 13.3 823 Families Species Rhyparobia maderae (Fabricius, 1781) BLATTODEA Blattidae Blatta orientalis Linnaeus, 1758 Neostylopyga rhombifolia (Ståll, 1861) Regime Native range First Invaded countries Record in Europe Unknown DE, ES, ES-CAN, FRCOR Habitat References Cochran (1999) Food stores indoors, outdoors prefers to live in sugarcane fields, as well as palms, guava, and bananas growing next to the fields; fond of bananas and grapes. A detritovorous Africa C detritovorous detritovorous Cryptogenic Unknown DK Africa AL, CH, CZ, DE, DK, J1 ES-CAN, FI, FR, GB, GR-SEG, GR, HU, IE, IL, IT-SAR, IT-SIC, IT, RO, SK Omnivorous, synanthropic, Chopard (1922), warm and dry habitats Ragge (1973), Rehn (1945), Šefrová and Laštůvka (2005) C detritovorous Cryptogenic 1500, CZ Omnivorous, synanthropic; Alexander et al. decaying organic matter (1991), Šefrová and (sewers, drains, damp Laštůvka (2005) basements, porches, and other damp locations), outdoors in bushes, under leaf groundcover and mulch C detritovorous Cryptogenic Unknown AL, AT, BA, BE, BG, J1, J6 CH, CY, CZ, DE, DK, EE, ES-CAN, FI, FR-COR, FR, GB, GR-SEG, GR, HR, AT, HU, IE, IL, IS, IT-SAR, IT-SIC, IT, LV, LT, LU, MT, NL, NO, PL, PTAZO, PT-MAD, PT, RO, SE, SI, SK, SE, UA CZ G, I2 A 1945, DE J, I1 Host J Princis (1947) Omnivorous, synanthropic, Šefrová and Laštůvka warm climate; not cold (2005) tolerant, moist conditions Jean-Yves Rasplus & Alain Roques / BioRisk 4(2): 807–831 (2010) BLATTODEA Blatellidae Nyctibora laevigata (Beauvois, 1805) Supella longipalpa (Fabricius, 1798) Status 824 Families Species Status Regime Native range First Invaded countries Record in Europe 1600, IT AL, AT, BE, BG, CH, CZ, DE, DK, EE, ESCAN, ES, FI, FR, GB, GR-CRE, GR-NEG, GR-SEG, GR, HR, AT, HU, IE, IL, IS, IT-SAR, IT-SIC, IT, LV, LT, LU, MT, NO, PL, PT-AZO, PT-MAD, PT, SI, SK, SE 1927, DE AT, CH, CZ, DE, DK, ES-CAN, FI, FR, GB, AT, IE, IS, IT-SAR, ITSIC, IT, PL, SK, SE Periplaneta americana (Linnaeus, 1758) A detritovorous Africa Periplaneta australasiae (Fabricius, 1775) A detritovorous/ phytophagous AsiaTropical Periplaneta brunnea Burmeister, 1838 A detritovorous Africa Shelfordella lateralis (Walker, 1868) A detritovorous Central Asia 2009, ITSAR IT-SAR Asia IT DERMAPTERA Anisolabididae Euborellia stali (Dohrn, A parasitic/ 1864) predator Unknown 2002, IT CZ, ES-CAN, PTMAD, SK, SE Habitat Host References J1, H1, Omnivorous, synanthropic, Princis (1966), Ragge J100 warm climate; not cold (1945), Šefrová and tolerant, moist conditions Laštůvka (2005) J1, J100 Omnivorous, synanthropic, Asshof and Coray warm climates, moist, eat (2003), Mileke plants outdoors (2001), Princis (1966), Ragge (1945), Šefrová and Laštůvka (2005) J1 Near human habitats Šefrová and Laštůvka in cold climate; mainly (2005), Stejskal outdoors, under the bark (1993) of trees and in sewers in native and warm J Herbaceous places near Fois et al. (2009) human habitats, along streets. I Sugarcane field in native range Vigna- Taglianti (2005) Dictyoptera (Blattodea, Isoptera), Orthoptera, Phasmatodea and Dermaptera. Chapter 13.3 825 Families Species Status Regime Native range First Record in Europe Invaded countries Habitat Host References parasitic/ predator Tropical, subtropical 1915, ITSIC ES-BAL, ES-CAN, ES, FR, IT-SAR, IT-SIC, IT, PT I, J Granivore, predator, Economic pest of agricultural crops; hosts: Beta vulgaris (beetroot), Glycine max (L.) (soybean), Glossipium sp.(cotton), Helianthus annuus L. (sunflower), Sorghum sp. (sorghum) Albouy and Caussanel (1990) ISOPTERA Kalotermitidae Cryptotermes brevis A (Walker, 1853) phytophagous C&S America 1993, DE DE, ES-CAN, GB, IT, PT-AZO, PT J Soil, buildings Becker and Kny (1977), Fontana and Buzzetti (2003), Gay (1969), Nunes et al. (2010), Raineri (2001), Scheffrahn et al. (2001) ISOPTERA Rhinotermitidae Reticulitermes flavipes A (Kollar, 1837) phytophagous North America 1934, DE AT, DE, FR J Soil, buildings Austin et al. (2005, 2006), Clément et al. (2001), Feytaud (1924), Weidner (1937) ORTHOPTERA Acrididae Dociostaurus tartarus A Shchelkanovtsev, 1921 Locusta migratoria (L, A 1758) phytophagous phytophagous Asia 1962, BG BG E Africa 1886, FR AL, BG, DK, FR, FRCOR, HU, LV, PT F3 Migration ? Hubenov et al. (1998) Budrys and Pakalniskis (2007), Presa et al. (2007), Rey (1936) Jean-Yves Rasplus & Alain Roques / BioRisk 4(2): 807–831 (2010) DERMAPTERA Labiduridae Nala lividipes (Dufour, A 1828) 826 Families Species Status Regime Notostaurus albicornis A phyto(Eversmann, 1848)) phagous Ramburiella turcomana A phyto(Fischer von Waldheim, phagous 1846) ORTHOPTERA Bradyporidae Ephippigerida A unknown nigromarginata (Lucas, 1849) ORTHOPTERA Gryllidae Gryllodes sigillatus Walker A detrito1869 vorous ORTHOPTERA Myrmecophilidae Myrmecophilus americanus C detritoSaussure 1877 vorous ORTHOPTERA Phaneropteridae Topana cincticornis (Stal, A detrito1873) vorous ORTHOPTERA Raphidophoridae Tachycines asynamorus A detritoAdelung, 1902 vorous ORTHOPTERA Tettigoniidae Copiphora brevirostris A Stäl, 1873 phytophagous Native range Asia First Invaded countries Record in Europe 1964, BG BG Habitat Host E Tomov et al. (2009) Asia 1962, BG BG, MK E Petkovski (2009) Africa 1953, FR FR, IT-SIC F6 Asia Unknown DE, GB, NL J100 Cryptogenic Unknown DE U C&S America 1991, FR FR U Asia 1892, DE AT, BG, CH, DE, DK, EE, FR, GB, AT, IE, IT, LV J100 Omnivorous, greenhouses and botanical gardens Asshoff and Coray (2003), Detzel (2001), Geiter et al. (2002), Weidner (1981) C&S America Unknown DE J100 Greenhouses Detzel (2001) ? References Morin (2007) Geiter et al. (2002), Weidner (1981) Ant nests Geiter et al. (2002), Wetterer and Hugel (2008) Morin (2001) Dictyoptera (Blattodea, Isoptera), Orthoptera, Phasmatodea and Dermaptera. Chapter 13.3 827 Families Species Status Regime Native range Habitat I2 Botanic garden Bramble, Eucalyptus, Cupressus Phlugiola dahlemica A Eichler, 1938 PHASMATODEA Phasmatidae Acanthoxyla geisovii A (Kaup, 1866) phytophagous C&S America phytophagous Australasia 1908, GB GB I2, E5 Acanthoxyla inermis Salmon, 1955 Carausius morosus (Sinéty, 1901) Clitarchus hookeri (White, 1846) phytophagous phytophagous phytophagous Australasia 1981, GB GB I2, E5 Asia Unknown DE, GB I2, E5 Australasia 1900, GB GB, IE I2, E5 A A A Host References Weidner (1938) Lee (1993), Turk (1985), Uvarov (1944) Rose, Bramble, Eucalyptus Lee (1993), Turk (1985) Privet, Ivy, Hawthorn, Lee (1993), Weidner Pyracantha, Bramble, Rose (1981) Bramble, Eucalyptus, Lee (1993) Guava Jean-Yves Rasplus & Alain Roques / BioRisk 4(2): 807–831 (2010) First Invaded countries Record in Europe 1924, DE DE 828 Families Species Table 13.3.2. Blattodea, Isoptera, Orthoptera, Phasmatodea and Dermaptera species alien in Europe. List and characteristics. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Last update 01/03/2010. Status Regime DERMAPTERA Anisolabidae Anisolabis maritima E parasitic/ (Bonelli, 1832) predator First Record in Europe Invaded countries Habitat Host References Mediterranean Unknown region ? (Cosmopolitan) DE, DK, GB, IL B, J1, J6 Waste, algae in coastal Albouy and Caussanel areas (1990) Mediterranean 1837, IT region ? (Cosmopolitan) CZ, DE, DK, ESB, J1, J6 BAL, ES-CAN, ES, FR, GB, GR-CRE, GR-SEG, GR, HR, IL, IT-SAR, IT-SIC, IT, MT, NL, PTAZO, PT-MAD, PT, UA Omnivorous, on plant Albouy and Caussanel and animal material; (1990) minor nuisance in gardens unknown Mediterranean 1882, FRregion COR FR-COR, FR F9, J6 Under plane bark, along Adour river Albouy and Caussanel (1990) phytophagous Mediterranean 2005, PTregion AZO PT-AZO G, J Dry wood, forests, buildings Borges and Myles (2007) phytophagous Mediterranean Unknown, region DE DE J Soil, buildings Becker (1970) phytophagous Mediterranean Unknown region AL, DE, DK F6 DERMAPTERA Carcinophoridae Euborellia annulipes E parasitic/ (Lucas, 1847) predator, phytophagous DERMAPTERA Labidae Forficula smyrnensis E Serville, 1838 ISOPTERA Kalotermitidae Kalotermes flavicollis E (Fabricius 1793) ISOPTERA Rhinotermitidae Reticulitermes lucifugus E (Rossi 1792) ORTHOPTERA Acrididae Anacridium aegyptium E (Linnaeus 1764) Native range Weidner (1981) Dictyoptera (Blattodea, Isoptera), Orthoptera, Phasmatodea and Dermaptera. Chapter 13.3 829 Families Species Status Regime ORTHOPTERA Meconematidae Meconema meridionale E phytoCosta, 1860 phagous First Record in Europe Invaded countries Habitat Host References Mediterranean 1900, AT region AT, BE, GB, AT, NL I2 Urban parks; highway Couvreur and Godeau parkings (2000), Decleer et al. (2000), Kleukers (2002) Southern Europe 1956, AT AT U Gardens Ebner (1958), Essl and Rabitsch (2002) Mediterranean Unknown region DE G3, G4 Cliffs in pine stands (pinus nigra) Geiter et al. (2002) Mediterranean 1998, CZ region CZ J6 Cave, cellars Šefrová and Laštůvka (2005) Mediterranean 1999, FR region FR J Slate quarry Nöel et al. (2002) Mediterranean Unknown region Mediterranean Unknown region GB I2, E5 Bramble, rose Lee (1993) GB I2, E5 Bramble, Broom Lee (1993) Jean-Yves Rasplus & Alain Roques / BioRisk 4(2): 807–831 (2010) ORTHOPTERA Phaneropteridae Leptophyes punctatissima E phyto(Bosc, 1792) phagous ORTHOPTERA Rhaphidophoridae Dolichopoda bormansi E detritoBrunner von Watt., vorous 1882 Troglophillus neglectus E detrito(Kraus, 1879) vorous ORTHOPTERA Tettigoniidae Antaxius spinibrachius E detrito(Fischer, 1853) vorous PHASMATODEA Bacillidae Bacillius rossius (Rossi, E phyto1788) phagous Clonopsis gallica E phyto(Charpentier, 1825) phagous Native range 830 Families Species Dictyoptera (Blattodea, Isoptera), Orthoptera, Phasmatodea and Dermaptera. Chapter 13.3 831 Table 13.3.3. Number of alien ‘polyneoptera’ per European country. Countries Germany mainland Denmark Spain Canary islands France mainland Great Britain Czech Republic Italy mainland Bulgaria Ireland Italy Sicily Switzerland Italy Sardinia Austria Portugal mainland Slovakia Albania Finland mainland Hungary Iceland Israel Latvia Poland Portugal Azores Portugal Madeira N 17 14 11 11 10 8 8 7 7 6 6 6 5 5 5 4 4 4 4 4 4 4 4 4 Countries Sweden Estonia Greece South Aegean Greece mainland Spain mainland Belgium Croatia France Corsica Lithuania Luxemburg Malta Norway mainland Netherlands Romania Slovenia Bosnia Cyprus Greece Crete Greece North Aegean Macedonia Serbia Spain Balearic islands Ukraine N 4 3 3 3 3 2 2 2 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 A peer reviewed open access journal BioRisk 4(2): 833–849 (2010) doi: 10.3897/biorisk.4.65 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Lice and Fleas (Phthiraptera and Siphonaptera) Chapter 13.4 Marc Kenis1, Alain Roques2 1 CABI Europe-Switzerland, 1, Rue des Grillons, CH- 2800, Delémont, Switzerland 2 Institut National de la Recherche Agronomique (INRA), UR 0633, Station de Zoologie Forestière, 2163 Av. Pomme de Pin, 45075 Orléans, France Corresponding authors: Marc Kenis (m.kenis@cabi.org), Alain Roques (alain.roques@orleans.inra.fr) Academic editor: David Roy | Received 26 March 2010 | Accepted 25 May 2010 | Published 6 July 2010 Citation: Kenis M, Roques A (2010) Lice and Fleas (Phthiraptera and Siphonaptera). Chapter 13.4. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 833–849. doi: 10.3897/biorisk.4.65 Abstract A total of 31 Phthiraptera species alien to Europe are listed. They include 24 chewing lice and seven sucking lice of 12 different families. The families Goniodidae (Ischnocera) and Menoponidae (Amblycera) largely dominate the alien entomofauna of chewing lice. Asia is the major supplier of alien Phthiraptera which are mostly associated with poultry farming, game birds, guinea pigs and invasive alien mammals. The recent period did not show any acceleration in alien arrival in Europe. Alien fleas include six species in the families Pulicidae and Ceratophyllidae. Three of them are primarily associated with rats and are capable of transmiting major human diseases such as the bubonic plague and the murine typhus. Keywords Phthiraptera, lice, flea, Siphonaptera, alien, Europe 13.4.1. Introduction Phthiraptera (lice) and Siphonaptera (fleas) are obligate ectoparasitic insects of birds and mammals, including humans. Some are of high importance for human and animal health because they cause itches and skin infection, and transmit serious diseases, e.g. the head louse (Pediculus capitis De Geer), the crab louse (Phtirus pubis (L.)), the cat flea (Ctenocephalides felis felis (Bouché)), the rat flea (Xenopsylla cheopis (Rothschild)) or the human flea (Pulex irritans L.). Although many of these are of unknown origin, they Copyright M. Kenis, A. Roques. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 834 Marc Kenis & Alain Roques / BioRisk 4(2): 833–849 (2010) are probably allochtonous in Europe, having arrived in ancient times with their hosts (Mey, 1988; Beaucournu and Launay, 1990). Thus, Pulex irritans was shown to have been present in Europe since the Bronze Age at least, having been found in remains of lake dwellings in the French Jura, dating back to 3100 B.C. (Yvinec et al 2000). Only the species considered as possibly neozoans*, i.e. 27 lice and six fleas, were originally included in the DAISIE database. Four further species have subsequently been added to the list of alien species and this review is therefore based on 31 species. Although a large part of these alien species were recorded in Europe for the first time at the end of the 19th century, many probably came much earlier; the exact date of arrival remaining unclear in nearly all cases. 13.4.2 Phthiraptera Lice are exopterygotes* of birds and mammals. Most species are host-specific but others are rather polyphagous. They spend their entire life on their host animal, feeding on epidermal tissue debris, parts of feathers, blood or sebaceous secretions. Until recently, they were divided into two orders, Anoplura (sucking lice) and Mallophaga (chewing lice), but they are presently grouped into a single order, Phthiraptera (Barker et al 2003; Price et al 2003). The order Phthiraptera comprises about 5,000 described species present in four sub-orders, Anoplura (543 spp. on mammals), Amblycera (ca. 1360 spp. on birds, mammals and marsupials), Ischnocera (ca. 3080 spp. on birds and mammals) and Rhynchophthirina (3 spp. on elephants and warthogs), this latter group being not present in Europe (Smith 2003). A total of 31 Phthiraptera species alien to Europe have been listed here, including 16 species known to be of exotic origin and 14 cryptogenic species, to be compared to the 691 species considered as native to Europe included in Fauna Europaea (Mey 2005). They include 24 chewing lice belonging to 8 different families and 7 sucking lice belonging to 4 different families (Table 13.4.1). Three of the families have no representatives in Europe (Gliricolidae, Gyropidae, Trimenoponidae; all in the Amblycera suborder). The families Goniodidae (Ischnocera) and Menoponidae largely dominate the alien entomofauna (Figure 13.4.1). In a number of families, the arrival of aliens has largely modified the composition of the total entomofauna currently present in Europe. In contrast to the trends reported in other arthropod groups, the majority of the alien lice were first observed in Europe during the 18th and 19th century (18 species out of 31- 58.1%), although they probably arrived much earlier with their animal host, in most cases a domestic species. The recent period did not show any acceleration in alien arrival in Europe with only 4 species (12.9 % of the total species) newly observed during the period 1975- 2007. Eight out the 17 alien species of known exotic origin came from Asia (47.0 %), with earlier arrival dates than those from North America (4 spp.; 23.5 %) or South America (4 spp.). Several chewing lice of cryptogenic origin are important pests of poultry farming, in particular Menopon gallinae, Goniocotes gallinae and Eomenacanthus stramineus Chapter 13.4: Lice and Fleas (Phthiraptera and Siphonaptera) 835 Figure 13.4.1. Relative importance of the Phthiraptera and Siphonaptera suborders and families in the alien and native fauna in Europe. Families of Phthiraptera are presented per suborder in a decreasing order based on the number of alien species. Species alien to Europe include cryptogenic species. The number over each bar indicates the number of species observed per family. (Sychra et al 2008). Other species parasitize pheasants (Phasianus spp.) and came with their host from Asia, such as Goniocotes chrysocephalus, Lagopoecus colchicus, Lipeurus maculosus, Uchida phasiani, Zlotorzyckella colchici (Kopocinski et al 1998). Chewing lice parasitising mammals in Europe are listed in Mey (1988). Some species are known to be of alien origin, such as the three South American species, Gyropus ovalis, Gliricola porcelli and Trimenopon hispidum, arriving in Europe with guinea pigs (Cavia porcellus L.) and causing scratching, loss of hair, and scabs to domestic and laboratory animals. Other species worth mentioning are the cryptogenic dog louse, Trichodectes canis, and the sheep louse, Bovicola ovis, which cause pruritus and skin infections such as eczema to their host animal. Finally, a few species are associated with invasive alien mammals, such as the South American Pitrufquenia coypus on coypu (Myocastor coypus (Molina)); (Laurie 1946; Newson and Holmes 1968) and the North American Trichodectes (Stachiella) octomaculatus on raccoon (Procyon lotor (L.)); (Hellenthal et al 2004). Only seven sucking lice of four families (Enderleinellidae, Hoplopleuridae, Linognathidae, and Polyplacidae) are considered Neozoans in Europe (Table 13.4.1). The Asian Polyplax spinulosa (spined rat louse) causes hair loss and pruritus to wild and domestic rats (Rattus spp.). The cryptogenic species Linognathus stenopsis and Haemodipsus lyriocephalus parasitize goats (Capra hircus L.) and hares (Lepus europaeus Pallas), respectively. According to Durden and Musser (1994), another Haemodipsus species, H. setoni Ewing associated with Lepus spp. in North America is possibly an introduced species in Eurasia (this species has not been included here). Three species have been introduced to Europe with their Sciuridae host from either North America (Enderleinellus longiceps and Hoplopeura sciuricola with grey squirel, Sciurus carolinensis Gmelin; Britt and Molineux 1979) or Asia (Enderleinellus tamiasis with Siberian chipmunk, Tamia sibiricus (Laxmann); Beaucornu et al 2008). Solenopotes muntiacus has also been 836 Marc Kenis & Alain Roques / BioRisk 4(2): 833–849 (2010) Figure 13.4.2. Alien Phthiraptera (Anoplura). Solenopotes muntiacus female from Muntjac deer, Muntiacus muntjak (Credit: British Museum of Natural History, London) introduced from Asia to Great Britain with muntjac deers, Muntiacus reevesi (Ogilby) (Dansie et al 1983). In addition, Haemodipsus ventricosus (Denny) which lives on rabbits (Oryctolagus cuniculus L.) can be considered as alien in Europe, originating, as its host, from the Iberic pensinsula (Durden and Musser 1994). 13.4.3 Siphonaptera Fleas are holometabolous insects whose adults must feed on blood of mammals and birds in order to reproduce. Larvae feed on organic matter, often in the host’s nest. In the DAISIE database, six fleas are listed as alien to Europe, including 5 species known to be of exotic origin and 1 cryptogenic species, in comparison to the 260 species considered as native to Europe (Soledad Gomez Lopez 2005) (Table 13.4.1). The aliens belong equally to two families, Pulicidae and Ceratophyllidae, whereas the latter family largely dominates the native entomofauna. Three of these fleas have rats as their main host (Beaucornu and Launay, 1990). The tropical rat flea, Xenopsylla cheopis, probably originates from the Nile area (Beaucornu 1999). It became synanthropic in most of Southern Europe where it could not survive before because of large temperature variations between summer and winter within human habitats (Beaucornu 1999). X. brasiliensis, originates from tropical Africa and invaded the Canary islands (Beaucornu and Launay, 1990); it has also been Chapter 13.4: Lice and Fleas (Phthiraptera and Siphonaptera) 837 Figure 13.4.3. Alien Phthiraptera (Amblycera). Gliricola porcelli male from guinea pig, Cavia porcellus (Credit: British Museum of Natural History, London) found sporadically in port areas and elsewhere, e.g. it was recorded from Wales in the 1950s (Hopkins and Rotschild 1953). The third species, Nosopsyllus fasciatus, is a temperate species from Asia. Rat fleas are also able to feed on other mammals, including humans, to which they can transmit the bubonic plague by carrying the bacteria Yersinia pestis (Audouin-Rouzeau, 2003). Xenopsylla cheopis is also a vector of another human disease, the murine typhus fever caused by the bacteria Rickettsia typhi (Beaucournu and Launay, 1990). The North American species Orchopeas howardi is found on the grey squirrel (Sciurus carolinensis), an invasive rodent in Europe (Keymer, 1983). In addition, a rabbit flea, Spilopsyllus cuniculi (Dale), can be considered as alien in Europe, probably originating with its host from the Iberian Peninsula. It has invaded a large part of Western and Central Europe (Soledad Gomez Lopez 2005). 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List and characteristics of the Phthiraptera and Siphonatera species alien to Europe. Status: A Alien to Europe C cryptogenic species. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Last update 27/03/2010; Polyplacidae Haemodipsus lyriocephalus (Burmeister, 1839) C parasitic/ Cryptopredator genic 1st record Invaded countries Habitat in Europe Host References 1979 GB G, I2 Sciurus carolinensis Britt and Molyneux (1979), O’Connor et al (2005) 1916 DE, FR G, I2 Tamias sibiricus Beaucournu et al (2008), Durden and Musser (1994) 1976 GB, IE G, I2 Sciurus carolinensis Britt and Molyneux (1979), O’Connor et al (2005) 1838 BG, CH, CZ, DE, J FR, GR, IT 1983 GB 1839 BG, CH, CZ, DE, E FI, FR, GB, IT, NL, PL G Goat (Capra) Fauna Italia (2003), Himonas and Liakos (1989), Piaget (1880), Šefrová and Laštùvka (2005), Séguy (1924, 1944) Touleshkov(1954) Muntjac Dansie et al (1983), Durden and deer Musser (1994) (Muntiacus reevesi) Hares Broekhuizen (1971), Fauna Italia (Oryctolagus) (2003), Geiter et al (2002), Kenis (2005), Piaget (1880), Séguy (1924, 1944), Thompson (1939), Touleshkov(1954), Wegner (1966), Wegner and Eichler (1968), Büttiker and Mahnert (1978), Šefrová and Laštùvka (2005), Silfverbeg (1984) Chapter 13.4: Lice and Fleas (Phthiraptera and Siphonaptera) Order Species Status Regime Native Suborder range Family Phthiraptera Anoplura Enderleinellidae Enderleinellus A parasitic/ North longiceps Kellogg predator America & Ferris, 1915 Enderleinellidae Enderleinellus A parasitic/ Asia tamiasis predator (Korea) Fahrenholz, 1916 Hoplopleuridae Hoplopleura A parasitic/ North sciuricola Ferris, predator America 1921 Linognathidae Linognathus C parasitic/ Cryptostenopsis predator genic (Burmeister, 1838) Linognathidae Solenopotes A parasitic/ Asia muntiacus predator Thompson, 1938 843 Species Status Regime Native range 1st record Invaded countries Habitat in Europe Host A parasitic/ Asia predator 1839 BG, CH, CZ, DE, J ES, FI, FR, HR, PL Rats(Rattus spp.) Gliricola porcelli (Schrank 1781) A parasitic/ C & S predator America 1781 AT, BG, CH?, DE, J ES, FI, FR, HU, IT, PL, RO, SI Guinea pigs (Cavia porcellus) Gyropidae Gyropus ovalis Burmeister, 1838 A parasitic/ C & S predator America 1838 AT, BG, CH?, DE, J ES, FI, FR, HR, HU, IT, PL Gliricolidae Pitrufquenia coypus Marelli 1932 Eomenacanthus stramineus (Nitzsch 1818) A parasitic/ C & S predator America 1932 AT, BE, CH? , DE, GB C2 C parasitic/ Cryptopredator genic 1818 BG, DE, ES, FI, FR, IT, PL, RS, UA E, J Phthiraptera Amblycera Gliricolidae Menoponidae Geiter et al (2002), Gomez et al (1987), Kenis (2005), Šefrová and Laštùvka (2005), Séguy (1944), Silfverbeg (1984), Stojcevic et al (2004), Touleshkov (1954) Bordeaul (2008), Fauna Italia (2003), Geiter et al (2002), Kenis (2005), Mouchet and Morel (1957), Paradiznik (1989), Piaget (1880), Schrank (1781), Séguy (1924, 1944) Touleshkov (1955a) Guinea Bordeau (2008), Fauna Italia pigs (Cavia (2003), Geiter et al (2002), Kenis porcellus) (2005), Mouchet and Morel (1957), Piaget (1880), Séguy (1924, 1944), Stojcevic et al (2004),Touleshkov (1955a) Coypu Hellenthal et al (2004), Kenis (Myocastor (2005), Laurie (1946), Newson and coypus) Holmes (1968) Pheasant Geiter et al (2002), Ilieva (2009), (Phasianus), Mouchet and Morel (1957), Nitzsch Domestic (1818), Pavlovic and Nesic (1991), fowl (Gallus Prelezov and Koinarski (2006), gallus Séguy (1924, 1944), domesticus), Turkey (Meleagris) Marc Kenis & Alain Roques / BioRisk 4(2): 833–849 (2010) Polyplax spinulosa (Burmeister, 1839) References 844 Order Suborder Family Polyplacidae Order Suborder Family Menoponidae Species Status Regime Native range 1st record Invaded countries Habitat in Europe parasitic/ Cryptopredator genic 1880 C parasitic/ Cryptopredator genic 1781 Menoponidae Myrsidea quadrifasciata (Piaget, 1880) A parasitic/ Asia predator Menoponidae Neocolpocephalum turbinatum (Denny 1842) Uchida phasiani (Modrzejewska & Zlotorzycka, 1977) Trimenopon hispidum Burmeister, 1838 C Bovicola (Bovicola) ovis (Schrank, 1781) Menoponidae Menoponidae Trimenoponidae Phthiraptera Ischnocera Bovicoliidae BE, BG, DE, ES, FI, FR, HU, PL, RO BE, BG, DE, ES, FI, FR, GB, HU, IT, PL, RO, RS, UA J 1880 BE, CZ, DE, FR, HU, IT J, J1 parasitic/ Cryptopredator genic 1842 G, J A parasitic/ Asia predator 1998 BG, DE, ES, FR, GB, HU, IT, ITSAR, PL, RO CZ, DE, PL A parasitic/ C & S predator America 1966 AT, CH?, DE, FR, J FI, HU, PL Guinea pigs (Cavia porcellus) C parasitic/ Cryptopredator genic 1916 BE, BG, CZ, ES, FI, FR, GB, HU, IT, LT, NL, PL, RO Sheep (Ovis) Cummings (1916), Hellenthal et al (2004), Šefrová and Laštùvka (2005), Séguy (1944), Silfverbeg (1984), Touleshkov (1955b) J E, J J Columba Hellenthal et al (2004), Ilieva (2009), Piaget (1880), Séguy (1924, 1944), Touleshkov (1974) Denny (1842), Geiter et al (2002), Domestic fowl (Gallus Hellenthal et al (2004), Ilieva (2009), Mouchet and Morel (1957), gallus domesticus), Pavlovic and Nesic (1991), Pigate Turkey (1880), Prelezov and Koinarski (Meleagris) (2006), Schrank (1781), Séguy (1924, 1944), Silfverbeg (1984), Touleshkov (1955a) House Hellenthal et al (2004), Piaget sparrow (1880), Šefrová and Laštùvka (Passer (2005), Séguy (1924, 1944), domesticus) Falcons Denny (1842), Geiter et al (2002), (Falco), Ilieva (2009), Piaget (1880), Séguy Columba (1944), Touleshkov (1957) Pheasant Šefrová and Laštùvka (2005) (Phasianus) Geiter et al (2002), Kenis (2005), Mouchet and Morel (1957) 845 C References Chapter 13.4: Lice and Fleas (Phthiraptera and Siphonaptera) Hohorstiella gigantea lata (Piaget 1880) Menopon gallinae (L. 1758) Host Species Status Regime Native range 1st record Invaded countries Habitat in Europe A parasitic/ North predator America 1877 CZ, DE, ES, FI, FR, HU, IT, NL, PL, PT, RO Goniocotes chrysocephalus Giebel 1874 Goniocotes gallinae (De Geer 1778) C parasitic/ Cryptopredator genic 1874 C parasitic/ Cryptopredator genic 1880 BE, DE, FR, HU, E, J IT, NL, PL, RO, ES BE, BG, DE, ES, J FI, FR, HU, IT, PL, RS, UA Goniodidae Goniodes pavonis (Linnaeus, 1758) C parasitic/ Cryptopredator genic 1892 BG, DE, FI, FR, HU, IT, PL, RO J Goniodidae Goniocotes rectangulatus Nitzsch, 1818 C parasitic/ Cryptopredator genic 1818 DE, HU, RO J Goniodidae Stenocrotaphus gigas (Taschenberg 1879) A parasitic/ Tropical, predator subtropical 1924 BE, BG, DE, ES, FR, GB, IT, PL J1 Goniodidae Goniodidae J1, G References Wild and domesticated Turkey (Meleagris) Pheasant (Phasianus spp.) Domestic fowl (Gallus gallus domesticus) Geiter et al (2002), Mouchet and Morel (1957), Piaget (1880), Šefrová and Laštùvka (2005), Séguy (1924, 1944), Fauna Italia (2003), Geiter et al (2002), Hellenthal et al (2004), Piaget (1880), Séguy (1924, 1944), Geiter et al (2002), Hellenthal et al (2004), Mouchet and Morel (1957), Pavlovic and Nesic (1991), Piaget (1880), Prelezov and Koinarski (2006), Séguy (1944), Touleshkov (1955a) Geiter et al (2002), Fauna Italia (2003), Séguy (1924, 1944), Touleshkov (1955a) Indian Peafowl (Pavo cristatus) Helmeted Guinea Fowl (Numida meleagris), Indian Peafowl (Pavo cristatus) Domestic fowl (Gallus gallus domesticus), Turkey (Meleagris) Geiter et al (2002), Nitzsch (1818), Piaget (1880) Geiter et al (2002), Hellenthal et al (2004), Ilieva (2009), Séguy (1924), Touleshkov (1955a) Marc Kenis & Alain Roques / BioRisk 4(2): 833–849 (2010) Chelopistes meleagridis (Linnaeus, 1758) Host 846 Order Suborder Family Goniodidae Order Suborder Family Goniodidae Philopteridae Philopteridae Philopteridae Trichodectidae Trichodectidae Zlotorzyckella colchici (Denny, 1842) Cuclotogaster heterographa (Nitzsch in Giebel 1866) Lagopoecus colchicus Emerson, 1949 Lipeurus maculosus Clay, 1938 Native range 1st record Invaded countries Habitat in Europe Host References A parasitic/ Asia predator 1977 BE, CZ, DE, ES, IT, PL, RO G, I2 Pheasant (Phasianus) Dlabola (1977), Hellenthal et al (2004) C parasitic/ Cryptopredator genic 1876 BE, BG, DE, ES, FI, FR, HU, IT, NL, PL, RO,UA J Domestic fowl (Gallus gallus domesticus) A parasitic/ Asia predator 1989 BE, CZ, DE, PL G, I2 A parasitic/ Asia predator 1938 BE, CZ, DE, GB, G, I2 HU, IT, PL, RO C parasitic/ Cryptopredator genic 1880 BG, DE, FR, IT Pheasant (Phasianus colchicus) Pheasant (Phasianus colchicus), Partridge (Perdrix perdrix) Turkey (Meleagris) Fauna Italia (2003), Geiter et al (2002), Hellenthal et al (2004), Mouchet and Morel (1957), Piaget (1880), Séguy (1924, 1944), Touleshkov (1955a) Geiter et al (2002), Hellenthal et al (2004), Šefrová and Laštùvka (2005) A parasitic/ North predator America Unknown AT, BE, CH?, DE F9 Raccoon (Procyon lotor) C parasitic/ Cryptopredator genic <1880 Dogs (Canis Fauna Italia (2003), Hellenthal et al domesticus) (2004), Mouchet and Morel (1957), Piaget (1880), Séguy (1924, 1944), Touleshkov (1955b) BE, BG, DE, ES, FI, FR, IT, PL J J Clay (1938), Dlabola (1977), Fauna Italia (2003), Geiter et al (2002), Hellenthal et al (2004) Fauna Italia (2003) Geiter et al (2002), Mouchet and Morel (1957), Piaget (1880), Séguy (1944) Geiter et al (2002), Hellenthal et al (2004), Kenis (2005) 847 Reticulipeurus (=Oxylipeurus) polytrapezius (Burmeister 1838) Trichodectes (Stachiella) octomaculatus Paine 1912 Trichodectes (Trichodectes) canis (De Geer 1778) Status Regime Chapter 13.4: Lice and Fleas (Phthiraptera and Siphonaptera) Philopteridae Species A parasitic/ Asiapredator Temperate 1985 CH 1811 AL, AD, AT, BA, J1, J2 BE, BG, CH, CY, CZ, DE, DK, EE, ES, ES-BAL, ES-CAN, FI, FÖ, FR, FR-COR, GR, GR-CRE, GR_NEG, GRSEG, GB, HR, HU, IE, IS, IT, IT-SAR, IT-SIC, LI, LT, LU, MD, MK, MT, NL, PL, PT, PT-AZO, PTMAD, RO, SK AT, BE, CH, CZ, E,J DE, DK, ES, ES-BAL, FÖ, FR, FR-COR, GB, GR, HU, IE, IT, IT-SAR, IT-SIC, LT, LU, ME, MT, NL, PL, PT, PTAZO, PT-MAD, RO, RS, SK 1900 J1 Host References Man (Colomba livia in the native range) Mus domesticus, Rattus rattus and other Muridae (fur fleas) Beaucornu and Aeschlimann (1985) Rattus spp., Apodemus spp., Mus spp. and other Muridae Beaucornu (1972, 1976, 1978), Beaucornu and Alcover (1984), Beaucornu and Launay (1990), Beaucornu and Pascal (1998), GalliValerio (1900), Krause (1911), Mifsud et al (2008), Peus (1963), Smit (1957, 1966), Soledad Gomez Lopez (2009) Beaucornu and Launay (1990), Dale (1878), Rotschild (1899), Soledad Gomez Lopez (2009), Stojcevic et al (2004) Marc Kenis & Alain Roques / BioRisk 4(2): 833–849 (2010) Ceratophyllidae Nosopsyllus (Nosopsyllus) fasciatus (Bosc dAntic, 1800) 1st record Invaded countries Habitat in Europe 848 Order Species Status Regime Native Suborder range Family Siphonaptera Ceratophyllidae Callopsylla A parasitic/ Asia (Geminopsylla) predator gemina (Ioff, 1946) Ceratophyllidae Leptopsylla C parasitic/ Crypto(Leptopsylla) predator genic segnis (Schönherr, 1811) Order Species Suborder Family Ceratophyllidae Orchopeas howardi Baker 1895 Status Regime Native range 1st record Invaded countries Habitat in Europe A parasitic/ North predator America 1800 GB, IE G, X11 Euhoplopsyllus glacialis affinis (Baker, 1904) A parasitic/ North predator America 1977 FR, IT E, F, G Pulicidae Xenopsylla brasiliensis (Baker, 1904) A parasitic/ Africa predator 1942 ES-CAN, GB J1 Pulicidae Xenopsylla cheopis cheopis (Rothschild, 1903) A parasitic/ Africa( predator Nile region) 1904 DE, ES, ES-CAN, J1 FR, FR-COR, GB, GR, HU, IE, IT-SIC, IT, MT, PL, PT-AZO, PTMAD, PT, RU Anonymous (1994),Donisthorpe (1925) Beaucornu and Launay (1977), Beaucornu et al (1981), Fauna Italia (2003) Beaucornu and Launay (1990), Hopkins and Rotschild (1953), Najera (1942), Smit (1957), Bernard et al (1947), Beaucornu and Launay (1990), Cartana Castella and Gil-Collado (1934), Giles (1905), Ilvento (1913), Lavier (1921), Najera (1942), Séguy (1924), Tanon (1923), Tiraboschi (1904), Zapatero-Ramos et al (1982), 849 Sciurus carolinensis (grey squirrel), Clethrionomys glareolus, Glis glis, Dama dama, Vulpes vulpes, Oryctolagus cuniculus cottontail, rabbit Sylvilagus floridanus, Oryctolagus cuniculus Rattus spp., vector of plague and murine typhus Rattus norvegicus, R. rattus, humans, Mus musculus; vector of plague References Chapter 13.4: Lice and Fleas (Phthiraptera and Siphonaptera) Pulicidae Host A peer reviewed open access journal BioRisk 4(2): 851–854 (2010) doi: 10.3897/biorisk.4.47 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Springtails and Silverfishes (Apterygota) Chapter 13.5 Jürg Zettel University of Bern, Institute of Ecology and Evolution Baltzerstrasse 6, CH-3012 Bern, Switzerland Corresponding author: Jürg Zettel (juersi.zettel@bluewin.ch) Academic editor: David Roy | Received 27 January 2010 | Accepted 25 May 2010 | Published 6 July 2010 Citation: Zettel J (2010) Springtails and Silverfishes (Apterygota). Chapter 13.5. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(2): 851–854. doi: 10.3897/biorisk.4.47 Abstract The alien fauna of Apterygota is still limited in Europe. Springtails (Collembolla) alien to Europe include only three species to which add a cryptogenic one. Two nowadays cosmopolitan species of silverfishes may originate from Central America. The reasons of this limited colonisation of Europe are briefly discussed. Keywords Apterygota, Collembola, springtails, Zygentoma, silverfishes 13.5.1. Diplura alien to Europe No introductions of alien species into Europe are known. 13.5.2. Collembola (Springtails) alien to Europe Worldwide ca. 6500 collembolan species are listed, belonging to 18 families (Hopkin 1997). For Europe, there are estimated to be ca. 1500 species, belonging to 16 families (taxonomic work is still progressing). Collembola are the most abundant terrestrial arthropods, colonising all soil habitats that provide enough humidity and food, such as organic matter or microorganisms. Example habitats include root rosettes of high alpine plants, plant debris on the shore, natural soils, as well as microhabitats such as flower pots. Most species are soil or Copyright Jürg Zettel. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 852 Jürg Zettel / BioRisk 4(2): 851–854 (2010) litter dwellers, whilst only few species live on the surface or in the vegetation (mainly Entomobryidae and Symphypleona). In mature soil, abundances may attain values of 50–100‘000 individuals/m2. Local gradations in abundance are a well known phenomenon in many Collembola. As detritivores, Collembola are not generally considered as pest species. Exceptions are two species of Symphypleona living above ground in the vegetation layer: the European Sminthurus viridis which became a severe pest in Australia on alfalfa, clover etc, and the ubiquitous Bourletiella hortensis is known to feed on vegetable seedlings when natural food (weed seedlings) is absent. The ubiquitous onychiurid Protaphorura armata also switches food source in the absence of weeds, but only as a secondary pest when feeding on wounds infected by microorganisms. In Europe, no Collembola are declared as agronomic pests (e.g. in CABI Crop Protection Compendium) (CABI 2009). A 100 or more Collembola species may occur in the same soil habitat, and through occupying all available niches are believed to preclude establishment of alien species. To date, alien Collembola have only been observed to become invasive and replace indigenous species in isolated microhabitats and in extreme climates such as two species on sub-Antarctic islands (Convey et al. 1999, Greenslade 2002). Identifying alien Collembola is difficult due to the limited number of specialists investigating soil fauna. No intentional introductions to Europe have occurred. Unintentional transport within soil of ornamental plants, with vegetables, dirty equipment and vehicles easily moves Collembola over large distances. Short life cycles and parthenogenetic development of a number of species may also increase the chances to colonise new sites. Therefore the distribution ranges of alien species within Europe may increase steadily. Only three records of alien Collembola introduced to Europe have been published. These concern two species in the family Isotomidae, Proisotoma filifera, originating from Central America but found in Dutch greenhouses (Ellis 1970), and Desoria trispinata, that originates from North America but has appeared in anthropogenic habitats, mainly towns (Christian 1987, Christian and Kindl-Stomatopolos 1999, Kindl- Stomatopoulos). A third species in the family Onychiuridae, Onychiurus folsomi, originating from Australia, is restricted to earthworm cultures in Spain (Arbea and Jordana 1988). In addition, we considered a cryptogenic species, Sminthurinus trinotatus (Katiannidae), which presents a very disjunct known distribution (southern Europe, eastern Asia). 13.5.3. Zygentoma (Silverfishes) alien to Europe Zygentoma or silverfishes comprise five families (in Europe Lepismatidae only) with 12 genera (three in Europe) and ca. 370 species (ten in Europe). The two (today) cosmopolitan species Ctenolepisma longicaudata and Thermobia domestica (both Lepismatidae), may originate from central parts of America. Once moved from western Mediterranean regions to central and northern Europe, they mainly colonise anthropogenic habitats, where they may become pests by destroying paper or stored products. Springtails and Silverfishes (Apterygota). Chapter 13.5 853 References Arbea JL, Jordana R (1988) Nota sobre la presencia masiva de Onychiurus folsomi Schaeffer (Collembola, Onychiuridae) en lechos de Eisenia andrei (Oligochaeta, Lumbricidae). Boletin de Sanidad Vegetal Plagas 14: 535–540. CABI Crop Protection Compendium. http://www.cabicompendium.org/cpc. Christian E (1987) Collembola (Springschwänze). Catalogus Faunae Austriae XIIa, Wien: Österreichische Akademie der Wissenschaften. 80 pp. Christian E, Kindl-Stamatopolos L (1999) Arthropods on plastered riverbanks in the builtup area of Vienna. In: Tajovský K, Pižl V (Eds) Soil Zoology in Central Europe. Proceedings of the 5th Central European Workshop on Soil Zoology, April 1999. České Budějovice: Institute of Soil Biology, 27–30. Convey P, Greenslade P, Arnold RJ, Block W (1999) Collembola of subantarctic South Georgia. Polar Biology 22: 1–6. Ellis WN (1970) Proisotoma filifera Denis in Holland, with a note on its classification (Collembola, Isotomidae). Entomologische Berichten 30: 18–24. Essl F, Rabitsch W (2002). Neobiota in Österreich. Wien: Umweltbundesamt. 432 pp. 8-Greenslade P (2002) Assessing the risk of exotic Collembola invading subantarctic is lands: priorising quarantine management. Pedobiologia 46: 338–344. Hopkin SP (1997). Biology of the springtails (Insecta: Collembola). Oxford: Oxford University Press. 330 pp. Hopkin SP. The Natural World in Close Up. available at [www.stevehopkin.co.uk]. Kindl-Stamotopolos L (2001) Arthropoden des Wienflussufers im dicht bebauten Stadtgebiet Wiens. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Österreich 138: 1–15. Collembola- Katiannidae Sminthurinus trinotatus C (Axelson, 1905) Zygentoma- Lepismatidae Ctenolepisma C longicaudata (Escherich, 1905) Thermobia domestica C (Packard, 1873) 1st record in Europe Invaded countries Habitat References 1988 ?, ES ES, GB I (Vermiculture), Arbea and Jordana (1988), J100 Hopkin (2009) 1968, NL NL J100 Ellis (1970) ca 1900, AT AT, DE, IT, NO, PT, RU J Christian (1987), Christian and KindlStomatopolos (1999), Kindl-Stamotopolos (2001) detritivorous South Europe, East Asia? 1925, GB AT, DE, FR, GB, IT I2, J100 Essl and Rabitsch (2002), Hopkin (2009) detritivorous Central America? Unknown CY, FR, IT, MT, PT J1 (stored products) Essl and Rabitsch (2002) detritivorous Central America? Unknown CY, DE, DK, J1 (stored FR, GB, IT, products) PT Essl and Rabitsch (2002) Jürg Zettel / BioRisk 4(2): 851–854 (2010) Order-Family Status Regime Native range Species Collembola- Onychiuridae Onychiurus folsomi A detritivorous Australia (Schäffer, 1900) Collembola- Isotomidae Proisotoma filifera A detritivorous Central America (Denis, 1931) Desoria trispinata A detritivorous North America (MacGillivray, 1896) 854 Table 13.5.1. List and characteristics of the Collembola and Zygentoma species alien to Europe. Status: A Alien to Europe C cryptogenic species. Country codes abbreviations refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Last update 11/01/2010; A peer reviewed open access journal BioRisk 4(2): 855–1021 (2010) doi: 10.3897/biorisk.4.69 RESEARCH ARTICLE BioRisk www.pensoftonline.net/biorisk Introductory notes to factsheets Chapter 14 Alain Roques1, David Lees2 1 Institut National de la Recherche Agronomique (INRA), UR 0633, Station de Zoologie Forestière, 2163 Av. Pomme de Pin, 45075 Orléans, France 2 INRA UR633 Zoologie Forestière, 2163 Av. Pomme de pin, 45075 Orléans, France Corresponding author: Alain Roques (alain.roques@orleans.inra.fr) Academic editor: Alain Roques | Received 14 May 2010 | Accepted 26 May 2010 | Published 6 July 2010 Citation: Roques A, Lees D (Eds) (2010) Factsheets for 80 representative alien species. Chapter 14. In: Roques A et al. (Eds) Arthropod invasions in Europe. BioRisk 4(2): 855–1021. doi: 10.3897/biorisk.4.69 Among the 1590 terrestrial arthropod species alien to Europe identified in this book, 78 were selected to produce specific factsheets in order to provide more information on their biology, distribution and impact. We included two more species which are alien in Europe, the horse-chestnut leaf miner (Cameraria ohridella) and the African cotton leafworm (Spodoptera littoralis) because of their importance. These 80 species are perhaps not the most important alien invaders, but they are rather representatives of the main taxonomic groups of alien terrestrial arthropods. They were selected so as to represent different pathways of introduction and diverse impacts on ecosystems, economic activities and human and animal health. These species include two myriapods, one spider, one mite, 18 coleopterans, seven dipterans, 23 hemipterans, 10 hymenopterans, one termite, 14 lepidopterans, and three thrips. Each factsheet includes information on the following aspects: Description and biological cycle: A brief description of adults and immature stages is given, whenever possible illustrated by a photograph, to help the reader identify the species. Further information details the general characteristics of the biological cycle in the invaded area, especially the species’ potential to reproduce and the hosts it has colonized. Native habitat: The factsheet includes the habitat type where the species is found in its native range. In order to make habitat types comparable among taxa, we adopted the classification of the European Nature Information System (EUNIS) database Copyright A. Roques, D. Lees This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 856 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) (http://eunis.eea.europa.eu). The habitat type codes are detailed in Appendix II. When information was available, we included specific habitat requirements which may help understand the potential of the species to establish and spread in Europe. Habitat occupied in invaded range: The different habitats colonized by the alien species are described as for native habitats. Native range: The native distribution of the species is described. For some species, there is very precise information available, but for others, only brief details of a region or even continent can be given. Introduced range: The date of the first record in Europe and the location of this record is given, as well as details of the process of dispersion in the continent when available. A distribution map is supplied for all species. For most of them, presence/ absence data have been obtained only at country level, but for a few species, more detailed maps are given to show the distribution at regional scale. However, the missing occurrence of species from some countries does not always mean that these countries are not colonized, but may rather result from a lack of data for the country concerned. The map also indicates eradication records where relevant. Pathways: We included information on the routes of introduction to Europe, and the potential of the species to disperse within the continent once it has established. Impact and management: This section details the importance of the species’ impacts in the colonized habitats. Both ecological and economical impacts are detailed when known. Practical advice where known is given regarding mechanical, chemical and biological control methods. Selected references: Three of the most relevant references to the history of the species’ introduction and spread in Europe are given. Factsheets for 80 representative alien species. Chapter 14 857 List of the species 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. Lamyctes emarginatus Oxidus gracilis Mermessus trilobatus Varroa destructor Agrilus planipennis Anoplophora chinensis Anoplophora glabripennis Diabrotica virgifera Epitrix hirtipennis Leptinotarsa decemlineata Harmonia axyridis Gonipterus scutellatus Rhopalapion longirostre Trogoderma granarium Diocalandra frumenti Rhynchophorus ferrugineus Megaplatypus mutatus Gnathotrichus materiarius Phloeosinus rudis Xylosandrus crassiusculus Xylosandrus germanus Tribolium confusum Liriomyza huidobrensis Liriomyza trifolii Dasineura gleditchiae Obolodiplosis robiniae Aedes albopictus Ceratitis capitata Rhagoletis completa Adelges (Dreyfusia) nordmannianae Bemisia tabaci Trialeurodes vaporarium Aphis gossypi Cinara curvipes Macrosiphum euphorbiae Myzocallis walshii Myzus persicae Prociphilus fraxiniifolii Toxoptera citricidus Scaphoideus titanus Pulvinaria regalis 862 864 866 868 870 872 874 876 878 880 882 884 886 888 890 892 894 896 898 900 902 904 906 908 910 912 914 916 918 920 922 924 926 928 930 932 934 936 938 940 942 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. Leptoglossus occidentalis Aspidiotus nerii Diaspidiotus perniciosus Pseudaulacaspis pentagona Metcalfa pruinosa Nysius huttoni Stictocephala bisonia Halyomorpha halys Viteus vitifoliae Corythucha arcuata Corythucha ciliata Cales noacki Lysiphlebus testaceipes Dryocosmus kuriphilus Ophelimus maskelli Lasius neglectus Linepithema humile Nematus tibialis Megastigmus spermotrophus Sceliphron curvatum, S. caementarium and S. deforme Vespa velutina Reticulitermes flavipes Hyphantria cunea Paysandisia archon Diplopseustis perieresalis Phthorimaea operculella Cameraria ohridella Parectopa robiniella Phyllonorycter issikii Phyllonorycter platani Phyllonorycter robiniella Cacyreus marshalli Spodoptera littoralis Epichoristodes acerbella Grapholita molesta Argyresthia thuiella Frankliniella occidentalis Pseudodendrothrips mori Thrips palmi 944 946 948 950 952 954 956 958 960 962 964 966 968 970 972 974 976 978 980 982 984 986 988 990 992 994 996 998 1000 1002 1004 1006 1008 1010 1012 1014 1016 1018 1020 858 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) Adresses of factsheets authors Sylvie Augustin (#68) INRA UR633 Zoologie Forestiere, 2163 Av. Pomme de Pin, 45075 Orléans, France; (sylvie.augustin@orleans.inra.fr) Yuri Baranchikov (#5) Department of Forest Zoology, V.N.Sukachev Institute of Forest, Siberian Branch, Russian Academy of Science,50 Akademgorodok, Krasnoyarsk 660036, Russia (baranchikov_yuri@yahoo.com) Ejup Çota (#24, #36, #39) Plant Protection Department, Faculty of Agriculture and Environment, Agriculture University of Tirana, Albania; (ejupcota@gmail.com) Jurate De Prins (#66, #73) Entomology Section, Royal Museum for Central Africa, Leuvensesteenweg 13, B-3080 Tervuren, Belgium; (jurate.de.prins@africamuseum.be) Willy De Prins (#66, #73) Entomology Section, Royal Museum for Central Africa, Leuvensesteenweg 13, B-3080 Tervuren, Belgium; (willy.deprins@gmail.com) Massimo Faccoli (#17, #18, # 20, #21) Universita di Padova, Department of Environmental Agronomy and Crop Sciences, Agripolis, Viale dell’Università 16, 35020 Legnaro (PD), Italy; (massimo.faccoli@unipd.it) Milka M. Glavendekić (#25, #46, #55, #59) University of Belgrade, Faculty of Forestry, Kneza Viseslava 1, 11030 Belgrade, Serbia; (milka.glavendekic@nadlanu.com) Stanislav Gomboc (#75) Siskovo naselje 19, SI-4000 Kranj, Slovenia; (stane.gomboc@gov.si) Marc Kenis (#29, #41, #44, #50, #51, #63) CABI Europe-Switzerland, 1, Rue des Grillons, CH- 2800, Delémont, Switzerland; (m.kenis@ cabi.org) Zoltán Korsós (#2) Zoological Department, Hungarian Natural History Museum H-1088 Budapest, Hungary (korsos@zoo.zoo.nhmus.hu) Factsheets for 80 representative alien species. Chapter 14 859 Ferenc Lakatos (#13, #14, #22, #43, #64, #77) University of West-Hungary, Institute of Sylviculture and Forest Protection, Bajcsy-Zs. u. 4., H-9400 Sopron, Hungary; (flakatos@emk.nyme.hu) Zdeněk Laštůvka (#76) Department of Zoology, Fisheries, Hydrobiology and Apidology, Faculty of Agronomy, Mendel University in Brno, Zemědělská 1, CZ-613 00 Brno, Czech Republic; (last@mendelu.cz) Yves Le Conte (#4) Institut National de la Recherche Agronomique (INRA), UMR0406 AE Abeilles et Environnement, Domaine Saint-Paul - Site Agroparc 84914 Avignon, France; (yves.leconte@avignon.inra.fr) David Lees (Introductory notes, #65, #69, #70, #71, #72) INRA UR633 Zoologie Forestiere, 2163 Av. Pomme de Pin, 45075 Orléans, France; (david.lees@ orleans.inra.fr) Carlos Lopez-Vaamonde (#10, #65, #74) INRA UR633 Zoologie Forestiere, 2163 Av. Pomme de Pin, 45075 Orléans, France; (carlos.lopezvaamonde@orleans.inra.fr) Ljubodrag Mihajlović (#25) University of Belgrade, Faculty of Forestry, Kneza Viseslava 1, 11030 Belgrade, Serbia; (mljuba@ EUnet.rs) Leen Moraal (#19) Alterra, Wageningen UR, Centre Ecosystems, PO Box 47, NL-6700 AA Wageningen, The Netherlands; (Leen.Moraal@wur.nl) Franck Muller (#62) Museum National d’Histoire Naturelle Entomologie CP50, 45 rue Buffon, 75005 Paris, France Maria Navajas (#4) Institut National de la Recherche Agronomique, UMR CBGP (INRA/IRD/Cirad/Montpellier SupAgro), Campus International de Baillarguet, CS 30016, F-34988, France; (navajas@supagro.inra.fr) Olivera Petrović-Obradović (#34, #35, #37, #38) University of Belgrade, Faculty of Agriculture, Nemanjina 6, SER-11000, Belgrade, Serbia; (petrovic@agrif.bg.ac.rs) 860 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) Wolfgang Rabitsch (#8, #15, #16, #40, #42, #47, #48, #49, #51, #52, #57, #58) Environment Agency Austria, Dept. Biodiversity & Nature Conservation, Spittelauer Lände 5, 1090 Vienna, Austria; (wolfgang.rabitsch@umweltbundesamt.at) Jean-Yves Rasplus (#53, #54, #56, #61) Institut National de la Recherche Argonomique, UMR Centre de Biologie et de Gestion des Populations, CBGP, (INRA/IRD/CIRAD/Montpellier SupAgro), Campus international de Baillarguet, CS 30016, 34988 Montferrier-sur Lez, France; (rasplus@supagro.inra.fr) Hans Peter Ravn (#30) Forest & Landscape Denmark, University of Copenhagen, Hoersholm Kongevej 11, DK-2970 Hoersholm, Denmark; (hpr@life.ku.dk) Philippe Reynaud (#79, #80) Laboratoire national de la protection des végétaux, Station d’Angers, 7 rue Jean Dixméras, 49044 Angers Cedex 01, France; (philippe.reynaud@agriculture.gouv.fr) Quentin Rome (#62) Museum National d’Histoire Naturelle Entomologie CP50, 45 rue Buffon, 75005 Paris, France (vespa.velutina@gmail.com) Alain Roques (Introductory notes, #23, #27, #28, #31, # 32, #33, #55, #60, #78) Institut National de la Recherche Agronomique (INRA), UR 0633, Station de Zoologie Forestiere, 2163 Av. Pomme de Pin, 45075 Orléans, France; (alain.roques@orleans.inra.fr) David B. Roy (#11) Centre for Ecology & Hydrology, Crowmarsh Gifford, Oxfordshire, OX10 8BB, United Kindgom (dbr@ceh.ac.uk) Helen Roy (#11) NERC Centre for Ecology & Hydrology, Biological Records Centre, Crowmarsh Gifford, Oxfordshire, OX10 8BB, United Kindgom (hele@ceh.ac.uk) Daniel Sauvard (#6, #7, #12) INRA UR633 Zoologie Forestiere, 2163 Av. Pomme de Pin, 45075 Orléans, France; (daniel.sauvard@orleans.inra.fr) Martin H Schmidt-Entling (#3) University of Bern, Institute of Ecology and Evolution, Community Ecology, CH-3012 Switzerland (martin.schmidt@zos.unibe.ch) Marcela Skuhravá (#26) Bítovská 1227/9, 140 00 Praha 4, Czech Republic; (skuhrava@quick.cz) Factsheets for 80 representative alien species. Chapter 14 861 Pavel Stoev (#2) National Museum of Natural History, Tsar Osvoboditel Blvd. 1, 1000 Sofi a, Bulgaria (pavel.e.stoev@gmail.com) Jean-Claude Streito (#52) Laboratoire national de la protection des végétaux, CBGP Campus international de Baillarguet, CS 30016, FR-34988 Montferrier-sur-Lez cedex, France; (streito@supagro.inra.fr) Rumen Tomov (#9) University of Forestry, 10 Kliment Ohridski blvd., 1756 Sofia, Bulgaria (rtomov@yahoo.com) Georgyi Trenchev (#67) University of Forestry, 10 Kliment Ohridski blvd., 1756 Sofia, Bulgaria (k_trencheva@yahoo. com) Katia Trencheva (#45, #67) University of Forestry, 10 Kliment Ohridski blvd., 1756 Sofia, Bulgaria; (k_trencheva@yahoo. com) Katalin Tuba (#13, #14, #22, #43) University of West-Hungary, Institute of Silviculture and Forest Protection, Sopron, Bajcsy-Zs. u. 4. 9400, Hungary (tubak@emk.nyme.hu) Claire Villemant (#62) Museum National d’Histoire Naturelle, UMR Origine, Structure et Evolution de la Biodiversite, OSEB, (MNHN/CNRS) CP50, 45 rue Buff on, 75005 Paris, France; (villeman@mnhn.fr) Marzio Zapparoli (#1) Universita degli Studi della Tuscia, Dipartimento di Protezione delle Piante, via S. Camillo de Lellis s.n.c., I-01100 Viterbo, Italy (zapparol@unitus.it) 862 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.1 – Lamyctes emarginatus (Newport, 1844) (Chilopoda, Henicopidae) Marzio Zapparoli Description and biological cycle: Body length 6.0–10.5 mm, shape slender, feebly fusiform, first tergite distinctly narrower than head and than tergite 3. Chestnut brown to dark brown, with leg tips and antennae yellow (Photo). One ocellus only at each side of the head. Antennae one-third to two-fifths of body length, with 25 segments. 15 pairs of legs, slender and without spines. Tibia of 1st-12th segment with a sharp projection on the anterior edge of its distal extremity. Reliable identification possible only after light microscope examination. L. emarginatus is a soil dwelling predator of small invertebrates, apparently hygrophilous and generally solitary. It is a pioneer of disturbed habitats (e.g. mine sites). This centipede develops through five anamorphic larval and five post-larval stadia. The species reproduces by thelytokous parthenogenesis* (producing only females) in most of its range. Bisexual populations are known in the Azores and Canary Islands, as well as in New Zealand, Tasmania and Hawaii. Native habitat (EUNIS code): Often collected at banks of creeks and rivers, and under stones where there is moisture. Some records from grasslands (E). In its native range Lamyctes emarginatus also commonly colonizes disturbed urban and suburban habitats. Habitat occupied in invaded range (EUNIS code): In Europe recorded in a wide range of habitats, from open or semi-open habitats (E); cultivated lands (I, X6, X7); city parks (X23); gardens (I1, I2); more or less intensively built-up areas of villages and cities (J1, J2, J4); waste dumps (J6); plant nurseries (I1); mine sites (J3); artificial banks of streams and lakes more or less temporary flooded (J5); also in natural or semi-natural habitats such as woodlands (G1, G3); heathlands (F4); riversides (F9) ; bogs (D) and coastal environments (B1) (British Isles, Faroe Isl., Northern Italy). Almost the same range of habitats is known for North America, New Zealand and Tasmania. Native range: Australasian species (western and southern Australia). Credit: Massimo Vollaro Factsheets for 80 representative alien species. Chapter 14 863 Introduced range: Present in almost all European countries and Atlantic islands (Mapnot recorded elsewhere probably due to lack of research), Southwestern Asia (Georgia, Iran, Turkey), Africa (Morocco, Ethiopia, Kenya, Tanzania, Somalia), North America (Canada, USA), Central America (Cuba, Guadeloupe), South America (Galapagos Islands, Brazil), Hawaii. Probably introduced also in Tasmania (proposed common name: Domestic Woodrunner), New Caledonia, New Zealand, Chatham Islands, Fiji and Kermadec Islands. Described as a new species by G. Newport from New Zealand in 1844, it has been recorded in Europe for the first time in 1868 in Denmark (Øerne) under the name of Lamyctes fulvicornis Meinert, 1868. The species may have been introduced into Europe even earlier, e.g. immediately after the British colonization of the Australian continent (18th century). Pathways: Mode of introduction and spread of this species is unknown, but probably occurred/occurs by passive transfer in rootballs of transplanted plants in which soil this small species temporarily settles. Parthenogenesis can then help the successful establishment of viable populations. Local-scale dispersal is probably achieved actively. Impact and management: Unknown, but effects on trophic chains are possible.This species is presently economically unimportant, thus monitoring, chemical or biological control are considered unnecessary. Selected references Andersson G (2006) Habitat preferences and seasonal distribution of developmental stadia in Lamyctes emarginatus (Newport, 1844) (L. fulvicornis Meinert, 1868) and comparisons with some Lithobius species (Chilopoda, Lithobiomorpha). Norwegian Journal of Entomology 53: 311–320. Bocher J, Enghoff H (1984) A centipede in Groenland: Lamyctes fulvicornis Meinert, 1868 (Chilopoda, Lithobiomorpha, Henicopidae). Entomologiske Meddelelser 52: 49–50. Zerm M (1997) Distribution and phenology of Lamyctes fulvicornis and other lithobiomorph centipedes in the floodplain of the Lower Oder Valley, Germany (Chilopoda, Henicopidae: Lithobiidae). Entomologica Scandinavica suppl. 51: 125–132. 864 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.2 – Oxidus gracilis (C.L. Koch, 1847) (Diplopoda, Polydesmida, Paradoxosomatidae) Pavel Stoev and Zoltán Korsós Description and biological cycle: Typical “flat-backed” millipede (Order Polydesmida), body length 16–21 mm (males), 17–23 mm (females). Dorsal surface smooth, chestnut to dark reddish or blackish brown, with lighter, pale yellowish paranota* (“lateral wings”) (Photo). Ventral surface and legs, head and antennae also dark brown. Immature stages light amber to cream. Further taxonomic traits and good illustrations of gonopods can be found in Blower (1985). Native habitat (EUNIS code): From grasslands (E1, E2, E3, E5, E7) to all kinds of woody habitats (G1, G2, G3, G4), miscellaneous inland habitats (e.g. H5), occasionally entering subterranean localities (H1), common in agricultural areas and other places under different degrees of disturbance (I1-I2, J2, J3, J4, J6, X7, X10, X11, X13, X14, X15, X16, X22, X23). Habitat occupied in invaded range: (EUNIS code): In Europe the'greenhouse millipede' has been recorded most often in hothouses (J100), city parks (X23) and gardens (I1, I2). The only alien millipede to have invaded some natural ecosystems and at least partially acclimatized in Europe. Earliest records in Europe are from the 19th century from Hungary, Austria, Germany, and the Netherlands from greenhouses and cities. Koch’s (1847) type locality ‘Puloloz’ may refer to a locality in former Czechoslovakia or in ‘Ostindien’. In North Europe, it was found for the first time in Finland in 1900. In the British Isles, first recorded from Edinburgh (1898) and Kew Gardens. Native range: East or Southeast Asia. Two other congeners (O. avia and O. riukiaria) are recently confirmed as validly occurring in the Ryukyu Archipelago, both confined to undisturbed, natural forest habitats. This suggests that the genus might have had a centre of origin in that region. Credit: Zoltán Korsós Factsheets for 80 representative alien species. Chapter 14 865 Introduced range: Recorded from 33 European countries, including several Mediterranean islands (Map). Also introduced to numerous temperate and tropical countries: USA, Canada, Puerto Rico, Asian part of Russia, Australia, New Zealand, South Africa, Brazil, archipelagos and isolated islands in Indian and Pacific Oceans: Hawaii, New Caledonia, Madagascar, Sandwich Islands, Seychelles, etc. Pathways: Must have arrived in Europe with goods from East Asia although when is unknown. Oxidus gracilis is spreading within Europe mainly with expanding trade and greenhouse tropical plant cultivation. Some populations in Central and South Europe have established themselves naturally, most often close to suburban/urban areas, but also in woodlands in nature reserves and in caves. Impact and management: A plant-damaging millipede regarded as a pest in several European countries. In some places, its density may exceed 2500 ind./m2. It is known for attacking vegetable and fruit crops such as sugar beet, potatoes, strawberries, cucumbers, orchard fruits, peanut seedlings, roots of wheat, and flowers in outdoor cultivated areas. No data on impact of O. gracilis on native species and its interactions with local invertebrate communities. So far no evidence for wild populations taking over new habitats. The species dies when subjected for two hours to a temperature of -4°C. Thus in Northern Europe, it can survive only in hothouse conditions. Various different chemicals such as Methodyl, Carbaryl and Propuxur proved to be effective for dealing with O. gracilis. Another effective method for decreasing the numbers of the species in buildings is the removal of all excess damp organic matter and debris from gardens and associated areas. Selected references Blower JG (1985) Millipedes. Keys and notes for the identification of the species. Synopses of the British Fauna 35: 1–242. Chornyi NG, Golovatch SI (1993) Dvuparnonogie mnogonozhky ravninnych territorij Ukrainy. [Millipedes (Diplopoda) of the plain areas of the Ukraine]. Kiev: Kiev University. 56 pp. Vicente M, Enghoff H (1999) The millipedes of the Canary Islands. Vieraea 27: 183–204. 866 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.3 – Mermessus trilobatus (Emerton, 1882) (Araneae, Linyphiidae) Martin H Schmidt-Entling Description and biological cycle: Small, 1.5–2.1 mm long sheetweb spider with orangebrown prosoma* and legs, gray to black opisthosoma*and no markings (Photo). In contrast to most native European sheetweb spiders, the female epigyne* is frequently covered with a dark reddish-brown, hemispherical mating plug. Egg sacs light pink with a smooth surface and 3 mm diameter. Individual females produce up to 11 egg sacs. Adults are found year-round, and several generations per year are likely. The species builds small horizontal webs low in the vegetation or at the ground surface. Long distance dispersal is by ballooning (also shown by adults), and possibly by passive transport on road, rail or by aircraft. Native habitat (EUNIS code): Inhabits various types of E - Grassland and tall forb habitats; I1 - Arable land and market gardens; D – wetlands; B - coastal habitats; G - Woodland and forest habitats and other wooded land. Habitat occupied in invaded range (EUNIS code): The species can reach high densities (exceeding 10 adults per m2) in E- Grassland and tall forb habitat, and in ruderal habitats in both rural and urban areas. It is frequently among the ten most abundant spider species in these habitats in Northern Switzerland. It also occurs on I1 - Arable land and market gardens and in G - Woodland and forest habitats and other wooded land. Native range: Widely distributed in North America from the Gulf coast (Florida, Mexico) to boreal climate (Northwest Territories, Newfoundland) and from the east to the west coast including the Great Plains. Credit: Martin H Schmidt Entling Factsheets for 80 representative alien species. Chapter 14 867 Introduced range: First recorded in 1981 near Karlsruhe, southwestern Germany, M. trilobatus spread in all directions with recent records as far as the Netherlands and Italy (near Venice) (Map). Pathways: Presumably transported unintentionally by aircraft, with no evidence for multiple introductions. Impact and management: Negative impacts on the native fauna are unknown, but are only starting to be explored. Given the wide distribution and high abundance, prey or other predators may be affected through predation or intraguild interference. M. trilobatus was the dominant sheetweb spider in nine protected E3.5 - fen meadows in Switzerland, indicating a potential threat to biodiversity. The species can be effectively recorded with vacuum sampling. It is underrepresented in pitfall traps relative to other spiders. Once established, successful control of M. trilobatus appears very unlikely. Specialist natural enemies are not known. Selected references Dumpert K, Platen R (1985) Zur Biologie eines Buchenwaldbodens, 4. Die Spinnenfauna. Carolinea 42: 75–106. Eichenberger B, Siegenthaler E, Schmidt-Entling MH (2009) Body size determines the outcome of competition for webs among alien and native sheetweb spiders (Araneae: Linyphiidae). Ecological Entomology 34: 363–368. Schmidt MH, Rocker S, Hanafi J, Gigon A (2008) Rotational fallows as overwintering habitat for grassland arthropods: the case of spiders in fen meadows. Biodiversity and Conservation 17: 3003–3012. van Helsdingen PJ, IJland S (2007) Mermessus species in the Netherlands (Araneae, Linyphiidae). Nieuwsbrief Spined 23: 27–29. 868 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.4 – Varroa destructor Anderson & Trueman, 2000 - Bee Varroa (Acari, Varroidae) Maria Navajas and Yves Le Conte Description and biological cycle: Varroa destructor is today’s most important honey bee parasite. As an ectoparasite, the mite causes serious damage to honey bee colonies almost worldwide. Varroa feeds on the hemolymph of adult, larval and pupal bees. Adult females are reddishbrown, 1.00–1.77 mm long and 1.50–1.99 mm wide (Photo- female Varroa on abdomen of Apis mellifera). Adult males are yellowish with lightly tanned legs and spherical body shape measuring 0.75–0.98 mm long and 0.70–0.88 wide. The male chelicerae are modified for transferring sperm. Varroa destructor reproduces by pseudoarrenotoky*. The female lays eggs in bee brood cells. Developing mites feed on developing honey bee larvae and pupae. Males and females copulate in capped brood cells. The male dies, but fertilized females emerge from the cell along with their bee host and seek a nurse bee on which it feeds for a few days and then repeat the cycle. In temperate climates, V. destructor populations complete an average of 10 generations per year. Native habitat (EUNIS code): J1- Hives. Habitat occupied in invaded range (EUNIS code): J1- Hives. Native range: South East Asia, where it was originally confined on its original host, the Asian honey bee, Apis cerana. Importation of commercial A. mellifera colonies into areas with A. cerana brought this previously allopatric bee species into contact and allowed V. destructor to switch to the new host. While the populations of the parasite reach only a small size within colonies of A. cerana and do not damage the colony, infested A. mellifera colonies die. Introduced range: practically worldwide except Australia, the state of Hawaii and some parts of Africa remain free of this pest (see lower Map for known spread routes of Varroa). First reported in the Eastern Europe in the 70s, it spread rapidly all over the continent (see upperMap). Two different genotypes (characterized by mitochondrial DNA sequences) have spread as independent clonal populations: the Korean and the Japanese haplotypes, the later having been found, besides Asia, in The Americas only. International travel and commerce has facilitated the varroa dispersal. Credit: Alain Migeon Factsheets for 80 representative alien species. Chapter 14 869 Pathways: Once established in a new region, the mite spread using drifting, robbing, and swarming behaviour of the host. Human mediated varroa dispersion also occurs by apicultural practices. The importation of honey bees from infested areas, and the movement of infested colonies for pollination or hives transhumance* led to the rapid spread of this mite. Impact and management: Although several chemicals are applied against honey bee larvae and adults, pesticide-resistant varroa populations occur. In addition, there is much concern about chemical residues in hive products. Alternative varroa control methods are attempted, including the use of organic acids, as formic acid, but they are temperature dependent and can be dangerous to humans. Another efficient varroa control is the use of plant volatile essential oil extracts. These different methods used in combination with an integrated pest management (IPM) plan, including bee colony management techniques (e.g. removal of the infested brood) may be helpful. The recent detection of varroa-resistant honeybee stocks is a promising avenue for honeybee breeding. Selected references De Rycke PH, Joubert JJ, Hossein Hosseinian S, Jacobs FJ (2002) “The possible role of Varroa destructor in the spreading of American foulbrood among apiaries.” Experimental and Applied Acarology 27: 313–318. Le Conte Y, de Vaublanc G, Crauser D, Jeanne F, Rousselle JC, Bécard JM (2007) Honey bee colonies that have survived Varroa destructor. Apidologie 38: 566–572. Martin C, Provost E, Roux M, Bruchoux C, Clément JL, Le Conte Y (2002) Potential mechanism for detection by Apis mellifera of the parasitic mite Varroa destructor inside sealed brood cells by Apis mellifera. Physiological Entomology, 27: 175–188. 870 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.5 – Agrilus planipennis Fairmaire, 1888 - Emerald Ash Borer (Coleoptera, Buprestidae) Yuri Baranchikov Description and biological cycle: Adults 7–15 mm long and 3–3.4 mm wide. Body narrow, elongate and metallic coppery-green (Photo left). Eyes large, black- or copper-coloured. Emerald Ash Borer has four larval instars. Mature larvae 26–32 mm long , creamy white with head brown (Photo right). Pre-pupal body characteristically J-shaped. Pupa exarate*, initially beige, later darkening. Adults emerge from ash trees through distinct D-shaped exit holes chewed through the bark. Adults found in June-early August, flying near the host trees and feeding on leaves. After several weeks of maturation, feeding and mating, female deposits up to 270 eggs, singly or in small clusters in bark crevices. Larvae form distinct S-shaped galleries, widening as they grow. Larvae mostly found under the bark during summer and some may be present all year around. Full development takes 1–2 years. All European and most American ash species highly susceptible to infestation. Up to three years’s attack can kill a middle-sized ash tree. All species of Fraxinus can be used as hosts. Reportedly, Agrilus planipennis colonize Ulmus, Juglans and Pterocarya in Asia; in the USA it can lay eggs on Ulmus americana, Celtis occidentalis, Carya ovata and Syringa reticulata, but larvae die in early instars. Native habitat (EUNIS code): G1 - Broadleaved deciduous woodland; G4 - Mixed deciduous and coniferous woodland. Habitat occupied in invaded range (EUNIS code): G1 - Broadleaved deciduous woodland; G5 - Lines of trees, small anthropogenic woodlands, recently felled woodland, early-stage woodland and coppice; I2 - Cultivated areas of gardens and parks; X24 Domestic gardens of city and town centres; X25 Domestic gardens of villages and urban peripheries. Native range: North Eastern China, Korea Japan, Taiwan, Eastern Mongolia and the Southern part of the Russian Far East. Introduced range: First discovered in Michigan, USA, in June 2002 and in Ontario, Canada, in August 2002 (Cappaert et al., 2005). Agrilus planipennis then established in 11 states in North-Eastern USA and adjacent provinces of Canada, expanding very fast. In the early 2000s, introduced to European Russia (see upper Map) where ash trees were killed within Credit: David Cappaert, Michigan State University, Bugwood.com Factsheets for 80 representative alien species. Chapter 14 871 a 100 km radius of Moscow; since has spread westwards. In approximately 10 years the beetle spread 95 km to the West, 90 km to the South, 30 km to the East and 20 km to the North. (see lower Map- red points detailing the outer spots of infestation in Moscow area in 2009). Pathways: transportation with wood for industry and firewood. Adults are active flyers. Impact and management : In < 10 years, this buprestid killed an estimated over 20 million forest and ornamental trees in North America. In Europe, the ecological impact manifests in fast dieback of all ashes in cities and forests, irrespective of their previous condition. Only Asian species of ashes are relatively resistant to the pest, so the beetle poses one of the most important economic threats to trees in Western Europe. In Asia, controlled by a few hymenopteran parasitoid species: eggs are attacked by Oobius agrili Zhang and Huang (Encyrtidae); larvae are infested by the gregarious endo-parasitoid Tetrastichus planipennisi Yang (Eulophidae) and the gregarious idiobiont ectoparasitoid Spathius agrili Yang (Braconidae). Two other braconids (Spathius depressithorax Belokobylskiy and S. generosus Wilkinson) have been found on larvae in Far Eastern Russia. Woodpeckers are active predators of larvae and pupae. Introduction of some parasitoid species is successfully implemented in the US. Traps with attractants are actively used to monitor spread. A dozen systemic insecticide formulations are used to protect individual ash trees in settlements and historical places. Selected references Anulewicz AC, McCullough DG, Cappaert DL, Poland TM (2008) Host range of the emerald ash borer (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae) in North America: results of multiple-choice field experiment. Environmental Entomology 37: 230–241. Baranchikov Y, Mozolevskaya E, Yurchenko G, Kenis M (2008) Occurrence of the emerald ash borer, Agrilus planipennis in Russia and its potential impact on European forestry. Bulletin OEPP/EPPO Bulletin 38: 233–238. Cappaert D, McCullough DG, Poland TM, Siegert NW (2005) Emerald ash borer in North America; a research and regulatory challenge. American Entomologist 51: 152–165. 872 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.6 – Anoplophora chinensis (Förster, 1848) (=A. malasiaca (Thompson, 1865) - Citrus longhorn beetle (Coleoptera, Cerambycidae) Daniel Sauvard Description and biological cycle: Large, 21–37 mm long, stout beetle with shiny black elytra marked with 10–12 white round spots (Photo left). Antennae long, basally marked with white or light blue bands. The larva is a legless grub creamy white in colour, up to 50 mm long when fully grown (Photo right). Polyphagous insect attacking over 100 species of broadleaved trees and shrubs (Acer, Betula, Carpinus, Citrus, Corylus, Rosa and deciduous shrubs). Adults can fly up to 1.5 km from their emergence place. Human-mediated longdistance dispersal is possible via infested wood movement or adults hitch-hiking on vehicles. Females lay eggs throughout their lifespan from spring to late summer. Fecundity varies from tens to more than a hundred eggs per female. Full development is achieved in one or two years depending on climate and egg-laying date. Larvae and pupae overwinter inside their tunnels in wood. Native habitat (EUNIS code): G1- Broadleaved deciduous woodland; G5- Lines of trees, small anthropogenic woodlands. Habitat occupied in invaded range (EUNIS code): G5- Lines of trees, small anthropogenic woodlands. Prefers subtropical to temperate climate; can survive in a large part of Europe. Native range: East Asia (China, Taiwan, Korea, Japan, Myanmar, Vietnam). Introduced range: Italy and a spot in the Netherlands (Map). First recorded in Lombardia, near Milano, Italy in 2000 but probably arrived several years earlier. Increasing frequency of interceptions during the last ten years in Europe. Eradicated in France and Great Britain. Italian populations from Lombardia recently spread in the peninsula, including the Roma area. Credit: Franck Hérard Factsheets for 80 representative alien species. Chapter 14 873 Pathways: Introduced with infested woody materials, especially bonsai plants. Impact and management: Citrus longhorn beetle may disturb broadleaved forest ecosystems by selective tree killing or via direct/indirect competition with native xylophagous insects, including protected ones. Social impact occurs because in urban areas (streets, private and public gardens) the species kills trees and Rosa shrubs. This is one of the most destructive cerambycid pests of fruit orchards in its native range, especially on Citrus trees. Larval tunnels also depreciate harvested wood. This longhorn beetle is difficult to trap; surveys are generally based on visual detection of damage. Mechanical control involves destruction of infested trees by chipping or burning; trees can also be protected with fine wire meshes to prevent oviposition. Chemical control is of limited effect because the insects are deep within the tree, but systemic insecticides might be used. Biological control using natural enemies (parasitoid insects, entomopathogenic nematodes, fungi or bacteria) is under investigation but not yet being used. Selected references Colombo M, Limonta L (2001) Anoplophora malasiaca Thomson (Coleoptera Cerambycidae Lamiinae Lamiini) in Europe. Bollettino di Zoologia agraria e di Bachicoltura II Ser 33: 65–68. EPPO (2010) Anoplophora chinensis found again in the Netherlands. EPPO Reporting Services 2: 2010/025. Hérard F, Ciampitti M, Maspero M, Krehan H, Benker U, Boegel C, Schrage R, BouhotDelduc L, Bialooki P (2006) Anoplophora species in Europe: infestations and management processes. Bulletin OEPP/EPPO Bulletin 36: 470–474. 874 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.7 – Anoplophora glabripennis (Motschulsky, 1853) - Asian longhorned beetle (Coleoptera, Cerambycidae) Daniel Sauvard Description and biological cycle: Large, stout beetle, 20–35 mm long with jet-black body and white spotted elytra. Antennae longer than body, black with blue rings at segment base (Photo up). The larva is a legless grub up to 50 mm long when fully grown. It is creamy white in colour, with a chitinized brown mark on the prothorax (Photo down). Xylophagous* species, feeding on a wide range of deciduous trees, mostly species with soft wood such as Acer or Populus where the larvae live inside the wood, in tree boles or large branches. Adults also eat bark on small branches. Adults fly up to 1.5 km from the emergence place. Possible human-mediated longdistance dispersal by infested wood movement or adults hitchhiking on vehicles. Eggs are laid throughout female life from spring to late summer; fecundity is variable from tens to more than a 100 eggs per female. Full development is achieved in one or two years depending on climate and oviposition date. Larvae and pupae overwinter inside wood tunnels. Native habitat (EUNIS code): G- Broadleaved deciduous woodland; G5- Lines of trees, small anthropogenic woodlands. Habitat occupied in invaded range (EUNIS code): G5- Lines of trees, small anthropogenic woodlands. Prefers subtropical to temperate climate; can survive in a large part of Europe up to S Sweden. Native range: East Asia (China, Taiwan, Korea, Japan) Credit: F. Hérard (above), Alain Roques (below) Factsheets for 80 representative alien species. Chapter 14 875 Introduced range: USA, Canada, Austria, France, Germany, Italy (Map). Increasing frequency of interceptions and introductions in Europe during the last ten years. Where the species has been introduced, always in urban areas, eradication attempts have been undertaken. Pathways: Introduced repeatedly with infested woody materials, especially wood packaging, pallets and waste materials. Impact and management: May disturb European broadleaved ecosystems by selective tree killing or direct/indirect competition with native xylophagous insects, including protected ones. Social impact occurs because primary introduction is always in urban areas where the beetle weakens or kills trees in streets, private and public gardens. One of the most destructive cerambycid forest pests in its native range, inducing heavy damage in broadleaved stands, including poplar plantations. Larval tunnels also depreciate harvested wood. Difficult to trap; surveys generally based on visual detection of damage. Mechanical control involves destruction of infested trees by chipping or burning; trees can also be protected with fine wire mesh to prevent oviposition. Chemical control is of limited effect because the insects live deep within the tree; systemic insecticides may be tried. Biological control using natural enemies (parasitoid insects, entomopathogenic nematodes, fungi or bacteria) is under investigation but not yet used. Selected references Dauber D, Mitter H (2001) Das erstmalige Auftreten von Anoplophora glabripennis Motschulsky 1853 auf dem europaïschen Festland (Coleoptera: Cerambycidae: Lamiinae). Beiträge zur Naturkunde Oberösterreichs 10: 503–508. Haack RA, Hérard F, Sun JH, Turgeon JJ (2010) Managing invasive populations of Asian longhorned beetle and Citrus longhorn beetle: A worldwide perspective. Annual Review of Entomology 55: 521–546. Hérard F, Maspero M, Ramualde N, Jucker C, Colombo M, Ciampitti M, Cavagna B (2009) Anoplophora glabripennis - Eradication programme in Italy (April 2009). http://www.eppo. org/QUARANTINE/anoplophora_glabripennis/ANOLGL_IT.htm. 876 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.8 – Diabrotica virgifera virgifera LeConte, 1868 - Western corn rootworm (Coleoptera, Chrysomelidae) Wolfgang Rabitsch Description and biological cycle: Small beetle, 5–6 mm long, with pale greenish-yellow body (Photo). Larvae are wrinkled, yellowish-white, with a brown head capsule. The Western corn rootworm is a major crop pest on maize (Zea mays), repeatedly introduced to Europe from North America in the early 1990s, which spreads rapidly. Adult flight dispersal is 20–100 km per year; intercontinental dispersal occurs via the transfer of goods. Up to 1000 eggs are produced per female during lifespan, laid preferentially in the soil at the base of maize plants. Larvae develop in and on roots of the food plant and adults move upwards and feed on the plant. The species develops as one generation per year. Eggs overwinter in diapause. Native habitat (EUNIS code): E- Grassland. Habitat occupied in invaded range (EUNIS code): I: Regularly or recently cultivated agricultural, horticultural and domestic habitats; I1- Arable land and market gardens. Temperature not only influences larval development, but also triggers flight activity which governs the rate of dispersal. Increased habitat diversity slows the rate of spread. Native range: Probably in the tropics and subtropics of Mexico and Central America. Introduced range: North America, Europe (Serbia 1992; Croatia, Hungary 1995; Romania 1996; Bosnia and Herzegovina 1997; Bulgaria, Italy, Montenegro 1998; Slovakia, Switzerland 2000; Ukraine 2001; Austria, Czech Republic, France 2002; Belgium, Netherlands, Slovenia, UK 2003; Poland 2005; Germany 2007; Map). Genetic data provides evidence for Credit: Margarita Auer Factsheets for 80 representative alien species. Chapter 14 877 repeated introductions from America (Ciosi et al 2008). Spread in Europe continues and the species is expected to colonize all maize producing countries in Eurasia. Pathways: Repeatedly transported via vehicles (airplanes, railways, ships).redit : Margarita Auer Impact and management: Ecosystem impacts including side-effects on non-target species as a consequence of insecticide treatment or biological control are possible, but not demonstrated. This species is regarded as one of the most serious pest species of corn in the USA and its damage to crops and chemical control amounts to 1 billion US$ per year. Current economic damage in Europe is restricted to some countries, but there is clearly a time lag of several years between the first record and economic damage. Predictive models forecast an economic impact of about 500 million €/year in Europe. Crop rotation is the most feasible preventative measure, although crop rotation resistant rootworm variants are known in the USA. Monitoring the spread of adults via pheromone-traps is used as a predictor of damage and further treatment is applied in the following season. Chemical control involves several toxicants applied as granular soil insecticides against the larvae and by aerial spraying against adults (spraying is not permitted in most European countries). Biological control using natural enemies (tachinid flies, nematodes, entomopathogenic fungi) is currently under investigation. Selected references Ciosi M, Miller NJ, Kim KS, Giordano R, Estoup A, Guillemaud T (2008) Invasion of Europe by the western corn rootworm, Diabrotica virgifera virgifera: multiple transatlantic introductions with various reductions of genetic diversity. Molecular Ecology 17: 3614–3627. Kuhlmann U, van der Burgt W (1998) Possibilities for biological control of the western corn rootworm, Diabrotica virgifera virgifera LeConte, in Central Europe. BioControl 19, 59–68. Vidal S, Kuhlmann U, Edwards CR (2004) Western Corn Rootworm: Ecology and Management. CABI Bioscience, Delémont, 320 pp. 878 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.9 – Epitrix hirtipennis (Melsheimer, 1847) - Tobacco flea beetle (Coleoptera, Chrysomelidae) Rumen Tomov Description and biological cycle: Adult about 1.5–2 mm long, brown (Photo left). Elytra the same colour as pronotum except on their central areas which are almost black; with rows of fine, distinct punctures, and scattered thin, relatively short brown-yellow setae. Eggs tiny, elongate and white. Fully developed larvae about 4–5 mm long with slender, cylindrical, whitish body and brown head. Pupae white turning darker with anterior end curved downwards. The tobacco leaf beetle is oligophagous on plants of the family Solanaceae: tobacco (Nicotiana tabacum), potato (Solanum tuberosum), tomato (Solanum lycopersicum), aubergine (Solanum melongena), pepper (Capsicum annuum), ground cherry (Physalis pruinosa), jimson weed (Datura stramonium) and horsenettle (Solanum carolinense). Overwinters as adult among debris around fields of host plants or in tobacco seedbeds. Activity resumes in spring feeding on weedy hosts until crop hosts are available. Females lay eggs in the soil near the host plants, which hatch after a week. Larval development lasts 3–4 weeks. Pupal period is 7–10 days, before new generation of beetles emerges. The species develops 3–4 generations per year. Damage is caused by both larvae and adults. Adult beetles feed mainly on leaves in which they produce small round holes (Photo left). The larvae burrow into the soil, living in the roots or tunneling into the stalk (Photo right). Native habitat (EUNIS code): E- Grassland and tall forb habitats. Habitat occupied in invaded range (EUNIS code): I- Regularly or recently cultivated agricultural and domestic habitats. The species can also feed on weedy Solanaceae and, thus, is able to invade various types of habitats. Native range: North America, Canada, Continental US, Mexico Introduced range Recorded for the first time in 1984 in southern Italy; then spread in the Balkans (Map) and Turkey. Also present in the archipelago of the Azores and of Pacific Ocean (French Polynesia - Tahiti, Hawaii). Credit: Rumen Tomov Factsheets for 80 representative alien species. Chapter 14 879 Pathways : The main mode of introduction and spread is the transport of infested plant material. Adults fly actively between fields. Impact and management: No ecological impact has been reported so far. The species is mainly known as a serious pest of tobacco. It is most dangerous early in the growing season for young seedlings in plant beds. The plants often die after being transplanted into the field. Numerous holes on leaves of mature plants and the adults’ excrement delay the growth of plants and reduce the quality and value of tobacco leaves. In heavy infestations, the plant may be totally defoliated like lacework. Both types of attack reduce plant vigour and value, and may ruin an entire plant bed. The pest has been suggested as a vector of Tobacco ringspot nepovirus, which causes significant disease in tobacco. It may also damage potato, aubergine and tomato. Monitoring seedlings is important for early detection of flea beetles. Chemical control involves a number of insecticides for plant bed and the field situations. Biological control using Bacillus thuringiensis ssp. tenebrionis on adults may represent a suitable alternative. Possible cultural control practices include the sterilization or fumigation of top soil before planting and removal of weedy vegetation and excess organic debris in surrounding areas. Selected references Deseo KV, Balbiani A, Sannino L, Zampelli G (1993) Zur Biologie und biologischen Bekampfung des Tabakkafers, Epitrix hirtipennis Melsh. (Col., Chrysomelidae) in Italien. Anzeiger fuer Schaedlingskunde Pflanzenschutz Umweltschutz 66: 26–29. Lykouressis DP, Mentzos G, Parentis A (1994) The phenology of Epitrix hirtipennis (Mels.) (Col., Chrysomelidae) and damage to tobacco in Greece. Journal of Applied Entomology 118: 245–252. Paparatti B, Scubla P (1994) A contribution to the study of the bioethology of Epitrix hirtipennis (Melsheimer) (Coleoptera, Chrysomelidae). Frustula Entomologica 17: 175–184. 880 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.10 – Leptinotarsa decemlineata Say, 1824 - Colorado potato beetle (Coleoptera, Chrysomelidae) Carlos Lopez-Vaamonde Description and biological cycle: Adults up to 11 mm long; elytra yellow with ten characteristic black longitudinal bands (Photo). Main natural spread of beetle over large areas is by windborne migration. Females usually deposit eggs on the underside surface of host plant leaves. An egg mass may contain 10–40 eggs. Most adult females deposit over 300 eggs during 4–5 weeks, but they can lay up to 800 eggs. Potatoes are the preferred host, but the Colorado potato beetle (Colorado beetle) may feed and survive on a number of other Solanaceae: eggplant, tomato, pepper, tobacco, ground cherry, horse-nettle, common nightshade, Belladonna, thorn apple, henbane, and its first recorded host plant: buffalo-bur, Solanum rostratum. Larvae are hardy and resistant to unfavourable weather. Native habitat (EUNIS code): G1- Broadleaved deciduous woodland. Habitat occupied in invaded range (EUNIS code): I1- Arable land and market gardens; I2Cultivated areas of gardens and parks. Beetles are sensitive to cold temperatures. They need at least 60 days of temperature over 15 °C in summer and winter temperatures not falling below 8 °C. Native range: Mexico, where beetles are still present and feed on wild Solanaceae such as Solanum rostratum. Introduced range: beetles were accidentally introduced into USA. In 1922, the species was introduced to France from where it expanded almost throughout the European continent (Map) and to parts of Asia in about 30 years. Capable of adapting to different climatic Credit: György Csóka Factsheets for 80 representative alien species. Chapter 14 881 conditions and different host plants, this beetle is constantly moving to new areas. Its distribution is limited by temperature and therefore climate warming could further expand its distribution range. Pathways: International trade appears to be the most likely pathway for introduction on imported commodities such as fresh vegetables from infested areas. Beetles can also be spread through wind and attachment to all forms of packaging and transport. Impact and management: Serious pest of potatoes. Both adults and larvae feed on potato leaves and damage can greatly reduce potato yields. Beetles can also be a pest of other solanaceous plants such as tomato, aubergine, tobacco and peppers. This beetle may be managed culturally by crop rotation. Mechanical control involving destruction of crop debris is very effective at reducing population levels. Chemical control commonly involves insecticides, but resistance to them develops rapidly. Biological control includes a long list of natural enemies. Bacillus thuringiensis and some species of nematodes have particularly been used as control agents. Selected references CABI/EPPO (1997) Quarantine pests for Europe, 2nd Ed. Wallingford, UK: CAB International. Grapputo A, Boman S, Lindström L, Lyytinen A, Mappes J (2005) The voyage of an invasive species across continents: genetic diversity of North American and European Colorado potato beetle populations. Molecular Ecology 14: 4207–4219. Pucci C, Dominici M, Forcina A (1991) Population dynamic and economic threshold of Leptinotarsa decemlineata (Say) (Col., Chrysomelidae) in Central Italy. Zeitschrift für Angewandte Entomologie 111: 311–317. 882 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.11 – Harmonia axyridis (Pallas, 1773) - Harlequin ladybird (Coleoptera, Coccinellidae) Helen Roy and David Roy Description and biological cycle: Polyphagous predatory ladybird, 5–8 mm long, variable in colour pattern (yellow to orange to black) with a variable number of spots (0–21) (Photoadults mating). Highly dispersive, flying readily between host plants during breeding periods. This species migrates long distances in Asia and America. 20–50 eggs are produced per day, 1000–4000 in their lifetime. Adults typically live for a year, reproducing for three months. The Harlequin ladybird is generally bivoltine but can produce four generations per year in favourable conditions (Majerus et al. 2006). Native habitat (EUNIS code): G- Woodland, forest habitats and other wooded land. Habitat occupied in invaded range (EUNIS code): Same range of habitats as in the native range as well as G3- Coniferous woodland; G5- Lines of trees, small anthropogenic woodlands, recently felled woodland, early-stage woodland and coppice; I- Regularly or recently cultivated agricultural, horticultural and domestic habitats; I1- Arable land and market gardens; I2- Cultivated areas of gardens and parks; and, J1- Buildings of cities, towns and villages. The wide native range in Asia shows that the species can reproduce in both warm and cool climates and is well adapted to temperature extremes. Native range: Central and Eastern Asia. Introduced range: America, South Africa, Egypt, Europe (Brown et al., 2008). Increasing trend (Map; from Brown et al 2008, modified). Pathways: Harmonia axyridis was introduced intentionally as a biocontrol agent of aphids and unintentionally in horticultural/ornamental material. Impact and management: Potential to effect biodiversity, particularly that of other aphidophages and non-pest insects, through resource competition, intraguild predation and direct intra-specific competition. This beetle is also a pest of orchard crops (apples and pears) because as aphids become scarce in autumn, H. axyridis feeds on soft fruit, causing blemishing Credit: Helen Roy Factsheets for 80 representative alien species. Chapter 14 883 and an associated reduction in the market value. A tendency to aggregate in bunches of grapes prior to harvest makes the ladybirds difficult to separate from the fruit and so they are sometimes processed during wine making. Alkaloids contained within these beetles adversely affect the taste of the vintage. The beetle’s propensity to swarm and its large winter indoors aggregations are regarded as a nuisance. Economic impact is mainly on the wine industry, with reduction in fruit quality and management measures required in domestic dwellings (Kenis et al., 2008). Preventing the use of H. axyridis as a biocontrol agent and ensuring that fruit and cut flower imports are clean will reduce introduction events. Invasion into households can be limited by covering entrances with fine mesh. Adults and late instar larvae can be removed from unwanted locations manually, e.g., using a vacuum cleaner. Light traps can attract adults but the efficiency of these is not yet quantified. Chemical control in field situations such as orchards and vineyards is not applicable because of the impact of insecticides on other aphidophages and beneficial insects (Kenis et al., 2008). Selected references Brown PMJ, Adriaens T, Bathon H, Cuppen J, Goldarazena A, Hagg T, Kenis M, Klausnitzer BEM, Kovar I, Loomans AJ, Majerus MEN, Nedved O, Pedersen J, Rabitsch W, Roy HE, Ternois V, Zakharov I, Roy DB (2008) Harmonia axyridis in Europe: spread and distribution of a non-native coccinellid. BioControl, 53: 5–22. Kenis M, Roy HE, Zindel R, Majerus MEN (2008) Current and potential management strategies against Harmonia axyridis. BioControl, 53: 235–252. Majerus MEN, Strawson V, Roy HE (2006) The potential impacts of the arrival of the Harlequin ladybird, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), in Britain. Ecological Entomology, 31: 207–215. 884 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.12 – Gonipterus scutellatus Gyllenhal 1833 - Eucalyptus snout beetle (Coleoptera, Curculionidae, Cyclominae) Daniel Sauvard Description and biological cycle: Medium-sized weevil species (12–14 mm), grey to brown with a light transverse band on the elytra and pale brown hairs (Photo left- adult on eucalyptus branch). This species is morphologically very similar to another Australian eucalyptus weevil, G. gibberus, invasive too but not present in Europe at this time. Hosts are different Eucalyptus species. The weevil has several generations per year (generally two in southern Europe). The adults emerge from the soil and feed on leaves (see Figure 8.2.6 in Chapter 8) and growing shoots. Throughout their life, females lay several egg batches protected by brown capsules on surfaces of young leaves (overall fecundity of a female is about 150–300 eggs). Yellowish-green larvae feed on leaves (Photo right- larval damage) and twigs, then fall on the ground and pupate in the soil. Overwintering occurs in the adult stage. Native habitat (EUNIS code): G2- Broadleaved evergreen woodland. Habitat occupied in invaded range: G2- Broadleaved evergreen woodland; G5- Lines of trees, small anthropogenic woodlands; I2- Cultivated areas of gardens and parks; X- parks and gardens. Native range: Southeastern Australia. Introduced range: Progressively introduced in all places where eucalyptus have been introduced: USA, South America, Western Australia, New Zealand, China, South and East Africa. In Europe, the Eucalyptus snout beetle was first recorded in Italy in 1990 and then in other Mediterranean countries (Map). Credit: Alain Roques Factsheets for 80 representative alien species. Chapter 14 885 Pathways: Adults, eggs and larvae can be transported with live eucalyptus; larvae and pupae can be transported with soil. The adults can fly to disperse locally; adult may hitch-hike, e.g. on vehicles. Impact and management: This weevil is an important eucalyptus pest in all areas where it has been introduced. Adults and especially larvae damage eucalyptus leaves, mainly young ones. Larvae characteristically damage only one surface of leaves, while adults chew the edge. Defoliation causes growth reduction, and even tree mortality in case of successive severe damage. Young trees are generally the most damaged. Susceptibility depends of Eucalyptus species; in Europe, the commonly planted E. globulus is one preferred host. Chemical control is not recommended due to side effects on honey bees often visiting eucalytus flowers. Biological control has been successfully achieved in several world and European countries using the Australian chalcid Anaphes nitens (Girault 1928) (Hymenoptera, Mymaridae), an egg parasitoid. Selected references Arzone A (1976) Un nemico dell’Eucalipto nuovo per l’Italia. Apicoltore Moderno 67: 173–177. Mansilla JP, Pérez Otero R (1996) El defoliador del eucalipto Gonipterus scutellatus. Phytoma España 81: 36–42. Rabasse JM, Perrin H (1979) Introduction en France du charançon de l‘eucalyptus, Gonipterus scutellatus Gyll. (Col., Curculionidae). Annales de Zoologie, Écologie Animale 11: 336–345. 886 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.13 – Rhopalapion longirostre (Olivier 1807) (= Apion longirostre) - Hollyhock weevil (Coleoptera, Apionidae) Katalin Tuba and Ferenc Lakatos Description and biological cycle: Small weevil species distinguished from other Apion and Aspidapion species by its orange legs and enormously narrow rostrum* (Photo- adults mating on Alcea leaf) The females have one of the longest rostra among all Middle-European Apionidae, longer than half the remaining body length which is 3–3.5 mm. Their black or rarely yellowish antennae are located halfway along this snout. The greyish-black body is densely hairy. Hosts are different Alcea species, especially A. rosae (Malvaceae). It develops on hollyhock with other host specialist Apionidae species, like Aspidapion aeneum, A. radiolus and Alocentron curvirostre. The hollyhock weevil has one generation per year. The adults overwinter under fallen leaves or in the soil around the stem. In spring the adults come out and chew the leaves, the petioles and the stem. The females lay eggs at the bottom of the buds or on young ornamental or herbaceous plants. The larvae develop by chewing seeds and sometimes leaf tissue. Adults emerge from the seeds in August-September. Native habitat (EUNIS code): G- Woodland and forest habitats and other wooded land; I: Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): I2 - Cultivated areas of gardens and parks. Credit: Ferenc Lakatos Factsheets for 80 representative alien species. Chapter 14 887 Native range: Temperate Asia. Introduced range: First detected in 1875 in Romania. Then, observed in most of the Mediterranean and Central-European countries (Map). Also introduced in North America since 1914 in Georgia. Pathways: Probably trade of ornamentals. Impact and management: This species can spread very quickly due to its distinct habit and special host plant especially in urban gardens. It can cause serious ecological damage in hollyhock cultivation as ornamentals or herbs. Both chemical and biological control is possible against this insect. However application is recommended mainly in production circumstances rather than garden situations. Selected references Ehret JM (1983) Apion (Rhopalapion) longirostre, espèce nouvelle pour la France (Coleoptera, Curculionidae). L‘Entomologiste 39: 42. Kozłowski MW, Knutelski S (2003) First evidence of an occurrence of Rhopalapion longirostre in Poland. Weevil News 13: 4 pp. http://www.curci.de/Inhalt.html. Perrin H (1984) Présence en France d‘Apion (Rhopalapion) longirostre (Olivier) (Coleoptera, Curculionidae, Apioninae) et répartition dans la région paléarctique. L‘Entomologiste 40: 269–274. 888 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.14 – Trogoderma granarium (Everts, 1898) (= T. khapra (Arrow, 1917)) (= T. afrum (Priesner, 1951)) - Khapra beetle (Coleoptera, Dermestidae) Katalin Tuba and Ferenc Lakatos Description and biological cycle: Adults of this tiny beetle species are ovate and densely haired (Photo- Adult, larva, larval skins and damage to wheat grains). Males are 1.4–2.3 mm long and 0.75–1.1 mm wide, while females are slightly larger and lighter in colour. Head and pronotum are dark reddish-brown, while elytra are reddish brown with two or three lighter, indistinct bands. Legs are yellowish-brown. Antennae consist of 10–11 segments; the last 3–4 segments at females and the last 5 segments at males forming a club. Antennae are yellowish-brown. Eggs are 0.7 mm long and 0.25 mm wide, cylindrical, milky white, turning pale yellowish with age. At the end of the embryonic development, six brown bands are visible in the eggs. Generally there are 6–8 larval stages, but under unfavourable development conditions, up to 10–12. A tail on the last abdominal segment is half of the whole larval body length. Larval body is yellowish-white with brown head and setae. Mature larva is 4–6 mm long and 1.5 mm wide. Male pupa is 3.5 mm long and female 5 mm, both sexes yellowish-brown and hairy. The khapra beetle has one to nine generations per year depending on nutritional resources, temperature, humidity, light and season. Each female lays a total of 50–100 eggs on host material. Development time varies between 26–220 days. Egg stage takes 3–14 days, while larvae live 4–6 weeks. Larvae can enter diapause if the temperature falls below 25°C and may remain in this condition for many years. The pupa stage takes 2–5 days. Adult khapra beetles have wings, but are not known to fly and feed very little. Mated females live from 4–7 days, unmated females from 20–30 days, and males from 7–12 days. Habitat occupied in invaded range (EUNIS code): J- Constructed, industrial and other artificial habitats. Credit: Ministry of Agriculture and Regional Development Archives of Hungary, bugwood.org Factsheets for 80 representative alien species. Chapter 14 889 Native range: India. Introduced range: all over the word except for Australia and New Zealand. Warmer climates are preferred, living in stores further north. The khapra beetle first moved into Europe in 1908. At first the eradication was successful but during the First World War, the species was introduced again and became established. Pathways: The wide geographical distribution of this pest is to a certain extent due to human activities. In the absence of flight, its spread is dependent on movement of infected goods. The species can spread with containers in which it diapauses. Impact and management: This polyphagous species is one of the most important and dangerous insect pest of the stored products. The gregarious larvae damage both quality and quantity of stored foodstuff, chewing seeds, while adults cause negligible damage. Food produce attacked includes grain and cereal (wheat, barley, oats, rye, maize, rice, flour, pasta) as well as beans, herbs, chocolates, nuts and many other products. Both physical and chemical control can be used against this insect but it is one of the most difficult detritivorous pests to manage. Established infestations are difficult to control and eradicate because the larva can live without food for long time and it can crawl and slip into tiny cracks and crevices, and is resistant towards many insecticides. Selected references Harris DL (2006) Trogoderma granarium Everts (Insecta: Coleoptera: Dermestidae). http://entnemdept.ufl.edu/creatures/urban/beetles/khapra_beetle.htm. [accessed 25 February 2010]. Jenser G., Mészáros Z, Sáringer Gy (Eds) (1998) A szántóföldi és a kertészeti növények kártevői. Mezőgazda Kiadó Budapest: 157–158. Szőnyegi S, Kalmár K (1999) Szemestermény tárolók károsítói és az ellenük való védekezés. Agroinform Kiadó Budapest: 49–50. 890 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.15 – Diocalandra frumenti (Fabricius, 1801) - Four-spotted coconut weevil (Coleoptera, Dryophthoridae) Wolfgang Rabitsch Description and biological cycle: Curculionid beetle, shiny black with four large red to brown spots on the elytra. Length of adults is 6–8 mm. Larvae bore galleries in roots, petioles, infloresences and fruits of palms, where they pupate (Photo left- pupal chambers in a palm frond). Adult emergence is noticed by round holes (Photo right- Adult emerging from a palm tree and emergence holes) The whole life-cycle takes 10–12 weeks. Larvae cause premature yellowing and collapse of palm fronds as well as shedding of fruits and may cause the death of the tree. This weevil causes damage to many palm species, including economically important and ornamental species (Cocos nucifera, Phoenix dactylifera, P. canariensis and Elaeis guineensis). Native habitat (EUNIS code): G2.5- Palm groves. Habitat occupied in invaded range (EUNIS code): G2.5- Palm groves; I- Regularly or recently cultivated agricultural, horticultural and domestic habitats. Native range: Southeast Asia. Introduced range: In Europe, it was found in the south of Gran Canaria, the Canary Islands, in 1998, on the endemic palm Phoenix canariensis. It has also established on the islands of Tenerife, Lanzarote and Fuerteventura, where it occurs in several protected areas. Also intro- Credit: EPPO Factsheets for 80 representative alien species. Chapter 14 891 duced in South Asia (Bangladesh, India, Indonesia, Malaysia, Philippines, Sri Lanka, Taiwan, Thailand), Japan (Ryukyu), East-Africa, Madagascar, Seychelles, Oceania (Australia, Guam, Palau, Papua New Guinea, Samoa, Solomon Islands, Vanuatu), and South America (Ecuador). Pathways: Probably with ornamental trade Impact and management: Introduction occurs with infested ornamental palm trees. D. frumentii is a threat to native Canary Islands Date Palm and it may change the fire regime and the structure, abundance and succession of invaded habitats. Detection and control is difficult due to the secretive life habit. Selected references González Núñez M, Jiménez Álvarez A, Salomones F, Carnero A, Del Estal P, Esteban Durán JR (2002) Diocalandra frumenti (Fabricius) (Coleoptera: Curculionidae), nueva plaga de palmeras introducida en Gran Canaria. Primeros estudios de su biología y cría en laboratorio. Boletín de Sanidad Vegetal Plagas, 28(3): 347–355. Samarin Bello CR (2008) Diocalandra frumenti (Fabricius, 1801). In: Silva L, Ojeda Land E, Rodríguez Luengo JL (Eds) Invasive Terrestrial Flora & Fauna of Macaronesia. Top 100 in Azores, Madeira and Canaries. ARENA, Ponta Delgada, pp. 418–420. 892 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.16 – Rhynchophorus ferrugineus (Olivier, 1790) - Red Palm Weevil (Coleoptera, Dryophthoridae) Wolfgang Rabitsch Description and biological cycle: Curculionid beetle, reddish-brown, with dark spots on elytra (Photo- see also Figure 8.2.7a in Chapter 8). Length of adults is 3.5 cm, length of larvae 5 cm. Females bore small holes in the trunk of a palm tree and lay up to 300 eggs. Grub-like larvae (see Figure 8.2.7b in Chapter 8) feed on soft tissues inside the host plant, producing long tunnels up to 1 m inside the trunk. Pupation takes place at the base of the palm. Completion of the life-cycle takes four months. Feeding activity of larvae can completely destroy palms. The diurnal adults search for new palms and can fly distances up to 1 km. Long-distance dispersal happens via infested palm trees. Host plants are different palm tree species. According to EUDecision 2007/365/EC, susceptible host plants, other than fruit and seeds, having a diameter of the stem at the base of over 5 cm are Areca catechu, Arenga pinnata, Borassus flaellifer, Calamus merillii, Caryota maxima, C. cumingii, Cocos nucifera, Corypha gebanga, C. elata, Elaeis guineensis, Livistonia decipiens, Metroxylon sagu, Oreodoxa regia, Phoenix canariensis, P. dactylifera, P. theophrasti, P. sylvestris, Sabal umbraculifera, Trachycarpus fortunei and Washingtonia spp. The weevil was also found in Agave americana, Brahea armata, Butia capitata, Howea firsteriana and Saccharum officinarum. Native habitat (EUNIS code): I- Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): I- Regularly or recently cultivated agricultural, horticultural and domestic habitats; X24- Domestic gardens of city and town centres; X25- Domestic gardens of villages and urban peripheries. Credit: Olivier Denux Factsheets for 80 representative alien species. Chapter 14 893 Native range: South(east)-Asia Introduced range: R. ferrugineus has been introduced in most of Mediterranean countries and islands of Europe (Map). Since its first discovery in Spain (Andalucía and Valenciana, 1994) it colonized Italy (2004, incl. Sardegna, Sicilia), the Canary Islands (2005), Mallorca (2006), France (incl. Corsica), Greece, Cyprus (2006), Malta, Portugal (2007), Albania (2009) and Ceuta (2009). It also occurs in the Eastern Mediterranean region (Turkey, Syria, Israel, Jordan, Egypt) and North Africa (Morocco). Also introduced to Oceania (Australia?, Papua New Guinea, Solomon Islands), China, the Near and Middle East, and the Caribbean (Netherlands Antilles). Pathways: Probably ornamental trade of palm trees Impact and management: Symptoms are visible only late after colonization of the beetle and usually palms do not recover when symptoms become visible (see Figure 8.2.7c in Chapter 8). Early detection is difficult, as symptoms of infested palms remain unnoticed for several years. EU-Decision 2007/365/EU requires phytosanitary certificates for palm imports and eradication measures at infested areas. Imported palms needs to be kept in quarantine for inspection. This also is recommended for another Rhynchophorus species, R. palmarum, listed by EPPO as A1 species, not yet introduced to Europe, but presenting a similar phytosanitary risk to palms. Infected palms must be cut and burned or buried deeply. IPM employs pheromone traps to monitor and collect beetles. Removal of offshoots, which are preferred oviposition sites, is not recommended, because wounds attract females to oviposit. Also pruning of palm leaves should be carried out in winter, as pruning attracts beetles and facilitates egg laying. It is estimated that more than 30.000 palm trees have been destroyed in Spain within three years. Selected references Barranco P, de la Pena J, Cabello T (1996) El picudo rojo de las palmeras, Rhynchophorus ferrugineus (Olivier), nueva plaga en Europa (Col., Curculionidae). Phytoma-Espana 76: 36–40. Faleiro JR (2006) A review on the issues and management of red palm weevil Rhynchophorus ferrugineus (Coleoptera: Rhynchophoridae) in coconut and date palm during the last one hundred years. International Journal of Tropical Insect Science 26: 135–154. Liu G, Peng ZQ, Fu YG (2002) Research advances on the red palm weevil Rhynchophorus ferrugineus. Journal of Tropical Agricultural Science 22: 73–77. 894 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.17 – Megaplatypus mutatus (Chapuis, 1865) - The grand forest borer (Coleoptera, Curculionidae, Platypodinae) Massimo Faccoli Description and biological cycle: Adult females 8–9 mm long, males slightly smaller (7.5 mm). Body brown above and reddish-yellow below, with reddish tarsi and antennae (Photo). Elytrae of males with sulcate* striae and characteristic spiniform processes on the declivity*; female elytral declivity rounded and without processes. Mature larvae about 7.2 mm long. M. mutatus bores 3 mm-wide holes in the trunk, approximately 4 m above ground level. Adults excavate long and sinuous galleries that become covered by the black mycelium of symbiotic fungi, which nourish larval instars. Unlike other ambrosia beetles, M. mutatus attacks standing and vigorous trees. The stem attack does not kill the plant immediately, and the same tree may be re-infested several times by subsequent generations. M. mutatus infests mainly poplars (Populus spp.), willows (Salix spp.) and important fruit trees species such as apples (Malus spp.), walnuts (Juglans spp.) and avocados (Persea spp.). The species has been recorded also on Acer, Citrus, Eucalyptus, Fraxinus, Laurus nobilis, Magnolia grandiflora, Platanus, Prunus, Quercus, Robinia pseudacacia, Tilia and Ulmus. Credit: G. Allegro, CRA Istituto di Sperimentazione per la Pioppicoltura, Casale Monferrato, eppo.org Factsheets for 80 representative alien species. Chapter 14 895 Native habitat (EUNIS code): G1 - Broadleaved deciduous woodland; G2 - Broadleaved evergreen woodland. Habitat occupied in invaded range (EUNIS code): G1 - Broadleaved deciduous woodland; I2 - Cultivated areas of gardens and parks. Native range: subtropical and tropical areas of South America. The weevil has extended its range into temperate regions, reaching as far south as Neuquén in Argentinean Patagonia. Introduced range: recently introduced and acclimatized in the Napoli region, Italy (2000) (Map). Adult flight can ensure species dispersal over short distances. Pathways: Man-mediated long-distance dispersal is possible by international trade of infested woody plants and woody materials. Impact and management: M. mutatus represents a threat for many woody species widely cultivated in Europe for commercial or ornamental purpose. It is a primary pest in commercial poplar plantations, especially those of P. deltoides. In Italy, damage has been recorded also on fruit trees (Corylus avellana, Prunus cerasus, Pyrus communis and Malus domestica). The damage is caused by adults, which bore large gallery systems into living host-trees. The galleries and associated fungi degrade the lumber and weaken the tree stems, which often then break during windstorms. As most of the life cycle is accomplished within wood tissues, this species is difficult to detect and to control, although some chemicals are available. Recent applications of the mating disruption technique are giving promising results. Selected references Tremblay E, Espinosa B, Mancini D, Caprio G (2000) Un coleottero proveniente dal Sudamerica minaccia i pioppi. L‘informatore Agrario 48: 89–90. 896 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.18 – Gnathotrichus materiarius (Fitch, 1858) American Utilizable Wood Bark Beetle (Coleoptera, Curculionidae, Scolytinae) Massimo Faccoli Description and biological cycle: Small species, 3.2–3.5 mm long, with a cylindrical, narrow and elongated body (Photo). Elytra dark reddish, generally smooth with only few short bristles on the declivity. Pronotum black and very fine punctured, except at the front where there are several granules. Head blackish and smooth, antennae with funicle*5-segmented and club with two round sutures. Anterior coxae fused. This weevil is polyphagous on conifer trees, in Europe being recorded on Pinus, Abies, Picea, Larix and Pseudotsuga. Gnathotrichus materiarius is an ambrosia beetle excavating timber galleries 1mm in diameter and 10–15 cm long. The galleries host the black fungus Endomycopsis fasciculata Batra, which nourishes the larvae. In Central Europe, the adults fly between April and the middle of June, but it is possible that a second flight takes place at the end of the summer. The species is monogamous, but males are very rare. Mature larvae or young adults overwinter in their galleries. Native habitat (EUNIS code): G3- Coniferous forests. Habitat occupied in invaded range (EUNIS code): G3- Coniferous forests. Credit: Louis Michel Nageilesen Factsheets for 80 representative alien species. Chapter 14 897 Native range: North America Introduced range: Since its first discovery in 1933 in France, this weevil has invaded a large part of Central, Western, Central and Northern Europe (Map). Man-mediated longdistance dispersal is possible by infested wood movement. Pathways: Trade of trees or timber. Impact and management: This species may disturb forest ecosystems by direct/indirect competition with native xylophagous insects. Typically a secondary species, recorded only on trees already felled or killed by other bark beetles. However, infestations reduce timber value because of damage from adult galleries and black discolouration caused by associated ambrosia fungi. Control is usually not required. Population monitoring may be based on visual detection of damage and by pheromone traps. Mechanical control may consist in the destruction of infested trees by chipping or burning. Natural enemies (parasitoid insects, entomopathogenic nematodes, fungi or bacteria) for possible biological control are under investigation but not yet being used. Chemical control has limited effect because the insects live deep within wood. Selected references Balachowsky A (1949) Coléoptères Scolytides. Faune de France 50. Paris: Librairie de la faculté des sciences. 320 pp. Faccoli M (1998) The North American Gnathotrichus materiarius (Fitch) (Coleoptera Scolytidae): an ambrosia beetle new to Italy. Redia 81: 151–154. Valkama H, Martikainen P, Raty M (1997) First record of North American ambrosia beetle Gnathotrichus materiarius (Fitch) (Coleoptera, scolytidae) in Finland - a new potential forest pest? Entomologica fennica 8: 193–195. 898 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.19 – Phloeosinus rudis Blandford, Japanese Thuja Bark Beetle (Coleoptera, Curculionidae, Scolytinae) Leen Moraal Description and biological cycle: Scolitid beetle, dark-brown, length of adults 2.5–3.0 mm (Photo). Females bore into the trunks of shrubs and trees weakened by drought and other stress factors. They excavate 2–3 armed egg galleries with a length varying from 5–16 cm. Feeding activity of the larvae kills the hosts. The diameter of emergence holes varies from 1.1–1.9 mm. The beetle overwinters, with a small percentage of adults, since the larval stage predominates. This weevil produces one generation per year. The photo shows an adult and also larval galleries in a 60-year-old tree of Thuja occidentalis. Host plants are Thuja, Chamaecyparis and Juniperus. Native habitat (EUNIS code): G3- Coniferous forests. Habitat occupied in invaded range (EUNIS code): X24- Domestic gardens of city and town centres; X25- Domestic gardens of villages and urban peripheries. Native range: Japan Introduced range: First found outside Japan in Southern France near St-Tropez, in June 1940, in dying branches of a Thuja japonica plantation. However, the beetle has not been Credit: Leen Moraal Factsheets for 80 representative alien species. Chapter 14 899 recorded in France since then. During the summer of 2004, hundreds of conifers, old solitary trees as well complete hedges, died in several cities around Rotterdam. Between 2004–2008, few infestations were found. This is probably due to the return to normal summer precipitation of the years following 2004, leading to more vigorous plants with less infestation. Because small beetle populations may survive in weakened trees, a new drought period may result in a new weevil population build-up. Pathways: All Dutch locations were situated within 30 km of the harbour of Rotterdam. It is suspected that P. rudis may have escaped from imported material from this harbour, but this could not be verified. The beetle was intercepted several times in the USA in wood off-cuts integrated in steel products from Asia, but there are no records of establishment in the USA. Impact and management: Symptoms become visible in summer when the needles turn brown and the hosts are dying. Removal and destruction of larval infested plants is recommended to control populations. Selected references Balachowsky A (1949) Coléoptères scolytides. Faune de France 50. Paris : Lechevallier, 124–126. Hoffmann A (1942) Description d ‘un genre nouveau et observations diverses sur plusieurs espèces de Scolytidae (Col.) de faune Francaise. Bulletin de la Société Entomologique de France 47: 72–74. Pfeffer A (1995) Zentral- und westpaläarktische Borken- und Kernkäfer (Coleoptera: Scolytidae, Platypididae). Pro Entomologia, Naturhistorisches Museum, Basel: 69. 900 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.20 – Xylosandrus crassiusculus (Motschulsky, 1866) - Granulate Ambrosia Beetle (Coleoptera, Curculionidae, Scolytinae) Massimo Faccoli Description and biological cycle: The adult is about 2.1–2.9 mm long, with stout body, elytra 1.15 times longer than pronotum and reddish (Photo left- female). Elytra with vestiture of long setae in irregular rows. Elytral declivity dull, with dense, numerous, uniformly distributed granules, allowing easy distinction from other Xylosandrus species occurring in Europe. Male smaller than female, rare. Xylosandrus crassiusculus is an ambrosia beetle developing within the wood and feeding on the mycelium of ambrosia fungi. Mated females bore small chambers and lay eggs in groups. Larvae develop together feeding on the fungus growing on the chamber walls. During gallery formation, the female compacts and pushes out the frass, which extends from the entrance hole forming a long, easily visible cylinder (Photo right). The adults usually overwinter in galleries at the base of the trees. X. crassiusculus develops in Europe on Carob tree, Ceratonia siliqua, but is highly polyphagous in the native range on Pinus spp. and broadleaved trees. Native habitat (EUNIS code): G - Woodland and forest habitats and other wooded land Habitat occupied in invaded range (EUNIS code): G2 - Broadleaved evergreen woodland; J100- Greenhouses. Native range: Paleotropical species (Africa and Asia) Credit: meta.arsia.toscana.it (left), EPPO (right) Factsheets for 80 representative alien species. Chapter 14 901 Introduced range: since 2003 recorded in Europe (Italy: Tuscany and Liguria- Map), probably introduced with infested trees or timber. Also introduced in North America. Pathways: Long-distance dispersal is possible by trade of infested timber. Impact and management: This beetle may disturb forest ecosystems by direct and indirect competition with native xylophagous insects. It may attack trees from about 2 cm stem diameter upwards in both stressed plants and is found in harvested timber. At high population density, X. crassiusculus may attack and kill healthy trees causing significant economic loss. Infested timber has reduced value because of adult galleries and black discolouration due to associated ambrosia fungi. Population monitoring and control may be performed using pheromone traps and the trees themselves as traps. Mechanical control is effected by destruction of infested trees via chipping or burning. Chemical control has limited effect because the insects develop deep within wood. Prevention is achieved via timber debarking before insect infestation and by keeping trees in good physiological condition. Selected references Pennacchio F, Roversi PF, Francardi V, Gatti E (2003) Xylosandrus crassiusculus (Motschulsky) a bark beetle new to Europe. Redia 86: 77–80. 902 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.21 – Xylosandrus germanus (Blandford, 1894) (= Xyleborus germanus) - Black stem borer (Coleoptera, Curculionidae, Scolytinae) Massimo Faccoli Description and biological cycle: Ambrosia beetle, 2.0–2.4 mm long, with shiny black elytra, surface of elytral declivity* shining, anterior margin of pronotum with 8–10 asperities* (Photo). This species is highly polyphagous on conifers and broadleaves, attacking a wide range of both living plants and timber. Adults fly during early to mid May, infesting timber of the lower part of the trunk of stressed trees. By specialized organs (tegumental mycangia*), during tunnel excavation, the female introduces the pathogenic fungus Ambrosiella hartigii (Batra) (= Monilia candida Hartig) into the host plants. The associated fungus causes cankered areas in the stem and treetop as well as branch dieback and suckering. During gallery formation, frass ejected by the female often protrudes as a long and conspicuous cylinder. Although in Europe generally considered to be monovoltine, two generations per year have been observed in Germany and Italy. Adults of X. germanus usually overwinter in galleries at the base of attacked trees. Hosts in the invaded range include both broadleaved species (Fagus, Castanea, Buxus, Ficus, Carpinus, Quercus and Juglans) and conifers (Picea and Pinus). Native habitat (EUNIS code): G - Woodland and forest habitats and other wooded land. Habitat occupied in invaded range (EUNIS code): G - Woodland and forest habitats and other wooded land. Native range: Asia. Credit: Christoph Benisch, kerbtier.de Factsheets for 80 representative alien species. Chapter 14 903 Introduced range: First detected in Germany in 1950. Then, the weevil spread in most countries of Western and Central Europe as far as the European part of Russia (Map). Increasing frequency of interceptions has been reported during recent years in Europe. Also introduced in North America. Pathways: Man-mediated long-distance dispersal is possible by movement of infested timber. Impact and management: This weevil may disturb forest ecosystems by direct/indirect competition with native xylophagous insects. Such secondary* species have been recorded on stressed living trees or harvested timber. Water stress is one of the main causes inducing stem colonization of living trees. Infestation kills the host plant and reduces timber value because of damage from adult galleries and black discolouration due to associated ambrosia fungi. Population monitoring and control may be performed by pheromone traps and trees used themselves as traps. Mechanical control involves destruction of infested trees by chipping or burning. Chemical control is of limited effect because the insects live deep within wood. Damage reduction and prevention may rely on timber debarking before insect infestation and keeping trees in good physiological condition. Selected references Bruge H (1995) Xylosandrus germanus (Blandford, 1894) [Belg. sp.nov.] (Coleoptera Scolytidae). Bulletin et annales de la Société Royale Belge d’Entomologie 131: 249–264. Graf E, Manser P (1996) Der Schwarze Nutzholzborkenkäfer, Xylosandrus germanus. Wald und Holz 2: 24–27. Henin JM, Versteirt V (2004) Abundance and distribution of Xylosandrus germanus (Blandford 1894) (Coleoptera, Scolytidae) in Belgium: new observations and an attempt to outline its range. Journal of Pest Science 77: 57–63. 904 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.22 – Tribolium confusum (Du Val, 1868) - Confused flour beetle (Coleoptera, Tenebrionidae) Katalin Tuba and Ferenc Lakatos Description and biological cycle: Small beetle species. Adults are 2.6–4.4 mm long, shiny, reddish-brown or chestnut-brown (Photo). The first antennal segments are obscured by the forehead. Antennae widen from the 5–6th segment. The head and the pronotum are finely dotted. The elytra are patterned with lines also consisting of fine dots. Adults may live more than three years. Eggs are 0.4 mm long and white. Larvae have six larval stages. Their length is 6–7 mm in the final larval stage. Young larvae are white, aging yellowish. The body of larvae is cylindrical and slight hairy. On the ninth abdominal segment there are two hook-like projections. Larvae have three pairs of legs. The length of the yellowish brown pupa is 3–4 mm. There are three or five generations per year. Development time is about 40–45 days under optimal circumstances depending on sex, temperature, humidity and nutrition. Each female lays a total of 450–900 eggs. Females lay eggs one by one and thus the oviposition period is long. Eggs adhere well to the crop surface with a glue-like material. Larvae live for 3–4 weeks chewing crops. The pupa stage lasts a maximum of two weeks. The adults mature after 2–7 days. Adults overwinter in stores. Native habitat (EUNIS code): I- Regularly or recently cultivated agricultural, horticultural and domestic habitats; J- Constructed, industrial and other artificial habitats. Habitat occupied in invaded range (EUNIS code): J- Constructed, industrial and other artificial habitats. Native range: Africa. Credit: Christoph Benisch, kerbtier.de Factsheets for 80 representative alien species. Chapter 14 905 Introduced range: All over the world. Tribolium confusum can withstand cooler climates than red flour beetle (T. castaneum), which is found in more temperate areas. The confused flour beetle moved into Europe from America at the end of the 19th century. Its range limit in Europe is now Scandinavia (Map). Pathways: The wide geographical distribution of this pest is to a certain extent due to dissemination of infested stored products. Impact and management: The confused flour beetle is one of the most important pests of the stored products in homes, groceries and granaries. This is a highly polyphagous species. Both adults and larvae cause damage, but the main pests are larvae. These attack flour, cereals, meal, crackers, pasta, cake, beans, peas, spices, dried pet food, chocolates, nuts, seeds and even dried museum specimens. Crops contaminated by larval skins, excrement and chewed residues are smelly and rendered inedible by both humans and animals. Physical and chemical as well as biological control can be used against this insect. Interestingly, cannibalistic interactions among certain life stages (eggs and pupae by adults, and eggs by larvae) constitute a natural control mechanism of confused flour beetles. Selected references Benoit HP, McCauley E, Post JR (1998) Testing the demographic consequences of cannibalism in Tribolium confusum. Ecology 79: 2839–2851. Baldwin R, Fasulo TR (2003): Tribolium confusum Jacqulin du Val (Insecta: Coleoptera: Tenebrionidae). http://www.entnemdept.ufl.edu/creatures/urban/beetles/red_flour_beetle. htm [accessed 25 February 2010]. Jenser G, Mészáros Z, Sáringer Gy (Eds) (1998) A szántóföldi és a kertészeti növények kártevői. Mezőgazda Kiadó Budapest: 177–178. 906 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.23 – Liriomyza huidobrensis (Blanchard, 1926) - Serpentine leaf miner (Diptera, Agromyzidae) Alain Roques Description and biological cycle: Small fly, adult 1.3–2.3 mm long, compact-bodied, greyishblack (Photo left); maggot appears headless, up to 3.3 mm in length, yellow-orange at maturity. Larvae are leaf miners on a wide range of hosts, especially economically important vegetables and ornamental plants in both glasshouses and outdoors. Adult flight range is limited. Longrange dispersal (eggs, larvae) occurs with human-transported infested plant material, including cut flowers. The vase-life of chrysanthemums is sufficient to allow completion of the life-cycle. Under laboratory conditions, a female lays about 100–130 eggs but up to 250 eggs have been observed. Eggs are laid into the leaf tissue. Larvae tunnel within the leaf tissue forming characteristic mines (Photo right- mines with a puparium), then cut a semi-circular opening in the tissue and drop to the soil to pupate. The life cycle can be as short as 14 d at 30 °C or as long as 64 d at 14 °C. Generations follow in quick succession as long as the growing conditions of the host plant provide suitable food. Optimal temperatures for feeding and egg laying range between 21–32°C. Egg-laying is reduced at temperatures below 10 °C. All stages are killed within a few weeks by cold storage at 0 °C and above 40 °C. Native habitat (EUNIS code): F5- semi-arid and subtropical habitats Habitat occupied in invaded range (EUNIS code): I1- Arable land and market gardens; I2- Cultivated areas of gardens and parks; J100- glasshouses. Native range: South America. Introduced range: First recorded from France in 1989, spreading with imported ornamentals; now outdoors in southern Europe (including Sicily and the Canary islands- Map), but mainly a glasshouse pest in northern Europe. Also introduced in Central America, most of Asia (China, Taiwan, India, Thailand, Singapore, Indonesia), Asia Minor, Africa (Kenya), and Indian Ocean (Reunion, Mauritius, Seychelles). Credit: Jean Yves Rasplus (left), Michel Martinez (right) Factsheets for 80 representative alien species. Chapter 14 907 Pathways: Passive transport with plant trade including vegetables, cut flowers and nursery stock. Impact and management: A serious pest for the floricultural industry, where leaf-miner damage directly affects the marketable portion, or in vegetable crops where the leaves are sold as the edible part. Sticky traps can be used to monitor adult flies. Crop rotation is an effective pest management tool, as is avoiding varieties more highly susceptible to leaf-miner infestations in glasshouses. There is little information about the leaf-miner tolerance of field vegetables. In this case, cultivation of crop debris or removal of infected plant material is recommended. L. huidobrensis adults are resistant to conventional insecticides. At present, the only effective insecticides are translaminar insecticides (abamectin, cyromazine, neem and spinosad), which penetrate the leaves to affect the leaf-miner larvae. Parasitoid wasps (e.g., Diglyphus isaea and Dacnusa sibirica)are available for control in glasshouse crops. These parasitoids will not be effective for vegetables growing in the field. However, there may be natural parasites present that can reduce the population. Selected references Maseti A, Luchetti A, Mantovani B, Burgio G (2006) Polymerase Chain Reaction-restriction fragment length polymorphism assays to distinguish Liriomyza huidobrensis (Diptera: Agromyzidae) from associated species lettuce cropping systems in Italy. Journal of Economic Entomology 99(4): 1268–1272. Phalip M, Martinez M (1994) Liriomyza huidobrensis : ses plantes hôtes et les confusions possibles avec d’ autres espèces. PHM Rev Hort 353: 24–28. Scheffer SJ, Lewis ML (2001) Two nuclear genes confirm mitochondrial evidence of cryptic species within Liriomyza huidobrensis (Diptera: Agromyzidae). Annals of the Entomological Society of America 94: 648–653. 908 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.24 – Liriomyza trifolii (Burgess, 1880) - Chrysanthemum leaf miner (Diptera, Agromyzidae) Ejup Çota Description and biological cycle: The adult fly is small, greyish-black, compact-bodied, 1–1.3 mm in length, up to 1.7 mm in the female, with wings 1.3–1.7 mm (Photo left). Eggs are 0.2–0.3 mm x 0.10–0.15 mm, off-white and slightly translucent. The larva is a “headless” maggot up to 3 mm in length when fully grown. First instar larvae are colourless on hatching, turning pale yellow-orange. Later instars are yellow-orange. Female flies puncture the leaves of the host plants causing wounds, which serve as sites for feeding or oviposition. Eggs are inserted just below the leaf surface. Hatching occurs 2–5 days later and the three larval instars make serpentine mines in the leaves (Photo right). The larvae develop in a few days and leave the mine to pupate in the soil or in crop debris. There are many generations per year. The life cycle from oviposition to adult emergence can be as short as 12 d at 35°C or as long as 54 d at 15°C. Adult flies are capable of limited flight. Native habitat (EUNIS code): I- Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): I1- Arable land and market gardens; I2- Cultivated areas of gardens and parks; J100- glasshouses. Native range: North America. Introduced range: First detected in France in 1976, now occurring in most European countries (Map) but unable to overwinter outdoors in Northern and Central Europe, and found only in glasshouses in these regions. Pathways: Trade of plant material, e.g. cut flowers, plants for planting out, and vegetables. Credit: Rémy Coutin/ OPIE (left), Jean Pierre Lyon/ INRA (right) Factsheets for 80 representative alien species. Chapter 14 909 Impact and management: Feeding and oviposition punctures of adults affect the value of ornamentals. However, damage is mainly done by larvae mining into the leaves and petioles, which reduces photosynthesis and may result in leaf drop. Mines are typically serpentine, tightly coiled and of irregular shape. Liriomyza trifolii is a major pest of various Asteraceae worldwide, both in outdoor crops and in glasshouses. It is particularly serious on Chrysanthemum, but also celery, onion, tomato, Gerbera, etc. In addition to the impact on yield, mines and feeding punctures also reduce the commercial value of ornamental plants and vegetables. Control by insecticides is feasible, although resistance is a problem. In glasshouses, the leaf miner is best controlled using natural enemies, such as parasitoids or nematodes. In field vegetables, cultivation of crop debris or removal of infected plant material is recommended. To prevent the introduction and establishment of L. trifolii and other leaf miner species, it is recommended that propagating material of susceptible plants from countries where the pests occur should be inspected at least every month for three months and verified free from the pests. Selected references Aguilar J d’, Martínez M (1979) Sur la présence en France de Liriomyza trifolii Burgess. Bulletin de la Société Entomologique de France 84, 143–146. Arzone A (1979) L’agromizide neartico Liriomyza trifolii (Burgess) nuovo nemico di Gerbera in Italia. Informatore Fitopatologico 29: 3–6. Sher RB, Parrella MP (1996) Integrated biological control of leafminers, Liriomyza trifolii, on greenhouse chrysanthemums. Proceedings of the meeting Integrated control in glasshouses, held in Vienna, Austria, 20–25 May 1996. Bulletin OILB/SROP 1996: 147–150. 910 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.25 – Dasineura gleditchiae (Osten Sacken, 1866) - Honey locust pod midge (Diptera, Cecidomyiidae) Ljubodrag Mihajlović and Milka M. Glavendekić Description and biological cycle: Approximately 2.0–3.0 mm long, antennae, long, moniliform* antennae with 12 flagellar segments, compound eyes holoptic* with no ocelli. Thorax grey with two prominent black longitudinal stripes. First tarsomere considerably shorter than second segment out of five and tarsal claws with large, basal teeth. Mouthparts reduced. Unsculptured eggs elongate-ovoid and opaque-white, turning opaque-red. Larvae elongate and dorso-ventrally flattened with pebble-like integument, varying from white to orange. First-third larval instars 0.57–2.44 mm long. Pupae approximately 2.43 mm long, obtect* with horn-like spines located at antennal base. Pupae sexually dimorphic; females with red and males with grey abdomen. D. gleditchiae is monophagous, living on Gleditsia triacanthos L. Generation time ranges from 21–30 days with several overlapping generations per year, overwintering as pupae or late instar larvae in cocoons in the soil. First appearance of the gall midge is in April, with males appearing first. Females deposit eggs on new foliage along the rachis or on edges of developing leaf buds. Eggs usualy hatch in two days. Young larvae crawl along the leaf to begin feeding. Only one larva is required to induce galling of the leaf. Leaf galls may be folded, partially pod-like, or the entire leaf may form a pod (Photo). The leaf gall dies and drops once the larvae pupate and emerge. Native habitat (EUNIS code): G1 - Broadleaved deciduous woodland; G5 - Lines of trees, small anthropogenic woodlands, recently felled woodland, early-stage woodland and coppice. Habitat occupied in invaded range (EUNIS code): FA- Hedgerows; I2 - Cultivated areas of gardens and parks and landscape; X24- Domestic gardens of city and town centres. Native range: Nearctic species widespread in North America. Credit: Milka M. Glavendekić Factsheets for 80 representative alien species. Chapter 14 911 Introduced range: First discovered in Europe in 1980 in the Netherlands. Since that time, galls of this gall midge were recorded in several other countries of Central and Southern Europe (Map). Also introduced in Turkey. Pathways: The main mode of introduction and spread is passive transport of coccons in soil with nursery stock or directly with infested young plants. Dispersal on a local scale is realized by active flight of adults and favoured by wind. Impact and management: Honey locust pod gall midge is a major pest of honey locust. Feeding by midge larvae causes leaflets of new growth to form pod like galls in which the larvae pupate. After the adult midge emerges from the pod, the leaf tissue dies and drops prematurely. Much of new growth can be affected, reducing the aestetic quality of trees in nurseries and landscapes. Monitoring of honey locust trees in nursery and landscape sites should begin in early spring and throughout the growing season, noting the appearance of eggs deposited on buds and new foliage by overwintering and first generation adults. Clusters of red midge eggs on honey locust buds can be observed with a hand lens. Effective chemical control is achieved by using various organophosphates, pyrethroids, carbamates and neonicotinoids. Oil applications in a narrow date-range targeting the first two egg depositions in April should facilitate midge population suppression. Biological control can be achieved using the Nearctic parasitoid Zatropis catalpae Craw. (Hymenoptera., Pteromalidae). Selected references Bolchi Serini G, Volonté L (1985) Dasineura gleditchiae (Osten Sacken), Cecidomide nuovo per l´Italia (Diptera, Cecidomyiidae). Bollettino di Zoologia agraria e di Bachicoltura Ser. II 18: 185–189. Dauphin P (1991) Sur la présence en France de Dasineura gleditchiae (Diptera, Cecidomyiidae), gallicole sur Gleditsia triacanthos (Fabacées, Caesalpinoidae). Bulletin de la Société Linnéenne de Bordeaux 19: 126. Estal P, Soria S, Vinuela E (1998) Nota de la presencia en España de Dasineura gleditchiae (Osten Sacken), sobre acacia de tres espinas. Boletín de Sanidad Vegetal Plagas 24: 225–230. 912 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.26 – Obolodiplosis robiniae (Haldeman, 1847) - Black locust gall midge (Cecidomyiidae, Diptera) Marcela Skuhravá Description and biological cycle: Adult small, body 2.5–3.2 mm long, reddish-brown, with long antennae, hairy wings and long densely haired legs (Photo left). Larvae at first whitish, fully grown larvae pale yellow, with typical sclerotized organ - spatula sternalis* – on ventral surface of prothoracic segments. Larvae are monophagous, inducing galls on leaves of black locust (Robinia pseudoacacia), a Fabaceae tree originating in North America. Larvae are gregarious and develop in galls formed of downwards rolled leaflet margins (Photo right). Several overlapping generations develop during one vegetative season. Pupation takes place in galls. In autumn, fully grown larvae leave galls and drop to the soil, where they hibernate till the spring of the next year. The population density is high. Native habitat (EUNIS code): G1 - broadleaved and deciduous woodland, native in north-eastern part of USA. Habitat occupied in invaded range (EUNIS code): G5- Lines of trees, small anthropogenic woodlands, recently felled woodland, early-stage woodland and coppice; I2 - Cultivated areas of gardens and parks; FB - Shrub plantations Native range: North America. Introduced range: Galls appeared suddendly in north-eastern Italy at Paese, Treviso Province in 2003, but the source of the infestation remains unknown. In 2004, the galls were found in northern Italy and in the Czech Republic at high infestation levels and in Slovenia. In the course of four years, the species spread very quickly in several countries of Europe and at present it occupies a large distribution area from England to Ukraine (Doneck) and from northern Germany to southern Italy (Map). The galls of O. robiniae appeared also suddendly in Korea and Japan in 2002. O. robiniae has a strong tendency to spread and can quickly become abundant in newly colonized areas. Pathways: Black locust gall midge probably arrived in Europe with plant materials imported from the USA. The source of its rapid spread in Europe may be international traffic along Credit: György Csóka (left),Vaclav Skuhravý (right) Factsheets for 80 representative alien species. Chapter 14 913 roads. Larvae drop from the galls and may be transported in vehicles over large distances. Adult midges, due to their small size, may be transferred by the wind. Young seedlings in forest nurseries or nurseries raising ornamental shrubs and trees may be transported to new places hidden in their indistinct galls. High fecundity of females and exponential growth of populations in the course of one vegetative season has contributed to the rapid spread of this species in Europe. Impact and management: Black locust is a tree with continual growth during the vegetative season. Nearly all leaflets of young shoots may be attacked by gall midges. Attacked leaflets dry up and fall off precociously after larval exit. The aesthetic value of damaged trees and shrubs is reduced. Monitoring may be achieved by visual detection of galls on trees and shrubs. Until now, insecticides have not been used to reduce populations. Natural enemies have been found in Europe, but surprisingly not in North America where O. robiniae is native. The endoparasitoid Platygaster robiniae (Hymenoptera: Proctotrupoidea: Platygastridae) has potential to reduce the gall midge population future biological control. The fungus Beauveria bassiana (Entomophthoraceae) was found during a study of gall midge larvae. Mechanical control is effected by cutting off infested parts with galls and burning is also recommended. Selected references Bathon H (2007) Die Robinien-Gallmücke Obolodiplosis robiniae (Haldeman) (Diptera: Cecidomyiidae) in Deutschland. Hessische Faunistische Briefe 26: 51–55. Duso C, Fontana P, Tirello P (2005) Diffusione in Italia e in Europa di Obolodiplosis robiniae (Haldeman), dittero cecidomiide neartici dannoso a Robinia pseudoacacia. Informatore fitopatologico 5: 30–33. Glavendekić M, Roques A, Mihajlović L (2009) An ALARM Case study: The Rapid Colonization of an Introduced Tree, Black Locust by an Invasive North-American Midge and Its Parasitoids. In: Settele J et al. (Eds) Atlas of Biodiversity Risks - from Europe to the globe, from stories to maps. Pensoft, Sofia & Moscow (www.pensoftonline.net/alarm-atlas-info, in press. 914 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.27 – Aedes albopictus (Skuse, 1894) - Asian tiger mosquito (Diptera, Culicidae) Alain Roques Description and biological cycle: Mosquito with black adult body and conspicuous white stripes on body and legs. Males (Photo left) have plumose antennae, whereas females have sparse short hairs (Photo right- female on human skin). Females are active during the day and are bloodfeeders on vertebrates, including humans. Adult flight range is limited (200–400 m). Long-distance dispersal (eggs, larvae) mediated by human activity. Average fecundity of 150–250 eggs, up to 5 generations per year. Eggs are laid in the water in tree holes and domestic containers. Breeding populations are present from March to November; overwintering at egg stage. Eggs are resistant to desiccation and cold. Larvae require only 6 mm of water depth to complete life cycle. Areas at risk have mean winter temperatures higher than 0 °C, at least 500 mm precipitation and a warm-month mean temperature higher than 20 °C. Native habitat (EUNIS code): G- Woodland and forest habitats and other wooded land; J6: Waste deposits. Typically breeds in tree holes and others small water collections surrounded by vegetation but also in peri-domestic containers filled with water. Habitat occupied in invaded range (EUNIS code): J6: Waste deposits. Mostly opportunistic container breeder capable of using any type of artificial water container, especially discarded tyres, but also saucers under flower pots, bird baths, tin cans and plastic buckets. It can establish in non-urbanised areas lacking artificial containers. Native range: Southeast Asia. Introduced range: Continuous spread all over the world since the late 1970s. First recorded in Europe in 1979 in Albania. Then, accelerated expansion was observed in Southern Europe since 2000, mostly along the Mediterranean Coast (Map). Some spots were detected in northwestern Europe, where it was tentatively eradicated. Also introduced in the Middle East, Africa, the Caribbean and North and South America. Credit: Susan Ellis, Bugwood.org (left), James Gathany, Centres for Disease Control and Prevention, USA (right) Factsheets for 80 representative alien species. Chapter 14 915 Pathways: Stowaway. Passive transport as dormant eggs via the international tire trade (due to the rainwater retained in the tires when stored outside), aircraft, boats and terrestrial vehicles and as larvae in “lucky bamboo” Dracaena spp., and other phytotelmata* shipped with standing water. Impact and management: Interspecific larval competition causes displacement of native mosquito species. Considerable health risk and economic costs result from the biting nuisance and the potential as vector for at least 22 arboviruses (including dengue, chikungunya, Ross River, West Nile virus, Japanese encephalitis, eastern equine encephalitis), avian plasmodia and dog heartworm filariasis Dirofilaria. For monitoring, ovitraps are used: artificial breeding containers (e.g., tyres) baited with frozen CO2 from dry ice. Mechanical control: removal of discarded tyres. All sources of standing water should be emptied every 3 d in areas at risk; water reserves that cannot be dumped can be treated with a spoonful of vegetable oil to suffocate mosquito larvae. To control larvae, spray water with derivates of Bacillus thuringiensis israelensis or larval growth inhibitors (diflubenzuron). To control adults, spray with deltamethrine. To control adults, spray with deltamethrine. Cyclopoid copepod predators (e.g., Macrocyclops, Mesocyclops) can be used for container-breeding larvae, and fishes and dragonflies in other situations. Selected references Eritja R, Escosa R, Lucientes J, Marquès E, Roiz D, Ruiz S (2005) Worldwide invasion of vector mosquitoes: present European distribution and challenges for Spain. Biological Invasions 7: 87–97. Gratz NG (2004) Critical review of the vector status of Aedes albopictus. Medical and Veterinary Entomology 18: 215–227. Urbanelli S, Bellini R, Carrieri M, Sallicandro P, Celli G (2000) Population structure of Aedes albopictus (Skuse): the mosquito which is colonizing Mediterranean countries. Heredity 84: 331–337. 916 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.28 – Ceratitis capitata (Wiedemann, 1824) - Mediterranean fruit fly (Diptera, Tephritidae) Alain Roques Description and biological cycle: Small fly, 4–5 mm long. Adults with yellowish body, brown abdomen and legs, and yellow-banded wings (Photo). Larva 6–8 mm long at maturity, elongate, cream coloured, and of cylindrical maggot shape. Phytophagous on a wide range of temperate and subtropical fruits. Adult flight range up to 20 km but winds can carry flying adults over longer distances; intercontinental dispersal (eggs, larvae) via infested fruits transported by humans. Before reaching sexual maturation, adults feed 6–8 d on fruit juices. Females lay up to 22 eggs per day and 300–800 eggs during lifetime, under the skin of a fruit just beginning to ripen. Under tropical conditions, overall life cycle is completed in 21–30 d. Adults may survive for up to six months. Native habitat (EUNIS code): G- Woodland and forest habitats and other wooded land. Habitat occupied in invaded range (EUNIS code): I- Regularly or recently cultivated agricultural, horticultural and domestic habitats; I1- Arable land and market gardens. Native range: Tropical Eastern Africa. Introduced range: Observed in Europe since 1873 in Italy. Present all over southern Europe (Map); regularly observed but not established in other parts of Europe; global warming may allow populations to establish at higher latitudes than at present. It has also been introduced in Africa, Middle East, Central and South America, the Caribbean, Hawaii, Australia. Eradicated in USA except Hawaii. Credit: Michel Martinez/ INRA Factsheets for 80 representative alien species. Chapter 14 917 Pathways: Imported with fruit trade but also with passengers transporting infested fruits during trips. Impact and management: Probably the most important fruit fly pest, inducing large damage in fruit crops, especially citrus fruits and peach. Fly damage results from both oviposition in fruit, feeding by the larvae, and decomposition of plant tissue by invading secondary microorganisms (bacteria, fungi) that cause fruit rot. Their presence often requires host crops to undergo quarantine treatments, other disinfestation procedures or certification of fly-free areas. The costs of such activities and phytosanitary regulatory compliance can be significant and definitely affect global trade. To ensure early detection, traps baited with chemical attractants (especially trimedlure) can be used. Larvae can be killed by soaking, freezing, cooking or pureeing infested fruits. Fruits can be bagged to prevent egg laying. Field sanitation needs to destroy all unmarketable and infested fruits; harvesting fruit weekly also reduces food sources by keeping the quantity of ripe fruit on the trees to a minimum. Chemical sprays are not completely effective. It is better to use foliage baits combining a source of protein with an insecticide to attract both males and females. Biological control involves use of sterile insects and release of parasitoids. Selected references Copeland RS, Wharton RA, Luke Q, De Meyer M (2002) Indigenous Hosts of Ceratitis capitata (Diptera: Tephritidae) in Kenya. Annals of the Entomological Society of America 95: 672–694. Liebhold AM, Work TT, McCullough DG, Cavey JF (2006) Airline Baggage as a Pathway for Alien Insect Species Invading the United States. American Entomologist 52: 48–56. Malacrida AR, Marinoni F, Torti C, et al (1998) Genetic aspects of the worldwide colonization process of Ceratitis capitata. Heredity 89: 501–507. 918 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.29 – Rhagoletis completa Cresson, 1929 - Walnut husk fly (Diptera: Tephritidae) Marc Kenis Description and biological cycle: Adults are typical tephritid yellow-orange flies with black stripes on wings, 4–8 mm long (Photo left). Adults fly in summer, and can live up to 40 days. Breeds in the husks of walnuts (Juglans spp.). Eggs are laid under the skin of the host fruit and hatch after 3–7 days. Larvae feed for 2–5 weeks, usually in the mesocarp (Photo right- larva emerged from a walnut). Overwinters in its puparium in the soil. There is only one generation per year. Native habitat (EUNIS code): G1 - Broadleaved deciduous woodlands; I- Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): G1 Broadleaved deciduous woodland; I1- Arable land and market gardens. I2- Cultivated areas of gardens and parks. Native range: North America. Introduced range: First found in Switzerland and Italy in the 1980s, from where it spread to several European countries, including France, Germany, Slovenia and Croatia (Map). Its distribution is closely linked to that of walnut species. Pathways: The main mode of dispersal is probably human-mediated transport through larval infested fruits. Adults can fly, but only a short distance. Impact and management: Attacked walnut fruits are pitted by oviposition punctures around which discolouration usually occurs. Larvae usually feed on the mesocarp, but Credit: Erwin Mani, eppo.org Factsheets for 80 representative alien species. Chapter 14 919 at high density, larvae also damage the pericarp and the nut itself. Walnuts attacked by the fly become unfit for sale, because of the discolouration of the nut. Walnut husk fly is a major pest of walnut in the USA. Since its introduction into Europe, populations are increasing, and severe damage has been observed, with up to 100% of harvested walnuts infested in some orchards. Various chemical treatments are effective against R. completa. Attacked fruits should be removed and destroyed before the larva emerges. Covering the soil under trees may prevent the larvae from entering the soil and pupating. Yellow sticky traps baited with ammonia can be used as a monitoring method, but are not efficient as a control method. Selected references Duso C, Lago G dal (2006) Life cycle, phenology and economic importance of the walnut husk fly Rhagoletis completa Cresson (Diptera: Tephritidae) in northern Italy. Annales de la Société Entomologique de France 42: 245–254. Mani E, Merz B, Brunetti R, Schaub L, Jermini M, Schwaller F (1994) Zum Auftreten der beiden amerikanischen Fruchtfliegenarten Rhagoletis completa Cresson und Rhagoletis indifferens Curran in der Schweiz (Diptera: Tephritidae). Mitteilungen der Schweizerischen Entomologischen Gesellschaft 67: 177–182. Romani M (1998) Gravi attacchi di Rhagoletis completa nei noceti lombardi. Informatore Fitopatologico 48: 13–16. 920 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.30 – Adelges nordmannianae (Eckstein, 1890) (= Dreyfusia nordmannianae, = D. nüsslini Börner) - Silver fir woolly aphid (Hemiptera, Adelgidae) Hans Peter Ravn Description and biological cycle: Winged female adult aphids (emigrants) from the primary host have a body length of 1.1–2.3 mm and wing span of about 4,6 mm. They are greenish just after the moult, turning darker. Winged female adults from the secondary host (remigrants) are grey-green and have a body length of 0.8–1.2 mm. Body length of parthenogenetic females is 0.7–1.5 mm; they are black-brown or black-violet. The body is covered with wax-wool. The small, turtle-shaped nymphs usually have only a peripheral fringe of wax round their body. In the native range, the aphid has a two-year life cycle with sexual reproduction on a primary host, Oriental Spruce, Picea orientalis (or P. omorica), and a parthenogenetic reproduction on a secondary host, Caucasian fir, Abies nordmanniana, which is replaced by European silver fir, Abies alba, in the introduced range in Europe. On Picea orientalis, aphids induce a 6- 8 mm gall (Photo left) growing from the short side-branches and also consisting of thickened needles. Galls are not induced on Abies species. The overwintering stage on the secondary host is 2nd-3rd instar larvae, situated on the shoot axis of the previous year’s shoot. In early spring, they develop into egg-producing females. Each female produces 110–500 eggs in a rosette-shaped heap. After hatching, young larvae will move to the new shoots and suck either on the new shoot axis or on the needles (Photo right). Some of the larvae develop into winged adults that will try to re-migrate to P. orientalis. Needle-feeding larvae and some shoot feeding larvae develop into females producing 10–30 eggs, from which the larvae move to the shoot axis for overwintering. Credit: L. Goudzwaard Factsheets for 80 representative alien species. Chapter 14 921 Native habitat (EUNIS code): G3 - Coniferous woodland. Habitat occupied in invaded range (EUNIS code): G3 - Coniferous woodland; I2 Cultivated areas of gardens and parks; X24 - Domestic gardens of city and town centres; i.e. Christmas tree plantations (A. nordmannianae) in forests and on arable field land. Native range: Mountain areas of Caucasus, Northeastern Turkey (Pontus) and Crimea. Introduced range: First detected in 1840 in Germany, then spreading to stands of native Abies alba throughout the distribution range of this tree species in Europe (Map). However native Norway spruce, Picea abies, has not been accepted as a primary host. Therefore, the sexual life cycle rarely occurs in Europe. Pathways: Forestry plantations of exotic conifers and trade of ornamental trees. Impact and management: Aphid suction curls needles on new twigs. At severe attack levels, honeydew production may cause formation of sooty mould, loss of needles and even death of the leading shoot. Attacks are more abundant and severe in the region of introduction than in the region of origin. Silver fir woolly aphid has developed into the severest pest problem for Christmas tree production in Europe. Silver fir woolly aphid is responsible for the major part of insecticides used in Christmas trees. Selected references Eichhorn O (1991) On the generation cycle of Dreyfusia nordmannianae Eckst. (Hom., Adelgidae). Journal of Applied Entomology 112: 217–219. Schneider-Orelli O, Schaeferrer C, Wiesmann R (1929) Untersuchungen über die Weisstannenlaus Dreyfusia nüsslini C.B. in der Schweiz. Mitteilungen der schweizerischen Cebtralanstalt für das forstliche Versuchswesen 15: 191–242. Varty IW (1956) Adelges Insects of Silver Firs. Edinburgh: Her Majesty’s Stationery Office. 75 pp. 922 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.31 – Bemisia tabaci (Gennadius, 1889) - Cotton whitefly (Hemiptera, Aleyrodidae) Alain Roques Description and biological cycle: Small, about 1 mm long, sap-sucking whitefly with two pairs of white wings and a white to light yellow body, covered with waxy powdery material (Photo left). Larvae also sap-sucking, feeding on > 900 plant species. This taxon corresponds to a species complex that comprises a large number of genetically variable populations, some of which are discernible owing to distinct phenotypes. Well-studied B.tabaci populations that have been differentiated are referred to as races or biotypes. The B biotype is a particularly aggressive variant. One female produces 80–300 eggs per lifetime. Unmated females produce parthenogenetically only male progeny. Development needs 15–70 d from egg to adult depending on temperature (10–32 °C, 27 °C is optimal), while 11–15 generations per year are possible (Photo right- empty exuviae). Native habitat (EUNIS code): Unknown. Habitat occupied in invaded range (EUNIS code): I- Regularly or recently cultivated agricultural, horticultural and domestic habitats; I1- Arable land and market gardens; glasshouses. Native range: Asia -Pacific region. Cotton whitefly appears to be a species complex. Recent genetic data indicate as many as ten morphologically indistinguishable species indigenous to the Asia-Pacific region. Introduced range: Widely spread in the last 15 years. Reported at present from all continents; present in the field in most of Southern Europe but restricted to glasshouses in Western, Central and Northern Europe (Map). Apparently eradicated in Finland, Ireland and the United Kingdom. Pathways : Intercontinental dispersal of eggs, nymphs and adults occurs with plant trade. Directional adult flight is limited but winds may carry flying adults over long distances due to their small size. Credit: Jean Yves Rasplus (left), Jean Claude Streito (right) Factsheets for 80 representative alien species. Chapter 14 923 Impact and management: Heavy infestations cause important yield losses, ranging from 20–100% depending on the crop and season, to both field and glasshouse agricultural crops and ornamental plants. Three types of damage are observed. Direct feeding damage by adults and larvae may reduce host vigour and growth, cause chlorosis and uneven ripening, and induce physiological disorders. Indirect damage results from accumulation of honeydew produced by nymphs, which serves as a substrate for the growth of black sooty mould on leaves and fruit. The mould reduces photosynthesis and lessens market value of the plant or yields it unmarketable. Finally, it is the most important vector of plant viruses worldwide. As vector of over 100 plant viruses, a small population of whiteflies is sufficient to cause considerable damage. Avoid importations from infested areas. Sequential plantings, avoiding the establishment of affected crops near infested fields, can be used. Adult activity and abundance can be monitored using yellow sticky traps. Chemical control: a number of insecticides provided effective control in the past, but resistance has developed rapidly. Biological control: the use of natural enemies such as chalcids (e.g., Encarsia formosa, Eretmocerus spp.) and the entomopathogenic fungus Verticillium lecanii is moderately efficient, but cannot sufficiently decrease infestations to stop virus transmission. Selected references De Barro PJ (2005) Genetic structure of the whitefly Bemisia tabaci in the Asia-Pacific region revealed using microsatellite markers. Molecular Ecology 14: 3695–3718. Martin JH, Rapisarda C, Mifsud D (2000) The whiteflies (Hemiptera: Aleyrodidae) of Europe and the Mediterranean Basin. Bulletin of Entomological Research 90: 407–448. Moya A, Guirao P, Cifuentes D, Beitia F and Cenis JL (2001) Genetic diversity Genetic diversity of Iberian populations of Bemisia tabaci (Hemiptera: Aleyrodidae) based on random amplified polymorphic DNA-polymerase chain reaction. Molecular Ecology 10: 891–897. 924 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.32 – Trialeurodes vaporariorum (Westwood, 1856) - Glasshouse whitefly (Hemiptera, Aleyrodidae) Alain Roques Description and biological cycle: Adult small, white to pale yellow, about 1mm long; the wings held relatively flat when at rest and coated with powdery wax (Photo- adult male). While single whitefly can be difficult to see, large numbers clustered on the underside of leaves are very obvious. They tend to fly rapidly when the plant is disturbed. The female may lay more than 500 eggs during its 3–6 weeks- long life. Eggs are laid in a circle on smooth leaves; on hairy leaves, they are more dispersed and less regularly situated. The eggs hatch about 9 days after egg-laying at 21°C. Newly emerged nymphs are mobile for a short period before settling to feed, their stylets inserted in leaf tissue, passing through three instars. Then, they stop feeding, moult and remain in a pupa for about 18 days. Reproduction is essentially parthenogenetic. Overwintering occurs at all instars. In northern climates, this whitefly usually lives in glasshouses on wild plants, or in summer on adjacent plants outside. Further south, adults may also overwinter on wild plants growing outdoors if the climatic conditions are not too severe. Reproduction occurs throughout the year when conditions are favourable, with several generations overlapping. Under optimum conditions at 21–24°C, the development from egg to adult takes about 3–4 weeks. Highly polyphagous, this species is capable of attacking 249 genera of plants. It attacks mainly vegetables, especially tomatoes, cucumbers and several other economic plants especially when they are grown in greenhouses. It can also be found on a wide selection of ornamentals, with a prediliction for Asteraceae, and of weeds, including sow thistles (Sonchus spp.), milkweed (Euphorbia peplus), and mallows (Malva spp.). Native habitat (EUNIS code): Unknown. Habitat occupied in invaded range (EUNIS code): J100- glasshouses; I- Regularly or recently cultivated agricultural, horticultural and domestic habitats; I1- Arable land and market gardens. Native range: Central America, essentially tropical and subtropical. Credit: LNPV/ Montpellier Station Factsheets for 80 representative alien species. Chapter 14 925 Introduced range: First recorded in Europe in Great Britain in 1856. Nowadays, present in the major part of Western, Central and Southern Europe (Map). In cold regions, this whitefly is found only in heated glasshouses whilst it may occur outdoors in southern Europe on both wild and cultivated plants. Pathways: Intercontinental dispersal of eggs, nymphs and adults occurs with plant trade. Directional adult flight is limited but winds may carry flying adults over long distances due to their small size. Impact and management: A major pest in glasshouses. The whitefly is responsible for very severe damage on vegetables through both sap sucking, and the production of honeydew and the consequent formation of sooty moulds. Up to 2,000 nymphs may be found on a single bean leaf, each being capable of producing 20 drops of honeydew in an hour. Affected tomatoes cannot be sold. The species may also transmit viruses.A certain resistence to synthetic insecticides has been observed, particularly amongst parthenogenetic strains. Populations are controlled by the action of entomophagous species such as fungi, ladybirds, Neuropterae, and hymenopteran chalcids. Biological control is widely used in commercial glasshouses, by introduction of a small endoparasitic wasp, Encarsia Formosa Gahan, which attacks and kills the whiteflies. Other biological control agents becoming available to gardeners include a small black ladybird, Delphastus sp., and a small predatory bug, Macrolophus sp. Selected references Kirk AA, Lacey LA, Roditakis, N Brown JK (1993) The status of Bemisia tabaci (Hom.: Aleyrodidae), Trialeurodes vaporariorum (Hom.: Aleyrodidae) and their natural enemies in Crete. Entomophaga 38: 405–410. Martin JH, Rapisarda C, Mifsud D (2000) The whiteflies (Hemiptera: Aleyrodidae) of Europe and the Mediterranean Basin. Bulletin of Entomological Research 90: 407–448. Van Dorst HJM, Huijberts N, Bos L (1983) Yellows of glasshouse vegetables, transmitted by Trialeurodes vaporariorum. European Journal of Plant Pathology 89: 171–184. 926 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.33 – Aphis gossypii Glover, 1877 - Cotton aphid, melon aphid (Hemiptera, Aphididae) Alain Roques Description and biological cycle: Small aphid, about 2 mm long, phloem-feeding with two virginiparous forms. Winged and wingless, highly variable in colour from yellowish green to partly black; immature stages pale yellow to pale green (Photo- wingless female and immatures). Highly polyphagous species, a major pest of cultivated plants in the families Cucurbitaceae, Rutaceae, Malvaceae and of Citrus trees. Flight range of winged adults is limited. Long-range dispersal of eggs, immature stages and adults is human-mediated with the transport of infested plant material. In Europe, it reproduces by apomictic parthenogenesis, and can produce nearly sixty generations a year. The optimal temperature is 21–27 °C. Viviparous females produce 70–80 offspring at a rate of 4.3 per day. Developmental periods of immature stages vary from 21 d at 10°C to 4 d at 30°C. Good resistance to summer heat. Dry weather conditions are favourable and heavy rainfall decreases population sizes. Native habitat (EUNIS code): Unknown. Habitat occupied in invaded range (EUNIS code): I1- Arable land and market gardens; I2- Cultivated areas of gardens and parks; J100- glasshouses. Native range: Unknown. Introduced range: Found in tropical and temperate regions throughout the world except northern areas. Common in Africa, Australia, Brazil, East Indies, Mexico and Hawaii, Present in most of Europe (Map) but it can develop outdoors only in Southern Europe, surviving in glasshouses in Northern Europe. Pathways: Passive transport with plant trade including vegetables, fruits, cut flowers, ornamental plants, bonsai, and nursery stock. Credit: Jérôme Carletto Factsheets for 80 representative alien species. Chapter 14 927 Impact and management: Economically important because nymphs and adults feed on the underside of leaves, or on growing tip of vines, sucking nutrients from the plant. The foliage may become chlorotic and die prematurely. Feeding also causes distortion and leaf curling, hindering photosynthetic capacity of the plant. In addition, honeydew production fosters growth of sooty moulds, resulting in a decrease of fruit/vegetable quantity and quality. Vector of crinkle, mosaic, rosette, Tristeza citrus fruit (CTV) and other virus diseases. Impact is especially high on courgette, melon, cucumber, aubergine, strawberry, cotton, mallow and citrus. Resistance has arisen to many pesticides. Insecticides should be used sparingly and in conjunction with other non-chemical control methods. Parasitoid aphidiid wasps (e.g., Aphidius colemanior, Lysiphlebus testaceipes), aphelinid wasps (e.g., Aphelinus gossypii), predatory midges (e.g., Aphidoletes aphidimyza), predatory anthocorid bugs (e.g., Anthocoris spp.), predatory coccinelids, and entomopathogenic fungi (e.g., Neozygites fresenii) are efficient and available for biocontrol in glasshouse crops. Selected references Fuller SJ, Chavigny P, Lapchin L, Vanlerberghe-Masutti F (1999) Variation in clonal diversity in glasshouse infestations of the aphid, Aphis gossypii Glover in southern France. Molecular Ecology 8: 1867–77. Margaritopoulos JT, Tzortzi M, Zarpas KD, Tsitsipis JA, Blackman RL (2006) Morphological discrimination of Aphis gossypii (Hemiptera: Aphididae) populations feeding on Compositae. Bulletin of Entomological Research 96: 153–165. Martin B, Rahbé Y, Fereres A (2003) Blockage of stylet tips as the mechanism of resistance to virus transmission by Aphis gossypii in melon lines bearing the Vat gene. Annals of Applied Biology 142: 245–250. 928 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.34 – Cinara curvipes (Patch, 1912) - Bow-legged fir aphid (Hemiptera, Aphididae) Olivera Petrović-Obradović Description and biological cycle: Wingless viviparous females are pearlike, 4–6 mm long. Body is dark brown, almost black, glossy, with two long white wax lines, extending dorsally from head to end of abdomen (Photo). Cornicles are short, on an oval sclerotised plate. Cauda are short and rounded. Rostrum is very long and may exceed the length of the body. Winged viviparous females are somewhat finer, with well developed wings. Monoecious species (host alternation does not occur) on Abies spp., Cedrus athlantica and Cedrus deodora. In America, develops a sexual generation in autumn, but in Europe, males have not been observed and it seems that it has anholocyclic* development. Native habitat (EUNIS code): G- Woodland and forest habitats and other wooded land. Habitat occupied in invaded range (EUNIS code): G3- Coniferous woodland; G3F- Highly artificial coniferous woodland; G5- Lines of trees, small anthropogenic woodlands, recently felled woodland, early-stage woodland; I2- Cultivated areas of gardens and parks; X24- Domestic gardens of city and town centres; X25- Domestic gardens of villages and urban peripheries. Native range: North America and Mexico Introduced range: First recorded in Germany in 2000, later found in Serbia (2001), Switzerland (2007), Slovakia (2007) and Czech Republic (2008). Credit: Armin Spürgin Factsheets for 80 representative alien species. Chapter 14 929 Pathways: Introduced with infested coniferous host plants. Spread in Europe continues by transport of the host plants and by active and passive flight of winged viviparous females. Impact and management: Economically one of the most important aphids as it is a pest of many crops (peach, potato, tobacco, sugar beet, vegetables, ornamental plants). Also, among aphids, it is the most efficient vector of plant viruses, transmitting more than 100 nonpersistent and many important persistent viruses, including Potato leaf roll (PLRV), Bean leaf roll (BLRV), Pea enation mosaic (PEMV) and Beet yellow net (BYNV). For monitoring flight activity, yellow water traps and suction traps are used. For chemical control, since resistance to insecticide is easily developed, only a few new insecticides are sufficiently effective. Many predators act as biological controls in colonies of the pest, especialy Coccinelidae, Syrphidae, Chrysopidae, Miridae and Cecidomyiidae (Aphidoletes aphidomyza Rond.). A very rich parasitoid complex includes 18 species of Aphidiidae wasp. Two of them are used in control in glasshouses: Aphididus colemani Vier. and Aphidius ervi Hal. Selected references Balachowsky A, Mesnil L (1935) Les insectes nuisibles aux plantes cultivées. Paris, France: Mery L. 1921 pp. Blackman RL, Eastop VF (2000) Aphids on the World’s Crops - an Identification and Information Guide. 2nd edn. Chichester UK: John Wiley & Sons. 476 pp. Theobald FV (1926)The plant lice or Aphididae of Great Britain, Vol I., London, UK: Headley Brothers. 372pp. 930 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.35 – Macrosiphum euphorbiae (Thomas, 1878) - Potato aphid (Hemiptera, Aphididae) Olivera Petrović-Obradović Description and biological cycle: Medium-sized to large aphid (1.7–3.5 mm), spindle-shaped, green (Photo) or pink. Adults are rather shiny and larvae have a light dusting of greyish-white wax. Mainly anholocyclic*, usually with only winged and wingless forms present in colonies. Sexual morphs are produced on primary host (Rosa spp.) in North America and only rarely in other parts of the world. Highly polyphagous on secondary hosts, feeding on plant species in more than 20 different plant families. In Europe, develops usually without sexual generation. During winter, regularly found in glasshouses. Native habitat (EUNIS code): Unknown. Habitat occupied in invaded range (EUNIS code): : I1 - Arable land and market gardens; I2- Cultivated areas of gardens and parks; glasshouses; X7 - Intensively-farmed crops interspersed with strips of spontaneous vegetation. X24 - Domestic gardens of city and town centres; X25: Domestic gardens of villages and urban peripheries. Native range: North America. Credit: Rémi Coutin/ OPIE Factsheets for 80 representative alien species. Chapter 14 931 Introduced range: Cosmopolitan species. In Europe, first found in 1917 in Great Britain. Then, the potato aphid colonized most of Europe (Map). Pathways: Trade of ornamentals. Impact and management: Serious pest of many crops (potato, vegetable, flowers), causing direct damage by sucking nutrients and indirect damage as a vector of viruses. This aphid can transmit more than 40 non-persistent viruses and five persistent viruses (potato leaf roll, beet yellow net, bean leaf roll, zucchini yellow mosaic and sweet potato leaf-speckling virus). Monitoring can be effected using yellow water traps and suction traps. Chemical control involves use of selective insecticide and is often both necessary and effective. Many specific predators and parasitoids can be used for biological control, epecially in glasshouses. Selected references Blackman RL, Eastop VF (2000) Aphids on the World’s Crops - an Identification and Information Guide. 2nd edn. Chichester UK: John Wiley & Sons. 476 pp. Eastop VF (1958) The history of Macrosiphum euphorbiae (Thomas) in Europe. The Entomologist 91: 198–201. 932 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.36 – Myzocallis walshii (Monell, 1879) (Hemiptera, Aphididae) Ejup Çota Description and biological cycle: Small (1.5–2.0 mm), delicate, usually yellowish aphid with a knobbed cauda and bilobed anal plate. Nymphs usually have capitate dorsal hairs. Adult viviparous females (viviparae) are all alate. The life cycle is monoecious and holocyclic. Not antattended. Alate viviparae of M. (Agriomyzus) castanicola have a distinct dark medial stripe on head and thorax, black spots on abdomen, dark siphunculi* and dark 2nd antennal segment. The dark pigmentation is less distinct in spring forms. Sides of pronotum and mesonotum of both species bear a black band extending from the eye to the base of hind wings. The late-summer form of M. (Lineomyzocallis) walshii has a broad foreground band of black pigment from the costal vein in the forewing, extending well past the stigma to the wing apex (Photo left- alate viviparous female of summer form; right- ovipara in aautumn). Mainly associated with Quercus rubra, the American red oak, but attacks other oaks of North American origin (Q. coccinea, Q. palustris) and one native species (Q. robur). Over-winters in the egg stage. Eggs hatch in the spring and give rise to the first of several asexual generations in which winged (alate) parthenogenic females give rise to wingless (apterous) nymphs that develop into alate parthenogenic females. In late fall, the sexual generation begins with production of apterous females (oviparae) and alate males. When mature and mated, the oviparae lay from 4–6 eggs/female in cracks and crevices among the bark, shortly before the leaves begin to fall. Credit: Jan Havelka Factsheets for 80 representative alien species. Chapter 14 933 Native habitat (EUNIS code): G- Woodland and forest habitats and other wooded land. Habitat occupied in invaded range (EUNIS code): I1- Arable land and market gardens; I2 - Cultivated areas of gardens and parks. Native range: North America Introduced range: Myzocallis walshii was detected for the first time in Europe in 1988 (France), and subsequently in several other European countries (Switzerland, Spain, Andorra, Italy, Belgium and Germany- Map). Pathways: Accidental introduction with trade of ornamental plants. Impact and management: Monitoring can be carried out using yellow sticky traps. As a mechanical means these exert limited control on populations. A number of aphicides can be used for chemical control as well as biological control agents, such as Aphidoletes spp, and Aphidius spp. Selected references Hullé M, Renoust M, Turpeau E (1998) New aphid species detected by permanent aerial sampling programmes in France. In: Nieto Nafria JM, Dixon AFG (Eds) Aphids in Natural and Managed Ecosystems. León, Spain: Universidad de León, Secretariado de Publicaciones. 365–369. Remaudière G (1989) Découverte en France de l’espèce américaine Myzocallis (Lineomyzocallis) walshii (Monell) (Hom.Aphididae). Revue Francaise d’Entomologie: 14: 172. 934 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.37 – Myzus persicae (Sulzer, 1776) - Peach potato aphid (Hemiptera, Aphididae) Olivera Petrović-Obradović Description and biological cycle: Small to medium-sized aphid (1.2–2.1 mm), yellow-green, grey-green, pink or red, not shiny. The aphid on tobacco is usually red, as well as specimens kept in cold conditions. Winged forms have a black central dorsal patch on the abdomen. Both winged and wingless forms are present in colonies (Photo left- Colony on tomato). Situated on the underside of leaves, aphids excrete honeydew. They curl leaves of peach in spring (Photo right) and migrate on to many secondary hosts in summer. Many generations can be produced a year with very rapid development under favorable conditions. Highly polyphagous species. The sexual phase occurs on the primary host, Prunus persica. In glasshouses and where outdoor conditions are good, parthenogenetic development occurs all year round on secondary hosts. Secondary hosts are very numerous, feeding on plants in over 40 different families. Populations colonizing tobacco are recognized as subspecies Myzus persicae nicotianae (Blackman, 1987). Native habitat (EUNIS code): Unknown. Habitat occupied in invaded range (EUNIS code): I1- Arable land and market gardens; I2- Cultivated areas of gardens and parks; glasshouses; X7- Intensively-farmed crops interspersed with strips of spontaneous vegetation. X24- Domestic gardens of city and town centres; X25- Domestic gardens of villages and urban peripheries. Native range: Unknown, possibly Asia. Credit: Rémi Coutin/ OPIE Factsheets for 80 representative alien species. Chapter 14 935 Introduced range: Cosmopolitan. Present since a very long time (since <1758) in Europe. Probably introduced repeatedly with infested plants. Pathways: Plant trade. Impact and management: Economically one of the most important aphids as it is a pest of many crops (peach, potato, tobacco, sugar beet, vegetables, ornamental plants). Also, among aphids, it is the most efficient vector of plant viruses, transmitting more than 100 nonpersistent and many important persistent viruses, including Potato leaf roll (PLRV), Bean leaf roll (BLRV), Pea enation mosaic (PEMV) and Beet yellow net (BYNV). For monitoring flight activity, yellow water traps and suction traps are used. For chemical control, since resistance to insecticide is easily developed, only a few new insecticides are sufficiently effective. Many predators act as biological controls in colonies of the pest, especialy Coccinelidae, Syrphidae, Chrysopidae, Miridae and Cecidomyiidae (Aphidoletes aphidomyza Rond.). A very rich parasitoid complex includes 18 species of Aphidiidae wasp. Two of them are used in control in glasshouses: Aphididus colemani Vier. and Aphidius ervi Hal. Selected references Balachowsky A, Mesnil L (1935) Les insectes nuisibles aux plantes cultivées. Paris, France: Mery L. 1921 pp. Blackman RL, Eastop VF (2000) Aphids on the World’s Crops - an Identification and Information Guide. 2nd edn. Chichester UK: John Wiley & Sons. 476 pp. Theobald FV (1926)The plant lice or Aphididae of Great Britain, Vol I., London, UK: Headley Brothers. 372pp. 936 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.38 – Prociphilus fraxinifolii Riley ex Riley & Monell, 1879 - Woolly Ash Aphid (Hemiptera, Aphididae) Olivera Petrović-Obradović Description and biological cycle: Aphids 2.0–2.5 mm long, soft bodied, with well-developed wax glands, producing enormous quantities of wax rendering a snow-white appearance. Siphuncular* pores are absent. Both winged (Photo right) and wingless forms have yellow-green to pale green bodies. Compact colonies inhabit curled leaves at twig tips throughout the vegetative period (Photo left). Host plant is red ash (Fraxinus pennsylvanica) and some other American species of Fraxinus. In North America, P. fraxinifolii is holocyclic* but overwinters as parthenogenetic females in Europe. Host alternation does not occur. Native habitat (EUNIS code): G- Woodland and forest habitats and other wooded land. Habitat occupied in invaded range (EUNIS code): G1- Broadleaved decidous woodland; G5- Lines of trees, small anthropogenic woodlands, recently felled woodland, early-stage woodland; I2- Cultivated areas of gardens and parks; X24- Domestic gardens of city and town centres; X25- Domestic gardens of villages and urban peripheries. Native range: North America and Mexico. Introduced range: First recorded in Europe in 2003 in Hungary, then in Serbia and Bulgaria (Map). Arrived three hundred years after introduction of its host plants, the American species of Fraxinus. Also introduced into Chile and South Africa. Credit: Claude Pilon Factsheets for 80 representative alien species. Chapter 14 937 Pathways: Trade of ornamental plants. Impact and management: Very destructive aphid because deformation of curled leaves and twigs make trees much less attractive. Colonies also occur on ash roots, where overwintering occurs in Europe. Key pest in nursery production of Fraxinus. For monitoring, it is important to make inspections of ash in early spring and to use systemic insecticides as soon as colonies appear. Biological control involves Aphelinus prociphili, the parasitoid in North America; no parasitoids are found in Europe. The natural enemy complex fails to keep plant damage below an acceptable level. Selected references Petrović-Obradović O, Tomanović Ž, Poljaković-Pajnik L, Vučetić A (2007) An invasive species of aphid, Prociphilus fraxinifolii (Hemiptera, Aphididae, Eriosomatinae), found in Serbia. Archives of Biological Sciences, Belgrade 59: 9–10. Remaudière G, Ripka G (2003) Arrival in Europe (Budapest, Hungary) of American ash aphid, Prociphilus (Meliarhizophagus) fraxinifolii (Hemiptera, Aphididae, Eriosomatinae, Pemphigini). Revue Francaise d’Entomologie 25: 152. 938 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.39 – Toxoptera citricida (Kirkaldy, 1906) - Tropical citrus aphid, oriental black citrus aphid, brown citrus aphid (Hemiptera, Aphididae) Ejup Çota Description and biological cycle: Aphid with medium-sized body, 1.5–2.4 mm long, shiny, reddish-brown to black. Alates (Photo left) can be identified, using a pocket lens, by the wholly black third antennal segment which is succeeded by a pale fourth segment. Median nervure of forewings normally forked twice. Siphunculi* of alates about l/6 body length and strongly sculptured, while cauda rather bulbously rounded at apex. Apterous forms should be examined microscopically to observe the very long, fine and erect hairs on the legs and body margins. Siphunculi* as in alates but relatively shorter (Photo- right). Cauda thick and bluntly rounded at the apex. Immature stages brown A useful character to distinguish T. citricidus from T. aurantii is that a distinct scraping sound produced by disturbed colonies of the latter, audible up to 45 cm away from the leaf, while T. citricidus are silent. Females are parthenogenetic and a single generation develops in 6–8 days. Tropical citrus aphids attack solely Citrus spp. Reproductive potential depends on the abundance of plant sap. About 30 generations are produced annually, depending on temperature. Winged females give rise to new infestations. Dark-brown to black colonies develop on young growths and are usually visited by ants. Native habitat (EUNIS code): I - Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): I1- Arable land and market gardens; I2 - Cultivated areas of gardens and parks; glasshouses. Credit: Aphidweb.com Factsheets for 80 representative alien species. Chapter 14 939 Native range: Occurs predominantly in humid tropical regions and presumably originated in south-east Asia. Introduced range: First detected in Madeira in 1994, later observed in continental Portugal and Spain (Map). Pathways: T. citricidus can spread locally by flight, but is very unlikely to be introduced into the region by natural means. Introduction occurs on potted plants and associated transportation materials. Impact and management: Growth of shoots is greatly impaired and they become distorted; leaves become brittle and wrinkled and curl downwards. Attacked flowers fail to open or do so abortively since the ovaries are deformed. T. citricidus is an efficient vector of important virus diseases of citrus: citrus tristeza closterovirus, stem-pitting and seedling yellows strains. Control measures are intended to prevent damage to young shoots and fruits, and especially to suppress the formation of alates. Young trees are treated preventively with systemic insecticides. Many natural enemies are known (e.g. predators and entomopathogenic fungi). Some are being considered for use in integrated control programmes. Selected references Aguiar AMF, Fernandez A, Ilharco FA (1994) On the sudden appearance and spread of the black citrus aphid Toxoptera citricidus (Kirkaldy) (Homoptera Aphidoidea) on the island of Madeira. Bocagiana 168: 1–7. Doncaster JP, Eastop JF (1956) The tropical citrus aphid. FAO Plant Protection Bulletin 4: 109–110. Ilharco FA, Sousa-Silva CR, Alvarez Alvarez A (2005) First report on Toxoptera citricidus (Kirkaldy) in Spain and continental Portugal (Homoptera, Aphidoidea). Agronomia Lusitana 51: 19–21. 940 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.40 – Scaphoideus titanus Ball, 1932 - Vine leafhopper (Hemiptera: Cicadellidae) Wolfgang Rabitsch Description and biological cycle: Small leafhopper, average adult body size 5 mm, ochrebrown to mottled dark brown (Photo), nymphs yellowish-white with two dark brown spots on abdomen. Females lay clusters of 10–12 eggs in late summer in crevices in the bark of one- or two-year-old grapevine wood. Eggs overwinter, and development from first instar to adult takes 35–40 days. Adults of the new generation appear in late spring with one generation per year. Larvae and adults live ampelophagously*, ie. monophagous on grapevine (Vitis vinifera). Native habitat (EUNIS code): I1- Arable land and market gardens. Habitat occupied in invaded range (EUNIS code): I1- Arable land and market gardens. Native range: Nearctic species, originally present in the northeastern parts of USA and South Canada. Introduced range: Unintentionally introduced to south-western France, presumably in the 1950s (first record in 1958), from where it subsequently spread to neighbouring countries: Italy (1963), western and southern Switzerland (1968), Slovenia (1983), Croatia (1987), northern Spain (1995) and northern Portugal (1999). Later, the species extended its range Credit: Gernot Kunz Factsheets for 80 representative alien species. Chapter 14 941 north- and eastward and was also found in Serbia (2003), southern Austria (2004), Bulgaria (2006), and southwestern Hungary (2006) (Map) Pathways: Studies of the genetic structure of European populations has revealed longdistance translocations, most probably of eggs with grapevine propagation material. In addition the species spreads naturally, probably favoured by current climatic conditions. Impact and management: Scaphoideus titanus is vector of “Flavescence dorée” (FD), a serious disease of grapevine, caused by the phytoplasma Candidatus Phytoplasma vitis, belonging to the elm yellow group 16Sr-V subgroups C and D. Larvae acquire phytoplasmas by feeding on infected plants and after 4–5 weeks (in the third larval stage), they are able to transmit the disease to healthy plants. FD phytoplasma is reported from France, Italy, Portugal, Serbia, Slovenia, Spain, and Switzerland. Productivity of infected plants is greatly reduced by discolouration (yellowing) and desiccation. Selected references Bertin S, Guglielmino CR, Karam N, Gomulski LM, Malacrida A, Gasperi G (2006) Diffusion of the Nearctic leafhopper Scaphoides titanus Ball in Europe: a consequence of human trading activity. Genetica 131: 275–285. Seljak G (2008) Distribution of Scaphoides titanus in Slovenia: its new significance after the first occurrence of grapevine “flavescence dorée”. Bulletin of Insectology 61: 201–202. Steffek R, Reisenzein H, Zeisner N (2007) Analysis of the pest risk from Grapevine flavescence dorée phytoplasma to Austrian viticulture. EPPO Bulletin 37: 191–203. 942 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.41 – Pulvinaria regalis Canard, 1968 - Horse chestnut scale (Hemiptera, Coccidae) Marc Kenis Description and biological cycle: Adult scales are dark brown, flattish, round and about 4 mm in diameter. They are found on the edge of white egg masses on bark of trunks and branches (Photo). Nymphs on foliage are pale yellow and oval in shape. At outbreak density, P. regalis can be recognized by their white egg masses covering the trunk and the main branches in spring and in summer. This scale is univoltine. Crawlers hatch in May and June and move to leaves of the host tree. Nymphs feed on leaves until September/October and then migrate to twigs where they overwinter in the third instar. In spring, newly emerged females first feed, then move to the main branches and the trunk to lay eggs. Crawlers can be transported by wind. Host plant transportation is probably another important mode of dispersal. Although P. regalis is known to attack a high number of woody plants, heavy infestations occur mainly on Aesculus, Tilia and Acer in urban and suburban areas or along roads. Native habitat (EUNIS code): Unknown. Habitat occupied in invaded range (EUNIS code): G1 - Broadleaved deciduous woodland; I2 - Cultivated areas of gardens and parks. Native range: Unknown Credit: Chris Malumphy Factsheets for 80 representative alien species. Chapter 14 943 Introduced range: First found in London in 1964 and subsequently observed in France, Belgium, Netherlands, Germany, Ireland, Switzerland, Austria and Denmark (Map). Pathways: Trade of ornamental plants. Impact and management: Horse-chestnut scale does not kill trees, but outbreaks have a considerable impact on growth, particularly of young trees. This scale also causes aesthetic damage to ornamental trees. Additionally, it produces high quantities of honeydew that may become a nuisance in urban areas. Occurrence and incidence of the scale in natural habitats is unclear, and its interaction with native fauna is not known. Use of insecticides is possible, but difficult in urban areas. In spring, egg masses on trunks and branches can be washed off with water using a high-pressure cleaner. On small plants, mature scales and their eggs can be scraped or wiped from the stems. Selected references Hippe C, Frey JE (1999) Biology of the horse chestnut scale, Pulvinaria regalis Canard (Hemiptera: Coccoidea: Coccidae), in Switzerland. Entomologica 33: 305–309. Jansen MGM (2000) The species of Pulvinaria in the Netherlands (Hemiptera: Coccidae). Entomologische Berichten 60: 1–11. Speight MR. The impact of leaf-feeding by nymphs of the horse chestnut scale, Pulvinaria regalis Canard (Hom.: Coccidae), on young host trees. Journal of Applied Entomology 112: 389–399. 944 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.42 – Leptoglossus occidentalis Heidemann, 1910 - Western Conifer Seed Bug (Heteroptera: Coreidae) Wolfgang Rabitsch Description and biological cycle: Large coreid true bug, reddish brown with a white zig-zag band on the forewing and a characteristic leaf-like dilation on the hind tibia (Photo), average size of adults 18 mm. Adults emerge from overwintering sites in late spring. Females lay up to 80 eggs in chains on conifer needles. Nymphs develop into new generation in late summer, one generation per year. Feeds on the young seeds or flowers of conifer species, with a preference for Pinaceae (Plinus spp., Pseudotsuga menziesii), but it was also observed on Picea, Cedrus, Abies and Juniperus. eptoglossus occidentalis overwinters in crevices or secret places under bark or other structures. Native habitat (EUNIS code): G3 - Coniferous woodland; G3F - Highly artificial coniferous plantations. Habitat occupied in invaded range (EUNIS code): G3 - Coniferous woodland; G3F Highly artificial coniferous plantations; I2 - Cultivated areas of gardens and parks. Native range: Presumed to be west of the Rocky Mountains in North America, from British Columbia to Mexico. Introduced range: Since the 1950s, the species spread eastward and reached the east coast of North America in the 1990s. First European records date from 1999 near Vicenza (northern Italy). Western conifer seed bug then spread rapidly in Europe and is known from Switzerland (2002), Slovenia, Spain (2003), Croatia, Hungary (2004), Austria (2005), Czech Republic, France, Germany, Serbia (2006), United Kingdom, Belgium, Netherlands, Slovak Credit:Wolfgang Rabitsch Factsheets for 80 representative alien species. Chapter 14 945 Republic, Poland (2007), Bulgaria, Montenegro and Greece (2008) (Map). In most countries, rapid within-country spread and increasing abundance has been observed. Recently, it was also introduced to Japan. Pathways: The species is capable of flying over long distances, but also is translocated as egg, nymph or adult with its host plant (conifers). Impact and management: Enters buildings in large numbers in autumn and so becomes a nuisance. Feeding on conifers causes reduction of seed fertility, and the species is regarded as pest in the native range. Although no economic impact has yet been measured in Europe, first observations tend to show that it may largely decrease the potential of regeneration of conifers in both seed orchards and natural pine stands. Mechanical exclusion is recommended to avoid public nuisance. Selected references Bernardinelli I, Zandigiacomo P (2001) Leptoglossus occidentalis Heidemann (Heteroptera, Coreidae): a conifer seed bug recently found in northern Italy. Journal of Forestry Science, 47: 56–58. Ishikawa T, Kikuhara Y (2009) Leptoglossus occidentalis Heidemann (Hemiptera: Coreidae), a presumable recent invader to Japan. Japanese Journal of Entomology (New Series) 12: 115– 116. Lis JA, Lis B, Gubernator J (2008) Will the invasive western conifer seed bug Leptoglossus occidentalis Heidemann (Hemiptera: Heteroptera: Coreidae) seize all of Europe? Zootaxa 1740: 66–68. 946 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.43 – Aspidiotus nerii (Bouché, 1833) (= A. hederae (Vallot, 1829)) - Oleander scale (Hemiptera, Diaspididae) Katalin Tuba and Ferenc Lakatos Description and biological cycle: Adult female covered with a scale that is 1.5–2.0 mm in diameter, nearly circular and flat, yellowish white with a yellow or gold central part. Female body bright yellow. Wings, legs, and eyes absent. Scale cover of male white, oval, translucent and smaller, and more elongate than female. Adult males winged. Highly polyphagous species; > 200 host species recorded including Nerium oleander, Acer spp., Olea europaea, Populus spp. Ribes spp. and Vitis vinifera. Attacks and can wholly cover leaves, bark and the fruits (Photo- Colony on a palm leaf). Reproduction is either sexual (A. nerii nerii) or parthenogenetic (A. nerii unisexualis). The sexual population has higher fecundity and faster development than the parthenogenetic one. There are two or three generations per year depending on climatic conditions. Development time is about 30–35 d influenced by the sex, temperature, humidity, and rainfall. Each female lays a total of 100–150 eggs under the scale of the female, where they develop. The settled female nymph moults twice, the males four times. Adult females remain under scale throughout their life. Males became winged after the second moult, but their flight ability is limited. Male lifespan is only a few hours. Native habitat (EUNIS code): G- Woodland and forest habitats and other wooded land; I- Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): G1 - Broadleaved deciduous woodland; I8: Part of agricultural land and artificial landscapes. Native range: Afrotropical region. Credit: Claude Bénassy/INRA Factsheets for 80 representative alien species. Chapter 14 947 Introduced range: Nowadays with worldwide distribution, occurring especially in tropical and subtropical zones. First record in Europe from Italy in 1829. At present, the oleander scale is observed in most of Europe (Map) but in cold areas, occurs only in greenhouses and indoors. Pathways: Trade of ornamental plants. The wide geographical distribution of this pest is primarily due to human activities. The first instar, the only nymphal stage with legs, is active and responsible for short-distance dispersal. Impact and management: : Aspidiotus nerii is particularly important where aesthetic value of the crop is high, like cut flowers, ornamentals in gardens, nurseries, under glass and indoors. After heavy infestation in olive orchards, quality and quantity reduces. Economically important on other mediterranean forest tree species too. Both adults and nymphs cause damage. Mechanical, chemical and biological control is used to reduce damage. Nowadays, biological control plays the most important role, especially in greenhouses. Natural enemies have already adapted to the species: parasitoids, e. g. Aphytis chilensis (attacking nymphs and adults in Europe, the Middle East, Africa, America and Australia), Encarsia aurantii (South America), and also predators, e.g. Aleurodothrips fasciapennis (attacking eggs, nymphs, adults), Chilocorus circumdatus (attacking nymphs and adults) and Hemisarcoptes coccophagus (attacking all stages, except eggs). Selected references Alexandrakis V, Bénassy C (1981) Experiment in biological control on olive in Crete using Aphytis melinus DeBach (Hym. Aphelinidae), parasite of Aspidiotus nerii Bouché (Hom. Diaspididae). Acta Oecologica, Oecologia Applicata 2: 13–25. Gerson U, Hazan A (1979) A biosystematic study of Aspidiotus nerii Bouché (Homoptera: Diaspididae), with the description of one new species. Journal of Natural History 13: 275 – 284. Longo S, Marotta S, Pellizzari G, Russo A, Tranfaglia A (1995) An annotated list of the scale insects (Homoptera: Coccoidea) of Italy. Israel Journal of Entomology 29: 113–130. 948 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.44 – Diaspidiotus perniciosus (Comstock, 1881) - San José scale (Hemiptera: Diaspididae) Marc Kenis Description and biological cycle: Female is grey, circular and about 2 mm in diameter (Photo left- female with scale turned upside down to show the body colour). Male has only forewings present. Larvae highly variable, depending on stage and sex, white to black, round to elongate, and fixed scales or little mobile yellow organisms (Photo left- young nymphs pointed with arrow). In Europe, two to four generations per year, depending on climatic conditions. In cold climates, the winter is usually spent in the first larval stage. Development starts in early spring. Females become adult after the second moult and gradually increase in size. Males have two larval instars, a prepupal and a pupal stage. Males are winged and fly, but lack mouthparts, whereas females remain stationary and feed. Females are viviparous and produce about 100 larvae, 30–40 d after copulation. First instar crawls to find new host tissues. Then, it attaches itself and secretes a waxy substance forming the scale cover. Diaspidiotus perniciosus is a highly polyphagous species. The main hosts are apples, peaches, pears (Photo right- change in epidermis colour of a damaged pear), plums and Rubus. Native habitat (EUNIS code): G1- Broadleaved deciduous woodland; I- Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): G1- Broadleaved deciduous woodland; I1- Arable land and market gardens; I2- Cultivated areas of gardens and parks; J100glasshouses. Native range: East Asia. Introduced range: First introduced into California in the 19th century, from whence it spread to the whole North American continent. It is also present in many Asian, African and South American countries, as well as in New Zealand and Australia. First discovered in Hun- Credit: Rémi Coutin/ OPIE (left), Claude Bénassy/ INRA (right) Factsheets for 80 representative alien species. Chapter 14 949 gary and Italy in 1928 and now present in most European countries (Map) although in many of them, it has not yet reached its potential distribution. Pathways: International spread probably occurs through human-mediated transport of planting material of trees and shrubs, or fruits. The crawling first instar larvae are the main dispersal stage and can be carried a few kilometres by wind. Adult males, but not females, can also be carried by wind. Impact and Management: Various young host plant tissues are affected. Attacks occur on wood mainly, but also on leaves and fruits. The insect injects toxic saliva, causing localized discolouration. San José scale can kill a young tree in 2–3 years in the absence of control. Older trees are weakened and growth is reduced, as well as fruit production and quality. This is considered a serious orchard pest in several European countries, reducing growth, fruit quality and marketability. Mineral oil can be applied in winter against overwintering stages, whereas pesticides during the growing season. Sex pheromone traps are used to monitor the timing and level of attack. Biological control with the aphelinid wasp Encarsia perniciosi has been carried out in several regions, with varying degrees of success. Selected references Kosztarab M, Kozár F (1988) Scale insects of Central Europe. Budapest, Hungary: Dr. W. Junk Publishers. 456 pp. Mani E, Schwaller F, Baroffio C, Hippe C (1995) Die San-José-Schildlaus in der deutschen Schweiz: Wo stehen wir heute? Schweizerische Zeitschrift für Obst- und Weinbau 131: 264–267. Melis A (1943) Contributo alla conoscenza dell’Aspidiotus pernicious. Redia 29: 1–170. 950 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.45 – Pseudaulacaspis pentagona (Targioni-Tozzetti, 1886) - Mulberry scale (Hemiptera: Diaspididae) Katia Trencheva Description and biological cycle: Adult female cover is convex, circular white (Photo- Encrustation on a branch of peach); shed skins usually sub-central, yellowish orange. Male cover smaller, felted, white, elongate, sometimes with slight median carina completely enclosing developing male; shed skin white, sometimes tinged with yellow. Body of adult female light yellow, eggs of male white, that of females yellow or pink. Mulberry scale reproduces sexually, with two to five generations per year depending on climate. It has three generations per year in Bulgaria, where overwintering occurs as a fertile female. In the USA, it can also overwinter as adult females or as eggs. Females each lay about 100 eggs, which hatch 3–5 days after oviposition. Native habitat (EUNIS code): G1- Broadleaved deciduous woodland; I- Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): G1- Broadleaved deciduous woodland; I1- Arable land and market gardens; I2- Cultivated areas of gardens and parks; J100glasshouses. Native range: East Asia. Introduced range: Accidentally introduced to Italy in 1886, then recorded in most countries of Southern and Central Europe and in the Atlantic islands (Map). Nowadays, it is one of the best examples of the northward expansion of insects in central Europe where it has colonized both cultivated and natural habitats, primarily occurring on bark and fruit of various trees and shrubs, occasionally on leaves. Also introduced in Africa, Australia, New Zealand, southern Central America and many Pacific Islands. Credit: ACTA/ INRA Factsheets for 80 representative alien species. Chapter 14 951 Pathways: Trade in plants and plant products. Mulberry scale can also be transported by wind and by birds. Impact and management: Most serious problems are caused in areas of accidental introduction in the absence of its natural regulators. The efficiency of natural enemies is reduced in urban areas by pollution. Consequently, P. pentagona can cause severe damage to ornamental plants in towns and cities. It is particularly destructive on flowering cherry, mulberry, peach and other deciduous fruit trees. In Europe, outbreaks have occurred in many countries, including Hungary, Switzerland, France, Greece and Bulgaria. Scale insects are difficult to control because the waxy or cottony covering serves as a protective barrier to traditional contact insecticides. However, a pest management program that incorporates natural, mechanical, and/or chemical controls should provide satisfactory control of most scales. Pheromone traps are used for detection in newly infested regions, especially in Europe. Colour and sticky traps have also been developed to monitor the flight and dispersal of males. Natural enemies, particularly the parasitoid Encarsia berlesei, can be effective control agents. Chemical control may not be advisable for orchards, since the natural enemies of P. pentagona can be killed, causing local outbreaks. Selected references Kozár F, Sheble DAF, Fowjhan MA (1995) Study on the further spread of Pseudaulacaspis pentagona (Homoptera: Coccoidea: Diaspididae) in Central Europe. Israel Journal of Entomology 29: 161–164. Targioni Tozzetti A (1886) (1885) Sull’insetto che danneggia I gelsi. Rivista di bachicoltura 18: 1–3. Watson GW (2002) Pseudaulacaspis pentagona. In: Arthropods of Economic Importance. Diaspididae of the world. World Biodiversity database. http://nlbif.eti.uva.nl/bis/diaspididae. php. 952 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.46 – Metcalfa pruinosa (Say, 1830) - Citrus Flatid Planthopper (Hemiptera: Flatidae) Milka M. Glavendekić Description and biological cycle: Adult usually 5.5–8 mm long and 2–3 mm wide. Adults vary from brown to grey. Forewings and the body are covered with a soft white powder, giving them a bluish tone (Photo left). Larvae white, less than twice as long as wide, when mature about 4 mm long (Photo right). The species is univoltine. Highly polyphagous, recorded on 330 woody and herbaceous plant species in 78 plant families in Europe. Eggs laid singly in splits under bark of host plant during late summer and early autumn, where they overwinter and hatch during spring of the following year between late May and early June to mid-July. First adults of the new generation appear from mid-July and live until early November. Native habitat (EUNIS code): F5 - semi-arid and subtropical habitats Habitat occupied in invaded range (EUNIS code): G1 - Broadleaved deciduous woodland; I1 - Arable land and market gardens; I2 - Cultivated areas of gardens and parks; X16 Land sparsely wooded with mixed broadleaved and coniferous trees; X25 - Domestic gardens of villages and urban peripheries. Native range: Eastern North America. Introduced range: Since it was first recorded in north-eastern Italy in 1979, citrus flatid planthopper has spread in the Mediterranean region, as well as to Central- and South-East Europe (Map). Pathways: Trade appears to be the most likely pathway for introduction, on imported commodities such as nursery stock, both ornamentals and vegetables from infested areas. Credit: LNPV/ Montpellier Station Factsheets for 80 representative alien species. Chapter 14 953 Spread mostly passive through eggs laid into the bark of plants. Adults can also spread occasionally, attached to commodities or via transport by human activity. Impact and management: Considered as one of the most prolific pests for its ability to infest a wide variety of plant species in agricultural, forest and urban ecosystems. Metcalfa pruinosa sucks the sap from small diameter stems, but the damage is usually minor. There is evidence in Italy that M. pruinosa is infected by phytoplasma*, which could induce diseases on fruit trees. Oviposition injuries sometimes kill seedlings. Buds with deposited eggs could be frozen during winter. The most severe damage in Europe is caused by the secretion of honeydew which is colonized by sooty-mould fungus thus hindering photosynthetic capacity of the plant. High population could have a nuisance effect on tourism in some places. Mechanical control is effective on young plants by pruning and destroying shoots that contain oviposition punctures. Chemical control is possible in juvenile stages, but less effective against adults. Biological control includes the use of a dryinid wasp predator-parasitoid Neodryinus typhlocybae (Ashmead) (Hymenoptera: Dryinidae), which is introduced in Italy in 1994. Selected references Lucchi A (Ed) (2000) La Metcalfa negli ecosistemi italiana. Firenze: Agenzia Regionale per lo Sviluppo e l’Innovazione nel settore Agriocolo-forestale.163 pp. Orosz A, Der Z (2004) Beware of the spread of the leafhopper species Metcalfa pruinosa (Say, 1830). Novenyvedelem 40: 137–141. Trenchev G, Ivanova I, Nicolov P, Trencheva K (2007) Metcalfa pruinosa (Say, 1830) (Homoptera, Flatidae) a species new to the Bulgarian fauna. Plant Science 34: 195–198. 954 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.47 – Nysius huttoni White, 1878 - Wheat bug (Hemiptera: Lygaeidae) Wolfgang Rabitsch Description and biological cycle: Small lygaeid true bug, brown, with characteristic dorsal erect pubescence (Photo- adults mating). Average adult size 3.5 mm. Wing morphs comprise macropters, submacropters and brachypters; macropters are capable of flight over some distances; adults hibernate and two generations are developed per year in western Europe. Polyphytophagous species feeding on different weeds and crops (e.g. Brassica, Capsella, Chenopodium, Hieracium, Medicago, Polygonum, Rumex, Silene, Senecio, Trifolium and Triticum), attaining pest status in its native area. Native habitat (EUNIS code): B - Coastal habitats; E - Grassland and tall forb habitats; I - Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): I - Regularly or recently cultivated agricultural, horticultural and domestic habitats. In Europe, found in dry and warm sites, waste grounds, roadsides, sparsely vegetated sandy soils, and abandoned fields. The presence of acrocarpous mosses seems necessary. Native range: New Zealand. Introduced range: First recorded 2002 in the Netherlands, 2003 in Belgium and subsequently found at the French/Belgian border. In 2007 and 2009 it was found in Great Britain Credit: R. Kleukers Factsheets for 80 representative alien species. Chapter 14 955 (East Suffolk, North Essex) (Map). Nysius species are well known for their high abundances and effective dispersal strategies and it is expected that this species will further spread across Europe. Nysius huttoni currently is included in the EPPO Alert List. It has been intercepted on fruits in Australia and the United States. Pathways: Unintentional introduction, probably with shipments Impact and Management: N. huttoni is an economically important pest species in New Zealand, particularly when feeding on wheat and degrading gluten thus diminishing baking quality. Insecticides may be used, but no effective control treatment is known. Selected references Aukema B, Bruers JM, Viskens G (2005) A New Zealand endemic Nysius established in the Netherlands and Belgium (Heteroptera: Lygaeidae). Belgian Journal of Entomology 7: 37–43. He XZ, Wang Q, Carpenter A (2003) Thermal requirements for the development and reproduction of Nysius huttoni White (Heteroptera: Lygaeidae). Journal of Economic Entomology 96: 1119–1125. Smit JT, Reemer M, Aukema B (2007) Een invasie van de nieuw-zeelandse tarwewants Nysius huttoni in Nederland (Heteroptera: Lygaeidae). Nederlandse Faunistische Mededelingen 27: 51–70. 956 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.48 – Stictocephala bisonia Kopp & Yonke, 1977 - Buffalo treehopper (Hemiptera: Membracidae) Wolfgang Rabitsch Description and biological cycle: Medium-sized membracid treehopper, average adult body size 8–10 mm, adults bright green (Photo), larvae light grey-green with longitudinal row of spines and conical abdomen. Characteristic pronotum with buffalo-horn-like protrusions. Females lay batches of 5–12 yellow eggs in the bark of host plants by cutting small punctures. Overwinters as eggs, with adults of the new generation appearing in July, and one generation per year. Stictocephala bisonia is polyphagous on different herbs, shrubs and trees (e.g. Rosa, Malus, Pyrus, Prunus, Cornus, Crataegus, Populus, Ulmus, Coronilla, Melilotus, Solidago and Medicago). Native habitat (EUNIS code): F9 - Riverine and fen scrubs; I - Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): F9 - Riverine and fen scrubs; I - Regularly or recently cultivated agricultural, horticultural and domestic habitats; FA - Hedgerows; FB - Shrub plantations; E - Grassland and tall forb habitats. A preference for moist and wet riverine habitats, woody margins and tall herb stands, but it also can be found in dry meadows and agricultural land. Native range: North America, widely distributed from Canada to Mexico. Introduced range: accidentally introduced to Europe in the 20th century (first documented record: 1912 in former Hungary) and is present now in almost all countries (Map). It also spread to North Africa and Central Asia. Credit: Wolfgang Rabitsch Factsheets for 80 representative alien species. Chapter 14 957 Pathways: Long-distance dispersal of eggs with infected tree-seedlings; adults capable of flight. Impact and management: Females deposit their eggs in shrub and tree stems, which usually die above the insertion slits and also provide entry for pathogens; this may cause economic loss in orchards and vine cultures. Feeding activity on fruit trees and in vineyards may cause discolouration and wrinkling of the leaves and malignant growth of twigs; severely scarred branches should be pruned out. Application of insecticides usually is not effective; release of an introduced Nearctic parasitoid braconid (Polynema striaticorne) successfully controlled Stictocephala populations in Italy, where the braconid established and spread subsequently. Selected references Alma A, Arno C, Vidano C (1987) Particularities on Polynema striaticorne as egg parasite of Stictocephala bisonia (Rhynchota, Auchenorryncha). Proceedings 6th Auchenorryncha Meeting, Turin, Italy, 7–11 September 1987, 597–603. Fursov VN (1994) New data on Polynema striaticorne (Hymenoptera, Mymaridae) and its cicada host Stictocephala bisonia (Homoptera, Membracidae). Vestnik Zoologii 2: 12–19. Holzinger W, Kammerlander I, Nickel H (2003) The Auchenorrhyncha of Central Europe. Vol. 1. Leiden: Brill. 674 pp. Schedl W (1991) Invasion der Amerikanischen Büffelzikade (Stictocephala bisonia Kopp & Yonke, 1977) nach Österreich (Homoptera, Auchenorrhyncha, Membracidae). Anzeiger für Schädlingskunde 64: 9–13. 958 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.49 – Halyomorpha halys (Stål, 1855) - Brown Marmorated Stink Bug (Heteroptera: Pentatomidae) Wolfgang Rabitsch Description and biological cycle: Large pentatomid true bug, shield-shaped, mottled brown, size of adults 12–17 mm; females lay egg clutches of up to 25 eggs on the underside of leaves. The species hibernates as adult, producing one generation per year and is regarded as a polyphagous horticultural pest, observed on over 60 host plants, including fruit and shade trees and other woody ornamentals (e.g. Acer, Buddleja, Citrus, Malus, Morus, Paulownia, Prunus, Pyrus and Rosa), vegetables and as an agricultural pest on various leguminous crops (e.g. soybean). Native habitat (EUNIS code): G1 - Broadleaved deciduous woodland; I - Regularly or recently cultivated agricultural, horticultural and domestic habitats; X25 - Domestic gardens of villages and urban peripheries. Habitat occupied in invaded range (EUNIS code): I2 - Cultivated areas of gardens and parks; X25 - Domestic gardens of villages and urban peripheries. Native range: Asia (Japan, Korea, China, Taiwan). Introduced range: Introduced to the east coast of USA since 1996, where it subsequently spread along the east coast south to South Carolina. In 2007, there were several records including nymphs from the area around Zürich in Switzerland, indicating established populations in Europe. Credit: Beate Wermelinger Factsheets for 80 representative alien species. Chapter 14 959 Pathways: Long-distance dispersal occurs as stowaways with goods and plant material. Adults are able to fly some distance. Impact and management: Detrimental impacts on ornamentals (necrosis on leaves and fruits) has been observed. Brown marmorated stink bug is also known to be a vector of witches’ broom, a phytoplasma* disease of Paulownia tomentosa, an East Asian ornamental tree introduced to Central Europe in 1834, which only recently established and spread in urban-industrial areas. In Asia and America, Halyomorpha halys causes a nuisance in households when seeking hibernation sites in large numbers in autumn. Mechanical exclusion and chemical control are suggested to control indoors pest problems. Selected references Hoebeke ER, Carter ME (2003) Halyomorpha halys (Stål), (Heteroptera: Pentatomidae) a polyphagous plant pest from Asia newly detected in North America. Proceedings of the Entomological Society of Washington 105: 225–237. Wermelinger B, Wyniger D, Forster B (2008) First records of an invasive bug in Europe: Halyomorpha halys Stål (Heteroptera: Pentatomidae), a new pest on woody ornamentals and fruit trees. Mitteilungen der Schweizerischen entomologischen Gesellschaft 81: 1–8. Gyeltshen J, Bernon G, Hodges A. Brown Marmorated Stink Bug, Halyomorpha halys Stål (Insecta: Hemiptera: Pentatomidae). http://edis.ifas.ufl.edu/in623. 960 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.50 – Viteus vitifoliae (Fitch, 1855), Grape phylloxera (Hemiptera: Phylloxeridae) Marc Kenis Description and biological cycle: Small, globular, 1–1.8 mm long (Photo). Complex life cycle, which depends both on vine species or cultivars and the environment. Grape phylloxera has several generations per year and alternates between sexual and asexual generations, and between an aerial form causing galls on leaves, named gallicolae, and a root-feeding form, radicicolae. Viteus vitifoliae is a monophagous species restricted to some vine species of the genus Vitis. On the European grapevine, Vitis vinifera, the gallicolae form is rare, the radicicolae form persists parthenogenetically. Native habitat (EUNIS code): G - Woodland and forest habitats and other wooded land; I- Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): I- Regularly or recently cultivated agricultural, horticultural and domestic habitats. Native range: Eastern North America. Introduced range: It was first recorded in Europe in 1860 in France. Nowadays, most grapevine-growing areas in Europe are invaded, except Cyprus, parts of Greece and some re- Credit: LNPV/ Montpellier Station Factsheets for 80 representative alien species. Chapter 14 961 stricted areas in a few other countries (Map). The pest also spread to most major wine-producing areas, in Western North America, South America, South Africa and New Zealand. Pathways: This aphid, particularly the radicicolae form, does not spread easily by itself and is mainly carried on grapevine plants. Impact and management: The main damage is caused by radicicolae that feed on grapevine roots and are associated with secondary pathogens. In susceptible vine species and cultivars, they cause root rot, decrease plant vigour and, ultimately, kill the vines within 3–10 years. In the 19th century, V. vitifoliae causes huge damage to vineyards and has endangered the European wine industry. In France alone, it caused the destruction of 1.2 million ha. The problem was solved by grafting European cultivars on less susceptible American rootstocks. Grape phylloxera is still the target of phytosanitary regulations in Europe and elsewhere, because some pest-free areas remain, where susceptible grape cultivars are cultivated on their own roots. In the last 30 years, the level of damage has occasionally increased in various countries, with the appearance of new biotypes that overcome the resistance of certain rootstock cultivars. Selected references CABI (2007) Crop Protection Compendium. CD-ROM. Wallingford, UK: CAB International. Forneck A, Walker MA, Blaich R (2001) An in vitro assessment of phylloxera (Daktulosphaira vitifoliae Fitch) (Hom., Phylloxeridae) life cycle. Journal of Applied Entomology, 125: 443–447. Granett J, Walker MA, Kocsis L, Omer AD (2001) Biology and management of grape phylloxera. Annual Review of Entomology, 46: 387–412. 962 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.51 – Corythucha arcuata (Say, 1832) - Oak lace bug (Heteroptera: Tingidae) Wolfgang Rabitsch and Marc Kenis Description and biological cycle: Small tingid true bug, adults greyish to whitish with lacelike forewings, nymphs black with spines, adult body size 3 mm. Differs from C. ciliata in forewing pigmentation. Adults hibernate in bark crevices. Females lay eggs on the leaf underside. Development from egg to adult takes 30–45 days, 2–4 generations per year. The species feeds on deciduous Quercus species and was also reported on Castanea sativa, and occasionally on Acer, Malus and Rosa species. Native habitat (EUNIS code): G1 - Broadleaved deciduous woodland. Habitat occupied in invaded range (EUNIS code): G1 - Broadleaved deciduous woodland; I2 - Cultivated areas of gardens and parks. Native range: North America, east of the Rocky Mountains. Introduced range: First record for Europe dates back to spring 2000, when the species was found in northern Italy (Lombardy, Piemont). In 2002, it was found in Turkey (Western Anatolia) and 2003 in southern Switzerland (Tessin). It subsequently expanded its range in Credit: Joseph Berger, insectimages.org Factsheets for 80 representative alien species. Chapter 14 963 Italy and Switzerland and particularly in Turkey, where it has so far invaded an area of 28,000 km2. The oak lace bug is expected to further spread in Europe, where host plants occur. Pathways: Long-distance dispersal occurs with human activity (introduction with oak plants); adults can fly and be spread by wind. Impact and management: Corythucha arcuata may cause damage to the host trees (chlorotic discolouration, desiccation, premature leaf-fall and reduced photosynthetic activity). C. arcuata is not considered an important pest species in North America, likely due to control by natural enemies. Since these are missing in Europe, the environmental and economic impact in Europe is unknown, but potentially high. Selected references Bernardinelli I (2000) Distribution of Corythucha arcuata (Say) in northern Italy (Heteroptera, Tingidae). Redia 83: 157–162. Bernardinelli I (2006) Potential host plants of Corythucha arcuata (Het., Tingidae) in Europe: a laboratory study. Journal of Applied Entomology 130: 480–484. Forster B, Giacalone I, Moretti M, Dioli P, Wermelinger B (2005) The American oak lace bug Corythucha arcuata (Say) new to southern Switzerland. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 78: 317–323. 964 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.52 – Corythucha ciliata (Say, 1832) - Sycamore lace bug (Heteroptera: Tingidae) Wolfgang Rabitsch and Jean-Claude Streito Description and biological cycle: Small tingid true bug, adults whitish with lacelike forewings, nymphs black with spines, average adult body size 3.5 mm. Adults hibernate under loose bark of their host trees. Females lay up to 350 eggs. Development from egg to adult takes 45 d, 1–3 generations per year. Sycamore lace bug feeds on different Platanus species (Platanaceae). In the introduced range, it is regularly found on P. occidentalis and P. orientalis and their hybrid P. acerifolia, used as an ornamental tree in cities. Native habitat (EUNIS code): G1 - Broadleaved deciduous woodland. Habitat occupied in invaded range (EUNIS code): I2 - Cultivated areas of gardens and parks; X22 - Small city centre non-domestic gardens; X24 - Domestic gardens of city and town centres. Native range: North America, east of the Rocky Mountains. Introduced range: First record for Europe dates back to 1964, when found in northern Italy (Padua). It is now distributed over much of Europe with records in the west (Portugal), north (United Kingdom) and east (Russia) (Map). Sycamore lace bug was also introduced to China, Korea, Japan, Australia and Chile. Credit:Wolfgang Rabitsch Factsheets for 80 representative alien species. Chapter 14 965 Pathways: Long-distance dispersal occurs with human activity (transport via vehicles or clothes). The species flies well and also drifts passively by wind. Impact and management: Corythucha ciliata may cause damage to host trees (chlorotic discolouration, desiccation, premature leaf-fall, reduced photosynthetic activity, and prompt secondary infections by fungi and pathogens). In addition, the species may become a nuisance to people in parks and gardens, but usually impacts are of aesthetic value only. Chemical treatment with insecticides is not recommended. Selected references Arzone A (1986) Spreading and importance of Corythucha ciliata (Say) in Italy twenty years later. Bulletin WPRS, Section Régionale Ouest Paléarctique 9: 5–10. Servadei A (1966) Un Tingide nearctico comparso in Italia (Corythucha ciliata Say). Bollettino della Societa Entomologica Italiana 96: 94–96. Stehlík JL (1997) Corythucha ciliata (Say), a pest of plane trees, now also in the Czech Republic (Tingidae, Het.). Acta Musei Moraviae, Scientiae Naturales 81: 299–306. 966 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.53 – Cales noacki Howard, 1907 (Hymenoptera, Aphelinidae) Jean-Yves Rasplus Description and biological cycle: Traditionally, the genus Cales has been placed in the family Aphelinidae but this position has changed several times. Indeed, within aphelinids, the genus exhibits four-segmented tarsi, straight fore tibial spur, narrow forewing, flagellum with three funicular segments, and unsegmented clava*. Furthermore, the genus does not group with other Aphelinidae in a molecular phylogeny of chalcid wasps. Cales is the only genus of the subfamily Calesinae and comprises three described species that are parasitic on whiteflies, including Cales noacki, an endoparasitoid of woolly whitefly, Aleurothrixus floccosus (Maskell), a serious pest of citrus trees worldwide. C. noacki has a preference for second stage nymphs of this whitefly (Photo- C. noacki laying eggs on A. floccosus). However, C. noacki is not species specific and can develop on several species of whiteflies. At 26°C, the biological cycle of C. noacki takes about 21–22 days to be completed. C. noacki has always been regarded as a single species, however recent molecular analyses suggest that at least three distinct haplotypes coexist in the biocontrol citrus grove at Riverside. These species have different biology and environmental preferences. Native habitat (EUNIS code): G1-Broadleaved deciduous woodlands; I- Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): I- Regularly or recently cultivated agricultural, horticultural and domestic habitats Native range: South America. Credit: Jean Pierre Onillon / INRA Antibes Factsheets for 80 representative alien species. Chapter 14 967 Introduced range: Cales noacki strains released in Europe came from Chile, where they were sampled through biological control projects in the 1970s. The species was thus introduced in France (1971), Spain (1973), Portugal (1977), Italy (1980) and Greece (1991) for inoculative biological control of the woolly whitefly, Aleurothrixus floccosus (Map). Pathways: Intentionnal introduction for biological control. Impact and management: C. noacki is now established in Europe and has proved very effective against the woolly whitefly. The parasitoid has rapidly achieved high rates of parasitization (> 90%) resulting in substantial mortality to populations of the invading whitefly. In some parts of the Mediterranean area, the introduction of Cales noacki as a classical biological control agent against the woolly whitefly, may have led to the partial or complete displacement of native parasitoids of the non-target whitefly species Aleurotuba jelinekii. Selected references Onillon JC (1973) Possibilités de régulation des populations d’Aleurothrixus floccosus Mask. (Homopt. Aleurodidae) sur agrumes par Cales noacki How. (Hymenopt. Aphelinidae). Bulletin de l’Organisation Européenne et Méditerranéenne pour la Protection des Plantes 3: 17–26. Viggiani G (1994) Recent cases of interspecific competition between parasitoids of the family Aphelinidae (Hymenoptera: Chalcidoidea). Norwegian Journal of Agricultural Sciences Supplement 16: 353–359. Spicciarelli R, Tranfaglia A, Battaglia D, Torraco R (1996) Biological control of Aleurothrixus floccosus with Cales noacki. Informatore Agrario 52: 67–70. 968 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.54 – Lysiphlebus testaceipes (Cresson, 1880) (Hymenoptera, Braconidae) Jean-Yves Rasplus Description and biological cycle: Small (<3 mm) dark greenish to black braconid wasp. Female lays an egg on an aphid (Photo- female laying eggs on rose aphid, Macrosiphum rosae (L.)) the endoparasitoid larva grows and transforms it into a dead brown mummy. Development takes about 14 d; the wasp exits from a hole cut in the top of the mummy. Lysiphlebus testaceipes has a 2 days adult lifespan. Females produce 1.8 offspring per aphid patch, spending relatively shorter time on larger groups, while distributing a total of ca. 200 eggs across many patches (Tentelier et al., 2009). L. testaceipes is a generalist parasitoid, exhibiting extremely broad host range (> 200 aphid species on various plants that host notable aphid pests such Aphis, Brachycaudus, Myzus). In natural habitats, Aphidiinae are the main components of parasitoid communities controling aphid populations. Several species have been used for biological control in greenhouses. Native habitat (EUNIS code): E- Grassland and tall forb habitats; I- Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): I- Regularly or recently cultivated agricultural, horticultural and domestic habitats. Native range : Possibly Cuba. Introduced range: Introduced into Europe (France) from Cuba in the 1970s to control Aphis spiraecola Patch, a pest of Citrus (Stary et al., 1988). This braconid soon became Credit: Peter J. Bryant Factsheets for 80 representative alien species. Chapter 14 969 established and subsequently spread in the Mediterranean basin, shifting to other native aphid species, and reaching Italy in 1977, Spain (1982–1984), Portugal (1985) and Greece (2002) (Map). Pathways: Intentionnal introduction for biological control Impact and management: In Europe, L. testaceipes is mass-reared, sold and released to control Aphis gossypii Clover. This braconid is a very efficient parasitoid that can reduce host infestations. However, recent studies have clearly shown that L. testaceipes outcompetes indigenous Aphidiinae. For example, on T. aurantii, it may have displaced two congeneric parasitoid species, L. fabarum (Marshall) and L. confuses Tremblay & Eady (Tremblay 1984). Such collateral effects on local faunas need more studies to estimate better the impact of this species on the parasitoid community foodweb associated with aphids. Selected references Stary P, Lyon JP, Leclant F (1988) Biocontrol of aphids by the introduced Lysiphlebus testaceipes (Cress.) (Hym., Aphidiidae) in Mediterranean France. Journal of Applied Entomology 105: 74–87. Tremblay E (1984) The parasitoid complex (Hymenoptera: Ichneumonoidea) of Toxoptera aurantii (Homoptera: Aphidoidea) in the Mediterranean area. Entomophaga 29: 203–209. Tentelier C, Lacroix MN, Fauvergue X (2009) Inflexible wasps: the aphid parasitoid Lysiphlebus testaceipes does not track multiple changes in habitat profitability. Animal Behaviour 77: 95–100. 970 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.55 – Dryocosmus kuriphilus (Yasumatsu, 1951) - Chestnut gall wasp (Hymenoptera, Cynipidae) Milka M. Glavendekić and Alain Roques Description and biological cycle: Female black, 2.5–3 mm long (Photo- left). Legs, antennal scapus and pedicel, apex of clypeus and mandibles yellow brown. Antennae 14-segmented with apical segments not expanded into a club. Head finely sculptured. Scutum, mesopleuron and gaster highly polished, smooth. Propodeum* with three distinct longitudinal carinae*; propodeum and pronotum strongly sculptured. Scutum* with two notaulices* converging posteriorly. Radial cell of forewing “open”. Eggs oval, milky white, 0.1–0.2 mm long, long-stalked. Full-grown larva 2.5 mm long, milky white, without eyes and legs. Pupa 2.5 mm long, dark brown. Monophagous on Castanea spp. and their hybrids, attacking Castanea crenata Sieb. et Zucc. (Japanese chestnut), C. dentata (Marsh.) (American chestnut), C. mollissima Blume (Chinese chestnut), C. sativa Mill. (European chestnut) and C. seguinii Dode (in China). Univoltine and thelytokous* parthenogenetic species. Adults emerge from galls from end of May until end of July. Lifetime short (about 10 d). Females lay 3–5 eggs per cluster inside buds. Each female can lay > 100 eggs. Some buds contain 20–30 eggs. Embryonic development lasts 30–40 d. Early instar larvae overwinter inside chestnut buds. At the time of bud burst in spring, gall wasps induce formation of a 5–20 mm diameter green (Photo right) or rose-coloured gall, containing 1–7 or 8 small cells where early instars develop. Galls develop in mid April on new shoots, leaves and twigs. Larvae feed 20–30 d within the galls before pupation from mid-May to mid-July. Native habitat: (EUNIS code): G- Woodland and forest habitats and other wooded land. Habitat occupied in invaded range: G1 - Broadleaved deciduous woodland; G5 - Lines of trees, small anthropogenic woodlands, recently felled woodland, early-stage woodland and coppice. Chestnut forests and monocultures within coppice deciduous forests, chestnut orchards, lines of chestnut trees, gardens, ornamental cultures. Native range: Asia (China). Introduced range: In Europe, first recorded in 2002 near Cuneo, Italy, then from Slovenia (2006), France (Alpes- Maritimes, 2007) Switzerland (2009), Hungary (2009) and elsewhere in Italy (Map). Also introduced in Japan, Korea, and USA. Credit: Milka Glavendekić Factsheets for 80 representative alien species. Chapter 14 971 Pathways: Passive transport with plants for planting and cut branches. Dispersal at a local scale is realized by adult flight. Impact and management: Chestnut gall wasp is the most severe worldwide insect pest on chestnuts. It disrupts twig growth and reduces fruiting, causing yield reduction up to 70%. Severe infestations may result in the decline and death of young chestnut trees and debilitate chestnut forests. Rapid recruitment of generalist parasitoids shared with oak cynipids suggests that chestnut gall wasp may have a negative impact on native cynipids through apparent competition. An effective measure would be to prohibit import of chestnut cut branches (or young plants) for grafting from China, Japan and America. In Italy, France and Slovenia, chestnut nurseries should be inspected annually to ensure trade of safe young plants. Infestations in small chestnut orchards may be reduced by pruning and destroying infested shoots. Treatment with systemic insecticides during the growing season at the place of production can be applied but is insufficient for control; as yet there are no efficient chemicals to control this pest. Torymus sinensis Kamijo was already introduced as a biological control agent in Italy from Japan. Several cultivars, prevalently belonging to the species Castanea crenata and its hybrids, are considered resistant; among them, Bouche de Bétizac (C. sativa x C. crenata) was reported. Larvae were found also in this cultivar but they die just at shooting time and do not develop galls. There are also new resistant Japanese and Korean chestnut cultivars. Selected references Csóka G, Wittmann F, Melika G (2009) The oriental sweet chestnut gall wasp (Dryocosmus kuriphilus Yasumatsu, 1951) in Hungary. Novenyvdelem 45: 359–360. Forster B, Castellazzi T, Colombi L, Furst E, Marazzi C, et al. (2009) First record of the chestnut gall wasp Dryocosmus kuriphilus (Yasumatsu) (Hymenoptera, Cynipidae) in Southern Switzerland. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 82: 271–279. Graziosi I, Santi F (2008) Chestnut gall wasp (Dryocosmus kuriphilus): spreading in Italy and new records in Bologna province. Bulletin of Insectology 61: 343–348. 972 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.56 – Ophelimus maskelli (Ashmead, 1900) - Eucalyptus gall wasp (Hymenoptera, Eulophidae) Jean-Yves Rasplus Description and biological cycle: Gall making wasp mostly attacking Eucalyptus species, with a preference for E. camaldulensis and E. tereticornis (= umbellata Smith.), but with a hostplant range encompassing 14 species of Eucalyptus belonging to three sections. Females can lay up to 100 eggs, usually in batches. They oviposit preferentially on the immature leaf blade close to the petiole, in the lower canopy. Each egg induces a small (about 1 mm diameter) pimple-like gall visible on both side of the leaf; galls are well separated (Photo). Gall density can reach 36 per cm2 in Israel. O. maskelli has three generations per year in Israel and probably also in other Mediterranean countries (Protasov et al 2007). Native habitat (EUNIS code): G1 Broadleaved deciduous woodland. Habitat occupied in invaded range (EUNIS code): G1 - Broadleaved deciduous woodland; G5 - Lines of trees, small anthropogenic woodlands, recently felled woodland, early-stage woodland and coppice; I2 - Cultivated areas of gardens and parks; X24 Domestic gardens of city and town centres. Native range: Australia. Introduced range: O. maskelli, erroneously reported as O. eucalypti (Gahan), has been introduced in the Mediterranean area since at least 2000. O. maskelli was first recorded in Italy (2000) (Arzone & Alma, 2000), then in Greece (2002), Spain (2003), UK (2004), France (2005), Portugal (2006) (Map). It also occurs in Israel and Turkey. Pathways: Trade of ornamental plants. Credit: Alain Roques Factsheets for 80 representative alien species. Chapter 14 973 Impact and management: Heavy leaf galling can lead to premature shedding of leaves and dessication of large parts of tree crowns, resulting a depreciated value. Some Eucalyptus are particularly affected, such as E. camaldulensis planted for forestry in the Mediterranean region and the Middle East. Repeated attacks can lead to loss of foliage from terminal branches. Heavy galling can damage two thirds of the entire leaf volume and results in premature shedding of the leaves. The impact of the wasp on E. camaldulensis is consequently serious and heavily infested trees exhibit strong desiccation of their crowns and premature leaf drop. Interestingly, O. maskelli has similar host range to Leptocybe invasa Fisher & LaSalle, another Eulophid wasp developing on Eucalyptus and introduced in the Mediterranean basin from Australia. Adults emerge en masse in large clouds that cause nuisance and health problems to humans (Protasov et al 2007). Closterocerus chamaeleon (Hymenoptera Eulophidae) has been used to successfully control O. maskelli in Israel and also in Portugal. This wasp exhibits several biological traits that favour population increase and spread, such as thelytoky*, high fecundity, short generation time, and high longevity that favours wind dispersion (Branco et al, 2009). Selected references Arzone A, Alma A (2000) A gall Eulophid of Eucalyptus in Italy. Informatore Fitopatologico 50: 43–46. Branco M, Boavida C, Durand N, Franco JC, Mendel Z (2009) Presence of the Eucalyptus gall wasp Ophelimus maskelli and its parasitoid Closterocerus chamaeleon in Portugal: First record, geographic distribution and host preference. Phytoparasitica 37: 51–54. Protasov A, La Salle J, Blumberg D, Brand D, Saphir N et al. (2007) Biology, revised taxonomy and impact on host plants of Ophelimus maskelli, an invasive gall inducer on eucalyptus spp. in the Mediterranean area. Phytoparasitica 35: 50–76. 974 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.57 – Lasius neglectus Van Loon, Boomsma & Andrásfalvy, 1990 - Garden Ant (Hymenoptera, Formicidae) Wolfgang Rabitsch Description and biological cycle: Ant with workers lacking erect hairs on the scape and extensor of hind tibiae, and with reduced mandibular dentition (Photo). Female immediately recognizable within European Lasius by its comparatively reduced size and proportionately smaller gaster, as compared with the thorax. Male the smallest of European Lasius. Sister species of Lasius turcicus Santschi. Ants are active throughout the entire day and aphid tending lasts for 24 h/d, from late April to late October, imposing a non-negligible cost on the energetic budget of individual trees. Nuptial flight seems to be absent. Nests are very difficult to delimit as they may coalesce and integrate a supercolony occupying enormous areas, as large as 16 ha. In urban areas, colonies are fragmented but may occupy a single tree. Finding many dealate* queens (polygyny) in a nest is a key diagnostic of this species, the single polygynous European Lasius (s.str.). The number of queens depends on colony size, but estimated from queens found under stones, was about 35,500 in the Seva supercolony. Using soil cores, worker number for that population in May 2002 was estimated as 1.12 x 108. This species is truly unicolonial., with inter-nest and inter-population relationships showing a typical unicolonial trait of reduced level of aggressiveness. Areas occupied furnish a wide array of possible nesting sites: under stones, temporal refuges with aphids at the base of herbs, amid rubbish, etc. The expansion process of a colony seems to be much helped by the progressive urbanization of lots. This development usually implies the cutting and burning of all natural vegetation but trees. The planting of grass and continuous irrigation that follows favours ant establishment. Native habitat (EUNIS code): Unknown. Habitat occupied in invaded range (EUNIS code): I2 - Cultivated areas of gardens and park; X24- Domestic gardens of city and town centres. Populations live in a wide range of conditions, from strictly urban habitats, streets with heavy traffic to semi-urban sites, mildly Credit: Jordan Wagenknecht, antbase.fr Factsheets for 80 representative alien species. Chapter 14 975 degraded habitats or seemingly undisturbed localities. A common feature to all such places is the presence of trees, on whose aphid populations the ants depend. Native range: Asia (possibly Turkey). Introduced range: The species was described in 1990 (Van Loon et al. 1990), although its presence in the garden of the Company for the Development of Fruit and Ornamental Production at Budapest, Hungary, was already known from the early seventies (Andrásfalvy, in litt.). Colonies were later observed in Western, Central and Southeastern Europe (Map). Pathways: Present distribution is likely to have been mediated by human intervention (commerce and transport of goods, soil, potted plants). Given the seeming absence of nuptial flight, dispersal capacity of this ant is very low. Local expansion is a very slow process and distances attained are two to five orders of magnitude smaller than minimum distances between known populations. Impact and management: In areas occupied by this species, other surface-foraging ant species have vanished or have reduced populations. Other arthropod groups also seem to be affected in positive (increased abundance; aphids), negative (lower density; lepidoptera larvae) or neutral ways. Occupation of electrical conduits in homes may cause nuisance to people. Selected references Cremer S, Ugelvig LV, Drijfhout FP, Schlick-Steiner BC, Steiner FM et al. (2008) The Evolution of Invasiveness in Garden Ants. PLoS One 3 (12): e3838. doi-10.1371/journal. pone.0003838 Espadaler X, Tartally A, Schulz R, Seifert B, Nagy C (2007) Regional trends and local expansion rate in the garden invasive ant Lasius neglectus (Hymenoptera, Formicidae). Insectes sociaux 54: 293–301. Van Loon AJ, Boomsma JJ, Andrásfalvy A (1990) A new polygynous Lasius species (Hymenoptera, Formicidae) from Central Europe. I. Description and general biology. Insectes Sociaux 37: 348–362. 976 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.58 – Linepithema humile (Mayr, 1868) - Argentine ant (Formicidae, Hymenoptera) Wolfgang Rabitsch Description and biological cycle: Light brown ant; females 4.5–4.9 mm long and workers 2.1–3.0 mm long (Photo) Omnivorous, feeding on honeydew, nectar, insects and carrion. Local dispersal by budding of large unicolonial nests (up to 150 m/year); long-distance dispersal human-mediated within the introduced ranges. Haplodiploid system with sterile workers; polygynous (multi-queened) nests; social organisation variable in its native range (from multicolonial to unicolonial), but entirely unicolonial in introduced range, with surface area covered by single supercolonies ranging from 2500 m² to many km². In the absence of queens, workers can lay unfertilised eggs, which develop into fully functional males. Prefers moderate temperature and moisture levels. Native habitat (EUNIS code): G- tropical and subtropical natural forests. Habitat occupied in invaded range (EUNIS code): I- Regularly or recently cultivated agricultural, horticultural and domestic habitats; G4- Mixed deciduous and coniferous woodland; preferably associated with disturbed, human-modified habitats in its introduced range, but may also invade natural habitats (e.g., oak and pine woodland in the Mediterranean basin). Native range: South America. Introduced range: The Argentine ant occurs throughout the world on all continents, especially in mediterranean-type climates, and many oceanic islands. First recorded in Europe in 1847 in Portugal, it invaded most of the Western Mediterranean Europe and Central Europe (Map). Ecological niche models predict that with changing climate, the species will expand at higher latitudes. Pathways: Transported with vehicles (airplanes, ships) together with goods and materials, soil, plants, etc. Credit: Alex Wild Factsheets for 80 representative alien species. Chapter 14 977 Impact and management: Supercolonies, by reducing costs associated with territoriality, allow high worker densities and interspecific dominance in invaded habitats. It has displaced native ant species in many parts of the world, even leading to species extinction in some cases. Also competes with other arthropod species for resources (e.g., for nectar with bees) and reduces local arthropod diversity. Taxa other than arthropods are also affected (e.g., causes nest failure of birds). Ecosystem level impacts such as reduction of seed dispersal capacity and disruption of mutualistic associations with other species are documented. Regarded as a nuisance for tourism at some places on the Mediterranean coast. Tending behaviour may increase homopteran populations, causing some crop loss. However, these costs are considered to be low. Several chemicals have been applied via ant baits, including insect growth regulators. Application needs supervision to optimise results and to minimise side-effects on non-target species. Since Argentine ants prefer disturbed sites, any extensification of land use or reduction in monoculture may help prevent high densities. Selected references Carpintero S, Reyes-López J, Arias de Reyna L (2005) Impact of Argentine Ants (Linepithema humile) on an arboreal ant community in Donana National Park, Spain. Biodiversity and Conservation 14: 151–163 Giraud T, Pedersen JS, Keller J (2002) Evolution of supercolonies: The Argentine ants of southern Europe. Proceedings of the National Academy of Sciences 99: 6075–6079 Way MJ, Cammell ME, Paiva MR, Colligwood CA (1997) Distribution and dynamics of the argentine ant Linepithema (Irydomirmex) humile (Mayr) in relation to vegetation, soil conditions, topography and native competitor ants in Portugal. Insectes Sociaux 44: 415–433. 978 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.59 – Nematus tibialis Newman, 1837 - Locust sawfly, false acacia sawfly (Hymenoptera, Tenthredinidae) Milka Glavendekić Description and biological cycle: Adult 6–7 mm long, head marked yellow above, thorax and abdomen marked with black; antennae black. Scutellum yellow. Legs yellow, exception of hind tibiae and tarsi, which are black and distinguish Nematus tibialis from the other sawflies. Larva green and shiny, 12 mm long with brownish-green head marked with black (Photo right). Feeds exclusively on black locust, Robinia pseudoacacia L., and its various ornamental cultivars, and on bristly locust Robinia hispida L. Adults emerge in May and June. Females deposit eggs in young leaflets and of the host plant. The young larvae feed on leaves, forming a small hole through the lamina (Photo left). Later, larvae consume more leaf area until maturity. Larval development last two to three weeks after which they enter the soil, forming tough dark brown cocoons, where they pupate. Adults emerge shortly afterwards. False Acacia sawfly develops a second generation in the late summer and sometimes a third brood in the autumn. Native habitat (EUNIS code): G1- Broadleaved deciduous woodland; G4- Mixed deciduous and coniferous woodland; G5- Lines of trees, small anthropogenic woodlands, recently felled woodland, early-stage woodland and coppice. Habitat occupied in invaded range (EUNIS code): G1- Broadleaved deciduous woodland; G5- Lines of trees and, small anthropogenic woodlands: plantations of black locust aimed to stop erosion, recently felled woodland, early-stage woodland and coppice, in the green belt down highways and at all sites where black locust is growing like a weed; I2 - Cultivated areas of gardens and parks; X6- Crops shaded by trees on sites where wind erosion is managed by planting black locust. Native range: North America (Pennsylvania). Credit: György Csóka Factsheets for 80 representative alien species. Chapter 14 979 Introduced range: First detected in Europe in 1825 in Germany. Then established in most countries of Western, Central and Southern Europe (Map). Pathways: Plant trade for ornamental purposes (parks, gardening, bonsai). Impact and management: Severe defoliation of leaves is common on black locust and bristly locust. Holed or partially devoured leaves on ornamental trees and nursery stock reduce ornamental value and health condition of young plants. Nematus also shares a leaf eating niche with several other invasive species, Parectopa robiniella, Phyllonorycter robiniella, Obolodiplosis robiniella, as well as with various aphids and mite species. Control measures are not needed in most cases. The survey of natural enemies revealed egg and larval parasitoids potentially available for biological control. The egg parasitoid Trichogramma aurosum Sugonjaev and Sorokina 1975 (Trichogrammatidae) is recorded from different locations in Central and Western Europe (Denmark, Netherlands, Austria, Luxemburg, Belgium and Germany). Impact is known on local fauna. A larval parasitoid Lathiponus bicolor (Brischke) (Ichneumonidae) has newly adapted to N. tibialis after switching from the congeneric native sawfly species, Nematus salicis L.and N. yokohamensis auct. Selected references Ermolenko VM, Sem’yanov VP (1981) Development of the fauna of sawflies (Hymenoptera, Symphyta) of man-made coenoses of cultivated lands in the south of the European part of the USSR. Noveishie dostizheniya sel’skokhozyaistvennoi entomologii (po materialam USh sëzda VEO, Vilnius, 9–13 October 1979.): 73–76. Markovic C, Stojanovic A (2008) Finding of locust sawfly Nematus tibialis (Newman) (Hymenoptera, Tenthredinidae) in Serbia. Biljni Lekar 36: 131–135. 980 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.60 – Megastigmus spermotrophus Wachtl, 1893 - The Douglas-fir seed chalcid (Hymenoptera, Torymidae) Alain Roques Description and biological cycle: Female 2.8–4.3 mm long, body entirely brownish-yellow to orange- yellow with a few darker spots and an ovipositor as long as body (Photo). Male 2.7–3.8 mm long, with body colour dark lemon yellow with distinct black patterns on head, thorax, propodeum and first two abdominal segments. Adults emerge from late April to mid-June, depending on location. Oviposition occurs after the host plant cone becomes pendant, when its water content is near its maximum. Egg laying begins when a red-brown or purple margin appears on cone scales and lasts until the cone scale turns entirely red-brown. In seed orchards, the oviposition period may last up to 7 weeks. Most oviposition punctures are made on scale margins, resulting in conspicuous resin droplets. Eggs are laid directly into the seed. The hatching larva feeds on archegonia*, then on cotyledons. The following larval instars progressively consume the megagametophyte* (endosperm), which is entirely destroyed by July. Larvae can successfully develop in unpollinated, unfertilized seeds where they prevent megagametophyte abortion. Larval diapause may extend up to four years, but most individuals emerge during the first two years. The proportion of individuals in prolonged diapause is highly correlated with cone abundance in the year following larval development. Sex ratio is highly variable with location and year, usually ranging from 1:0.5–1:1.5. In North America, Douglas-fir seed chalcid attacks both varieties of Douglas-fir, Pseudotsuga menziesii (var. glauca and var. menziesii). In Europe, it has been found in P. menziesii and on other introduced Pseudotsuga species such as P. macrocarpa and P. japonica. Native habitat (EUNIS code): G3 - Coniferous woodland. Habitat occupied in invaded range (EUNIS code): G3 - Coniferous woodland; G4 Mixed deciduous and coniferous woodland ; G5 - Lines of trees, small anthropogenic woodlands, recently felled woodland, early-stage woodland and coppice ; I2 - Cultivated areas of gardens and parks; X11- Large parks; X15- Land sparsely wooded with coniferous trees ; X24Domestic gardens of city and town centres. Credit: David Lees Factsheets for 80 representative alien species. Chapter 14 981 Native range: Western North America, from British Columbia to California and Mexico. Introduced range: First recorded in Europe in Austria in 1893. Then, observed wherever Douglas-fir has been planted, even in Mediterranean countries (Map). Pathways: Trade of tree seeds. The presence of larvae is is usually overlooked in traded seed lots, the infested seeds showing up only when X-rayed (see Figure 12.10 in Chapter 12). Impact and management: In Europe, this species has few indigenous competitors and parasitoids. Thus, the proportion of seeds infested in European seed orchards can reach up to 95%, especially during years of light cone crops. During years of moderate to heavy cone crops, seed infestation varies between 10%-50%. However, the true impact of this insect on seed production is difficult to assess because larvae can complete development in unfertilized seeds. For example, in the absence of fertilization, no viable offspring would be produced from seed, but seed damage would be estimated at 100 %, because only chalcid-infested seeds can be found. Monitoring can be carried out using yellow traps baited with terpinolete. Chemical control is possible, but effective only against adults, whereas systemic insecticides give contrasting results for larvae concealed in the seeds. The introduction of parasitoids from the native range, e.g. the pteromalids Mesopolobus spp., may constitute an alternative, biological control. Selected references Aderkas P Von, Rouault G, Wagner R, Rohr R, Roques A (2005) Seed parasitism redirects ovule development in Douglas-fir. Proceedings of the Royal Society of London B, 272: 1491–1496. Mailleux AC, Roques A, Molenberg JM, Grégoire JC (2008) A North American invasive seed pest, Megastigmus spermotrophus (Wachtl) (Hymenoptera: Torymidae): Its populations and parasitoids in a European introduction zone. Biological Control 44: 137–141. Roques A., Skrzypczynska M (2003) Seed-infesting chalcids of the genus Megastigmus Dalman (Hymenoptera: Torymidae) native and introduced to Europe: taxonomy, host specificity and distribution. Journal of Natural History 37: 127–238. 982 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.61 – Sceliphron curvatum (Smith, 1870), S. caementarium (Drury, 1773) and S. deforme (Smith, 1856) (Hymenoptera, Sphecidae) Jean-Yves Rasplus Description and biological cycle: The genus Sceliphron comprises four species native to Europe. These large black or dark brown wasps, banded with yellow, have thin waist and long legs. In summer, females are seen collecting mud to build their nests composed of several cells (Photo- nest of S. curvatum in Austria). The adult wasp preys on spiders that are packed into the cells, the female lay an egg in the cell and the larva develop at the expense of the stored spiders. During the last 100 years, three alien species of Sceliphron, S. caementarium, S. curvatum and S. deforme, have been introduced to Europe, which are treated together here. Native habitat (EUNIS code): C3- Littoral zone of inland surface waterbodies; G- Woodland and forest habitats and other wooded land. Habitat occupied in invaded range (EUNIS code): C3- Littoral zone of inland surface waterbodies; X25- Domestic gardens of villages and urban peripheries Native range: S. (S.) caementarium is originally native to North America, whilst the two other species belong to another subgenus (Hensenia) which is mostly Asiatic and Australasian. Introduced range: S. caementarium has been accidentally introduced several times into Europe during the 19th and 20th centuries. The species was first reported in 1945 from Versailles (but was never reported there again) and in 1949 from southern France. Since the 1970s, it Credit: Christine Sinhuber, www.aculeata.de Factsheets for 80 representative alien species. Chapter 14 983 is well established and occurs in several countries (France, Spain, Portugal and Ukraine). S. curvatum was first observed in Austria in the 1970s, where it was probably introduced by human activities. It subsequently spread all over central and southern Europe. The species is now reported from most of southern Europe (France, Italy, Greece), but has also reached northern countries (Netherland, Germany and Czech Republic) (Map). The species probably dispersed on its own following large river valleys, and has reached Mediterranean areas where it finds conditions ecologically similar to its native range. It is mostly associated with urban areas, where it constructs nests in different places of the houses. In southern areas, S. deforme may have several overlapping generations. S. deforme, a species naturally distributed in central and tropical Asia, is also reported from several east Mediterranean countries: Bulgaria, Greece, Italy and Montenegro. Pathways: Unknown. Impact and management: S. caementarium and S. curvatum probably threaten native Sceliphron species in France and Austria. However, the impact of S. curvatum on indigenous Sceliphron species is still poorly understood and needs further study. Selected references Bitsch J, Barbier Y (2006) Répartition de l’espèce invasive Sceliphron curvatum (F. Smith) en Europe et plus particulièrement en France (Hymenoptera, Sphecidae). Bulletin de la Société entomologique de France 111: 227–237. Cetkovic A, Radovic I, Dorovic L (2004) Further evidence of the Asian mud-daubing wasps in Europe (Hymenoptera: Sphecidae). Entomological Science 7: 225–229. Gepp J (2003) Verdrängt die eingeschleppte Mauerwespe Sceliphron curvatum autochthone Hymenopteren im Südosten Österreichs? Entomologica Austriaca 8. 984 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.62 – Vespa velutina nigrothorax du Buysson, 1905 - Asian yellow-legged hornet (Hymenoptera, Vespidae) Claire Villemant, Quentin Rome et Franck Muller Description and biological cycle: Black brown hornet, 20–35 mm long, with gastral segments bordered with a fine orange band, except the 4th that is almost entirely orange; front of head orange, extremity of legs yellow (Photo). This coloration corresponds to the variant nigrithorax du Buysson. While preying on a diverse range of insects, Asian yellow-legged hornet shows a strong preference for honeybees, waiting in flight for workers in front of hives. The large nest, often hooked high in tree tops, may contain several thousand individuals (see Figure 12.11 in Chapter 11). The colony founded in April always dies before the end of the year. Future founder queens only survive and overwinter in bark or ground shelters. Their long-distance dispersal is then possible through agricultural and forestry trade movements. The young queen flight capability is not yet assessed, but in general, a hornet adult is able to fly up to 2 km from its nest. Native habitat (EUNIS code): G- Woodland and forest habitats and other wooded land. Habitat occupied in invaded range: Mostly depends on the presence of high trees or buildings for nesting: X25- Domestic gardens of villages and urban peripheries; X10- Mixed landscapes with a woodland element (“bocages”); G- Woodland and forest habitats and other wooded land. Native range: Temperate Asia, probably from Yunnan (south-west China). Invaded range. Introduced in southwestern France (Lot-et-Garonne département) before 2004. Since then, has widely expanded in other parts of France (Map left). The invasion development has been annually traced (Map right). Pathways: Probably through international horticulture trade. Credit: Jean Haxaire Factsheets for 80 representative alien species. Chapter 14 985 Impact and management: As honeybees are its main prey, the Asian hornet represents a new threat to European beekeeping. It also feeds on ripe fruits, and may thus have detrimental effects on local fruit crops. However, economic impact needs to be accurately assessed. A significant public concern also exists because of the sting risk related to the increasing abundance of nests in invaded urban territories. Ecosystem impact includes threat to biodiversity due to the huge predatory pressure on insects (mainly pollinators), as well as potential side-effects on non target species as a consequence of uncontrolled mass trapping and colony destruction by beekeepers and general public. Traps and poisoned baits kill numerous other insects, notably the common yellow-jackets and the European hornet, while nests filled with insecticides and left on the spot threaten birds that fed intensively on brood of poisoned colonies. Nest distribution in France is mapped each year within the program Inventaire National du Patrimoine Naturel website (MNHN, Paris). Chemical control: mass destruction of founder queens in spring seems to have virtually no effect; the best control measure is to kill off a colony by spraying cypermethrine inside the nest after dark, when foraging activities have ceased. The nest is then removed and burnt. However, nests are often difficult to locate before leaf fall, when sexual progeny is already produced. The use of specific baited mass traps to protect hives is under investigation. Selected references Haxaire J, Bouquet JP, Tamisier JP (2006) Vespa velutina Lepeletier, 1836, une redoutable nouveauté pour la Faune de France (Hym., Vespidae). Bulletin de la Société entomologique de France 111: 194. Villemant C, Haxaire J, Streito JC (2006) Premier bilan de l’invasion de Vespa velutina Lepeletier en France (Hymenoptera, Vespidae). Bulletin de la Société entomologique de France 111: 535–538. 986 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.63 – Reticulitermes flavipes (Kollar, 1837) - Eastern subterranean termite (Isoptera, Rhinotermitidae) Marc Kenis Description and biological cycle: The genus Reticulitermes is represented by several species in Europe, America and Asia. A complex of closely related species with uncertain taxonomic status occurs in southern Europe. One species, R. flavipes, is of North American origin and has been introduced into Western France, where it had been first described as R. santonensis de Feytaud and subsequently synonymised with R. flavipes. The same species has also been accidentally introduced in some cities in Germany and Austria. In France, R. flavipes is expanding its range further north. In common with all termites, Reticulitermes spp. are social, living in colonies in the soil. These colonies contain various castes: workers, soldiers, alate reproductives and replacement reproductives. The latter are particularly numerous in Reticulitermes spp. and allow the species to build up colonies of millions of individuals. Nests are built in the ground, usually in a humid environment. Workers bore into wood in contact with the ground to feed the colony (Photo). Dry wood (e.g. building structures) as well as living trees or other sources of cellulose can be attacked. Native habitat (EUNIS code): G1- Broadleaved deciduous woodlands; I- Regularly or recently cultivated agricultural, horticultural and domestic habitats; J- Constructed, industrial and other artificial habitats. Habitat occupied in invaded range (EUNIS code): G1 Broadleaved deciduous woodlands, I: Regularly or recently cultivated agricultural, horticultural and domestic habitats, J: Constructed, industrial and other artificial habitats. Native range: North America. Credit: Gary Alpert, Harvard University, www.insectimages.org Factsheets for 80 representative alien species. Chapter 14 987 Introduced range: France, Germany and Austria (Map). Pathways: Unknown. Impact and management: Reticulitermes flavipes is particularly harmful to wooden elements in buildings but can also attack living trees, as observed with street trees in Paris recently. Reticultermes spp. have had huge economic impacts worldwide. In the USA, subterranean termites are believed to cause more than US$2 billion in damage each year. In France, the recent spread of R. flavipes and Southern European species has caused major concern. New regulations were therefore set up to limit the spread. In nature, indigenous termites are beneficial, by recycling dead trees and other wood material. The impact of Reticulitermes spp. on the soil fauna and flora in newly invaded areas in Europe has not been studied. Management options are numerous and include both prevention and control methods. Building chemical or physical barriers can achieve prevention before and after construction (e.g. chemical wood treatment or steel mesh). Preventing moisture in the soil and in construction structures is an alternative strategy. Curative methods include termiticide injections, baits, trapping methods, etc. Selected references Szalanski AL, Scheffrahn RH, Messenger MT, Dronnet S, Bagnères AG (2005) Genetic evidence for the synonymy of two Reticulitermes species: Reticulitermes flavipes and Reticulitermes santonensis. Annals of the Entomological Society of America 98: 395–401. Clément JL, Bagnères AG, Uva P, Wilfert l, Quintana A, Reinhard J, Dronnet S (2001) Biosystematics of Reticulitermes termites in Europe: morphological, chemical and molecular data. Insectes Sociaux 48: 202–215. Pearce MJ (1997) Termites. Biology and Pest management. Wallingford, UK: CAB Intenational. 172 pp. 988 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.64 – Hyphantria cunea Drury - Fall webworm (Lepidoptera, Arctiidae) Ferenc Lakatos Description and biological cycle: Large white moth species with a wingspan of 19–30mm (Photo left- see also Figure 11.6k- adult male and 11.6i- adult female in Chapter 11). Immature larvae feed gregariously, forming large webs on tree branches (Photo right). Mature larvae tend to be solitary feeders and consume the entire leaf leaving only the petiole. Highly polyphagous, with over 200 known host species including Acer negundo, Morus spp., Prunus spp., Malus spp. and even Populus and Quercus. Adults show remarkable dispersal powers. In Europe, there are up to three generations per year. In Japan, recent climate change has resulted in a shift from a bivoltine to a trivoltine life-cycle in at least a part of the range, together with significant changes in the length of the critical photoperiod for diapause induction. Overwinters as pupae. Adults emerge from mid April onwards and females lay 500 (spring generation) to 800 (summer generations) eggs, usually towards the top part of the host tree. Larvae produce 5–7 instars, feeding gregariously in a light web, except the first and last instars. Larval development takes 24–57 days, depending on climate and host nutrition conditions. Native habitat (EUNIS code): G- Woodland and forest habitats and other wooded land. Habitat occupied in invaded range (EUNIS code): G1 - Broadleaved deciduous woodland; G5- Lines of trees. Native range: North America. Introduced range: Two known introductions, the first one to Hungary during WW II (first individual found in 1940 in Budapest) and the second one in 1978 in Bordeaux, France. Credit: Zdenek Laštůvka Factsheets for 80 representative alien species. Chapter 14 989 Nowadays, present in most of Europe, except UK, Scandinavia and Iberian Peninsula (Map).; also introduced to Japan and China. Pathways: Probably trade of ornamental trees. Impact and management: Threat to orchards, ornamentals and forest trees in some regions of Central and Eastern Europe, as well as in Eastern Asia. Particularly damaging to ornamentals; however, severe damage has occurred only during the expansion phase after establishment. It was a serious pest in Bulgaria, Romania, Hungary, former Yugoslavia, Russia and northern Italy. Nowadays frequent along roadsides, urban forests, parks and gardens. Constantly present in orchards, where the usual plant protection practices keep the population low. Heavy feeding by the caterpillars over time can lead to defoliation (leaf loss) and limb and branch die-back. Trees/ plants are often totally defoliated by late-instar larvae, particularly in the second generation. Environmental impacts are likely, given the high polyphagy and impact on individual plants. Natural enemies have already adapted to the species as well (e.g. Trichogramma, Tachina and Chalcidoidea, and even birds). Previously both mechanical (elimination of webs) and chemical (insecticide) controls were widely used, but nowadays, biological control (at least in the native habitats) plays a much more important role. Selected references Gomi T, Nagasaka M, Fukuda T, Hagihara H (2007). Shifting of the life cycle and life-history traits of the fall webworm in relation to climate change. Entomologia Experimentalis and Applicata 125: 179–84. Rezbanyai-Reser L (1991) Hyphantria cunea Drury, 1773, und Noctua tirrenica Biebinger, Speidel & Hanigk, 1983, im Südtessin, neu für die Schweiz (Lep.: Arctiidae, Noctuidae). Entomologische Berichte Luzern 26: 94–96, 135–152. Torp R (1987) Ny dansk spinder: Hyphantria cunea Drury f. textor Harr. Lepidoptera 5: 83–86. 990 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.65 – Paysandisia archon (Burmeister, 1879) - The castniid palm borer (Lepidoptera, Castniidae) Carlos Lopez-Vaamonde & David Lees Description and biological cycle: Large dayflying moth with clubbed antennae, wingspan 75–120 mm, upperside forewing greenish brown in both sexes, hindwing bright orange with a black band postdiscal to white spots (Photo left). Forewing underside orange, excepting beige tips. Upright fusiform eggs, about 4.7 mm. long and 1.5 mm wide, laid by the female’s extensible ovipositor between mid-June and mid-October. Fertile eggs pink, laid among palm crown fibres, at the base of leaf rachis. Larvae hatch after 12–21 d, whitish and grub-like, up to 9 cm long, endophytic cannibals, forming galleries 20–30 cm long inside palm trunks, towards the crown (Photo right). 7–9 larval instars, overwintering as larva, in a false cocoon. Pupation occurs at the rachis base or between inflorescences, where larvae form a cryptic cocoon of palm fibres, pupating for 43–66 d. Pupae remain attached to the cocoon after adult emergence. Adults observed from mid May to late September, males especially exhibiting powerful territorial flight in hot sunshine. Males live about 24 d and females about 14 d. One generation per year (sometimes bivoltine) in Mediterranean locations. Larvae can live > 18 months and overall life cycle 13–22 months, exceptionally three years. Castniid palm borer infests a wide range of palm genera including Chamaerops, Latania, Livistona chinensis, Phoenix canariensis, Syagrus spp., Trithrinax campestris (probable import host), in the native area. Reported from Brahea, Butia, Chamaerops, Livistona spp. Phoenix spp., Sabal, Trachycarpus, Trithrinax campestris and Washingtonia, in the introduced area. Native habitat (EUNIS code): G2 - Broadleaved evergreen woodland. Habitat occupied in invaded range (EUNIS code): I2- Cultivated areas of gardens and parks; X24 Domestic gardens of city and town centres; J100- Greenhouses. Native range: Neotropical region: western Uruguay, northwest Argentina, Paraguay and southeastern Brasil. Introduced range: First introduced with its foodplant to Spain and France in the 1990’s, well established by 2001 when first reported from Catalonia in Northeastern Spain. Rapidly spread Credit: Laurence Olivier Factsheets for 80 representative alien species. Chapter 14 991 to coastal areas of the other Mediterranean regions where palms are widely used as ornamentals. Now common and widespread in Spain (along the Mediterranean coast from Girona to Alicante and the Balearic islands) southeastern France (Var and Hérault), Italy (Campania, Lazio, Toscana, Marche and Sicily), and in Greece mainland and Crete); also introduced in England (Sussex, one example in 2002) and Netherlands (one example in 2006) (Map). Spreading tendency. Pathways: Introduced with trade of palm trees as ornamentals. Impact and management: Pest species in parks and palm nurseries, causing severe damage (such as holes in leaves and deformation) and death of plants. Conservation concern exists for the native Mediterranean Fan Palm, Chamaerops humilis; numerous larvae may be found in one plant. Biological control in Europe is not yet achieved. As last resort, palms can be pulled up and burned. Chemical control of this species is also difficult since larvae are endophytes. Best control has been obtained by wetting crown and trunk with contact or systemic organophosphorus insecticides (Chlorpyrifos, Acephate and Dimethoate). Ostrinil (Beauveria bassiana147 strain) biological insectides normally used for the European Corn Borer cause egg and up to 80% larval mortality for crown treatment every two weeks during flight season, and can be used as a curative. Trials done with“glue” used as physical barrier (both preventing adult females from ovipositing and developing adults from emerging) have had positive results. Selected references Colazza S, Privitera S, Campo G, Peri E, Riolo P (2005) Paysandisia archon (Lepidoptera: Castniidae) a new record for Sicily. L’Informatore Fitopatologico 5: 56–57. Hollingsworth T (2004) Status of Paysandisia archon (Burmeister) (Lepidoptera: Castniidae) in southern Europe. British Journal of Entomology and Natural History 17: 33–34. Sarto i Monteys V (2002) The discovery, description and taxonomy of Paysandisia archon (Burmeister, 1880), a castniid species recently found in southwestern Europe (Castniidae). Nota Lepidopterologica 25: 3–16. 992 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.66 – Diplopseustis perieresalis (Walker, 1859) - The exotic pyraloid moth (Lepidoptera, Crambidae) Jurate De Prins & Willy De Prins Description and biological cycle: Small moth, wingspan 12–14 mm, forewing greyish with some whitish transverse lines, indicated by separate dots; submarginal line more conspicuous; in some specimens, darker greyish antemedian transverse line, most visible towards inner margin (Photo). Termen* sinuous. Labial palps porrect*, a little longer than eye diameter. Hindwings whitish with greyish suffusion, dark marking in anal corner, transversed by white line parallel to wing margin. Biology undescribed, but larva supposed to live on Carex spp. Probably hibernating in larval stage. Native habitat (EUNIS code): Unknown. Habitat occupied in invaded range (EUNIS code): E3 - Seasonally wet and wet grasslands. Native range: Oriental and Australasian regions: Australia, Brunei, China (coastal regions), Fiji, Hong Kong, India, Indonesia, Japan, Malaysia (Borneo, Sarawak), New Zealand, Taiwan. Credit: C. van den Berg Factsheets for 80 representative alien species. Chapter 14 993 Introduced range: First recorded from Portugal in 2000. Then observed in several countries of western and Southern Europe: Belgium, Netherlands, Portugal, Spain (mainland, Balearic and Canary islands), United Kingdom (Scilly Islands) (Map). Pathways: Long-distance dispersal probably human-mediated (plant trade between Australia, New Zealand, and Japan, and the Canary Islands), although the species has a strong dispersal power and could spread on its own. Spreading mode within Europe unknown. Impact and management: Infestation rate still very low, no economic damage (Carex spp. are not commercially valuable). Neither chemical nor biological measurements necessary. Selected references Mackay A, Fray R (2002) Diplopseustis periersalis [sic] (Walker) on Tresco, Isles of Scilly – the first record for Britain and the western Palearctic region. Atropos 16: 26. Speidel W, Nieukerken van EJ, Honey MR, Koster SJC (2007) The exotic pyraloid moth Diplopseustis perieresalis (Walker) in the west Palaearctic region (Crambidae, Spilomelinae). Nota Lepidopterologica 29: 185–192. 994 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.67 – Phthorimaea operculella (Zeller, 1873) - Potato tuber moth (Lepidoptera, Gelechiidae) Georgyi Trenchev and Katia Trencheva Description and biological cycle: Adult small, light grey moth, about 10 mm long with wingspan of about 12 mm (Photo left- adult on potato). Greyish-brown wings patterned with small, dark specks. Larvae light brown with a brown head. Mature larvae reach 15–20 mm long, pink or greenish. Multivoltine, developing four or more generations per year, depending on climate conditions. In field conditions, adult, larva or pupa overwinters under crop residues in the upper layer of ground. In temperate climates, overwintering occurs mainly in storehouses. Adult moth emerges in spring, and found until the end of October. Moths are active after sunset. They lay eggs in groups of 2–3 or individually on the lower sides of plant leaves, sometimes on leafstalks, stems, exposed potato tubers, lumps of soil in the field, on potato tuber buds and on bags in storehouses. Female fecundity is 150–200 eggs. Embryonic development lasts 3–10 days. Larval development lasts for 11–14 days and passes through four instars. Pupal stage lasts 6–8 d. In storehouses, the pest develops continuously throughout the year. Native habitat (EUNIS code): I- Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): I1- Arable land and market gardens; I2-Cultivated areas of gardens and parks; J100- glasshouses. Native range: Probably originates from South America. Introduced range: Widely distributed worldwide, recorded in more than 70 countries. Known in Europe since the beginning of the 20th century, where it is mainly found in potato fields and storehouses (Map). Credit: Katia Trencheva Factsheets for 80 representative alien species. Chapter 14 995 Pathways: Adults fly to potato fields over short distances and they can transported by wind. Over long distances, transportation occurs with infested tubers and re-infestation of fields then occurs from potato storehouses. Impact and management: Damage is most frequent on stored tubers after the spring growing season and on young plants in the autumn growing season. Larvae bore holes and galleries in the tubers (Photo right). Larval penetration holes are unsightly and induce soft rot. Phthorimaea operculella can be a very serious potato pest, especially in tropical and sub-tropical regions, including the Mediterranean region. The attack results in lowered market value and quality of the infested tubers. Infestation may start early in the field, up to 15 d before tuber maturity. By harvest time, a substantial number of tubers may already be infested. This harvest-time infestation is responsible for the further development of infestation in stores. Integrated pest management methods have been developed in various parts of the world. Control measures include the use of pesticides; cultural practices include use of healthy tubers, irrigation or early harvest. Biological control is achieved through the introduction of parasitoids but also the use of Bt or Baculovirus, and through use of resistant varieties. Pheromones can be used both for monitoring and for control trapping in storage. Selected references CABI (2007) Crop Protection Compendium. CD-ROM. CAB International, Wallingford, UK. Das GP, Magallona ED, Raman KV, Adalla CB, 1992. Effects of different components of IPM in the management of the potato tuber moth, in storage. Agriculture, Ecosystems & Environment 41: 321–325. Trivedi TP, Rajagopal D, 1992. Distribution, biology, ecology and management of potato tuber moth, Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae): a review. Tropical Pest Management, 38: 279–285. 996 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.68 – Cameraria ohridella Deschka & Dimić, 1986 - Horse chestnut leaf-miner (Lepidoptera, Gracillariidae) Sylvie Augustin Description and biological cycle: Tiny moth, 3–5mm long, forewing background orangebrown with basal white longitudinal streak and white v-marks bent towards costa and running straight across dorsum at rest, edged posteriorly in black, two of these fasciae continuous medially, one post-medial fascia interrupted and a final convex fascia towards wing apex, bisected by a diffuse blackish subapical streak (Photo left- see also Figure 11.7c in Chapter 11). Fringe forming a conspicuous orange tuft that is longest dorsally at rest. Head tufted with orange hair-like scales intermixed with white, scape and base of antenna silvery white. Antennae about 4/5 forewing length. Phytophagous larvae, mining leaves of white-flowered horse chestnut Aesculus hippocastanum; but can also develop on other Aesculus species and occasionally on maples, Acer pseudoplatanus and A. platanoides. Leaf mines from April onwards; an average of 75 eggs are laid per female on the upper epidermis of horse-chestnut leaves. Produces four (rarely five) mining and two spinning larval instars; usually three generations per year in W Europe, but up to five overlapping generations depending on weather conditions and climate. Pupae diapause in leaves. Native habitat (EUNIS code): G1 - Broadleaved deciduous woodland. Habitat occupied in invaded range (EUNIS code): G1 - Broadleaved deciduous woodland; I2 - Cultivated areas of gardens and parks; X13 - Land sparsely wooded with broadleaved deciduous trees; X11 - Large parks; J - Constructed, industrial and other artificial habitats. Native range: Southern Balkans. Introduced range: Most of Europe, except part of Northern Europe and Western Russia (Map). Increasing its distributional range and abundance in newly colonized areas. Credit: Sylvie Augustin Factsheets for 80 representative alien species. Chapter 14 997 Pathways: Adult moths are transported by wind. Anthropogenic transport occurs by vehicles, in infested leaf fragments or infested nursery stock. Impact and management: Severely defoliated trees produce smaller seeds with a lower fitness that affects tree regeneration and seriously impairs recruitment of horse chestnut in the last endemic forests in the Balkans. A single leaf can host up to 106 leaf-miners (Photo right). Parasitism rates low, as most parasitoids emerge when larvae or pupae are not yet available; this may have an important impact on native leaf-miners. There is significant public concern because of aesthetic impact. Main costs are caused by by removal or replacement of severely damaged horse chestnut trees planted in cities and villages. Complete removal of leaf litter, in which pupae hibernate, is the only effective measure available to lessen damage. The majority of adults can be prevented from emerging when leaves are properly composted (e.g., mulching of horse chestnut leaves with a layer of soil or uninfested plant material). Chemical control: aerial spraying with dimilin is efficient; spraying with Fenoxycarb combined with a surfactant has proved effective. Other “biological pesticides” with fewer non-target effects, such as neem, are also feasible, but their efficiency is considered to be lower. Stem injection is also efficient, but is not widely registered. This injures trees through necrosis and infections, and systemic insecticide may cause side effects on non-target species such as honey bees. Selected references Gilbert M, Grégoire J-C, Freise J, Heitland W (2004) Long-distance dispersal and human population density allow the prediction of invasive patterns in the horse-chestnut leafminer Cameraria ohridella. Journal of Applied Ecology 73: 459–468. Lees DC, Lopez-Vaamonde C, Augustin S (2009) Cameraria ohridella Deschka & Dimić 1986. http://eolspecies.lifedesks.org/pages/8675. Valade R, Kenis M, Hernandez-Lopez A, Augustin S, Mena NM, Magnoux E Rougerie R, Lakatos F, Roques A, Lopez-Vaamonde C (2009) Mitochondrial and microsatellite DNA markers reveal a Balkan origin for the highly invasive horse-chestnut leaf miner Cameraria ohridella (Lepidoptera, Gracillariidae). Molecular Ecology 18: 3458–3470. 998 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.69 – Parectopa robiniella (Clemens, 1863) - Locust Digitate Leafminer (Lepidoptera, Gracillariidae) David Lees Description and biological cycle: Small moth, wingspan 5.73–7.26 mm. Forewing background dark orange with four white curved flecks outlined in fuscous, running from costa and tergal edge of forewing in an interdigitate fashion towards middle of wing; white costal mark in between most apical flecks. Hindwing mid brownish with brownish cilia tipped apically in white (see Figure 11.6b in Chapter 11). Hindlegs conspicuously banded brown and white. Mine starts close to base of leaf with egg laid on underside, at the base of a fork made by the veins, where the larva bores through to upperside and forms a distinctive whitish digitate shape, straddling the midrib (Photo), unlike Phyllonorycter robiniella, which may precede it by about two weeks, and in U.S.A., Chrysaster ostensackenella Fitch. Larva greenish and solitary, leaf-miner on Black Locust (or False Acacia) trees Robinia pseudacacia and R. hispida, and on other Fabaceae including Amorpha fruticosa, Galactia volubilis and Desmodium sp. Leaf-mines and adult flight occurs from June to October (in two, sometimes overlapping, generations). Larva pupates in leaf litter on ground, in contrast to Phyllonorycter robiniella, and probably thus less susceptible to parasitism. Native habitat (EUNIS code): G- Woodland, forest and other wooded land. Habitat occupied in invaded range (EUNIS code): G5 - Lines of trees, small anthropogenic woodlands, recently felled woodland, early-stage woodland and coppice; I- Regularly or recently cultivated agricultural, horticultural and domestic habitats; I2- Cultivated areas of gardens and parks; X24- Domestic gardens of city and town centres. Native range: North America (Canada: Québec), U.S.A. (Florida, Kentucky, Maine, Maryland, Michigan, Missouri, New Orleans, New York, Pennsylvania, Vermont, Wisconsin) Credit: György Csóka Factsheets for 80 representative alien species. Chapter 14 999 Introduced range: First observed in Europe in Italy in 1970, Locust Digitate Leafminer has spread relatively quickly in various directions to Western and Central Europe and to the Balkans (Map). Pathways: Unknown, possibly via plant trade. Impact and management: Causes damage to false acacia trees including leaf drop as early as June in cases of severe infestation. Potential impact on ornamental trees and industrial plantations. Natural control includes at least 20 species of wasps in Braconidae (Pholetesor circumscriptus, P. nanus), Chalcidoidea Encyrtidae (Ageniaspis sp.), Eupelmidae (Eupelmus urozonusi), Eulophidae (Achrysocharoides cilla, Astichus trifasciatipennis, Chrysocharis nitetis, Cirrospilus viticola, Closterocerus cinctipennis, C. formosus, C. trifasciatus, Elachertus inunctus, Holcothorax testaceipes, Hyssopus benefactor, Minotetrastichus frontalis, Neochrysocharis formosa, Pediobius saulius, Pnigalio pectinicornis, P. soemius, Sympiesis acalle, S. marylandensis, S. sericeicornis and Ichneumonidae (Gelis acarorum, Diadegma sp.), but parasitism levels may be too low to have much impact. As for Phyllonorycter robiniella, parasitoids have easily shifted from other hosts, but this species has been less susceptible. Selected references Ivinskis P, Rimsaite J (2008) Records of Phyllonorycter robiniella (Clemens, 1859) and Parectopa robiniella Clemens, 1863 (Lepidoptera, Gracillariidae) in Lithuania. Acta Zoologica Lituanica 18: 130–133. Lakatos F, Kovács Z, Stauffer C, Kenis M, Tomov R, Davis DR (2003) The Genetic Background of Three Introduced Leaf Miner Moth Species - Parectopa robiniella Clemens 1863, Phyllonorycter robiniella Clemens 1859 and Cameraria ohridella Deschka et Dimic 1986. In: Proceedings: IUFRO Kanazawa (2003) “Forest Insect Population Dynamics and Host Influences”, 67–71. 1000 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.70 – Phyllonorycter issikii (Kumata, 1963) (Lepidoptera, Gracillariidae) David Lees Description and biological cycle: Small moth, wingspan 7.0–7.5 mm. Thorax and forewing ground colour golden to light brownish ochreous, with blackish stripes and three white stripes in summer form (see Figure 11.6d in Chapter 11), dusted with white scales in autumn form (see Figure 11.6c in Chapter 11), well camouflaged for hibernation; hindwings pale grey, cilia tinged with yellow. Adult resembles Phyllonorycter corylifoliella, but male genitalia highly asymmetric, right valve especially wide and left one slender. Eggs oblong, greenish, about 0.24 x 0.35 mm. Larva yellowish towards caudal end and white toward head. Leaf-miner on lower surface (Photo) of small-leaved Lime Tilia cordata, Tilia platyphyllos or various crosses such as Tilia x vulgaris (Tiliaceae), with adults flying in two generations at end of April and May and AugustSeptember (in Europe). Oligophagous on Tilia, apparently without strong preference. Feeds on T. maximowicziana, T. kiusiana and T. japonica (in Japan), T. amurensis (far eastern Russia) and T. mandshurica (in Korea). Development: egg, 4–8 d, larva in five instars, the last two tissue feeding, 13–40 d, pupa 10–15 d. Prefers trees in understory/shade. Mine when unfolded showing micro-ridges, elliptical to oblong, whitish, on underside of leaf, usually at fork of primary or secondary veins, with frass piled up at one end. Hibernates as adult. Native habitat (EUNIS code): G - Woodland, forest and other wooded land. Habitat occupied in invaded range (EUNIS code): G5 - Lines of trees, small anthropogenic woodlands, recently felled woodland, early-stage woodland and coppice; I - Regularly or recently cultivated agricultural, horticultural and domestic habitats; I2 - Cultivated areas of gardens and parks; X24 - Domestic gardens of city and town centres. Spreading quite rapidly westwards especially after 2000. Credit: Hana Šefrová Factsheets for 80 representative alien species. Chapter 14 1001 Native range: Japan (Hokkaidõ, Honshũ, Kyũshũ) and probably also in far eastern Russia eastern China and Korea (where first reported 1977). Introduced range: First reported from Moscow in 1985, Phyllonorycter issikii spread to the Baltic countries of the Baltic countries and most of Central Europe (Map and Šefrová (2002) for a review of spread in Europe). Pathways: Apparently spread by wind and possibly also by horticultural trade and passive spread of hibernating adults, since the distance between eastern and western Russia seems too large for possible long distance aerial transport. Impact and management: Causes damage including limited leaf folding to lime trees. Potential aesthetic impact to park and garden trees is relatively minor, since sunny branches are avoided, and no native Tilia populations are threatened. Natural controls include the chalcidoid eulophid wasp parasitoids Chrysocharis laomedon, Mischotetrastichus petiolatus, Pediobius saulius, Pleuroppopsis japonica, Sympiesis laevifrons and S. sericeicornis, but biological control seems unnecessary, since parasitoid levels attained 20% in some localities even the year after arrival in eastern Europe. Selected references Ermolaev IV, Motoshkova NV (2008) Biological invasion of the Lime Leafminer Lithocolletis issikii Kumata (Lepidoptera, Gracillariidae): interaction if the moth with the host plant. Entomological Review, 88: 1–9. Noreika R (1998) Phyllonorycter issikii (Kumata) (Lepidoptera, Gracillariidae) in Lithuania. Acta Zoologica Lituanica 8: 34–37. Šefrová H (2002) Phyllonorycter issikii (Kumata, 1963) - bionomics, ecological impact and spread in Europe (Lepidoptera, Gracillariidae). Acta Universitatis Agriculturae et Silviculturae Mendelianae Brunensis 50: 99–104. 1002 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.71 – Phyllonorycter platani (Staudinger, 1870) (Lepidoptera, Gracillariidae) David Lees Description and biological cycle: Small moth, wingspan 8–10 mm. Adult forewing light golden-orange, with silvery white markings: mediobasal stripe to a third of forewing length, longer white stripes running along costa and tergum; narrow transverse medial band at 2/3 that may be divided; apically two costal and two dorsal streaks that may meet in middle of wing and are basal to a small black apical eyespot (see Figure 11.6f in Chapter 11). Asymmetric male genitalia, left valve at least twice as broad as right. Larva yellowish white, up to 7 mm long. Leafminer on Platanus orientalis, P. racemosa, P. occidentalis, P. acerifolia, P. hispanica and commonly in urban areas on Platanus x hispanica (Platanaceae) (“London Plane”). Mine a large distinctive blotch commencing in a sinuous pattern, usually on underside of leaf, between veins, appears brownish underneath, mottled upperside, and semi-transparent against sky (Photo left- mine on Platanus leaf; Photo right- mine opened to show third- instar larva). Pupa light brown, pupates in a whitish cocoon within leaf on ground. Adults on wing between April and September, in two generations, with larvae of summer generation diapausing. Native habitat (EUNIS code): Unknown Habitat occupied in invaded range (EUNIS code): G5 - Lines of trees, small anthropogenic woodlands, recently felled woodland, early-stage woodland and coppice; I - Regularly or recently cultivated agricultural, horticultural and domestic habitats; I2 - Cultivated areas of gardens and parks; X24 - Domestic gardens of city and town centres. Native range: Unknown, but possibly Balkans; described from Italy. Credit: Hana Šefrová Factsheets for 80 representative alien species. Chapter 14 1003 Introduced range: Recorded in most European countries wherever Platanus trees have been planted (Map and Šefrová (2001) for a review of its dispersal history in Europe). Also introduced in central Asia (Kazakhstan, Tajikistan, Turkmenisatn), Asia Minor (Iran, Syria), and apparently in the USA (California). Pathways: Passive dispersal via fallen leaves has greatly facilitated the rapid spread of Phyllonorycter platani. Impact and management: Causes damage to plane trees, rarely unsightly, as leaves remain green, but sometimes reaching a density of 60 mines per leaf. Therefore of potential, but not usually severe, aesthetic impact. Leaves can be gathered up and burned. Chemical control not recommended nor necessary. Natural control by parasitoid wasps: at least 57 spp. recorded, including families Braconidae (genera Apanteles, Colstus, Pholeteor), Chalcididae (Conura), Encyrtidae (Ageniaspis); Eulophidae (Achrysocharoides, Aprostocetus,Chrysocharis, Cirrospilus, Clostocerus, Diglyphyus, Elachertus, Eulophus, Horismenus, Minotetrastichus, Pediobius, Pnigalio, Sympiesis); Ichneumonidae (Itoplectis, Pimpla, Scambus, Triclistus); Pteromalidae: Chlorocytus, Conomorium, Pteromalus); Torymidae (Torymus). Selected references Frankenhuyzen Av (1983) Phyllonorycter platani (Staudinger, 1870) (Lep.: Gracillariidae), een bladmineerder op plataan in Nederland. Entomologische Berichten 43: 19–25. Šefrová H (2001) Phyllonorycter platani (Staudinger) - a review of its dispersal history in Europe (Lepidoptera, Gracillariidae). Acta Universitatis Agriculturae et Silviculturae Mendelianae Brunensis 49: 71–75. 1004 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.72 – Phyllonorycter robiniella (Clemens, 1859) (Lepidoptera, Gracillariidae) David Lees Description and biological cycle: Small moth, wingspan 5.98–6.37 mm. Adult forewings light orange with four silvery-white diagonal striae running from costa towards tergal edge, but reflexed at middle of wing, at least the terminal one divided, apically a black eyespot; base of wings silvery white (see Figure 11.6g in Chapter 11). Larva in Europe a leaf-miner of Robinia pseudacacia (Fabaceae) (in North America using also R. hispida, R.viscosa and R. neomexicana, but not recorded on other genera), from June to October, in two or usually three generations from June to October in Europe. Diaphanous whitish, tentiform blotch mine that does not traverse midrib, but may occupy a large part of one side of the leaf, usually on underside (Photo), occasionally on leaf upperside, and which may sometimes merge to contain up to 15 larvae. Egg, light greenish grey, 6–10 d, larva, hypermetamorphic, in final two of five tissue-feeding instars, cylindrical, 20–50 d, pupa in oval white cocoon within the mine 7–20 d, 5–11 weeks for development. Hibernates as adult. Native habitat (EUNIS code): G - Woodland, forest and other wooded land. Habitat occupied in invaded range (EUNIS code): I - Regularly or recently cultivated agricultural, horticultural and domestic habitats; I2 - Cultivated areas of gardens and parks. Native range: Nearctic: Eastern and central USA (North America), throughout the native range of Robinia pseudacacia, including Canada (Québec), U.S.A (Connecticut, Florida, Illinois, Kentucky, Maine, Maryland, Massachusets, Michigan, New York, Texas, Vermont, and Wisconsin). Introduced range: First recorded in Europe in 1983 in Switzerland, it then invaded most of Western, Central and Northern Europe: Austria, Belgium (from 2000), Croatia, Czech Republic, France, Germany, Hungary, Italy (1988), The Netherlands (1999), Poland, Slovakia, Spain (Barcelona 2000), Switzerland (1983), Ukraine. Apparently spreading faster eastwards than westwards (Map). Credit: Hana Šefrová Factsheets for 80 representative alien species. Chapter 14 1005 Pathways: Passive wind dispersal may be unusually important for this species, as although leaves can be carried by cars, pupae hatch before leaf fall, making leaf transport more more unlikely than for some other gracillariid species. Impact and management: Causes premature leaf drop to false acacia trees and thus has potential aesthetic and physiological impact. Reported to have a higher surface area impact on industrial plantations than Parectopa robiniella. Damage must be weighed against considerations that false acacia is itself an undesirable alien in some European ecosystems. Chitin synthesis inhibitors applied in late May could cure leaf drop. Natural control includes at least 22 species of (polyphagous) braconid (Apanteles nanus, Colastes braconius, Pholetesor bicolor, P. circumscriptus, P. ornigis), and chalcidoid eupelmid and eulophid wasps (Achrysocharoides cilla, A. gahani, Astichus trifasciatipennis, Baryscapus nigroviolaceus, Chrysocharis nephereus, Closterocerus cinctipennis, C. trifasciatus, Elachertus inunctus, Horismenus fraternus, Minotetrastichus frontalis, M. platanellus, Pediobius liocephalatus, P. saulius, Pnigalio pectinicornis, P. soemius, Sympiesis acalle, S. marylandensis and S. sericeicornis). Parasitoids have easily shifted from other hosts. Selected references De Prins W, Groenen F (2001) Phyllonorycter robiniella, een nieuwe soort voor de Belgische fauna (Lepidoptera: Gracillariidae). Phegea 29: 159–160. Ivinskis P, Rimsaite J (2008) Records of Phyllonorycter robiniella (Clemens, 1859) and Parectopa robiniella Clemens, 1863 (Lepidoptera, Gracillariidae) in Lithuania. Acta Zoologica Lituanica 18: 130–133. Šefrová H (2002) Phyllonorycter robiniella (Clemens, 1859) - egg, larva, bionomics and its spread in Europe (Lepidoptera, Gracillariidae). Acta Universitatis Agriculturae et Silviculturae Mendelianae Brunensis 50: 7–12. 1006 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.73 – Cacyreus marshalli (Butler, 1898) - Geranium bronze (Lepidoptera, Lycaenidae) Jurate De Prins & Willy De Prins Description and biological cycle: Small butterfly, wingspan 15–27 mm, upperside forewing brown in both sexes, fringes checkered, hindwing with short tail and conspicuous dark spot close to tail (Photo left), upperside thus resembling Lampides boeticus (L.). Hindwing fringe pure white with sometimes a narrow, interrupted brown line in the middle. Underside unlikely to be confused with any other European lycaenid. Eggs are laid on the flowers buds or the underside of the leaves of Pelargonium spp. Larvae feed mainly on the flowers and flower buds, but also other parts of the foodplant are consumed. First two instars are obligate endophytes in the flower buds, young shoots or leaves; the last three instars are facultative endophytes in all plant tissues, except the roots, but they may also occur as external feeders. Pupation inside the stem or among leaf litter at the base of the foodplant (Photo right- pupae). No photoperioddriven diapause. The species cannot survive severe winters in Central Europe outdoors, but it can complete its life cycle during summer time in this region. Also, it often survives the winter season because geraniums are often put inside at this time, when development is slowed down. Five to six generations per year in Mediterranean locations. Closely associated to Pelargonium in the native range. Native habitat (EUNIS code): I2- Cultivated areas of gardens and parks. Habitat occupied in invaded range (EUNIS code): I1- Arable land and market gardens; I2- Cultivated areas of gardens and parks. Native range: Afrotropical region: South Africa. Introduced range: First introduced with its foodplant to the Balearic Island of Mallorca at the end of the 1980s. Spread fast to the other Balearic islands and the coastal regions of Credit: Paolo Mazzei Factsheets for 80 representative alien species. Chapter 14 1007 the West-Mediterranean, where geraniums (Pelargonium spp.) are widely used as ornamentals. At present common and widely spread in the Mediterranean basin (France common north to Lyon, Italy, Portugal, Spain mainland and Balearic Islands) but rare and isolated records in Central Europe (Belgium, Germany, Netherlands and Switzerland) and United Kingdom (southern coast) (Map). Also introduced in Morocco. Spreading tendency. Pathways: trade of ornamental geraniums (Pelargonium spp.) Impact and management: Pest species in geranium nurseries, causing severe damage and even death of plants. In laboratory conditions, oviposition has been observed on native European Geranium species (e.g. G. pratense, G. sylvaticum), and hence Cacyreus marshalli represents a potential threat for both native geraniums and for other Geranium-consuming lycaenids, such as Aricia nicias (Meigen) and Eumedonia eumedon (Esper). Trials with several insecticides on the island of Mallorca had positive results. Since 1995, no autochthonous parasitoids have been reared from Cacyreus marshalli. Selected references Holloway J (1998) Geranium bronze Cacyreus marshalli Atropos 4: 3–6. Quacchia A, Ferracini C, Bonelli S, Balletto E, Alma A (2008) Can the Geranium Bronze, Cacyreus marshalli, become a threat for European biodiversity? Biodiversity and Conservation 17: 1429–1437. Sarto i Monteys V (1992) Spread of the Southern African Lycaenid butterfly, Cacyreus marshalli Butler, 1898, (LEP: Lycaenidae) in the Balearic Archipelago (Spain) and considerations on its likely introduction to continental Europe. Journal of Research on the Lepidoptera 31: 24–34. 1008 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.74 – Spodoptera littoralis (Boisduval, 1833) - African cotton leaf worm (Lepidoptera, Noctuidae) Carlos Lopez-Vaamonde Description and biological cycle: Polyphagous moth, up to 2 cm long with wingspan of 4 cm (Photo left); eggs laid in batches covered with orange-brown hairs. The neonate larva is pale green with a brownish head; when fully developed, larvae 35–45 mm long, body colour varying from grey to reddish or yellowish, with a median dorsal line bordered on either side by two yellowish-red or greyish stripes, and small yellow dots on each segment (Photo right- mature larva on a tomato leaf). 1000–2000 eggs laid per female 2–5 d after emergence; egg masses of 100–300 on the lower leaf surface of host plants. Life cycle lasts 19–144 days. Larvae are extremely sensitive to climatic conditions, especially to combinations of high temperature and low humidity; temperatures above 40 °C or below 13 °C increase mortality. Native habitat (EUNIS code): F5 - semi-arid and subtropical habitats. Habitat occupied in invaded range (EUNIS code): F5 - Maquis, matorral and thermoMediterranean brushes; F6 - Garrigue; F8 - Thermo-Atlantic xerophytic habitats; H5 - Miscellaneous inland habitats with very sparse or no vegetation; I1- Arable land and market gardens; I2 - Cultivated areas of gardens and parks; J100- Glasshouses. Native range: Origin unclear, probably Egypt. Widespread in tropical and subtropical Africa and Southeastern Europe and Asia Minor. Introduced range: One of the most commonly intercepted species in Europe, for example on imported ornamentals. Present outdoors in Sicily, southern Italy, Corsica, Spain, southern Portugal, and in Madeira and the Canary Islands but only in glasshouses in northern Italy, Western and Central Europe (Map). Not established in Great Britain. Pathways: Trade appears to be the most likely pathway for introduction, through eggs and larvae present on imported commodities such as glasshouse crops, both ornamentals and vegetables from infested areas. Flight range of moths can be 1.5 km during a period of 4 h Credit: Paolo Mazzei (left), Jean Yves Rasplus/ INRA (right) Factsheets for 80 representative alien species. Chapter 14 1009 overnight. Adult moths can also be spread through wind, attached to or transported by another organism or through other natural means. Impact and management: Spodoptera littoralis is one of the most destructive agricultural lepidopteran pests within its subtropical and tropical range, attacking plants from 44 families including grasses, legumes, crucifers and deciduous fruit trees. In North Africa damages vegetables, in Egypt cotton, and in Southern Europe, plant and flower production in glasshouses or vegetables and fodder crops. It is important to seek assurance from suppliers that plants are free from this pest as part of any commercial contract. Avoid importing plant material from infested areas. Carefully inspect new plants on arrival, including any packaging material, to check for eggs and caterpillars and for signs of damage. As the adults are nocturnal, light or pheromone traps should be used for monitoring purposes. Mechanical control: physical destruction of insects and any plant material infested by this pest is recommended. Egg masses can be hand collected. Chemical control: there are many cases of resistance to insecticides. Biological control: includes the use of microbial pesticides, insect growth regulators and slow-release pheromone formulations for mating disruption. Selected references Abdel-Megeed MI (1975) Field observations on the vertical distribution of the cotton leafworm, Spodoptera littoralis on cotton plants. Zeitschrift für Angewandte Entomologie 78: 597–62. Brown ES, Dewhurst CF (1975) The genus Spodoptera (Lepidoptera, Noctuidae) in Africa and the Near East Bulletin of Entomological Research 65: 221–262. EPPO/OEPP (2003) Fiche informative sur les organismes de quarantaine. Spodoptera littoralis and Spodoptera litura. http://www.eppo.org/QUARANTINE/insects/Spodoptera_ littoralis/F-spodli.pdf. 1010 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.75 – Epichoristodes acerbella (Walker, 1864) - African Carnation Tortrix (Lepidoptera, Tortricidae) Stanislav Gomboc Description and biological cycle: Small moth, 14–24 mm wingspan, female larger than male. Forewing ochreous yellow to brownish yellow, often with a darker band towards the distal edge; hindwings greyish-white (Photo left). Number of generations depends on temperature. In Africa, the moth has several generations yearly, without diapause. In southern Europe, there are 3–4 generations (April - October) outdoors and 4–5 generations in glasshouses. Generations are difficult to distinguish since all stages are present for most of the year. Female lays 200–240 eggs in groups of 80–120 eggs in a period of three days, on the upper side of the leaf. Eggs hatch after about ten days. Lower threshold of their development is about 6°C, but they are able to withstand lower temperatures in the hibernation period. Larva variable in colour, green, yellowish or grey, with darker dorsal and subdorsal lines (Photo- right). Pupation occurs after about 30 d and the pupal stage lasts eight d. The development from egg to adult is influenced by temperature: 11°C - 170 d, 17°C – 70 d, 20°C - 40 d. Host plants mainly include Dianthus and Chrysanthemum but also Pelargonium, Medicago, Lupinus, Lycopersicon, Rosa, Capparis, Pyrus, Malus, Prunus, Rhamnus, and some weeds such as Sonchus, Rumex, Oxalis, Carex, Erigeron, Ornitogalum and others. Native habitat (EUNIS code): Unknown. Habitat occupied in invaded range (EUNIS code): Mainly J100- glasshouses ; but also I1- Arable land and market gardens; I2- Cultivated areas of gardens and parks ; FB- Shrub plantations; Shrub plantations for ornamental purposes ; F5- Maquis, arborescent matorral and thermo-Mediterranean brushes. Native range: South Africa, Eastern Africa (Kenya) and Madagascar. Introduced range: Firstly reported in Europe in mid-1960s from glasshouses in Scandinavian countries. At present, regional distribution in France, Italian mainland, Sardinia, Sicily, Credit: Stanislas Gomboc Factsheets for 80 representative alien species. Chapter 14 1011 Spanish mainland, south England, Serbia and also in plantations in Germany and Danish mainland (Map). Intercepted many times on cut flower shipments in other European countries, but not yet established there. Pathways: Passive international transport (airplanes, vehicles) of cut flowers and ornamental plants is the quickest means of spread. Adults fly only on short distances but the moth can disperse in any of its development stage, early stages being hidden on or inside plant tissue. Impact and management: Important indoor and outdoor pest of cultivated, mainly ornamental plants. In European carnation cultivars, African Carnation tortrix may attack up to 90% of the crop; an important pest of Chrysanthemum and some field crops. Larvae are polyphagous and feed first on the leaf, under a shelter of silk, later in buds, flowers or stems. Young leaves are perforated and wilt and, more typically, stems are mined; flower buds are also perforated, become desiccated and petals are often woven together by silk. Difficult to control, due to hidden lifestyle. Spraying or fumigation with insecticides is still the best control method. Avoid importing plant material from infested areas or inspection of plants on arrival. The adults are nocturnal, and can be monitored by pheromone traps or by light traps; eggs, larvae and pupae by observation of presence on plants or plant damage. Biological control is still under investigation: possibly by using mating disruption with pheromones or by parasitoids like trichograms (Trichogramma dendrolimi, T. voegelei, T. dendrolimi). Selected references Costa Seglar M, Vives Quadras JM (1976) Epichoristodes acerbella Walk, nuevo tortrícido, plaga de los claveles, en la Península Ibérica. SHILAP Revista de Lepidopterologia 4: 233–234. Zangheri S, Cavalloro R (1971) Sulla presenza in Italia di Epichoristodes acerbella (Walker) (Lepidoptera Tortricidae). Bolletino della Società Entomologica Italiana 103: 186–190. 1012 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.76 – Grapholita molesta (Busck, 1916) - Oriental fruit moth (Lepidoptera, Tortricidae) Zdeněk Laštůvka Description and biological cycle: Wingspan 11–13 mm, body length 6–7 mm; small tortricid moth, forewing dark, greyish black, more or less distinct black transverse lines, oblique strigulae on the costa, black spots along distal margin and distinct light spot in the middle of the distal half of the wing (Photo left); very similar to the native European plum fruit moth (Grapholita funebrana); reliable determination possible only after genitalic dissection; oligophagous on Prunus s.l. spp. (peach, nectarine, apricot, almond, plum, cherry), on apple (Malus), pear (Pyrus), occasionally on quince (Cydonia), medlar (Mespilus), hawthorn (Crataegus), loquat (Eriobotrya japonica), Cotoneaster, Eugenia and Photinia; the species develops 2–4 generations per year following climatic conditions and adults are on wing between May and October; female lies about 200 eggs during its life lasting 10–14 days; eggs are whitish, flattened, oval in shape, 0.7–0.8 mm long; they are laid usually on the leaf underside, less often on new shoots or on fruits; larva of the first generation bores tunnels in terminal parts of young shoots; of following generations it lives usually in fruits (Photo right- frass exiting from infested fruit); mature larvae of the last generation overwinter in cocoons in crevices under bark or in the soil litter and they pupate in early spring. Native habitat (EUNIS code): G- Woodland and forest habitats and other wooded land. Habitat occupied in invaded range (EUNIS code): I1 - Arable land and market gardens; I2 - Cultivated areas of gardens and parks (fruit orchards, lines of fruit trees, fruit gardens, ornamental cultures). Native range: East Asia (China, Korean Peninsula, Japan). Introduced range: Introduced over the world mostly during the first three decades of the 20th century. Described as new species by Busck from the introduced range (USA, Virginia) in 1916. In Europe recorded for the first time in 1920 in Italy and France. Today present in Western, Central, Southern and Southeastern Europe (Map); not known from Poland, but very probably present, occasionally imported with fruit into more northern countries such as Credit: Rémi Coutin/ OPIE Factsheets for 80 representative alien species. Chapter 14 1013 Great Britain, Belgium, Denmark, Sweden, Lithuania, Latvia, Byelorussia, but probably not naturalized there. Also recoreded from other temperate and partly subtropical regions of the world: Southwestern Asia (Armenia, Azerbaijan, Georgia, southern Kazakhstan - possibly native, Uzbekistan - possibly native), Africa (Morocco, Southern Africa), North America (USA, southern Canada – Ontario, North Mexico), southern parts of South America (Argentina, southern Brazil, Chile, Uruguay), eastern half of Australia, and New Zealand. Pathways: Mostly passive transport of cocoons on dormant fruit-tree nursery stock and in containers with fruits, or directly with infested fruits. Dispersal at a local scale is realized by active flight of adults. Impact and management: The oriental fruit moth is one of the most important pests of stone and other fruit trees (especially on peaches and nectarines) causing considerable economic damage. Ecological impact is not known, but an influence on native parasitoid abundance and their trophic chains is possible. Monitoring is possible using pheromone traps. A number of insecticides were used for chemical control during the last decades (various organophosphates, pyrethroids, carbamates, neonicotinoids, insect growth regulators). Biological control possibilities include various kinds of bioagents tested or applied locally as granuloviruses, Bacillus thuringiensis, entomoparasitic nematodes (Steinernema and Heterorhabditis spp.); also hymenopteran (several Ichneumonidae, Pteromalidae, Trichogramma spp., etc.) and dipteran (Tachinidae) parasitoids. Mating disruption by synthetic sexual pheromones was largely used during recent years. Selected references Kyparissoudas DS (1989) Control of Cydia molesta (Busck) by mating disruption using Isomate-M pheromone dispensers in northern Greece. Entomologia Hellenica 7: 3–6. Paoli G (1922) Un lepidottero nuovo per la fauna italiana (Laspeyresia molesta Busck). Bollettino della Società Entomologica Italiana 54: 122–126. 1014 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.77 – Argyresthia thuiella (Packard, 1871) - Arborvitae leaf miner (Lepidoptera, Yponomeutidae) Ferenc Lakatos Description and biological cycle: Tiny moth, adults with wingspan of 4–6mm. Forewings silvery, tip brownish. Larvae rosy with black head. Females lay eggs after mating on the foliage in June (Photo left - detail of an egg). After hatching, larvae enter the leaves, where they feed, overwinter and also pupate. Larva starts feeding at the tip of branch towards trunk. Branch tip becomes yellowish (Photo right), later brown. This species has one generation both in Europe and in the native area (Eastern North-America). Adults fly around the host trees, different Thuja species, during the daytime. Native habitat (EUNIS code): G- Woodland and forest habitats and other wooded land. Habitat occupied in invaded range (EUNIS code): G5- Lines of trees; I2- Cultivated areas of gardens and parks. Native range: North America. Introduced range: Supposedly introduced three times independently (1971: the Netherlands; 1975: Germany; 1976: Austria). Argyresthia thuiella expanded its distribution in the Credit: Hana Šefrová Factsheets for 80 representative alien species. Chapter 14 1015 last decades to most of continental Europe, except Scandinavia and Iberian Peninsula (Map). However damage caused by this species has decreased with this expansion. Pathways: Probably trade of ornamental Cupressaceae. Impact and management: Damage was important only during his expansion phase after establishment. At present, frequent in urban areas such as parks, gardens and urban forests. Several parasitoids were reared from different developmental stages already at the start of this moth’s presence in Europe (e.g. Pteromalidae, Eulopidae and Braconidae). After establishment, chemical suppressants were widely used, but as damage decreased so did the need for control. Attractants are known and available for the members of the genus Argyresthia, but not so far used for monitoring or mass trapping. Selected references Frankenhuyzen Av (1974) Argyresthia thuiella (Pack.) (Lep., Argyresthiidae). Entomologische Berichten 34: 106–111. Škerlavaj V, Munda A (1999) Argyresthia thuiella Packard - a new pest of Thuja in Slovenia. Zbornik predavanj in referatov 4. slovenskega posvetovanja o varstvu rastlin v Portorožu od 3. do 4. Marca 1999, Ljubljana: Društvo za varstvo rastlin Slovenije, p. 451. 1016 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.78 – Frankliniella occidentalis (Pergande, 1895) - Western flower thrips (Thripidae, Thysanoptera) Alain Roques Description and biological cycle: Tiny, slender thrips with narrow fringed wings (Photo). Males, 1.2–1.3 mm long, are pale yellow, females, 1.6–1.7 mm long, are yellow to brown; larvae are yellowish-white. Adults and larvae suck plant fluids from flowers and leaves of at least 244 plant species from 62 families. Western flower thrips reproduces in glasshouses with 12–15 generations/year. Overall life cycle lasts from 44 d at 15 °C to 15 d at 30 °C. A female can lay 20–40 eggs. Unmated females produce males. Different developmental stages are typically found in different parts of plants: eggs in leaves, flower tissue and fruits; nymphs on leaves, in buds and flowers; pupae in soil or in hiding places on host plants such as the bases of leaves; adults on leaves, in buds and flowers. Native habitat (EUNIS code): I - Regularly or recently cultivated agricultural, horticultural and domestic habitats. Habitat occupied in invaded range (EUNIS code): I1- Arable land and market gardens; J100 - glasshouses. Native range: North America. Introduced range: Reported from all continents; first record in Europe in 1983 in the Netherlands; continuous and rapid spread since the 1980s; present in glasshouses in North and central Europe, but already outdoors in Southern Europe (Map). Pathways: Intercontinental dispersal of eggs, larvae and adults is taking place with the trade of ornamental plants (e.g., cut flowers, potted plants). Adults can be easily carried by winds, but also by clothes, equipment and containers not properly cleaned. Credit: Philppe Reynaud Factsheets for 80 representative alien species. Chapter 14 1017 Impact and management: An outdoor pest as well as a glasshouse pest. Flowers and foliage of a great number of economically important crops are affected, in glasshouses as well as outdoors. On ornamental flower crops, feeding induces discolouration, indentation, distortion and silvering of the upper leaf surface as well as scarring and discolouration of petals and deformation of flower heads, largely reducing their economic value. In orchids, eggs laid in petal tissues cause a ‘pimpling’ effect on flowers. This thrips also kills or weakens terminal buds and blossoms in fruit trees (e.g., apricot, peach) and roses, and on most fruiting vegetables, especially cucumbers. In addition, nymphs are vectors of tobacco streak ilarvirus (TSV) and tomato spotted wilt virus (TSWV), which is inducing severe diseases in ornamental and vegetable crops in Europe. Blue sticky traps can be used to detect initial infestation and to monitor adult population levels. Chemical control is difficult because this thrips is resistant to most pesticides and feeds deep within the flower or on developing leaves. Biological predatory mites (e.g., Neoseiulus cucumeris, Amblyseius spp. and Hypoaspis spp.) and minute pirate bugs (e.g., Orius laevigatus, O. insidiosus) provide effective biological control, in glasshouses. Selected references Del Bene G, Gargani E (1989) [A contribution to the knowledge of Frankliniella occidentalis] (in Italian). Redia 72: 403–420. EPPO/CABI (1997). Frankliniella occidentalis. Quarantine Pests for Europe. Wallingford, United Kingdom: CAB International, 267–272. Mantel WP (1989) Bibliography of the western flower thrips, Frankliniella occidentalis. Bulletin SROP 12: 29–66. 1018 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.79 – Pseudodendrothrips mori (Niwa, 1908) - Mulberry thrips (Thripidae, Thysanoptera) Philippe Reynaud Description and biological cycle: Oligophagous thrips with small to minute and very pale bodies. Males 0.7–0.9 mm long; females 0.9–1.1 mm long. Mulberry thrips is a member of the Dendrothripinae subfamily distinguished from other Thripidae by the presence of a remarkably elongate metasternal endofurca* providing active jumping capacities (Photo left). Pseudodendrothrips mori commonly breeds on Morus alba and M. bombycis or on Ficus throughout the world and is a widespread although minor pest of Morus (Photo right - damage on leaf). Mulberry thrips reproduces outdoors with 6–10 generations per year in its native area. In mid-March, overwintering adults emerge to damage the leaves. The complete life cycle lasts 26–34 days in spring and autumn, and 16–23 days in summer, depending on conditions. Increased temperature (26–32˚C) directly inﬂuence the breeding activity of the thrips and thereby increases the population levels. July and August are peak months of occurrence; the ﬁnal stage of appearance is in late autumn. Adults overwinter after October, thus completing the annual life cycle. Seasonal population ﬂuctuations and the degree of damage caused to the host plant are inﬂuenced by various environmental factors, including climate, host plant variety, topography, soil type and management regimes. Native habitat (EUNIS code): Unknown. Habitat occupied in invaded range (EUNIS code): I2- Cultivated areas of gardens and parks. Native range: Probably originated from China or Japan. Introduced range: Recorded from several parts of the world, including USA, India, Iran and Australia. First recorded in 1974 in Italy; at present known only in few more countries (Spain and France). The rate of spread seems to be low. Credit: Georgi Trenchev and Katia Trencheva Factsheets for 80 representative alien species. Chapter 14 1019 Pathways: Probably trade of ornamental plants. Impact and management: Mulberry leaves are the exclusive food of the silkworm, Bombyx mori. Mulberry thrips have caused serious damage to sericulture in the southern states of India and China while damage to this industry is also reported in Sri Lanka and Vietnam. Feeding silkworms with mulberry leaves that have been damaged by mulberry thrips causes slower development, increases larval mortality and reduces cocoon yield. In other countries such as Japan, Iran and some countries in Europe and America, production of silk is very limited. Here mulberry is mainly an ornamental tree which is grown by roads because of its low need for water and nutrients, and P. mori can damage plants growing in these situations. Chemical control is the main method used to control P. mori in silk production areas. However, it is assumed that in countries where Morus are used as ornamental plants, damage by the pest could be mitigated using non-chemical methods which are economically or ecologically tolerable. Selected references Cappellozza L, Miotto F (1975) Pseudodendrothrips mori (Niwa) (Thysanoptera Terebrantia) specie nuova per la fauna Italiana. Redia 56: 387–389. Trenchev G, Trencheva K (2007)Pseudodendrothrips mori Niwa (Thysanoptera, Thripidae), a species new to the Bulgarian fauna. Zachita na Rastenija 18: 68–71. Vierbergen G, Cean M, Szeller IH, Jenser G, Masten T, Simala M (2006) Spread of two thrips pests in Europe: Echinothrips americanus and Microcephalothrips abdominalis (Thysanoptera: Thripidae). Acta Phytopathologica et Entomologica Hungarica 41: 287–296. 1020 Edited by Alain Roques & David Lees / BioRisk 4(2): 855–1021 (2010) 14.80 – Thrips palmi (Karny, 1925) - Melon thrips (Thripidae, Thysanoptera) Philippe Reynaud Description and biological cycle: Completely yellow thrips (Photo). Males 0.9–1.0 mm long, females 1.1–1.3 mm long. Identification is hampered by small size and a great similarity with other yellow species of Thrips. Melon thrips is a polyphagous feeder and an outdoor pest of aubergine (Solanum melongena), sweet pepper (Capsicum annuum), cotton (Gossypium spp.) cowpea (Vigna unguiculata), cucumber (Cucumis sativus), Cucurbita spp., melon (Cucumis melo), pea (Pisum sativum), Phaseolus vulgaris, potato (Solanum tuberosum), sesame (Sesamum indicum), soyabean (Glycine max), sunflower (Helianthus annus), tobacco (Nicotiana tabacum) and watermelon (Citrullus lanatus). In glasshouses, economically important hosts are aubergine, Capsicum annuum, Chrysanthemum, cucumber, Cyclamen, Ficus and Orchidaceae. To develop from egg to adult, Thrips palmi requires 194 day-degrees above a thermal threshold of 10.1°C, and takes between 10 days (at 30°C) to 40 days (at 15°C) to complete its life-cycle which is lengthened to 80 days when the insects are at 13°C. Melon thrips are able to multiply during any season that crops are cultivated, but are favoured by warm weather. Native habitat (EUNIS code): I1- Arable land and market gardens. Habitat occupied in invaded range (EUNIS code): I- Regularly or recently cultivated agricultural, horticultural and domestic habitats; J- Constructed, industrial and other artificial habitats. Native range: First described in 1925 in Sumatra but remained little known, often overlooked and the subject of taxonomic confusion until 1980. Introduced range: From the late 1970s, Melon thrips has spread across the Far East and in subsequent decades within South East Asia, and to Australia, the Paciﬁc, Florida, the Car- Credit: LNPV Factsheets for 80 representative alien species. Chapter 14 1021 ibbean, South America and West Africa. In Europe, numerous interceptions have been reported on cut ﬂowers and fruit vegetables from Thailand, Mauritius, India, etc. Several limited outbreaks were found in glasshouses in Netherlands and in Great Britain since 1988, but all these outbreaks were eradicated. May-be still present in glasshouses of Norway and the Czech Republic. T. palmi is considered to be absent outdoors in Europe although it was detected in ﬂowers of kiwi fruit (Actinidia deliciosa) plantations in Portugal in 2004; in later surveys the pest was no longer found. Pathways: Trade of plant material (ornamentals, vegetables, fruits). Impact and management: Melon thrips cause severe injury to infested plants. Leaves become yellow, white or brown, and then crinkle and die. Heavily infested fields sometimes acquire a bronze color. Terminal growth damage occurs via discolouration, stunting, or deformation. Fruits may also be damaged with scars, deformities and abortion. T. palmi has been shown to transmit plant viruses including Groundnut bud necrosis virus (GBNV), Melon yellow spot virus (MYSV), Watermelon silver mottle virus (WSMoV), Tomato spotted wilt virus (TSWV) and Capsicum chlorosis virus (CaCV). However, this list is questionable due to lack of consistent studies. Experience of controlling or eradicating T. palmi has been gained in a large number of countries as this pest has spread around the world. However, melon thrips requires frequent spraying of insecticides, so resistance to many chemicals has developed. It is now considered that control with insecticides alone is not adequate. Integrated pest management is necessary, including cultural practices and biological control. Selected references Anonymous (2004). First report of Thrips palmi in Portugal. EPPO Reporting Service 144: 2. Cannon RJC, Matthews L, Collins DW, Agallou E, Bartlett PW, Walters KFA, MacLeod A, Slawson DD, Gaunt A (2007) Eradication of an invasive alien pest, Thrips palmi. Crop Protection 26: 1303–1314. A peer reviewed open access journal BioRisk 4(2): 1023–1028 (2010) BioRisk RESEARCH ARTICLE doi: 10.3897/biorisk.4.71 www.pensoftonline.net/biorisk Abbreviations and glossary of technical terms used in the book Alain Roques1, David Lees2 1 Institut National de la Recherche Agronomique (INRA), UR 0633, Station de Zoologie Forestière, 2163 Av. Pomme de Pin, 45075 Orléans, France 2 INRA UR633 Zoologie Forestière, 2163 Av. Pomme de pin, 45075 Orléans, France Corresponding author: Alain Roques (alain.roques@orleans.inra.fr) Received 30 May 2010 | Accepted 7 June 2010 | Published 6 July 2010 Citation: Roques A (2010) Abbreviations and glossary of technical terms used in the book. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(2): 1023–1028. doi: 10.3897/biorisk.4.71 Appendix I Country codes abbreviations used in the book according to the International Organization for Standardization list ISO 3166. http://www.iso.org/iso/english_country_ names_and_code_elements. Abbreviation AD AL AT BA BE BG BY CH CY CZ DE DK EE ES ES-BAL ES-CAN FI Country/ Island Andorra Albania Autriche Bosnia and Herzegovina Belgium Bulgaria Belarus Switzerland Cyprus Czech Republic Germany Denmark Estonia Spain Spain - Baleares islands Spain - Canary islands Finland Abbreviation FI-ALN FÖ FR FR-COR GB GI GL GR GR-CRE GR-ION GR-NEG GR-SEG HR HU IE Country/ Island Finland - Aland Faroe islands France France - Corsica island United Kingdom Gibraltar Greenland Greece Greece - Crete Greece - Ionian islands Greece - North Aegean islands Greece - South Aegean islands Croatia Hungary Ireland Copyright A. Roques, D. Lees This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 1024 Alain Roques / BioRisk 4(2): 1023–1028 (2010) Abbreviation Country/ Island Israel Iceland Italy Italy - Sardinia island Italy - Sicily island Liechtenstein Lithuania Luxembourg Latvia Moldova Montenegro Macedonia Malta Netherlands IL IS IT IT-SAR IT-SIC LI LT LU LV MD ME MK MT NL Abbreviation NO NO-SVL PL PT PT-AZO PT-MAD RO RS RU SE SI SK UA YU Country/ Island Norway Norway - Svalbard Poland Portugal Portugal - Azores islands Portugal - Madeira island Romania Serbia Russia (European Part) Sweden Slovenia Slovakia Ukraine Former Yugoslavia Appendix II Habitat abbreviations used in the book according to the European Nature Information System (EUNIS) database. http://eunis.eea.europa.eu Code A Habitat Marine habitats B B1 B2 B3 Coastal habitats Coastal dune and sand habitats Coastal shingle habitats Rock cliffs, ledges and shores, including the supralittoral C C1 C2 C3 Inland surface water habitats Surface standing waters Surface running waters Littoral zone of inland surface waterbodies D D1 D2 D3 D4 D5 D6 Mire, bog and fen habitats Raised and blanket bogs Valley mires, poor fens and transition mires Aapa, palsa and polygon mires Base-rich fens Sedge and reedbeds, normally without free-standing water Inland saline and brackish marshes and reedbeds E E1 E2 E3 E4 E5 Grassland and tall forb habitats Dry grasslands Mesic grasslands Seasonally wet and wet grasslands Alpine and subalpine grasslands Woodland fringes and clearings and tall forb habitats Taxonomy, time and geographic patterns. Chapter 2 1025 Code E6 E7 Habitat Inland saline grass and herb-dominated habitats Sparsely wooded grasslands F F1 F2 F3 F4 F5 F6 F7 F8 F9 FA FB Heathland, scrub and tundra habitats Tundra Arctic, alpine and subalpine scrub habitats Temperate and mediterraneo-montane scrub habitats Temperate shrub heathland Maquis, matorral and thermo-Mediterranean brushes Garrigue Spiny Mediterranean heaths (phrygana, hedgehog-heaths and related coastal cliff vegetation) Thermo-Atlantic xerophytic habitats Riverine and fen scrubs Hedgerows Shrub plantations G G1 G2 G3 G4 G5 Woodland and forest habitats and other wooded land Broadleaved deciduous woodland Broadleaved evergreen woodland Coniferous woodland Mixed deciduous and coniferous woodland Lines of trees, small anthropogenic woodlands, recently felled woodland, early-stage woodland and coppice H H1 H2 H3 H4 H5 H6 Inland unvegetated or sparsely vegetated habitats Terrestrial underground caves, cave systems, passages and waterbodies Screes Inland cliffs, rock pavements and outcrops Snow or ice-dominated habitats Miscellaneous inland habitats with very sparse or no vegetation Recent volcanic features I I1 I2 Regularly or recently cultivated agricultural, horticultural and domestic habitats Arable land and market gardens Cultivated areas of gardens and parks J J1 J2 J3 J4 J5 J6 J100 Constructed, industrial and other artificial habitats Buildings of cities, towns and villages Low density buildings Extractive industrial sites Transport networks and other constructed hard-surfaced areas Highly artificial man-made waters and associated structures Waste deposits Greenhouses X X6 X7 Complex habitats Crops shaded by trees Intensively-farmed crops interspersed with strips of spontaneous vegetation 1026 Code X10 X11 X13 X14 X15 X16 X20 X22 X23 X24 X25 Alain Roques / BioRisk 4(2): 1023–1028 (2010) Habitat Mixed landscapes with a woodland element (bocages) Large parks Land sparsely wooded with broadleaved deciduous trees Land sparsely wooded with broadleaved evergreen trees Land sparsely wooded with coniferous trees Land sparsely wooded with mixed broadleaved and coniferous trees Treeline ecotones Small city centre non-domestic gardens Large non-domestic gardens Domestic gardens of city and town centres Domestic gardens of villages and urban peripheries Glossary of the technical terms used in the book (marked by *) Alatae: winged forms in aphids, adelgids, and other hemipterans. Ampelophagous: related to the grapevine. Anholocyclic: in cyclically parthenogenetic organisms, life cycles that do not include a sexual generation (e.g., in adelgids). Archegonia: female multicellular egg-producing organ occurring in mosses, ferns, and most gymnosperms. Archeozooan: an alien animal introduced to Europe since the beginning of the Neolithic agriculture but before the discovery of America by Columbus in 1492 (Daisie 2009). Arrhenotoky: a common form of sex-determination in Hymenoptera and some other invertebrates, in which progeny are produced by mated or unmated females, but fertilized eggs produce diploid female offspring, whereas unfertilized eggs produce haploid male offspring by parthenogenesis (only the females are biparental). Carina (sg.), Carinae (pl.): a ridgelike structure (e.g. antennal longitudinal ridge). Cercus (sg.), Cerci (pl.): paired sensory structures at the posterior end of some arthropods. Clava: apically differentiated region (sometimes club-like) of the antennal flagellum. Dealate: having lost its wings; used for ants and other insects that shed their wings after the mating flight. Declivity: posterior portion of the elytra that descends to its apex. Domestic: living in human habitats. Endofurca: the internal skeleton of the meso-and metathorax, that provides important muscle insertion points. In some thrips, the metasternal endofurca provides the insertion for powerful muscles that are associated with a remarkable jumping ability of adults. Endophytic (adj): living inside a plant. Endopterygote: insect that undergoes complete metamorphosis, with the larval and adult stages differing considerably in their structure and behaviour. Epigyne: the external female sex organ in arachnids. Exarate: for a pupa, having the appendages free and not attached to the body (as opposed to Obtect). Taxonomy, time and geographic patterns. Chapter 2 1027 Exopterygote: insect that undergoes incomplete metamorphosis. The young (called nymphs) resemble the adults but lack wings; these develop gradually and externally in a series of stages or instars until the final moult produces the adult insect. There is no pupal stage. Flagellum: the part of the antenna beyond the pedicel, which is differentiated into three regions, the anellus, funicle and clava. Frass: waste material produced by feeding insects, including excrement and partially chewed vegetation. Funicle: region of the antennal flagellum between the anellus and clava. Gallicolae: leaf gall making forms; e.g., in phylloxerans. Gnathosoma: anterior body region in mites. Halobiont: an organism that lives in a salty environment. Hemimetabolous: the type of insect development in which there is incomplete or partial metamorphosis, typically with successive immature stages increasingly resembling the adult; see Exopterygote. Holocyclic: in cyclically parthenogenetic organisms, life cycles that include a sexual generation (e.g., in adelgids). Holoptic: as in flies, with compound eyes meeting along the dorsal midline of the head. Hyperparasitoid: a parasitoid living on or in another parasitoid. Idiobiont parasitoid: a parasitoid which prevents further development of the host after initial parasitization. Idiosoma: abdomen of mites and ticks. Kleptoparasitoid: a parasitoid which preferentially attacks a host that is already parasitized by another species. Koinobiont parasitoid: a parasitoid which allows the host to continue its development and often does not kill or consume the host until the host is about to either pupate or become an adult. Ligula: the apical lobe of the labium. Megagametophyte: female haploid, gamete-producing tissue in conifers. Mesothorax: the second, and usually the largest, of the three primary subdivisions of the thorax in insects. Mesonotum: the dorsal part of the mesothorax. Metathorax: the third of the three primary subdivisions of the thorax in insects. Metanotum: the dorsal part of the metathorax. Moniliform: bead-like (as in antennae). Mycangium (sg.), mycangia (pl.): usually complex structures on the insect body that are adapted for the transport of symbiotic fungi, usually spores. Neozooan: an alien animal introduced to Europe after the discovery of America by Columbus in 1492 (Daisie 2009) . Notaulix (sg.), Notaulices (pl.): one of a pair of grooves on the mesoscutum, from the front margin to one side of the midline and extending backward; divides the mesoscutum into three parts. Obtect: for a pupa, having the legs and other appendages fused to the body. Oniscomorph: the state as in ‘pill’ millipedes of being able to roll up in a ball. Opisthosoma: posterior part of the body in spiders and mites. 1028 Alain Roques / BioRisk 4(2): 1023–1028 (2010) Paranota: lateral wings. Parthenogenesis, parthenogenetic (adj.): the production of offspring from unfertilized eggs. Special cases of this state are arrhenotoky, pseudo-arrhenotoky, and thelytoky. Phytoplasma: prokaryotes that are characterized by the lack of a cell wall, associated with plant diseases. Phytotelmatum (sg.), Phytotelmata (pl.): a small, water-filled cavity in a tree or any similar environment. Podosoma: anterior section of idiosoma in ticks; serving as connecting area for the four pairs of legs. Porrect: extended, especially forward; e.g., porrect mandibles. Proctiger: the reduced terminal segment of the abdomen which contains the anus. Prognathous: with the head more or less in the same horizontal plane as the body, and the mouthparts directed anteriorly. Pronotum: the dorsal part of the prothorax. Propodeum: the first abdominal segment. Prosoma: anterior part of the body in spiders and mites; also called cephalothorax. Prothorax: The first of the three primary subdivisions of the thorax in insects. Pseudo-arrhenotoky: A form of sex-determination (especially in some scale insects and mites) in which males and females arise from fertilized eggs and are diploid. However, males become haploid by inactivation of the paternal genomic complement. Puparium (sg.), puparia (pl.): the enclosing case of a pupa. Reticulate: net-like, anastomosing. Rostrum: beak-shaped projection on the head; e.g., in weevils. Scutellum: the middle region of the mesonotum or metanotum, behind the scutum. Scutum: the anterior part of the mesonotum or metanotum. Secondary pest: a pest that attacks only weakened plants. Sensorium: sensory structure present on antenna. Siphunculi, siphuncular (adj.): pair of protruding horn-shaped dorsal tubes in aphids which secrete a waxy fluid. Spatula sternalis: median cuticular sclerite, often bilobed, on the ventral side of the prothoracic segment of the last instars of some midge larvae; plays a role in larval locomotion. Stigma: conspicuous, usually melanised area at the apex of a vein of the forewing, generally at the leading wing edge. Sulcate: having narrow, deep furrows or grooves. Synanthropic: ecologically associated with humans. Tegula: Small, typically oval sclerite that covers the region of the mesothorax where the forewing and thorax articulate. Thelitoky: A form of sex-determination (especially in Hymenoptera: Symphyta and Cynipidae) in which only diploid female progeny are produced by parthenogenesis. Termen: distalmost edge of wing. Transhumance: in the case of hives, moving to new environments, according to the change in season. Xylophagous (adj.): feeding on wood. Index of the latin names of the arthropod species mentioned in the book Index of the latin names of the arthropod species mentioned in the book abdominale 362 abdominalis 770, 776, 778, 780, 783, 789 abietiperda 599 abietis 598, 762 abietorum 394 abnormis 719, 726 absoluta 608, 628, 633, 638, 643 acaciaebaileyanae 519, 542, 547 acaciella 642 acarisuga 559, 565, 567, 577, 587 acaudaleyrodis 729 acerbella 624, 640, 656, 757, 857, 1010, 1011 acerifoliae 455, 463 achaeae 757 acoreensis 147 acrenulatus 700, 726 acroxantha 628, 647 aculeata 144, 698, 986 aculeatus 215, 755 acuminata 385, 654 acuminatus 196, 198, 201, 204, 206, 212, 215, 375 392 acupunctatus 229, 234, 254 acuta 609, 646 acutangulus 352 acutellus 665 acuticollis 398 adansoni 146 adenocarpi 415, 432 admes 173, 187 adusta 579, 591 adustella 607, 625, 641 adustus 396 advena 374 aechmeae 501 aegeria 629, 663 aegyptium 165, 166, 175, 176, 182, 829 aenea 385 aeneopiceus 257 aenescens 557, 567, 580, 581, 593 aeneus 396 aequalis 321, 351 aequidentellus 658 aequinoctiale 333, 343 aestiva 395 afer 518, 540, 744 affine 250, 344 affinis 253, 352 africanus 366 agathidioides 353 agavium 478, 487, 505 agraensis 736 ainsliei 742 alashanica 672, 751 albiceps 555, 561, 578, 586 albicornis 754, 827 albida 201, 215 albifrons 446, 454, 455, 466 albipes 62, 71, 387, 750 albomaculatum 745 albopictus 12, 34, 41, 42, 58, 63, 65, 67, 71, 559, 561, 563, 563, 566, 567, 571, 573, 577, 579, 580, 581, 584, 589, 857, 918, 919 albosquamosa 258 albosquamosus 258 albothoracica 802 albus 504 algeriensis 624, 656 alienus 761 allotrichus 154, 162, 163, 179 alluaudi 84, 85, 87, 89, 96, 251, 749 alnivagrans 184 aloineae 745 alopecuri 472 alpestris 154, 161, 191 alpina 218, 762 alternecoloratus 751 amaranthi 181 amaryllidis 509 amasiella 647 ambrosiae 160, 161, 178 americana 34, 275, 282, 283, 292, 825 americanum 344 americanus 152, 183, 478, 494, 508, 777, 778, 779, 780, 782, 783, 786, 787, 812, 822, 827 americensis 742 amicula 398 amoena 573, 583, 590 ampellophaga 638, 668 ampelophaga 290 amphibola 765 amygdali 265 analis 398 anatolicum 166, 168, 183 ancylivorus 706, 734 andrei 714, 747, 853 andrenaeformis 619, 638, 665 andromedae 464 Alain Roques et al. (Eds) / BioRisk $$: @@–@@ (2010) angelicus 741 anguinus 262 angulata 765 angulipennis 648 angustatus 263, 387 angustiformis 403 angustifrons 489, 508, 738 angustisetulus 261 angustulus 384 angustum 358 angustus 340, 380 anneckei 738 annularis 196, 199, 200, 214 annulatus 736 annulipes 829 anthobia 385 antirrhini 393 antoinei 248 aonidiellae 588 aonidum 477, 486, 487, 499, 697, 732 apenninus 239, 243, 259 aphodioides 405 apicalis 388, 429 araucariae 478, 488, 505 arbuti 474 archon 16, 57, 61, 66, 71, 604, 608, 612, 615, 619, 624, 626, 626, 632, 634, 636, 642, 857, 994, 995 arcuata 414, 415, 423, 430, 857, 966, 967 arenarius 248 areolatus 754 argus 365 argus 294, 295, 302, 303, 313 aridicola 258 arietis 217 armadillo 259 armatus 238, 264 armatus 741 armillatus 266 aroidephagus 517, 540, 542, 545 arsona 559, 568, 576, 593 artemisiae 252 arundicolens 472 arundinariae 472 arvicola 218 ascalonicus 445, 450, 468 asellus 82 asiaticum 444, 452, 465 asparagi 276, 291, 436 asper 217 aspidistrae 481, 503 assimile 82, 85 assimilis 257 asynamorus 812, 818, 827 atedius 756 ater 263, 356 aterrima 764 atlanta 145 atomosella 655 atramentaria 398 atratus 238, 253, 516, 535, 544 atricapillus 386 atricapitella 617, 661 atropalpus 34, 42, 571, 580, 590 atropos 810, 814, 823 attelaboides 266 attemsi 104, 117, 118, 127 attenuatus 241, 253, 264 aubei 264, 265 aucubae 516, 518, 544 aulacaspidis 735 aulicus 385 aurantii 69, 446, 486, 499, 581, 588, 692, 697, 707, 711, 727, 729, 737, 951, 973 auratus 386 aurella 617, 661 aurifer 243, 259 australasiae 818, 820, 825 australicus 365 australiensis 745 australis 354, 790 autumnalis 144 axillaris 801 axyridis 12, 30, 32, 36, 39, 42, 62, 64, 65, 67, 70, 71, 71, 295, 297, 298, 300, 302, 303, 304, 305, 306, 307, 308, 309, 310, 312, 313, 857, 886, 887 azaleae 155, 181, 449, 464, 515, 546 azaleella 612, 644 azimi 729 bacoti 153, 155, 164, 169, 184 bagdasariani 155, 178 bakeri 453, 468 balssi 94 balteatus 321, 367 bambusae 188, 466, 479, 488, 496, 506 banksi 162, 171, 187 barbifrons 573, 595 beckii 481, 487, 502, 692, 710, 722, 727, 732 begini 743 bella 388, 596 berberidis 672, 759 bergrothi 394 berlesei 704, 717, 729, 955 Index of the latin names of the arthropod species mentioned in the book beszedesi 400 bicarinatum 707, 750 bicinctum 250 bicinctus 345, 788 biclavis 502, 786 bicolor 187, 250, 356, 393, 394, 676, 735, 802 bifasciata 364, 692, 707, 711, 737 bifasciatus 380, 390 bifenestratus 368 biformis 771, 790 bifoveolatus 347 biguttata 395 bihamata 400 bilimeki 707, 749 bilinealis 628, 652 bilineatus 376 billeni 784, 787 bilobatus 153, 155, 157, 164, 178 binotatus 385, 525 bioculatus 413, 425, 430 bipunctatus 361 bipustulata 397 bipustulatum 393 bischoffi 226, 230, 239, 248 bisonia 524, 526, 528, 529, 533, 534, 551, 857, 960, 961 bisselliella 610, 616, 618, 655 bituberculatum 252 bivari 795, 803 bivittatus 146, 724, 728 blanchardi 296, 503 bleusei 358 bodoanus 225, 235, 251 boisduvalii 500, 737 bondari 518, 546 boninsis 506 borgesi 416, 429 borinquenensis 564, 594 borriesi 756 borzi 598 bostrychoides 350 bostrychophila 798 bourbonica 748 brachypterus 404 brasiliensis 361, 440, 451, 462, 849 brassicae 30, 277, 283, 288, 687, 691, 765 brevicollis 290 brevicornis 258, 401, 741 brevicruris 509 brevilabiatus 105, 124 brevipennis 209, 212, 213, 218 brevipes 378, 506 brevirostris 423, 432, 812, 827 brevis 811, 817, 818, 819, 820, 821, 826 brevis 763 brevisetosa 155, 162, 163, 182 brevispinosa 746 briggsi 798, 800 brimblecombei 519, 543, 548 britannica 660 bromeliae 477, 500 brumata 633, 660 brunnea 194, 196, 198, 204, 210, 215, 802, 821, 825 brunneus 322, 331, 334, 366, 374, 390 buddleia 567, 568, 586 buoliana 668, 693, 743 buqueti 321, 351 bursa 153, 155, 157, 164, 168, 170, 171, 184 busckii 591 busseolae 707, 753 butleri 410, 415, 417, 431 buxi 503 byersi 178 cachectus 260 cacticans 508 caelatus 347 caerulescens 392 caesus 384 calandrae 693, 752 calceolariae 478, 486, 489, 507, 706, 721, 741 caldaria 123 caldarium 714, 762 caledonicus 600 caliberberis 178 calidella 627, 632, 664 californica 438, 464, 719, 753 californicus 149, 156, 159, 169, 186, 353, 690, 732, 754 californiensis 470 califraxini 161, 182 camelliae 170, 180, 503 cameroni 708, 719, 753 campestris 31, 43, 99, 189, 197, 200, 203, 410, 428, 615, 994 cancellatus 386 candefacta 634, 646 canestrinii 191 caniola 658 canis 835, 847 capense 336, 362, 531 capensis 393, 728 capitata 55, 558, 560, 563, 565, 567, 568, 569, 577, 579, 596, 699, 699, 713, 719, 857, 896, 920, 921 Alain Roques et al. (Eds) / BioRisk $$: @@–@@ (2010) caprai 413, 432 caprealis 664 caraganae 441, 460 cardinalis 59, 294, 296, 297, 298, 300, 304, 311, 479 carinatus 162, 180, 217 carinthiaca 290 carnaria 371 carnesi 732 carnica 759 carnivorus 356 carpophagata 659 carpophagus 226, 251 carus 351 caryae 467 caryella 451, 467 caspius 349 castanea 803 castaneum 320, 323, 328, 329, 331, 344, 381 castaneus 263 castanoptera 398 catalpae 461, 915 cataractae 83 cattleyae 560, 564, 567, 579, 586 cattleyana 322, 367 caucasicus 227, 244, 265, 354 caudatus 187 caularum 360 cautella 649 cavasolae 743 cavelli 324, 373 cavicollis 366 cayennensis 211, 216 cedri 440, 445, 447, 450, 463 cedricola 324, 626, 631, 656 cedrivora 624, 657 cedrobii 699, 714, 734 celarius 162, 174, 189 cellaria 82 cellaris 323, 352 celtis 162, 175, 181 cembrae 238, 264, 522 cementarium 754 centifoliella 617, 661 cephalonica 649 cephalotes 402, 753 cephi 734 ceratoniae 632, 664 cerealella 608, 631, 643 cerealium 780, 791 ceriferus 483, 494, 496 ceroplastae 728, 742 ceroplastis 736 cerricola 760 cervantesi 715, 737 chalcites 609, 662 chamaecypari 156, 158, 174, 185 cheopis 18, 833, 836, 837, 839, 849 chilensis 414, 430, 690, 707, 726, 951 chilonis 757 chinensis 21, 32, 57, 61, 65, 69, 71, 196, 203, 204, 205, 207, 208, 214, 270, 275, 276, 287, 732, 578, 585, 857, 876, 877 chionaspidis 725 chittendeni 517, 539, 545 chnumi 592 chrysanthemella 607, 624, 636, 642 chrysanthemi 163, 181, 561, 563, 589 chrysippus 606, 609, 638, 647 chrysocephalus 290, 835, 846 chrysomelinus 404 cidariae 743 ciliata 57, 414, 420, 425, 427, 430, 857, 966, 968, 969 cincticornis 812, 820, 827 cinctipennis 743, 1003, 1009 cinerea 810, 814, 823 cinereus 600 cingulata cingulata 101, 129, 575, 578, 583, 596 cinnamomeus 260 circellaris 404 circulatus 258 circumflexus 469 circumvagans 823 citrella 55, 608, 626, 632, 633, 644, 691, 692, 717, 720, 736, 743, 743 citri 55, 157, 162, 163, 169, 170, 171, 174, 177, 188, 296, 298, 308, 309, 310, 477, 479, 481, 486, 489, 507, 541, 545, 561, 562, 563, 568, 575, 576, 581, 586, 610, 618, 631, 657, 684, 708, 712, 721, 722, 731, 738 citricidus 446, 448, 452, 474, 857, 942, 943 citricola 717, 722, 736 citricolus 518, 547 citrifolii 162, 517, 540, 546 citrina 308, 477, 493, 499, 730 clarus 805 clauseni 740 clavicornis 375, 398 clavipes 345 clisiocampae 725, 744 coarctata 104, 127, 762 cocciferae 599 Index of the latin names of the arthropod species mentioned in the book coccineus 478, 485, 505 coccus 476, 494, 498 cockerelli 494, 502, 504 coeculum 82 coeculus 103, 104, 117, 125 coffeae 221, 225, 233, 246, 478, 481, 498, 506, 710 cognata 396 coheni 697, 726 colchici 835, 847 colchicus 835, 840, 847 colemani 683, 733, 933, 939 collaris 397 colon 371 coloratus 391 columba 723, 754 complanatus 349 completa 562, 568, 569, 574, 575, 579, 581, 596, 857, 922, 923 compressa 764 concinna 378 concinnus 402, 405 concolor 432, 735 confine 226, 230, 241, 247 confusum 329, 382, 733, 857, 908, 909 conica 524, 528, 534, 537, 551 conicirostris 262 conicola 562, 567, 589 conicus 263 conquisitor 693, 751 consobrinus 791 constricta 364 constrictus 410, 428 continua 410, 422, 424, 428, 595 contorta 373 contractus 358 convergens 297, 302, 307, 310, 312 convexus 386 cooki 745 cooleyi 522, 526, 532, 536, 548 coracipennella 658 corbetti 787 cordatus 388 coriaria 398 corni 709, 732 cornifrons 678, 706, 751 cornuta 467, 564, 593 cornutus 331, 333, 342, 344, 347, 381 coronatus 356 corrodens 802 corruptor 227, 237, 245, 259 corruptrix 760 corruscus 396 corticalis 522, 535, 550 corticinus 377 cosmtocki 508 costalis 317, 329, 359 costirostris 230, 231, 233, 239, 242, 249 costulata 365 costulatus 94 coweni 522, 526, 542, 549 cowperi 728 coypus 835, 840, 844 crassirostris 263 crataegi 259 crawi 505 crenatus 226, 228, 233, 254 cribratum 198, 207, 208, 216 cribricollis 259 crinitus 346 crisonalis 652 cristatus 95, 265, 387 crotonis 503 crudiae 252 cryptomeriae 801 cucullata 281, 290 cucumeris 177, 277, 285, 1021 cunea 57, 607, 615, 616, 618, 620, 635, 638, 641, 694, 719, 733, 733, 742, 857, 992, 993 cunicularius 264 cupressella 621, 657 cupressi 155, 156, 181 cuprifer 258 cupripennis 340, 349 curtum 252 curvatum 35, 42, 678, 693, 694, 695, 701, 714, 720, 755, 857, 986, 990 curvipes 445, 448, 452, 455, 457, 463, 857, 932 curvirostre 225, 246, 890 curvispina 563, 591 cyaneus 684, 703, 709, 754 cyanophylli 477, 499 cycadis 504 cymbalariae 468 cynodontis 560, 562, 563, 588 cynthia 29, 605, 654 czernohorskyi 346 daci 676, 712, 745, 752 dactyliferae 274, 281, 289 dactyliperda 274, 281, 289 dactylopii 684, 706, 707, 708, 709, 719, 721, 724, 738 dahlemica 812, 828 dalii 805 Alain Roques et al. (Eds) / BioRisk $$: @@–@@ (2010) dalmatica 129 danicus 83 darwinii 748 debachi 715, 732 decemlineata 33, 54, 268, 270, 271, 275, 277, 280, 281, 288, 413, 857, 857, 885 decemstriatus 360 decolor 803 decolorella 641 decora 348 defoliaria 659 deforme 678, 755, 857, 986, 987 degenerans 153, 156, 159, 165, 177, 192 delamarei 342, 364 delicata 734 dendrolimi 691, 723, 757, 1015 dentatus 389, 544 denticulatus 144 dentiger 83 depauperatus 248 depressus 323, 360, 380, 392, 405, 412, 431, 568, 593 dermestoides 406 deserticola 564, 590 desjardinsi 374 destructor 33, 63, 71, 149, 150, 153, 159, 160, 164, 165, 170, 173, 173, 190, 382, 499, 515, 546, 718, 723, 747, 763, 857, 872, 873 destruens 266 detorquens 750 deyrollei 368 dianthi 508 diaperinus 331, 380 diaphana 652 diaspidicola 730 diaspidis 726, 740 dichrous 391 dictyopsermi 500 dieckmanni 243 difflualis 650 difformis 559, 561, 569, 594 digitata 114, 116, 127 dilatatus 82 dilutipennis 376 dimidiatus 368, 392, 707, 734 diminutalis 652 diminutus 506 diplosidis 742 discoidea 371 discoideus 403 disparilis 804 dispersus 517, 530, 546, 710, 730 distincta 757 distinguendus 386, 389, 762 distylii 444, 470 ditella 666 diversepubescens 355 divisa 399 domestica 804, 852, 854 domesticus 164, 191, 388, 759 dominica 56, 319, 329, 331, 348 donisthorpei 145 doriae 104, 113, 119, 124 doriai 142, 146 dorrieni 85, 86, 87, 88, 89, 91, 92, 96 dorsalis 283, 287, 751 dorsata 349 dorsiplana 753 dozieri 152, 184 dracaenae 783, 789 dromedarii 166, 183 dromedarius 257 dubius 384, 397 duplicatus 227, 243, 264, 331, 362 eastopi 437, 440, 445, 469 echidninus 153, 155, 164, 183 echinocacti 338, 477, 501, 695, 715 ecphaea 608, 638, 646 edentata 563, 595 edmandsii 609, 630, 653 elaeagni 523, 551 elegans 432, 737, 753 elongata 364, 693 elutella 650 emarginatae 156, 181 emarginatus 99, 103, 104, 105, 107, 112, 113, 115, 125, 857, 866, 867, 871 emeryi 703, 717, 746 endogeus 105, 106, 128 entomophila 803 equestris 602 eremicus 684, 732 ergatandria 746 erichsonii 336, 391 erigerivagrans 182 erigeronense 474 erinea 154, 158, 160, 162, 178 eriosoma 608, 646 erosus 264 errabunda 324, 373 erratica 601 eruditus 252 ervi 291, 933, 939 erythreae 548 Index of the latin names of the arthropod species mentioned in the book erythrocephala 762 erythrogaster 762 escalerae 413, 424, 432 eucalypti 59, 519, 535, 538, 539, 548, 693, 695, 696, 716, 741, 976, 980 euonymi 481, 504 euphorbiae 55, 441, 445, 446, 450, 466, 485, 506, 718, 857, 934, 935 evansi 149, 150, 153, 159, 162, 164, 169, 171, 173, 185, 189 excavatum 168, 183, 402 excoriatus 262 excresens 497 exigua 749 exilis 257 exornatum 165, 167, 182 exotica 332, 339, 373 fabricii 274, 288 facilis 144 faini 152, 184 fallax 353 farinosus 258 fasciapennis 785, 951 fasciata 708, 730 fasciatus 58, 259, 355, 370, 787, 791, 837, 838 fasciculatus 126, 218 fauroti 145 fausta 743 femoralis 386, 778, 783, 788 fenestralis 364 fenestratus 403 fennahi 523, 526, 528, 533, 542, 550 ferrugatus 257 ferruginea 290 ferrugineus 12, 32, 33, 57, 61, 66, 71, 225, 229, 233, 234, 237, 237, 243, 254, 328, 329, 362, 857, 896, 897 festivus 391 feytaudi 57, 65, 67, 68, 479, 483, 490, 492, 510 ficicola 437, 439, 440, 444, 448, 454, 464 ficifolii 563, 567, 576, 583, 588 ficorum 769, 777, 778, 782, 785 ficus 264, 521, 723, 727, 737 figulilella 649 filiceti 672, 716, 759 filicivora 515, 544, 622, 655 filicornis 395 filifera 563, 571, 587, 852, 853, 854 filiformis 262 filum 321, 365 fimetarius 403 fioriniae 501, 786 flabella 465 flandersi 735 flava 471 flavicollis 117, 829 flavidus 319, 354 flavifrons 557, 564, 577, 592 flavilinea 413, 432 flavipennis 376 flavipes 354, 401, 748, 786, 811, 819, 822, 826, 857, 990, 991 flaviterminata 802 flavomaculata 396 flavoscutellum 728 flavus 566, 570, 599, 738, 761 fletcheri 481, 488, 497 flexivitta 823 floccifera 481, 485, 488, 497 floccissimus 517, 546, 710, 730 floccosus 55, 63, 530, 544, 684, 695, 697, 711, 718, 727, 732, 752, 752, 971 floralis 283, 322, 337, 346 floricola 717, 748 floridanum 147, 737 floridensis 496, 691, 728, 736, 740, 769, 785 fluctuosalis 652 folsomi 852, 853, 854 forbesi 448, 460 forestieri 294, 300, 308, 311 forficatus 100, 101, 129, 214 formicarius 384 formicarum 478, 505 formicola 127 formosa 30, 684, 695, 700, 706, 707, 716, 718, 722, 730, 789, 927, 931 formosana 448, 469 fornicata 283, 292 fovealis 631, 665 foveolatus 290 fracticornis 400 fragaefolii 448, 462 fratella 592 fraterna 823 fraxinifolii 444, 447, 455, 457, 470, 940, 941 freemani 368 frischi 319, 356 friuliensis 601 frumenti 225, 233, 244, 254, 857, 894, 895 fuchsiae 155, 160, 174, 179 fucicola 396 fulguralis 520, 535, 536, 540, 542, 547 fuliginosa 414, 415, 417, 433 fuliginosus 762, 383, 394 Alain Roques et al. (Eds) / BioRisk $$: @@–@@ (2010) fullawayi 734 fulva 365, 560, 563, 570, 571, 576, 586 fulvipenne 401 fulvipes 357 fulvus 728 fumatus 368 fungi 399 fur 91, 92, 152, 153, 155, 164, 345, 537, 579, 627, 848 furcisetus 187 fusca 373, 787 fuscata 389 fuscicollis 706, 710, 761 fuscicornis 584, 602, 736 fuscipennis 708, 721, 722, 752 fuscipes 389 fusciventris 721, 736 fuscum 402 fuscus 737 galbus 739 galilaeus 387 galili 696, 707, 725 gallinae 834, 845, 846 galloprovincialis 203, 207, 217 gayndahensis 379 geisovii 812, 818, 828 gemina 838, 848 geminata 371 geniculella 606, 660 gentilei 761 geographicus 257 germanica 328, 679, 765, 818 ghesquierei 592, 802 gibboides 384 gibsoni 562, 587 giffardii 713, 735 gigantea 98, 107, 108, 124, 845 gigas 405, 763, 846 glabripennis 18, 32, 48, 56, 65, 69, 196, 198, 200, 201, 203, 204, 204, 206, 207, 208, 209, 210, 212, 214, 857, 878, 879 glabrirostris 263 glabrum 358 glacialis 849 glauca 180, 764, 984 glaucescens 415, 417, 432 gleditchiae 562, 567, 568, 570, 571, 572, 574, 575, 576, 581, 582, 587, 587, 914, 915 gleditsiae 160, 162, 179 globulus 346, 390 gloverii 480, 487, 502, 708, 730, 731 gobicola 355 godmani 229, 230, 231, 234, 244, 249 goldamaryae 465 gossypariae 721, 728 gossypiella 608, 643 gossypii 21, 31, 55, 441, 446, 458, 460, 507, 684, 718, 930, 931, 934, 973, 977 gowdeyi 785 gracilis 99, 100, 103, 104, 110, 111, 113, 127, 129, 347, 377, 857, 868, 869, 873 graminis 488, 506, 700, 736, 740 granarium 319, 329, 358, 857, 892, 893 granarius 18, 32, 233, 384 grandis 62, 395 grassii 506 gratiosus 791 greeni 805 greenii 502 gregaria 399, 564, 569, 594 gressorius 260 griseus 218 grossa 120, 147 grossulariae 560, 562, 589, 760 grunertiana 606, 626, 627, 667 guadalupensis 377 guadeloupae 710, 730 guildingii 102, 107, 116, 123 guineensis 749, 894, 896 gularis 627, 651 gurneyi 729 gustavi 432 guttatus 360 guttula 387 haemorhoidalis 392 haemorrhoa 402 haemorrhoidalis 391, 717, 721, 770, 773, 778, 788 haldemani 732 halli 507 halys 413, 415, 427, 430, 857, 962, 963 hamatipennis 348 harwoodi 399 hasselti 59, 139, 141, 147 hederae 162, 178, 950 hedericola 156, 158, 161, 184 heeri 746, 799 hellenica 172, 479, 481, 510, 573 helvolus 691, 721, 739 hemerocallis 456, 467 hemipterus 323, 369 hendersoni 162, 163, 180 henrici 372 hercyniae 672, 760 Index of the latin names of the arthropod species mentioned in the book heringella 617, 661 herndoni 708, 730 hespericus 263 hesperidum 478, 479, 481, 483, 486, 496, 710, 721, 724, 728, 729 heterographa 847 heydeni 358 heymonsi 574, 593 hilleri 344 hilli 736 himalayensis 229, 251 hippocastani 154, 191 hippophaes 432 hirsutus 104, 126 hirtipennis 274, 275, 277, 284, 285, 857, 882, 883 hislopi 389 hispida 730, 982, 1002, 1008 hispidum 835, 845 hollisi 519, 536, 547 holoxanthus 697, 699, 727 hookeri 812, 828 horii 498 horni 357 hortensis 290, 852 hortorum 718, 759 hortulanus 96 howardi 837, 838, 849 huidobrensis 55, 555, 561, 564, 565, 567, 569, 572, 573, 575, 583, 585, 585, 910, 911 humboldti 474 humeralis 226, 234, 236, 248, 371 humile 35, 43, 61, 62, 64, 65, 66, 70, 71, 679, 684, 723, 723, 857, 980, 981 humilis 61, 62, 384, 525, 615, 619, 995 huttoni 223, 226, 229, 230, 234, 237, 239, 243, 244, 247, 412, 415, 416, 420, 423, 427, 429, 857, 958, 959 hyalinus 523, 528, 551 hydei 591 hydrangeae 478, 480, 485, 487, 498 hylaeformis 634, 665 hyphantriae 733 ilicis 175, 187 illinoisensis 439, 442, 457, 458, 461 illucens 558, 561, 567, 595 illutana 606, 667 illyricus 396 immigrans 591 immigrata 379 imperfectus 763 impexus 294 impressicollis 340, 378 inaequalis 248 inarmatum 252 incanus 396 inclusum 359 indianus 592 indica 234, 255, 276, 473, 560, 577, 588, 742 indicus 781, 789 indifferens 562, 568, 569, 571, 579, 596, 923 indigata 659 inermis 812, 828 inexpectus 322, 383 infelix 738 infirmus 400 infrequens 598 ingenuus 743 ingrata 634, 662 innumerabilis 476, 478, 480, 489, 495, 497 innuptum 796, 803 inquilinus 804 inquinata 632, 639, 659 inquinatus 361 inquirenda 730 inquisitor 218 insectella 666 insertum 471 insignis 307, 311, 479, 491, 505 insolens 714, 750 instabilis 384 insulare 359, 771 insularis 372 insularum 244, 634, 653 integer 321, 363 intermedia 601 interpunctella 616, 618, 653 intrudens 228, 239, 250 invasa 56, 705, 709, 713, 743, 977 inviscus 739 isabellae 665 isabellinus 249 ishidae 523, 528, 529, 533, 534, 551 isosomatis 755 issikii 615, 616, 620, 626, 633, 636, 645, 857, 1004, 1005 itoi 673, 755 jacarandae 564, 567, 572, 585 jaegerskioeldi 748 jamatonica 519, 531, 542, 543, 547 japonica 140, 143, 144, 318, 373, 374 japonicus 58, 69, 353, 478, 480, 487, 496, 571, 581, 590, 744 jaspidea 214 Alain Roques et al. (Eds) / BioRisk $$: @@–@@ (2010) javae 744 javanicus 104, 107, 108, 113, 120, 123 joannisi 660 johnsoni 373, 595 johnstoni 179, 758 josephi 372 juglandicola 438, 444, 463 juglandis 438, 439, 469 juncii 262 juvencus 762 kahawaluokalani 473 kanzawai 189 kelaarti 107, 127 kellneri 566, 570, 599 kellyanus 771, 777, 783, 789 kewi 790 kirkaldy 545 kirschii 321, 363 koehleri 702, 737 kollari 760 kondoi 447, 460, 733 kossuthi 82 kraatzii 265 kuehniella 56, 302, 307, 616, 651 kuricola 455, 474 kuriphilus 12, 32, 36, 56, 61, 676, 687, 691, 696, 699, 702, 735, 735, 974, 975 kutscherae 290 kuwanae 300, 309, 311, 693, 705, 715, 740 kwoni 495, 506 lachlani 804 lacticolella 607, 641 laeta 139, 142, 146 laevigata 397, 810, 824 laevigatella 668 laevigatus 38, 380, 410, 422, 431, 1021 laevis 82, 265, 677, 733 lafertei 362 lagoi 105, 125 lahorensis 712, 721, 722, 731 lambersi 452, 457, 465 lamimani 180 laminatus 361 lanigerum 30, 59, 438, 444, 446, 453, 464, 683, 726 lanuginosum 750 laportei 440, 445, 447, 450, 463, 699, 714 lardarius 319, 331, 356 laricella 617, 658 laricicola 598 lariciphila 693, 762 laricis 161, 187, 192 lasiocarpae 756 lataniae 462, 477, 489, 501 lateralis 388, 811, 819, 825 lateripunctatus 290 lateritius 375 lathrobioides 402 laticollis 404 latipennis 249 latro 345 latum 167, 182 latus 149, 152, 159, 160, 161, 163, 171, 174, 186, 508 latysiphon 444, 471 lauretorum 483, 510 lavandulae 600 lavaterae 412, 421, 425, 426, 427, 431 leautieri 606, 639, 662 lebasi 215 leechi 356 lemniscata 692, 700, 713, 737 lentis 291, 571, 598 lentus 83 lepidosaphes 692, 727 lepidus 260 leplastriana 668 leprosa 198, 201, 210, 216 leptocorisae 753 lespedezae 466 lethifera 405 leucodactylus 609, 648 leucographella 616, 621, 625, 637, 645 leucoloma 228, 231, 233, 241, 249 leucomelanellus 642 leucomelas 747 leucophthalmus 387 leucopus 765 leucotreta 627, 630, 656 levaillantii 201, 207, 215 lewisi 162, 186, 187, 352, 375, 749 leydigi 83 licarsisalis 651 lichenis 786 ligneus 369 lignicola 760 ligniperda 264 ligustica 40, 67, 705, 759 ligustri 179, 180 lilii 292 liminaris 234, 242, 253 lindbergi 250 linearis 95, 226, 234, 255, 260, 264, 404, 405 lineatus 260 Index of the latin names of the arthropod species mentioned in the book lingnanensis 691, 727 linguis 786 liriodendri 445, 447, 453, 465 littoralis 229, 245, 247, 662, 855, 857, 1012, 1013 litura 608, 647, 1013 liturata 659 lividimanus 291 lividipes 811, 818, 826 lividum 218 lividus 393 loewi 154, 158, 191 lohsei 389 longicaudata 852, 854 longiceps 412, 421, 423, 424, 426, 431, 835, 843 longicollis 226, 230, 245, 251, 395 longicornis 403, 684, 699, 703, 719, 748, 812, 822 longicorpus 745 longimanus 214 longipalpa 702, 737, 744, 824 longipalpis 401 longipennis 790, 800, 804 longipes 146 longirostre 225, 229, 230, 238, 241, 243, 247, 857, 890, 891 longirostris 502 longisetosum 359 longispinosus 750 longispinus 478, 481, 486, 488, 508 longiusculus 258 longiventris 404 longoi 59, 69, 707, 717, 736 longula 396 longulus 496 lophanthae 294, 296, 299, 300, 311 loricatus 144 lounsburyi 494, 497, 501, 691, 695, 710, 711, 712, 731, 739 lucifugus 99, 101, 102, 103, 104, 111, 113, 115, 118, 119, 128, 817, 817 lucipara 662 lucorum 759 luctuosus 751 luderti 361 lugens 361, 524 lunatus 386 lupini 159, 188 luridipennis 399 luteipes 374, 799 luteola 280, 292, 370 luteolus 739 lutescens 409, 413, 425, 431 luteum 445, 472 lymphaseus 145 lynx 355 lyoni 140, 145 lyriocephalus 835, 843 macedonica 432 macfarlanei 189 mackienziei 506 macularius 260 maculatus 282, 287, 357 maculipennis 739 maculosus 835, 847 madagascariensis 799 madeirensis 478, 507 maderae 745, 810, 814, 824 maidis 446, 471 maillardi 608, 620, 622, 623, 646 maindroni 801 mairei 250 major 84, 94, 392, 504 mali 30, 59, 240, 252, 683, 697, 726 malifoliae 180 malifoliella 617, 661 maligna 523, 535, 551 malinellus 602, 706 malinus 741 malloi 712, 737 mamillariae 485, 508 mandibulare 226, 241, 252 manicatus 791 manilensis 639, 650 marchii 404 marginalis 394 marginata 524, 528, 529, 534, 542, 551, 677, 760 marginatus 403 marginellus 369 marginepunctata 796, 803 marginiventris 733 marietti 401 maritima 376 marlatti 479, 505, 518 marmoratus 344 maroccanella 642 maroccanus 382 marshalli 61, 68, 71, 604, 605, 608, 612, 619, 621, 626, 628, 634, 634, 638, 646, 857, 1010, 1011 martini 794 maskelli 56, 59, 65, 695, 713, 715, 716, 744, 857, 976, 977 Alain Roques et al. (Eds) / BioRisk $$: @@–@@ (2010) Masonaphis 452, 457, 465 materiarius 234, 235, 238, 245, 251, 857, 900, 901 matsuyamensis 721, 729 matteiana 560, 562, 563, 588 mauli 324, 368 mauritanica 746 mauritanicus 214, 323, 383 maxillaris 102, 123 maxillosum 260 maxillosus 381, 400 mayeti 360 mcdanieli 175, 189 meeusei 94 megacephala 62, 71, 684, 707, 749 megalops 769, 773, 785 megatomoides 359 melagynalis 650 melaleucus 773, 786 melancholica 261 melania 598 melanocephalum 698, 716, 718, 723, 750 melanogaster 591 melanogrammum 261 melanopygius 140, 142, 144 melanura 396 melbae 557, 568, 579, 592 meleagridis 846 meleoides 234, 237, 249 meliloti 791 melinus 690, 704, 727, 951 mellifera 40, 67, 705, 712, 759 membranifera 749 memnonia 263 mendax 332, 347, 803 mercator 375 merceti 702, 737 meridionale 35, 830 meridionalis 266 meritoria 701, 731 mesembryanthemi 498 mesomelinus 404 messaniella 661 mexicana 678, 716, 754 micans 57, 263 microcarpae 707, 725 microcosmus 742 microps 592 migrator 318, 379 migratoria 826 milleniana 667 milleri 756, 805 millieridactylus 664 mimose 288 minei 547 minor 961, 1005 minozzii 791 minuta 195, 217 minutulus 387 minutum 758 minutus 328, 348, 366 mixticolor 433 modestus 752 molesta 857, 1016, 1017 mollis 320, 331, 343, 388 monachus 348 moneta 606, 662 monizianus 247 monodactyla 664 montandoni 350 montanus 763 montivaga 385 montrouzieri 294, 296, 298, 299, 304, 308, 311 mordvilkoi 440, 449, 472 mori 857 morio 328, 382 morosus 812, 828 morrisoni 465 morsitans 104, 107, 124 mortisaga 336, 405 moschata 217 mucronata 376, 405 multispinosus 153, 183 munda 389 mundus 718, 759 muntiacus 835, 836, 843 murariella 621, 666 murinus 391 murtfeldtae 728 musae 780, 787 museorum 391 mutatus 857, 898, 899 mutilatus 370 myersi 440, 454, 460 mymaripenne 757 myopaeformis 665 myricae 517, 542, 546, 715, 732 mytilaspidis 684, 727 naevana 668 nana 387 nanus 95, 1007, 1009 nasatum 82 nasicornis 397 natalensis 347 Index of the latin names of the arthropod species mentioned in the book nearcticus 566, 594 nefrax 647 neglecta 520, 548 neglectus 35, 62, 66, 71, 87, 92, 684, 696, 697, 698, 716, 747, 857, 974, 975, 978 negundivagrans 155, 156, 178 nemoralis 338, 386 neobrevipes 506 neocaledonicus 189 neocynarae 172, 179 nephrelepidis 464, 515, 545 nepos 370 nerii 477, 479, 481, 486, 489, 499, 690, 850, 946, 947, 950, 951 nervata 474 nicholsoni 146 nietneri 740 nigella 765 niger 754 nigra 399, 478, 484, 497, 692 nigricans 321, 338, 343, 361, 378 nigriceps 316, 317, 318, 321, 331, 338, 343, 350, 361, 378 nigricollis 401 nigricornis 392, 399 nigripennis 769, 788 nigrirostris 262 nigritulus 400 nigritus 300, 309, 311 nigroflavus 755 nigromarginata 811, 820, 827 nigronervosa 445, 449, 470 nigrovariegatus 756 nigrovittata 564, 580, 591 nipae 507 nipponicus 328, 335, 338, 354 nitens 674, 714, 751, 893 nitidifrons 376 nitidula 389 nitidulus 266, 398, 404 noacki 857, 970, 971 noctilio 80, 724, 763 noda 749 nodifer 321, 364 nodulosus 217 nonagrioides 662, 703, 707 nordmannianae 526, 528, 857, 924, 925 norvegica 340, 394 notata 558, 597 nova 744 novemmaculata 385 novimundi 658 novita 712, 751 nubivagus 349 numenius 412, 415, 416, 423, 425, 429 oberti 414, 423, 433 oblita 399 obliteralis 650 obliterata 128 obliteratoides 290 obliteratus 334, 364 oblongiusculus 387 oblongus 343 obovatus 159, 161, 186 obscura 400 obscuralis 652 obscuriceps 410, 426, 428 obscurior 746 obscuriscapa 710, 749 obscurus 264, 405 obsoleta 805 obsoletus 370, 393 obtectus 270, 273, 275, 276, 285, 519, 535, 547 occidentalis 31, 36, 40, 41, 55, 61, 68, 71, 165, 176, 411, 418, 419, 420, 421, 422, 423, 424, 425, 426, 427, 428, 519, 535, 547, 555, 566, 577, 580, 586, 707, 770, 771, 773, 776, 780, 784, 785, 788, 944, 945, 1016, 1017 oceanicus 324, 354, 798 ochracea 330, 401 ochraceus 99, 103, 104, 107, 109, 112, 126 octarticulatus 787 octomaculatus 835, 847 ocularis 371 oculatus 523, 542, 550 oenotherae 454 ohridella 12, 20, 26, 29, 35, 36, 39, 43, 48, 49, 57, 58, 60, 69, 71, 604, 605, 612, 617, 620, 623, 625, 627, 632, 636, 637, 639, 660, 855, 857, 996, 997, 999, 1000, 1001, 1003 oleae 478, 479, 481, 486, 498, 668, 691, 695, 709, 710, 711, 712, 714, 718, 721, 722, 728, 729, 736, 737, 739, 742, 743, 744, 745, 752, 761 oleastrella 668 olfersii 94 olgae 391 oligomacrochaeta 105, 123 oliveri 469 omoscopa 654 ondatrae 153, 184 ononidis 262 ononis 256 Alain Roques et al. (Eds) / BioRisk $$: @@–@@ (2010) opaca 397 operculella 857, 998, 999 opilionoides 145 orana 666 orbonalis 651 orchidaceus 787 orchidearum 440, 462, 745 orchidii 780, 787 orientalis 34, 95, 123, 171, 187, 322, 366, 522, 732, 739, 811, 812, 818, 824 ornatus 467 oryzae 225, 226, 229, 230, 232, 233, 247, 255, 381 oryzophilus 231, 233, 237, 255 osborni 500 oscinidis 742 ostreata 504 ovalis 321, 350, 835, 844 ovis 835 oxycoccana 567, 572, 587 pacifica 797 pacificus 156, 159, 161, 187, 413, 415, 416, 422, 429 packardi 517, 539, 546 pactolana 667 paeoniae 488, 504 paeta 803 paetula 803 pallens 316, 349 pallescentella 655 palliatus 264 pallida 804, 823 pallidipennis 274, 283, 284, 286 pallidulus 388 pallidus 92, 94, 145 pallipes 763 palmae 501, 789 palmarum 901 palmi 55, 67, 376, 777, 779, 780, 781, 790, 857, 1020, 1021 paludosa 400 palustris 399, 940 panici 561, 567, 569, 589 panporus 104, 105, 116, 126 pantherina 307, 311 pantographae 734 papuana 564, 594 parabolicus 823 paradianthi 179 parallelus 385 parasemus 159, 188 parcesetosum 301, 309, 310, 312 parcus 318, 377 parlatorioides 504 parumpunctatus 402 parva 379 parvulus 805 parvus 92, 94 pascuorum 258 patruelis 804 paulistus 732 pavonis 846 paykulli 146 pearmani 803 pecanis 448, 467 pechumani 558, 597 pectinicornis 322, 328, 335, 374, 1007, 1009 pelekassi 149, 155, 158, 162, 180 pellio 390 pellucidus 259 pennipes 573, 596 pentagona 57, 477, 480, 481, 488, 504, 706, 725, 728, 729, 730, 857, 950, 951, 954, 955 perditus 187 peregrina 548, 621, 657, 741 peregrinus 101, 105, 113, 115, 118, 124, 129, 804 perelegans 144 pergandiella 695, 731 pergandii 487, 503 pericarpius 257 perieresalis 650, 857, 992, 993 peringueyi 798 peritana 656 perkinsi 758 perminutus 737 perniciosi 31, 540, 731, 957 perniciosus 30, 55, 477, 480, 481, 488, 500, 857, 948, 952 pernyi 653 perplexus 789 perseae 149, 162, 188 perseaphagus 516, 544 persicae 449, 455, 459, 470, 857, 938 persimilis 102, 105, 124, 153, 155, 158, 165, 185, 575, 584 perspeciosus 725 perspectalis 604, 619, 622, 650 peruana 823 peruvianus 357 perversaria 660 petanovicae 154, 163, 179 petiolata 744 pfeilii 223, 253 Index of the latin names of the arthropod species mentioned in the book phaenota 563, 565, 592 phalaenopsidis 787 phalangioides 145 pharaonis 684, 748 phaseoli 287 phasiani 835, 845 philococcus 503 phoeniceata 606, 659 phoenicis 186 phyllocnistoides 743 phyllotretae 744 picata 402 piceaella 642 picipes 395 picirostris 258 picta 321, 360 pictus 765 pilosellus 370 pilosus 59, 353, 538, 693, 695, 708, 716, 741 pinastri 227, 239 pini 405, 661, 760 piniaria 632, 659 pinicola 501 pinicolus 791 pinnulifer 500 pinsapinis 756 pinus 723, 752, 756 pipiens 600 piriformis 226, 231, 243, 248 piscatorium 250 pisi 566, 570, 598 pisorum 276, 286, 714 pityocampa 62, 70, 605, 663 plagiatum 260 planicollis 367 planipennis 61, 64, 71, 315, 329, 330, 332, 349, 857, 870, 871, 874, 875 planiuscula 258 planiusculus 392 platani 57, 69, 605, 616, 617, 620, 626, 636, 645, 661, 1002, 1003, 1006, 1007 plexippus 305, 308, 606, 609, 632, 647 plicatus 256, 260 plumbea 401 plumbeomicans 256 plummistaria 659 podocarpi 437, 445, 469, 502 poligraphus 265 politus 372, 403 polydectalis 652 polyspila 288 polytrapezius 847 pomonella 667 populifolii 455, 462 porcatus 257 porcelli 835, 837, 839, 840, 844 porosum 470 porrectella 664 porteri 731 postica 262, 795 posticus 360 postvittana 610, 656 praecox 577, 602 prelli 522, 537, 549 pretiosum 696, 758 primita 258 primulae 460 pritchardi 188 procerulus 402 procnemis 590 procnemoides 572, 590 pronubana 55, 622, 667 propinquus 404, 768, 786 proteus 503 protransvena 701, 731 provisorius 83 proximus 237, 253 pruinosa 61, 66, 71, 514, 524, 528, 532, 533, 536, 537, 539, 543, 552, 677, 708, 735, 857, 882, 952, 953, 956, 957 pruinosus 82, 404 pseudoambrosiae 474 pseudococci 561, 562, 571, 587, 719, 761 pseudohesperidum 496 pseudoleucaspis 502 pseudomagnoliarum 478, 490, 497, 720 pseudomillotianus 123 pseudopopuleum 470 pseudotenera 376 psidii 498 psylloides 331, 333, 343 ptericolens 442, 445, 466 pteridicola 790 ptilinoides 384 puberula 376 puberulus 261 pubescens 256, 290, 365 pulchella 520, 547 pulchellus 377, 740 pulchripennis 796 pulicaria 400 pulsatorium 796, 804 pulverulentus 262 pumilio 318, 360 Alain Roques et al. (Eds) / BioRisk $$: @@–@@ (2010) pumilis 801 punctatissima 35, 684, 694, 697, 705, 716, 746, 830 punctatulus 386 punctatum 384 punctatus 405 puncticollis 261, 525 punctidorsa 180 punctithorax 389 punctum 392 pungens 485, 507 punicae 188 pupula 622, 651 purchasi 59, 296, 478, 480, 481, 486, 505 puritanus 144 purpurea 703, 736 purpureus 104, 109, 126 pusilla 389 pusillidactylus 609, 622, 648 pusillima 402 pusilloides 363 pusillum 396 pusillus 83, 85, 363, 377, 382, 401 pusio 557, 564, 592 puttleri 706, 743 pygmaea 595 pygmaeus 265 pyri 154, 158, 162, 181, 570, 599, 617, 662 pyriformis 478, 480, 484, 497, 501, 693, 739 pyrioides 414, 425, 427, 430 pyrivora 570, 599 quadraticeps 725 quadraticollis 257 quadrifasciata 845 quadrifoveolata 395 quadrimaculatus 390 quadripedes 155, 182 quadrisignatus 371, 395 quercuscalicis 760 quinquenotata 563, 570, 571, 586 quisquiliarius 403 quisquilius 393 rachipora 516, 530, 544 radiolus rafni 71, 756 ramburii 804 rancidella 624, 659 rapae 60, 71, 663 rapax 502 raptor 699, 716, 753 ratzeburgi 382 rebeli 642, 647 rectangulatus 846 rectangulus 318, 379 recticollis 376 recurva 199, 204, 206, 207, 210, 211, 215 recurvalis 653 reflexus 191 regalis 57, 480, 487, 498, 698, 942, 943, 947 regularis 504 remyi 378 repleta 591 reticulatus 804 reunioni 300, 311 rhodensis 180 rhododactylus 265 rhododendri 413, 421, 427, 429, 430, 599 rhois 466 rhombifolia 811, 824 richardsi 801 rileyae 801 rimariae 485, 509 riparius 102, 104, 115, 126, 110 rivnayi 786 rivulare 402 robiniae 447, 454, 461, 561, 562, 565, 566, 567, 568, 571, 572, 573, 574, 580, 582, 582, 588, 761, 857, 916, 917 robiniella 32, 616, 620, 625, 629, 633, 636, 638, 639, 644, 645, 717, 720, 743, 857, 983, 1002, 1003, 1007, 1008, 1009 rogeri 749 rosae 246, 247, 763, 890, 972 rosaecolana 667 roscipennella 617, 660 rossi 390, 502 rossicus 183 rostellum 489, 506 rotundata 678, 762 rubra 218, 467, 936 rubricollis 250 rubromaculatus 414, 430 rudis 225, 234, 242, 253, 857, 902, 903 rufa 805 rufescens 139, 140, 146 rufiabdominale 471 ruficollis 331, 351, 394 ruficorne 406 ruficornis 371 rufifrons 564, 566, 594 rufimanus 276, 286 rufinasus 229, 235, 238, 240, 255 rufipenne 197, 198, 200, 203, 206, 214 rufipennella 606, 660 Index of the latin names of the arthropod species mentioned in the book rufipes 147, 256, 291, 351, 753 rufocapillata 358 rufomarginatus 386 rufulus 383 rufum 226, 230, 242, 247 rufus 339, 372, 791 rugosostriatus 260 rugulosus 265 rumexicolens 461 russelli 742 russulus 170, 186 rusticus 217 sabalis 487, 491, 500 sabella 621, 635, 648 sacchari 55, 624, 654, 786 saginatus 390 saissetiae 729, 736 sakuntala 802 saliapterus 462 salicetum 696, 752 salomonis 748 saltans 456, 473 samayunkur 769, 782, 789 sanborni 465 sanctaehelenae 372 sanctivincentii 146 sanguinea 309, 757 sanguineus 151, 167, 169, 171, 176, 192 sangwani 700, 740 sarcitrella 663 sarothamni 410, 415, 417, 431 sartor 217 sawadai 736 sawatchense 179 saxesenii 266, 764 scaber 82, 406 scabricollis 252 scalaris 211, 564, 568, 571, 573, 574, 578, 584, 594 scammelli 463 scanicus 390 scarabaeoides 265, 393 schimitscheki 71, 688, 756 schineri 595 schmidti 390 schmitzi 555, 574, 586 schoblii 82, 87, 90, 92 schultzei 783, 787 sciuricola 835, 843 sculptus 402 scupense 166, 173, 191 scutatus 146 scutellaris 351, 698, 729, 761 scutellatus 56, 57, 67, 226, 230, 232, 233, 234, 241, 242, 243, 248, 248, 751, 857, 888, 889 secalis 386 secreta 503 segnis 848 sellatus 257 semiflavus 705, 720, 726, 742, 768 semifumipennis 758 semipunctata 56, 57, 59, 66, 69, 70, 204, 206, 207, 210, 211, 216, 216, 707, 717, 736 semistriata 385 semistriatus 392 semivittatum 256 senegalense 348 senilis 761 senoculata 140, 145 senticetella 659 sericorne 320, 323, 344 serrata 365 serraticorne 388 serratus 275, 287, 507 serripennis 366 sertifer 672, 713, 760 setariae 446, 464 seticosta 595 sexguttella 659 shakespearei 722, 743 sharpi 362 sheldoni 149, 155, 162, 179, 741 shoshone 147, 380 sibiricus 379, 632, 835, 843 sicarius 742 sigillatus 812, 818, 827 signaticornis 291 signatum 741 signatus 404 siliquastri 274, 275, 286 silphoides 377 silvestrii 729, 750 similaris 285 similis 144, 522, 526, 760 simillimum 714, 750 simplex 391, 628, 642, 781, 790 simsoni 318, 342, 350, 373 sinensis 366, 480, 481, 486, 489, 496 singularis 260 sinhai 159, 189 siskiyou 560, 562, 567, 573, 575, 582, 588 skuhravyorum 581, 600 smirnovi 355 smithi 732, 733 Alain Roques et al. (Eds) / BioRisk $$: @@–@@ (2010) smyrnensis 829 soccata 557, 584, 593 solani 493, 507 solanivora 633, 643 sophia 302, 310, 731 sophorae 709, 745 sordida 399 sordidus 225, 233, 254, 400 sorghicola 561, 563, 568, 569, 573, 578, 589, 713, 742 spadix 257 sparsa 398 spartii 363 spatulata 56, 520, 536, 540, 543, 548, 696 spatulatus 751 speciosa 606, 617, 662 speciosissima 728 speciosus 478, 480, 485, 508, 740 spectrum 445, 670, 763 specularis 691, 756 speculifrons 398 spermotrophus 684, 689, 708, 752, 757, 857, 984, 985 sphacelata 409, 414, 423, 430 spinalis 790 spinicollis 395 spiniferus 530, 531, 541, 544, 752 spinigera 558, 564, 565, 568, 578, 595 spinimanus 261 spinipes 283, 288, 334, 379 spiniventris 147 spinosus 95, 502 spinulosa 835, 844 spiraeae 763 spiraecola 63, 305, 446, 461, 972 spiraeella 617, 634, 658 spiraephaga 444, 461 spiraephila 461 splendana 667 squamiger 158, 185 squamosus 259 stali 811, 814, 825 stanleyi 691, 722, 740 stebbingi 83, 91, 199, 201, 204, 207, 212, 216 stenaspis 142, 145 stenopsis 835, 843 stephensi 258 stercorea 321, 368 strachani 504 straminea 580, 595 stramineus 834, 844 strandi 389 strauchi 801 striaticorne 674, 751, 961 striatulus 259 strigulatella 661 strobi 522, 566, 599 strobicus 160, 162, 184 strobilella 667 suavis 608, 622, 629, 644 subarctica 764 subcarinatus 382 subcinerea 659 subcribrosus 263 subdeplanata 352 subdepressus 381 subdiversus 563, 590 suberivora 606, 617, 662 suberosus 382 subfasciatus 273, 276, 279, 288 subflaviceps 717, 736 subfumatus 353 subnitescens 392 subnotatus 388 subspinipes 117, 119, 124, 129 subtile 362 subtilis 377 succisus 370 sudanensis 786 sulcata 401 sulcatus 260 sulphureus 292 suppressalis 626, 649 surinamensis 32, 56, 320, 329, 331, 375, 808, 810, 818, 823 suspectus 713, 723, 765 sutor 217 suturalis 272, 276, 279, 282, 283, 289 suzukii 565, 569, 575, 576, 591 swezeyi 745 swirskii 740 sylvanidis 677, 733 sylvestrana 667 syrphoides 601 tabaci 31, 55, 67, 160, 165, 791, 301, 310, 346, 514, 527, 529, 530, 535, 536, 536, 690, 700, 701, 706, 718, 731, 857, 926, 927, 929 tabulata 147 taiwana 472 takachihoensis 438, 473 takeyai 414, 420, 421, 422, 424, 427, 430 tamarisci 395 tamiasis 835, 838, 843 tamilnaduensis 561, 563, 568, 569, 594 Index of the latin names of the arthropod species mentioned in the book tapetzella 666 tardyi 257 tartarus 811, 826 taurus 397 taxus 499 taylorae 373 taylori 359 tectorum 802 tectus 346 teneriffana 749 tenuicornis 753 tenuipes 715, 744 tenuis 429 tepidariorum 147 teres 403 terminatus 386 terminella 658 territans 600 tessellatus 478, 497 testacea 389, 403, 414, 415, 417, 433 testaceipes 63, 71, 696, 718, 719, 734, 857, 931, 972, 973, 1003 testaceus 218 tetracolus 387 tetraphylla 381 thalictri 765 theae 162, 178, 503, 607, 641 thermarum 400 thuiella 57, 610, 616, 626, 637, 657, 857, 1018, 1019 thujae 265 thymus 743 tibialis 371, 684, 709, 857, 982, 983 tinerfensis 483, 488, 499, 510 titanus 32, 55, 514, 523, 526, 528, 532, 533, 534, 535, 551, 857, 857, 945 titschacki 801 tobias 347 tomentosa 84, 86, 91, 95, 96 tomentosum 207, 211, 217 tonkineus 273, 275, 282, 284, 287 topitotum 85, 87, 96 trachoides 515, 540, 545 traegardhi 240, 265 transitella 638, 652 transitionalis 180 translucens 655 transvaalensis 716, 757 transversus 266 triangulosa 147 triangulum 399 tricolor 357 trifasciata 617, 668 trifasciatipennis 742, 1003, 1009 trifasciatus 391, 1003, 1009 trifolii 55, 56, 68, 555, 561, 565, 567, 569, 571, 572, 579, 585, 585, 857, 912, 913 trilobatus 138, 139, 144, 857, 870, 871 trinotata 400 trinotatus 852, 854 trispinata 852, 854 tristis 785 tristriata 154, 160, 162, 179 tritaeniorhynchus 565, 571, 581, 590 trivialis 413, 431, 455 troglodytes 360 truncatum 166, 183 tryoni 734 tsigana 591 tubercularis 499 tuberculatus 583, 592 tumidellus 189 turanica 467 turbinatum 845 turcicus 363, 747, 978 turcomana 827 turgida 389 turki 105, 113, 124 typhlocybae 677, 708, 735, 957 ulmi 174, 684, 791 ulmiparvifoliae 443, 453, 455, 473 umbratilis 403 uncatoides 519, 547 undulana 656 undulatus 391 unicolor 218, 343, 355, 379, 786 unidentatus 397 unifasciatus 316, 349 unilachni 735 unisetiorbita 563, 565, 567, 568, 583, 585 univittatus 351 utilis 677, 728, 733 uvae 494, 500 uvida 401 vacca 397 vaccinii 567, 573, 589 vachoni 805 vagans 555, 561, 564, 590 valiachmedovi 102, 105, 125 validum 225, 246 validus 152, 184 vaneeckei 791 vaporariorum 55, 160, 174, 301, 517, 528, 529, 530, 546, 684, 695, 701, 701, 707, 716, 718, 721, 722, 752, 928, 929 Alain Roques et al. (Eds) / BioRisk $$: @@–@@ (2010) variabile 359 variabilis 182, 394 varians 468 variatus 146 variegana 666 variegata 521, 539, 764 variegatum 238, 256 variegatus 261 varii 665 variolosus 262 varipalpis 593 varius 282, 291, 387, 729 varus 515, 546 velox 145 velutina 35, 43, 63, 70, 71, 679, 689, 690, 703, 722, 758, 857, 857, 984, 988, 989 venatoria 146 venusta 146, 543 venustulus 786 vernalis 387 vernana 663 versicolor 147, 359, 658 verticalis 33, 42, 409, 411, 420, 425, 426, 427, 429 verticillata 707, 725 veruculata 147 vespiformis 769, 773, 776, 781, 785 vespulae 339, 357 vestitus 263, 393 vexans 600 viburni 478, 481, 508, 714 villosus 263 violacea 351 violae 448, 469 violicola 589 virens 256, 760 virescens 415, 432 virgata 506 virgifera virgifera 12, 39, 54, 64, 69, 268, 274, 277, 279, 280, 289, 880, 884 viridula 40, 413, 416, 423, 426, 433, 573 vishnui 563, 571, 590 vitegenella 636, 644 vitifoliae 32, 54, 522, 523, 532, 550, 857, 964, 965 vitreoradiata 520, 531, 532, 534, 540, 541, 548 vittata 592, 607 vittatae 735 vittella 609, 632, 647 vividula 748 vorax 319, 357 vulcanius 82 vulgare 82, 83, 86, 88, 90, 658 vulgaris 274, 288, 653, 679, 765, 826, 1004 vulnerata 523, 537, 550 wachtli 714, 765 wakibae 442, 463 walshii 438, 456, 467, 857, 936, 937 waterstoni 677, 699, 733 watsoni 363, 364 weldoni 187 wesmaeli 764 williamsi 515, 545 wirthi 565, 576, 590 wollastonii 251 wroughtoni 706, 746 xanthodera 277, 278, 281, 282, 285 xantholoma 400 yamamai 29, 615, 618, 624, 654 yanonensis 477, 486, 487, 505, 693, 727 yuccae 785 yusti 190 zamiae 477, 501 zealandicus 334, 377, 720, 741 zeamais 225, 229, 230, 232, 255 zeaphilus 370 zejana 596 zelkowae 473 ziziphi 477, 487, 493, 503

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Alien terrestrial arthropods of Europe

Alien terrestrial arthropods of Europe

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