2019 Book BiodiversityOfAngola

Brian J.
Huntley · Vladimir Russo

Fernanda Lages · Nuno Ferrand Editors
Biodiversity
of Angola
Science & Conservation:
A Modern Synthesis
Foreword by His Excellency
President of the Republic of Angola
João Manuel Gonçalves Lourenço
Biodiversity of Angola
Brian J. Huntley • Vladimir Russo
Fernanda Lages • Nuno Ferrand
Editors
Biodiversity of Angola
Science & Conservation: A Modern Synthesis
United Nations UNESCO Chair on Life on Land

Educational, Scientific and University of Porto
Cultural Organization Porto, Portugal
Editors
Brian J. Huntley Vladimir Russo
CIBIO-InBIO, Centro de Investigação em Fundação Kissama
Biodiversidade e Recursos Genéticos Luanda, Angola
Universidade do Porto
Vairão, Portugal Nuno Ferrand
CIBIO-InBIO, Centro de Investigação em
Fernanda Lages Biodiversidade e Recursos Genéticos
ISCED – Instituto Superior de Ciências da Laboratório Associado, Campus de Vairão
Educação da Huíla Universidade do Porto
Lubango, Angola Vairão, Portugal
Departamento de Biologia, Faculdade
de Ciências
Universidade do Porto
Porto, Portugal
Department of Zoology, Auckland Park
University of Johannesburg
Johannesburg, South Africa
ISBN 978-3-030-03082-7 ISBN 978-3-030-03083-4 (eBook)

https://doi.org/10.1007/978-3-030-03083-4
Library of Congress Control Number: 2018966550
© The Editor(s) (if applicable) and The Author(s) 2019. This book is an open access publication
Open Access This book is licensed under the terms of the Creative Commons Attribution 4.0
International License (http://creativecommons.org/licenses/by/4.0/), which permits use, sharing,
adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit
to the original author(s) and the source, provide a link to the Creative Commons licence and indicate if
changes were made.
The images or other third party material in this book are included in the book’s Creative Commons
licence, unless indicated otherwise in a credit line to the material. If material is not included in the book’s
Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the
permitted use, you will need to obtain permission directly from the copyright holder.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication
does not imply, even in the absence of a specific statement, that such names are exempt from the relevant
protective laws and regulations and therefore free for general use.
The publisher, the authors, and the editors are safe to assume that the advice and information in this book
are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the
editors give a warranty, express or implied, with respect to the material contained herein or for any errors
or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims
in published maps and institutional affiliations.
This Springer imprint is published by the registered company Springer Nature Switzerland AG.
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
To the new generation of Angolan
students of biodiversity. May they stand
on the shoulders of giants: the founders
of Angola’s biodiversity science
Friedrich Martin Josef Welwitsch (1806–1872)
José Vicente Barbosa du Bocage (1823–1907)
José Alberto de Oliveira Anchieta (1832–1897)
Johannes (John) Gossweiler (1873–1952)
Contributors to this volume record with
sadness the passing of William Roy (Bill)
Branch (1946–2018) - herpetologist,
indefatigable field researcher and mentor
of young Angolan scientists
Foreword
His Excellency the President of the Republic of Angola, João Manuel Gonçalves Lourenço
Angola occupies only four percent of the terrestrial area of the African continent,
yet it possesses the highest number of biomes of any African country. It is second in
terms of the number of ecoregions represented within its borders. It has ecosystems
as diverse as the rainforests of Maiombe of Cabinda to the vegetation-less dunes of
Namibe and the endless savannas and woodlands of the Cuando Cubango to the tiny
remnant forests of the highest valleys of Mount Moco in Huambo. It is the only
home to the most magnificent mammal in the world – the Giant Sable Antelope.
It was in Angola that one of the most extraordinary plant species Welwitschia
mirabilis was discovered and described – the enigmatic ‘living fossil’ of the desert.
It even puzzled Charles Darwin, who compared its evolutionary importance in the
plant kingdom to that of the Duck-billed Platypus in the animal kingdom.
Considering real fossils, Angola’s history goes back hundreds of millions of years,
to the earliest known living organisms, the bacterial stromatolites of the limestones
vii
viii Foreword
of Bembe and Humpata. Angola’s fossils range in size from microorganisms to the
gigantic dinosaur – Angolatitan adamastor – recently discovered in the sediments
of the Bengo coastline. Yet, despite this globally significant natural wealth, Angola
remains one of the least well-documented countries in the world in terms of its bio-
diversity. This situation is about to change.
Angolan scientists have collaborated with over 40 colleagues from 7 countries to
produce a new synthesis of knowledge of Angola’s remarkable biodiversity. They
have produced a magnificent volume that seeks to review all what is known about
Angola’s biodiversity, especially that which has been revealed through studies
undertaken in the twenty-first century. For several decades, field studies were ren-
dered nearly impossible because of the disruptions of war. But since peace was
achieved in 2002, a new generation of research has been made possible, bringing
many foreign specialists into partnerships with Angolan scientists and institutions
and introducing new technologies that have helped stimulate an unprecedented
wave of research activity.
Angola’s indigenous knowledge acquired over millennia provided the founda-
tions to the information documented and materials collected by visiting researchers
from the eighteenth to twenty-first centuries. Many detailed accounts have been
published over the past century and more, but often in scientific journals and official
reports that have been lost with the passage of time. More important, the knowledge
that does exist is fragmentary and largely inaccessible to Angolan students and
researchers. Much is published in foreign languages, and thus the inexistence of a
comprehensive synthesis of studies on Angola’s fauna, flora and ecosystems is a
challenge for young researchers. Angolan students, researchers and government
officials have very limited access to data sources where they can find a reliable,
science-based and up-to-date summary of that which has been recorded on the
country’s biodiversity.
It was the need for an integrated ‘state-of-knowledge’ summary, recognised dur-
ing the past decade by many Angolan university and government colleagues, that was
the catalyst that stimulated this important project. From humble beginnings by a few
Angolan and foreign partners, the effort expanded into the present volume of over
500 pages of authoritative accounts on our landscapes, seascapes, vegetation, flora
and fauna, its past and future. Most importantly, this work identifies the exciting
opportunities for research and conservation that Angolan scientists, conservationists,
government officials and the general public can embrace as the country moves for-
ward to an ever greater and more prosperous and environmentally sustainable future.
It is a pleasure to endorse this valuable contribution to the new wave of Angolan
scientific and conservation literature, a source of inspiration to our students and a
reminder to all our leaders, young and old, of our responsibility to treasure and
safeguard Angola’s unsurpassed, but vulnerable, biodiversity and natural resources.
President of the Republic of Angola João Manuel Gonçalves Lourenço

Luanda, Angola
Acknowledgements
This book was conceived as a collaborative and voluntary endeavour shared by

students of Angola’s biodiversity. Contributors were drawn from Angola, Britain,
Germany, Namibia, Portugal, Swaziland, South Africa, the Netherlands and the
United States. To all contributors to this synthesis, the Editors offer their thanks for
the unstinting efforts of the synthesis team to meet the strict demands of quality and
of timelines.
Over the past decades, biodiversity research in Angola has been encouraged by
successive leaders within the government, from academics and from the general
public of Angola. On behalf of all the contributors to this volume, the Editors wish
to thank the Ministers, past and present, of Science and Technology, Dr Cândida
Teixeira and Dr Maria do Rosário Bragança Sambo, and of Environment, Dr Fátima
Jardim and Dr Paula Francisco, for the support given to students of Angola’s biodi-
versity. Similarly, the encouragement and logistical support of Prof Liz Matos, Prof
Serôdio d’Almeida (Universidade Agostinho Neto), Dr Charles Skinner (De Beers,
Angola) and General João Traguedo (Lubango) are gratefully acknowledged.
Without their strong support, the scientific results described in this synthesis would
not have been possible.
Special thanks are due to Martim Melo, who provided his superb copy-editing,
proofreading and technical support during the final preparation of the manuscript
for submission to the publishers. Similarly, Pedro Tarroso and John Mendelsohn
offered their graphic design skills to greatly improve many figures and maps.
Photos were also generously offered by the chapter contributors and by Maans
Booysen, Merle Huntley, Tassos Leventis, Lars Petersson, Fiona Tweedie and
Alexandre Vaz.
The financial and logistic support of CIBIO and the UNESCO Chair Life on
Land (University of Porto, in association with Twinlab partners in Angola, Namibia,
Mozambique, South Africa and Zimbabwe) in this project was fundamental to its
success.
ix
x Acknowledgements
Finally, our thanks go to Margaret Deignan, Esther Rentmeester, Malini

Arumugam and Maadhuri Kandrakota of Springer (English Edition) and to Jorge
Reis-Sa of Arte e Ciência (Portuguese Edition), for their consistent and professional
support throughout the project.
Contents
Part I Introduction: Setting the Scene

1 Angolan Biodiversity: Towards a Modern Synthesis�� 3
Brian J. Huntley and Nuno Ferrand
2 Angola in Outline: Physiography, Climate and Patterns
of Biodiversity�� 15
Brian J. Huntley
3 Marine Biodiversity of Angola:
Biogeography and Conservation�� 43
Stephen P. Kirkman and Kumbi Kilongo Nsingi
4 The Fossil Record of Biodiversity in Angola Through
Time: A Paleontological Perspective�� 53
Octávio Mateus, Pedro M. Callapez, Michael J. Polcyn,
Anne S. Schulp, António Olímpio Gonçalves,
and Louis L. Jacobs
Part II Flora, Vegetation and Landscape Change

5 The Flora of Angola: Collectors, Richness and Endemism�� 79
David J. Goyder and Francisco Maiato P. Gonçalves
6 Vegetation Survey, Classification and Mapping in Angola�� 97
Rasmus Revermann and Manfred Finckh
7 Suffrutex Dominated Ecosystems in Angola�� 109
Paulina Zigelski, Amândio Gomes, and Manfred Finckh
8 Landscape Changes in Angola�� 123
John M. Mendelsohn
xi
xii Contents
Part III Invertebrate Diversity: Environmental Indicators

9 The Dragonflies and Damselflies of Angola:
An Updated Synthesis�� 141
Jens Kipping, Viola Clausnitzer, Sara R. F. Fernandes Elizalde,
and Klaas-Douwe B. Dijkstra
10 The Butterflies and Skippers (Lepidoptera: Papilionoidea)
of Angola: An Updated Checklist �� 167
Luís F. Mendes, A. Bivar-de-Sousa, and Mark C. Williams
Part IV Vertebrates: Distribution and Diversity

11 The Freshwater Fishes of Angola �� 207
Paul H. Skelton
12 The Amphibians of Angola: Early Studies and the Current
State of Knowledge�� 243
Ninda Baptista, Werner Conradie, Pedro Vaz Pinto,
and William R. Branch
13 The Reptiles of Angola: History, Diversity,
Endemism and Hotspots�� 283
William R. Branch, Pedro Vaz Pinto, Ninda Baptista,
and Werner Conradie
14 The Avifauna of Angola: Richness, Endemism and Rarity�� 335
W. Richard J. Dean, Martim Melo, and Michael S. L. Mills
15 The Mammals of Angola �� 357
Pedro Beja, Pedro Vaz Pinto, Luís Veríssimo, Elena Bersacola,
Ezequiel Fabiano, Jorge M. Palmeirim, Ara Monadjem,
Pedro Monterroso, Magdalena S. Svensson, and Peter John Taylor
16 The Cetaceans (Whales and Dolphins) of Angola �� 445
Caroline R. Weir
17 The Giant Sable Antelope: Angola’s National Icon�� 471
Pedro Vaz Pinto
Part V Research and Conservation Opportunities

18 Biodiversity Conservation: History, Protected Areas
and Hotspots �� 495
Brian J. Huntley, Pedro Beja, Pedro Vaz Pinto, Vladimir Russo,
Luís Veríssimo, and Miguel Morais
Contents xiii
19 Museum and Herbarium Collections for Biodiversity

Research in Angola�� 513
Rui Figueira and Fernanda Lages
20 Conclusions: Biodiversity Research
and Conservation Opportunities�� 543
Vladimir Russo, Brian J. Huntley, Fernanda Lages,
and Nuno Ferrand
Contributors
Ninda Baptista Instituto Superior de Ciências da Educaҫão da Huíla, Rua

Sarmento Rodrigues, Lubango, Angola
National Geographic Okavango Wilderness Project, Wild Bird Trust, Parktown,
Gauteng, South Africa
CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos,
Laboratório Associado, Universidade do Porto, Vairão, Portugal
Pedro Beja CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos
Genéticos, Universidade do Porto, Vairão, Portugal
CEABN-InBio, Centro de Ecologia Aplicada “Professor Baeta Neves”, Instituto
Superior de Agronomia, Universidade de Lisboa, Lisboa, Portugal
Elena Bersacola Nocturnal Primate Research Group, Faculty of Humanities and
Social Sciences, Oxford Brookes University, Oxford, UK
A. Bivar-de-Sousa Museu Nacional de História Natural e da Ciência, Universidade
de Lisboa, Lisboa, Portugal
Sociedade Portuguesa de Entomologia, Lisboa, Portugal
William R. Branch (deceased) National Geographic Okavango Wilderness
Project, Wild Bird Trust, Parktown, Gauteng, South Africa
Department of Zoology, Nelson Mandela University, Port Elizabeth, South Africa
Pedro M. Callapez CITEUC; Departamento de Ciências da Terra, Faculdade de
Ciências e Tecnologia, Universidade de Coimbra, Coimbra, Portugal
Viola Clausnitzer Senckenberg Museum for Natural History, Görlitz, Görlitz,
Germany
xv
xvi Contributors
Werner Conradie National Geographic Okavango Wilderness Project, Wild Bird

Trust, Hogsback, South Africa
School of Natural Resource Management, Nelson Mandela University, George,
South Africa
Port Elizabeth Museum (Bayworld), Humewood, South Africa
W. Richard J. Dean DST-NRF Centre of Excellence at the FitzPatrick Institute,
University of Cape Town, Rondebosch, South Africa
Klaas-Douwe B. Dijkstra Naturalis Biodiversity Center, Leiden, The Netherlands
Sara R. F. Fernandes Elizalde SASSCAL – BID GBIF, Instituto de Investigação
Agronómica, Huambo, Angola
Ezequiel Fabiano Department of Wildlife Management and Ecotourism, Katima
Mulilo Campus, Faculty of Agriculture and Natural Resources, University of
Namibia, Katima Mulilo, Namibia
Nuno Ferrand CIBIO-InBIO, Centro de Investigação em Biodiversidade e
Recursos Genéticos, Laboratório Associado, Campus de Vairão, Universidade do
Porto, Vairão, Portugal
Departamento de Biologia, Faculdade de Ciências, Universidade do Porto, Porto,
Portugal
Department of Zoology, Auckland Park, University of Johannesburg, Johannesburg,
South Africa
Rui Figueira CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos
Genéticos, Universidade do Porto, Vairão, Portugal
CEABN-InBio, Centro de Ecologia Aplicada “Professor Baeta Neves”, Instituto
Superior de Agronomia, Universidade de Lisboa, Lisboa, Portugal
Manfred Finckh Institute for Plant Science and Microbiology, University of
Hamburg, Hamburg, Germany
Amândio Gomes Institute for Plant Science and Microbiology, University of
Faculty of Sciences, Agostinho Neto University, Luanda, Angola
António Olímpio Gonçalves Departamento de Geologia, Faculdade de Ciências,
Universidade Agostinho Neto, Luanda, Angola
Francisco Maiato P. Gonçalves National Geographic Okavango Wilderness
Project, Wild Bird Trust, Parktown, Gauteng, South Africa
Instituto Superior de Ciências da Educaҫão da Huíla, Lubango, Angola
University of Hamburg, Institute for Plant Science and Microbiology, Hamburg,
Germany
Contributors xvii
David J. Goyder Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey,
UK
National Geographic Okavango Wilderness Project, Wild Bird Trust,
Parktown, Gauteng, South Africa
Brian J. Huntley CIBIO-InBIO, Centro de Investigação em Biodiversidade e
Recursos Genéticos, Universidade do Porto, Vairão, Portugal
Louis L. Jacobs Roy M. Huffington Department of Earth Sciences, Southern
Methodist University, Dallas, TX, USA
Jens Kipping BioCart Ökologische Gutachten, Taucha/Leipzig, Germany
Stephen P. Kirkman Department of Environmental Affairs, Oceans and Coasts
Research, Cape Town, South Africa
Fernanda Lages ISCED – Instituto Superior de Ciências da Educação da Huíla,
Lubango, Angola
Octávio Mateus GeoBioTec, Faculdade de Ciências e Tecnologia, Universidade
Nova de Lisboa, Lisbon, Portugal
Museu da Lourinhã, Rua João Luis de Moura, Lourinhã, Portugal
Martim Melo DST-NRF Centre of Excellence at the FitzPatrick Institute,
University of Cape Town, Rondebosch, South Africa
Instituto Superior de Ciências da Educaҫão de Huíla, Lubango, Angola
John M. Mendelsohn RAISON (Research & Information Services of Namibia),
Windhoek, Namibia
Luís F. Mendes Museu Nacional de História Natural e da Ciência, Universidade de
Lisboa, Lisboa, Portugal
CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Vairão,
Portugal
Michael S. L. Mills Instituto Superior de Ciências da Educaҫão de Huíla, Lubango,
Angola
A. P. Leventis Ornithological Research Institute, University of Jos, Jos, Plateau
State, Nigeria
Ara Monadjem Department of Biological Sciences, University of Swaziland,
Kwaluseni, Swaziland
Mammal Research Institute, Department of Zoology and Entomology, University of
Pretoria, Pretoria, South Africa
xviii Contributors
Pedro Monterroso CIBIO-InBIO, Centro de Investigação em Biodiversidade e

Recursos Genéticos, Universidade do Porto, Vairão, Portugal
Miguel Morais Faculdade de Ciências, Universidade Agostinho Neto, Luanda,
Angola
Kumbi Kilongo Nsingi Benguela Current Convention, Swakopmund, Namibia
Jorge M. Palmeirim Departamento de Biologia Animal, Faculdade de Ciências,
cE3c – Centre for Ecology, Evolution and Environmental Changes, Universidade de
Lisboa, Lisboa, Portugal
Pedro Vaz Pinto Fundação Kissama, Luanda, Angola
Universidade do Porto, Campus de Vairão, Vairão, Portugal
Michael J. Polcyn Roy M. Huffington Department of Earth Sciences, Southern
Methodist University, Dallas, TX, USA
Rasmus Revermann Institute for Plant Science and Microbiology, University of
Vladimir Russo Fundação Kissama, Luanda, Angola
Anne S. Schulp Naturalis Biodiversity Center, Leiden, The Netherlands
Faculty of Earth and Life Sciences, VU University Amsterdam, Amsterdam, The
Netherlands
Paul H. Skelton South African Institute for Aquatic Biodiversity (SAIAB),
Grahamstown, South Africa
Wild Bird Trust, National Geographic Okavango Wilderness Project, Hogsback,
South Africa
Magdalena S. Svensson Nocturnal Primate Research Group, Faculty of Humanities
and Social Sciences, Oxford Brookes University, Oxford, UK
Peter John Taylor School of Mathematical & Natural Sciences, University of
Venda, Thohoyandou, South Africa
Luís Veríssimo Fundação Kissama, Luanda, Angola
Caroline R. Weir Ketos Ecology, Kingsbridge, Devon, UK
Mark C. Williams Pretoria University, Pretoria, South Africa
Paulina Zigelski Institute for Plant Science and Microbiology, University of
Part I
Introduction: Setting the Scene
Chapter 1
Angolan Biodiversity: Towards a Modern
Synthesis
Brian J. Huntley and Nuno Ferrand
Abstract Angola possesses an unusually rich diversity of ecosystems and species,

but this natural wealth is poorly documented when compared with other countries in
the region. Both colonial history and extended wars challenged progress in biodiver-
sity research and conservation, but since peace was achieved in 2002 a rapidly
increasing level of collaboration between Angolan and visiting scientists and institu-
tions has seen a blossoming of biodiversity research. The absence of comprehensive
reviews and syntheses of existing knowledge, often published in extinct journals and
inaccessible official reports, necessitates a modern synthesis. This volume brings
together the existing body of scientific results from studies on Angola’s landscapes,
ecosystems, flora and fauna, and presents an outline of opportunities for biodiversity
discovery, understanding and conservation as well as collaborative research.
Keywords Africa · Biomes · Collaborative research · Conservation · Ecoregions
Background and Context
Angola is a country of unusually rich physiographic, climatic and biological diver-

sity. It occupies only 4% of the terrestrial area of Africa, yet it possesses the highest
diversity of biomes and is second only to mega-diverse South Africa in terms of the
number of ecoregions found within its borders. However, scientific literature on its
B. J. Huntley (*)
CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade
do Porto, Vairão, Portugal
e-mail: brianjhuntley@gmail.com
N. Ferrand
CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Laboratório
Associado, Campus de Vairão, Universidade do Porto, Vairão, Portugal
Departamento de Biologia, Faculdade de Ciências, Universidade do Porto, Porto, Portugal
Department of Zoology, Auckland Park, University of Johannesburg, Johannesburg,
South Africa
e-mail: nferrand@cibio.up.pt
© The Author(s) 2019 3

B. J. Huntley et al. (eds.), Biodiversity of Angola,
https://doi.org/10.1007/978-3-030-03083-4_1
4 B. J. Huntley and N. Ferrand
biodiversity is extremely limited when compared with most African countries.

Much of that which has been published is difficult to access or out of print. This
volume seeks to redress this situation.
Here we present a review of what is known about Angola’s biodiversity. Much of
the existing literature dates from the nineteenth and early to mid-twentieth centu-
ries. Following independence in 1975, field studies were curtailed by the instabili-
ties of an extended civil war. It was not until after the peace settlement of 2002 that
a new wave of research has been possible. Initial attempts to establish collaborative
field expeditions were frustrated by visa and permit restrictions, but these chal-
lenges were gradually overcome and by the 2010s a vibrant programme of joint
projects has evolved. Today many foreign specialists work in partnership with
Angolan researchers and institutions, producing a new flow of scientific results of
which many are presented in this volume.
For any comprehensive synthesis, both temporal depth and spatial breadth is
necessary. An historical perspective is presented in each chapter. Angolan indige-
nous knowledge has contributed to the insights and materials that have informed
visiting researchers from the eighteenth century to the present day. The pioneering
studies and exhaustive botanical collections of the Austrian botanist, Friedrich
Welwitsch (1806–1872), the zoological collections of the indefatigable Portuguese
naturalist José Anchieta (1832–1897) and the Swiss botanist John Gossweiler
(1873–1952) set benchmarks for later work (Swinscow 1972; de Andrade 1985).
Each succeeding student of Angola has added to the description of its biological
diversity. While botanists such as Romero Monteiro (1970) and zoologists such as
Crawford-Cabral (1983) have summarised available biogeographic information
within a national context, no comprehensive synthesis of studies on Angola’s fauna,
flora and ecosystems has yet been undertaken. The need for an integrated account
has become evident in the past decade, as increasing numbers of expeditions and
collaborative projects have evolved as part of the country’s ‘peace dividend’.
Approach and Purpose of This Synthesis
A modern synthesis is not easily achieved. Much of the early literature on Angola’s
biodiversity resides in publications and reports that are difficult to source. This
review attempts to reference these important but sometimes elusive accounts, in
order to provide students with access to what information is available. While focus-
ing on papers in peer-reviewed journals, some topics need to draw on unpublished
reports filed in government departments. It also seeks to bring together the findings
of recent, post-independence studies, many of which are still in progress or in press.
It is intended to serve the new generation of Angolan students by providing a com-
prehensive but focused synopsis of what is known on the biomes, landscapes, flora
and fauna of Angola. It should also bring Angola to the attention of researchers
across Africa and beyond, revealing the great diversity of life, and the multiple
questions on the structure and functioning of Angola’s biodiversity that await explo-
ration, examination and explanation.
1 Angolan Biodiversity: Towards a Modern Synthesis 5
In structuring this book, this introduction leads through synopses on the coun-
try’s terrestrial and marine biogeography, paleontological record, recent landscape
evolution and land transformation, to chapters on its flora and vegetation. The main
body of this volume is devoted to accounts of its fauna – selected invertebrate groups
that have promise as indicators of environmental stress, and all vertebrate groups. In
each treatment, the need for increased conservation measures for threatened taxa
and habitats is a recurrent theme, while research opportunities are highlighted.
While general inventories and checklists are progressing well, the state of ecologi-
cal knowledge remains rudimentary. Topics as fundamental as ecological processes
such as the flows of energy, water and nutrients; the ecological impacts of phenom-
ena such as fire, invasive species, herbivory, droughts and frosts; community struc-
ture, plant-animal interactions and the impacts of land-transformation and of climate
change are yet to be researched in Angola. This volume’s content is limited by the
availability of information. It is therefore opportunistic, covering those taxonomic
groups and those features and processes for which a critical mass of information is
available. The focus is primarily on the terrestrial ecosystems and biota of Angola,
but the importance of the marine environment is described in accounts on marine
biodiversity and ocean dynamics, and on the richness of the whale, dolphin and
marine turtle faunas of Angolan waters.
In comparison with similar reviews for other African countries with long and
strong traditions of research into their biodiversity and ecology, and for which com-
prehensive syntheses of the state of knowledge are available (e.g. Namibia: Barnard
1998; Southern Africa: Davis 1964; Werger and van Bruggen 1978; Huntley 1989;
Tanzania: Sinclair 2012), this account reveals both the strengths and weaknesses of
the research agenda of the colonial era, and the challenges of the recent past. While
institutions such as the Instituto de Investigação Científica de Angola and the
Instituto de Investigação Agronómica de Angola undertook very important studies
on many taxa, and on vegetation, soils and agronomy, and the Museu do Dundo
amassed and distributed a vast series of collections of the animal species of the
Lundas, the coverage of disciplines and of the remote regions of Angola was weak.
Biodiversity Surveys: Historical Synopsis
The history of scientific exploration and biological collection in Angola is relatively

modest. Whereas South Africa, by 1975, had over three million herbarium speci-
mens collected by some 2500 botanists since the late eighteenth century (Gunn and
Codd 1981), Angola had less than 300,000 specimens collected by just 300 bota-
nists during the same period (Figueiredo and Smith 2008). Despite the relatively
limited coverage of Angolan collections, the great botanist Francisco Mendonça
was occasioned to state in his preface to Gossweiler and Mendonça (1939):
We are happily able to confirm that the flora of Angola is the best known in tropical Africa,
due to the attention given by the state towards the botanical exploration of the colony, and
the great interest and zeal of scientists in its study.
The Swiss zoologist, Monard (1935), had been less sanguine:

A regrettable fact about the Natural History of Angola is the scarcity of concrete informa-
tion about the nature, distribution, and habits of the large game. The Boers … never com-
municated their observations. The Portuguese hunters … did not write reports on their
hunts, or if they did, did so in newspapers or magazines that never enter the scientific litera-
ture. The observations remain, in this way, lost to the naturalist who is not able to locate
such work.
In truth, during the colonial era, investment in research on the country’s biodiversity
was limited. The achievements of early pioneers such as Friedrich Welwitsch, José
Anchieta and John Gossweiler were quite remarkable, and those of more recent
agronomists, botanists and zoologists such as Castanheira Diniz, Romero Monteiro,
Grandvaux Barbosa, Brito Teixeira, Crawford-Cabral, Rosa Pinto, Barros Machado,
etc., were equally laudable, indeed amazing.
The war years from 1975 to 2002 saw very few researchers venturing into the
field. Most activities were limited to brief searches for remnant populations of giant
sable (Estes 1982), marine turtles (Carr and Carr 1991), birds (Günther and Feiler
1986a, b; Hawkins 1993), and a countrywide assessment of the state of wildlife
populations (Huntley and Matos 1992). The Southern African Botanical Diversity
Network (SABONET) project attempted to stimulate botanical studies in Angola
from the mid-1990s (Huntley et al. 2006), while the Kissama Foundation funded a
vegetation survey of the northern extreme of Quiçama (Jeffrey 1996) and intro-
duced a mixed assemblage of antelope and ostriches into the park in 2000 (Walker
2004). The last decades of the twentieth century were aptly described as a period of
confusão – confusion (Maier 2007). In brief, from Angola’s independence in 1975,
until the twenty-first century, cooperative field research over most of the country
was challenged by the impacts of war. But dramatic and positive change came with
the dawn of the new millennium.
Research Collaboration in the Twenty-First Century
From 2000, especially after the peace agreement of April 2002, field activities
expanded rapidly. Most notably, Vaz Pinto has focused on a long-term study on
giant sable in Cangandala (Walker 2004; Vaz Pinto 2018), Morais (2017) has led
surveys of marine turtles along the Angolan coast, and Mills (2010, 2018) has
undertaken field studies on birds across the country.
International support for environmental conservation and research strengthened
from 2001, when the Global Environment Facility, through the United Nations
Development Programme, initiated a multidisciplinary project to develop a trans-
boundary diagnostic analysis of hydro-environment threats within the Okavango
River Basin, known as the Environmental Protection and Sustainable Management
of the Okavango River Basin Project (EPSMO). The project’s aim was to facilitate
the protection of the Basin’s aquatic ecosystems and biological diversity (OKACOM
2009, 2011). The project included participation from Angola, Botswana and
Namibia and provided a strong impetus to future multi-national projects in the

Basin. A further initiative, the Integrated River Management Project, was funded by
USAID/Southern Africa between 2004 and 2009 and provided both institutional
and management planning support to the national partners (OKACOM 2009). The
EPSMO project was succeeded by the SAREP project described below.
The OKACOM projects were focused on major water management needs and
did not embrace detailed biodiversity surveys. Indeed, until 2009, biodiversity
research activities in Angola had been essentially individual efforts, with limited
funding. Difficulties experienced in obtaining visas to visit, and permits to collect
specimens in Angola were a continuing challenge faced by foreign scientists. With
the signing of an agreement between the South African National Biodiversity
Institute (SANBI), the Angolan Ministry of Environment and the Instituto Superior
de Ciências da Educação (ISCED), Lubango, in 2009, more ambitious cooperative
biodiversity projects became possible. Initially designed as training exercises, the
series of Rapid Biodiversity Assessments, in Huíla/Namibe (Huntley 2009), Lunda-
Norte (Huntley 2011; Huntley and Francisco 2015) and across western Angola
(Rejmánek et al. 2017), brought over 40 scientists from 14 countries to Angola to
work with local students and researchers.
By the early 2010s, a wide diversity of major cooperative programmes had
developed, including those of the Southern African Regional Environmental
Programme (SAREP), the Southern African Science Service Centre for Climate
Change and Adaptive Land Management (SASSCAL) (Revermann et al. 2018), the
National Geographic Okavango Wilderness Project (NGOWP 2018), and conserva-
tion initiatives of NGOs such as Elephants without Borders, Panthera, Peace Parks
Foundation, the Kavango-Zambezi Transfrontier Conservation Area (KAZA) proj-
ect and several others. Collaboration between foreign museums and universities and
Angolan counterparts stimulated additional specialist interests, collectively gaining
momentum until the present. In October 2012, CIBIO (Research Centre in
Biodiversity and Genetic Resources) at the University of Porto, Portugal, and
ISCED-Huíla (Lubango) established a collaborative research, capacity building and
advanced training project – the ISCED/CIBIO TwinLab initiative. The initiative
was soon replicated in South Africa, Mozambique, Namibia and Zimbabwe, and the
whole network of TwinLabs now forms a UNESCO Chair Life on Land, awarded at
the end of 2017.
For much of the post-independence period, biodiversity research efforts had been
un-coordinated and opportunistic. With the establishment of the Instituto Nacional
de Biodiversidade e Áreas de Conservação (INBAC) in 2011, the opportunity arose
for a greater level of coordination and priority setting. The Plano Estratégico da
Rede Nacional da Áreas de Conservação de Angola (GoA 2011), provided a stimu-
lus to studies in key biodiversity hotspots such as Mount Moco, Mount Namba,
Serra da Neve, Serra Pingano, Cumbira, Lagoa Carumbo, and to the vast and very
poorly researched catchments of the Cuando Cubango. While a greater level of
inter-institutional collaboration is still possible, the momentum developed over the
past decade has been unprecedented since 1975. The successes of the recent past are
presented in this volume, often drawing on work that is still in progress, is unpub-
lished, or is in press.
Chapter Outlines
Angola is a large country, and as emphasised throughout this volume, it has a rich
diversity of landscapes, seascapes and associated biomes and ecoregions. The his-
tory of biodiversity research in Angola stretches over 200 years. The spatial, tempo-
ral and taxonomic scales embraced in this book results in it being structured in five
parts. Part I, Chap. 1 (Huntley and Ferrand this chapter) provides an introduction to
the book and its content. Chap. 2 (Huntley 2019) outlines the country’s biogeogra-
phy, drawing on the long history of geomorphological and landscape analysis in
Angola, and describes the diversity of seven terrestrial biomes, 15 ecoregions and
32 vegetation types. In Chap. 3 Kirkman and Nsingi (2019) synthesise the findings
of recent multi-national research activities on the Benguela Current Large Marine
Ecosystem project and other studies on Angola’s coastal and marine systems. The
long history of the evolution of Angola’s biota is introduced by Mateus et al. (2019)
in Chap. 4, where the exciting recent discoveries in Angola’s fossil record, most
especially that of the Cretaceous, is described. A highlight was the discovery of the
sauropod dinosaur Angolatitan adamastor, the first dinosaur to be found in Angola
(Mateus et al. 2011). These authors emphasise the fact that for very long periods –
hundreds of millions of years – the absence of fossiliferous rocks in Angola excludes
the possibility of tracking animal and plant evolution in Angola over such long
periods.
Part II presents an historical and contemporary analysis of our understanding of
the country’s flora and vegetation and on curious patterns and evolutionary pro-
cesses in some typical Angolan plant communities. In Chap. 5 Goyder and Gonçalves
(2019) note that the vascular flora now totals 6850 species, with 14.8% of these
being endemic. The two early vegetation maps of Angola, prepared by the pioneers
Gossweiler and Mendonça (1939) and Barbosa (1970), having served the country
for many decades, now deserve renewed mapping efforts at a finer scale, using mod-
ern remote sensing and numerical analysis approaches, as recommended by
Revermann and Finckh (2019) in Chap. 6. Of the many intriguing features of
Angolan vegetation, the patterns of plant community/soil/animal associations, such
as the ‘fairy circles’ of the Namib (Juergens 2013; Cramer and Barger 2013), ‘fairy
forests’ of the miombo, and the influence of coastal fog on desert vegetation and
fauna; are of special interest to ecologists. Few of these phenomena have been ade-
quately interpreted, but Zigelski et al. (2019) in Chap. 7 presents recent studies on
the ‘underground forests’ of the chanas de ongote of the Angolan plateau. The land-
scapes of Angola are not static, being subject to multiple processes of transforma-
tion. In Chap. 8, Mendelsohn (2019) uses results from satellite technologies and
ground surveys to describe the dramatic impacts of deforestation, fires, mining and
agricultural activities on vegetation, soils and water quality at landscape scales.
Part III details the results of surveys that have advanced rapidly over the past two
decades on two invertebrate groups – dragonflies and butterflies. These colourful
and taxonomically distinctive insects are known to be sensitive to subtle changes in
environmental conditions, such as forest cover and water quality, and serve as effec-
tive indicators of change in environmental health. The chapters (9 and 10) on but-
terflies (Mendes et al. 2019) and on dragonflies (Kipping et al. 2019) have enriched
Angola’s knowledge of these important ecological groups. Prior to 2009, for exam-
ple, only 158 species of dragonflies and damselflies were known from Angola. By
2018 this number had increased to 260. The butterfly checklist now stands at 792
species and subspecies, up by over 220 species and subspecies since the turn of the
millennium. In contrast to the encouraging progress in these taxa, key environmen-
tal engineers – ants and termites – remain poorly documented and await study.
A major component of this volume has been devoted to the vertebrate taxa that
have enjoyed the attention of scientists active in Angola since the mid-nineteenth
century. Part IV presents detailed accounts of the pioneering work of such luminar-
ies as Anchieta, Bocage, Boulenger, Machado, Rosa Pinto and Crawford-Cabral,
but also of the very many other contributors to the inventory of Angola’s vertebrate
fauna. Skelton (2019), Chap. 11, provides a concise summary of what is known of
Angola’s ca. 358 species of freshwater fishes (of which 22% are endemic), and also
presents a model of post-Cretaceous biogeography of Angola and the roles of
regional tectonics and river capture on the speciation and distribution of the fish
fauna. Baptista et al. (2019), Chap. 12, cover the amphibian fauna, noting that the
group clearly deserves further survey given that thus far only 111 species have been
recorded (compared to 128 species for similar-sized but much drier and cooler
South Africa). In Chap. 13, Branch et al. (2019) offer a comprehensive account on
the 278 species of Angolan reptiles and their patterns of diversity and endemism,
documenting key reptile hotspots deserving further exploration. They predict that as
many as 75 new species of lizards await discovery in Angola. Both Branch and
Baptista demonstrate the value of molecular phylogenetics in clarifying taxonomic
complexes in reptiles and frogs.
Angola, with ca. 940 bird species recorded, has in recent years become a favoured
destination of ecotourists searching for the country’s 29 endemic bird species, and
Dean et al. (2019) provide in Chap. 14, a chronology of ornithological surveys, a
listing of endemics and near-endemics, and sites of special interest to bird enthusi-
asts, both professional and amateur. They emphasise, as highlighted by Hall (Hall
1960), the faunistic importance of the Angolan Escarpment, and also of the relict
Afromontane forests of the highlands (Vaz Silva 2015), as areas of critical impor-
tance to understanding the evolution of Africa’s avifauna. These isolated, frag-
mented and rapidly declining forests merit the highest level of protection to secure
their futures as evolutionary fingerprints of the past.
A team of ten mammal specialists, coordinated by Beja et al. (2019), present a
major synthesis (Chap. 15) on Angola’s 291 mammal species. This chapter fills a
need felt since Hill and Carter’s (1941) benchmark study, and the more recent
coverage of ungulates by Crawford-Cabral and Veríssimo (2005). With 73 species
of bats (one third of the bat species known for Africa) Angola has the highest num-
ber of species for southern Africa, despite the comparatively limited intensity of
surveys undertaken to date. The most diverse mammal group, rodents, has 85 spe-
cies listed for Angola, of which 13 are endemic or near-endemic. While the number
of endemic mammal species is modest, the vulnerability of many species to extinc-
tion within Angola is high and deserves urgent conservation measures. Less well
known to Angolans than the terrestrial mammals is the country’s unusually rich
marine mammal fauna. The 28 species of cetaceans (whales and dolphins) found off
Angola’s coast have been the subject of surveys and research undertaken by Weir
(2019) since 2003. As noted in Chap. 16, the possible presence of a further seven
species of cetaceans in Angolan waters makes the country globally important for
marine mammal conservation.
The Angolan mammal that has enjoyed national and international attention is the
Giant Sable Antelope, which has been the subject of an intense research and conser-
vation project since 2002 (Chap. 17), led by Vaz Pinto (2019). The successful rescue
and rehabilitation of this national icon, from the brink of extinction, is a conserva-
tion model for which Angola can justifiably be proud. The success of the Giant
Sable Project needs replication for the many mammal species that are known to
have been reduced to very low numbers, or have been hunted to extinction in Angola.
These include most large carnivores – Cheetah, Lion, Wild Dog, plus many herbi-
vores – Black-faced Impala, Red Hartebeest, Lichtenstein’s Hartebeest, Tsessebe,
Southern Lechwe, Puku, Forest Buffalo, Giraffe, Black Rhino, Western Gorilla,
Chimpanzee, Forest Elephant and Manatee.
The final section of this volume (Part V) presents an overview of the country’s
conservation history and current opportunities for action, Chap. 18 (Huntley et al.
2019); and an introduction to the importance of natural history museums and her-
baria in the country’s biodiversity science and conservation agenda, Chap. 19
(Figueira and Lages 2019). What is abundantly clear, as expressed in the concluding
chapter, (Russo et al. 2019), is that Angola is alive with research and conservation
opportunities, stimulated by recent initiatives led by the Angolan government and
supported by the international community.
This volume provides a first synthesis of what is known and published about
Angola’s diverse landscapes, biomes and ecosystems and the species that inhabit
them. It is a humble attempt by its 46 contributors to place this knowledge before
researchers and conservationists in Angola and beyond, especially those who might
be stimulated to strengthen the imperfect understanding and vulnerable state of
Angola’s biodiversity. It is the fervent hope of this book’s editors that this volume
will provide an entry point for many young Angolan students to study the literature,
be inspired by the dedication, tenacity and wisdom of the early pioneers and con-
temporary explorers, and enter careers in field-based biodiversity research and con-
servation in Angola.
References
Baptista N, Conradie W, Vaz Pinto P et al (2019) The amphibians of Angola: early studies and the
current state of knowledge. In: Huntley BJ, Russo V, Lages F, Ferrand N (eds) Biodiversity of
Angola. Science & conservation: a modern synthesis. Springer Nature, Cham
Barbosa LAG (1970) Carta Fitogeográfica da Angola. Instituto de Investigação Científica de
Angola, Luanda, 343 pp
Barnard P (1998) Biological diversity in Namibia: a country study. Namibian National Biodiversity
Task Force, Windhoek, 325 pp
Beja P, Vaz Pinto P, Veríssimo L et al (2019) The mammals of Angola. In: Huntley BJ, Russo V,
Lages F, Ferrand N (eds) Biodiversity of Angola. Science & conservation: a modern synthesis.
Springer Nature, Cham
Branch WR, Vaz Pinto P, Baptista N et al (2019) The reptiles of Angola: history, diversity, ende-
mism and hotspots. In: Huntley BJ, Russo V, Lages F, Ferrand N (eds) Biodiversity of Angola.
Science & conservation: a modern synthesis. Springer Nature, Cham
Carr T, Carr P (1991) Surveys of the sea turtles of Angola. Biol Conserv 58(1):19–29
Cramer MD, Barger NN (2013) Are Namibian “fairy circles” the consequence of self-organizing
spatial vegetation patterning? PLoS One 8(8):e70876
Crawford-Cabral J (1983) Esboço zoogeográfico de Angola. Unpublished manuscript. Instituto de
Investigação Científica Tropical, Lisboa, 50 pp + 13 maps
Crawford-Cabral J, Veríssimo LN (2005) The ungulate fauna of Angola: systematic list, distri-
bution maps, database report. Estudos, Ensaios e Documentos do Instituto de Investigação
Científica Tropical 163:1–277
Davis DHS (ed) (1964) Ecological studies in Southern Africa. Junk, The Hague, 415 pp
de Andrade AAB (1985) O Naturalista José de Anchieta. Instituto de Investigação Científica
Tropical, Lisboa, 187 pp
Dean WRJ, Melo M, Mills MSL (2019) The avifauna of Angola: richness, endemism and rarity.
In: Huntley BJ, Russo V, Lages F, Ferrand N (eds) Biodiversity of Angola. Science & conserva-
tion: a modern synthesis. Springer Nature, Cham
Estes RD (1982) The giant sable and wildlife conservation in Angola. Report to IUCN Species
Survival Commission. Gland, Switzerland
Figueira R, Lages F (2019) Museum and herbarium collections for biodiversity research in Angola.
Figueiredo E, Smith GF (eds) (2008) Plants of Angola / Plantas de Angola. Strelitzia 22:1–279
GoA (Government of Angola) (2011) Plano Estratégico da Rede Nacional de Áreas de Conservação
de Angola (PLENARCA). Ministry of Environment, Luanda
Gossweiler J, Mendonça FA (1939) Carta Fitogeográfica de Angola. Ministério das Colónias,
Lisboa, 242 pp
Goyder DJ, Gonçalves FMP (2019) The Flora of Angola: collectors, richness and endemism. In:
Huntley BJ, Russo V, Lages F, Ferrand N (eds) Biodiversity of Angola. Science & conserva-
Gunn M, Codd LE (1981) Botanical exploration of Southern Africa. AA Balkema, Cape Town,
400 pp
Günther R, Feiler A (1986a) Zur phänologie, ökologie und morphologie angolanischer Vögel
(Aves). Teil I: Non-Passeriformes Faunistische Abhandlungen aus dem Staatlichen Museum
für Tierkunde in Dresden 13:189–227
Günther R, Feiler A (1986b) Zur phänologie, ökologie und morphologie angolanischer Vögel
(Aves). Teil II: Passeriformes Faunistische Abhandlungen aus dem Staatlichen Museum für
Tierkunde in Dresden 14:1–29
Hall BP (1960) The faunistic importance of the scarp of Angola. Ibis 102:420–442
Hawkins F (1993) An integrated biodiversity conservation project under development: the ICBP
Angola Scarp Project. Proceedings of the VIII Pan-African Ornithological Congress: 279–284.
Kigali, Rwanda, 1992. Koninklijk Museum voor Midden-Afrika, Tervuren
Hill JE, Carter TD (1941) The mammals of Angola, Africa. Bulletin of the American Museum of
Natural History 78, 211 pp
Huntley BJ (ed) (1989) Biotic diversity in southern Africa: concepts and conservation. Oxford
University Press, Oxford, 380 pp
Huntley BJ (2009) SANBI/ISCED/UAN Angolan biodiversity assessment capacity building proj-
ect. Report on pilot project. Unpublished Report to Ministry of Environment, Luanda 97 pp,
27 figures
Huntley BJ (2011) Biodiversity rapid assessment of the Lagoa Carumbo area, Lunda-Norte,
Angola. Expedition report. Ministry of Environment, Luanda
Huntley BJ (2019) Angola in outline: physiography, climate and patterns of biodiversity. In:
Huntley BJ, Francisco P (eds) (2015) Avaliação Rápida da Biodiversidade de Região da Lagoa
Carumbo, Lunda-Norte – Angola / Rapid Biodiversity Assessment of the Carumbo Lagoon
Area, Lunda-Norte – Angola. Ministério do Ambiente, Luanda, 219 pp
Huntley BJ, Matos L (1992) Biodiversity: Angolan environmental status quo assessment report.
IUCN Regional Office for Southern Africa, Harare, 55 pp
Huntley BJ, Siebert SJ, Steenkamp Y, et al (2006) The achievements of the southern African botan-
ical diversity network (SABONET) – a southern African botanical capacity building project.
In: Ghazanfar SA, Beentje H (eds.) Taxonomy and ecology of African plants, their conserva-
tion and sustainable use: Proceedings of the 17th AETFAT Congress. Addis Ababa, Ethiopia,
2003. Royal Botanic Gardens, Kew, pp 531–543
Huntley BJ, Beja P, Vaz Pinto P et al (2019) Biodiversity conservation: history, protected areas and
hotspots. In: Huntley BJ, Russo V, Lages F, Ferrand N (eds) Biodiversity of Angola. Science &
conservation: a modern synthesis. Springer Nature, Cham
Jeffrey R (1996) A Phytosociological Survey of the Northern Sector of the Quicama National
Park in Angola. B.Sc. (Hons.) Dissertation. Faculty of Biological and Agricultural Sciences,
University of Pretoria, Pretoria
Juergens N (2013) The biological underpinnings of Namib Desert fairy circles. Science
339:1618–1621
Kipping J, Clausnitzer V, Fernandes Elizalde SRF et al (2019) The dragonflies and damselflies of
Angola. In: Huntley BJ, Russo V, Lages F, Ferrand N (eds) Biodiversity of Angola. Science &
Kirkman SP, Nsingi KK (2019) Marine biodiversity of Angola: biogeography and conservation. In:
Maier K (2007) Angola: promises and lies. Serif, London, 224 pp
Mateus O, Jacobs LL, Schulp AS et al (2011) Angolatitan adamastor, a new sauropod dinosaur
and the first record from Angola. An Acad Bras Cienc 83(1):221–233
Mateus O, Callapez P, Polcyn M et al (2019) Biodiversity in Angola through time: a paleonto-
logical perspective. In: Huntley BJ, Russo V, Lages F, Ferrand N (eds) Biodiversity of Angola.
Mendelsohn JM (2019) Landscape changes in Angola. In: Huntley BJ, Russo V, Lages F, Ferrand
N (eds) Biodiversity of Angola. Science & conservation: a modern synthesis. Springer Nature,
Cham
Mendes L, Bivar-de-Sousa A, Williams M (2019) The butterflies and skippers of Angola. In:
Mills MSL (2010) Angola’s central scarp forests: patterns of bird diversity and conservation
threats. Biodivers Conserv 19:1883–1903
Mills MSL (2018) The special birds of Angola / as Aves Especiais de Angola. Go-away-birding/
Kissama Foundation, Cape Town/Luanda
Monard A (1935) Contribution à la mammologie d’Angola et prodrome d’une faune d’Angola.
Arquivos do Museu Bocage 6:1–314
Monteiro RFR (1970) Estudo da Flora e da Vegetacão das Florestas Abertas do Planalto do Bié.
Instituto de Investigacão Cientifíca de Angola, Luanda, 352 pp
Morais M (2017) Projecto Kitabanga – Conservação de tartarugas marinhas. Relatório final da
temporada 2016/2017. Universidade Agostinho Neto / Faculdade de Ciências, Luanda
NGOWP (2018) National Geographic Okavango Wilderness Project (2018) Initial findings from
exploration of the upper catchments of the Cuito, Cuanavale and Cuando rivers in Central
and South-Eastern Angola (May 2015 to December 2016). National Geographic Okavango
Wilderness Project, 352 pp
OKACOM (2009) Final report: Okavango integrated river management project. The Permanent
Okavango River Basin Water Commission, Maun
OKACOM (2011) Cubango-Okavango river basin transboundary diagnostic analysis. The
Permanent Okavango River Basin Water Commission, Maun
Rejmánek M, Huntley BJ, le Roux JJ et al (2017) A rapid survey of the invasive plant species in
western Angola. Afr J Ecol 55:56–69
Revermann R, Finckh M (2019) Vegetation survey, classification and mapping in Angola. In:
Revermann R, Krewenka KM, Schmeidel U et al (eds) (2018) Climate change and adaptive land
management in Southern Africa – assessments, changes, challenges, and solutions. Biodivers
Ecol 6:1–497
Russo V, Huntley BJ, Ferrand N (2019) Biodiversity research and conservation opportunities. In:
Sinclair ARE (2012) Serengeti story: life and science in the world’s greatest wildlife region.
Oxford University Press, Oxford, 270 pp
Skelton PH (2019) The freshwater fishes of Angola. In: Huntley BJ, Russo V, Lages F, Ferrand N
(eds) Biodiversity of Angola. Science & conservation: a modern synthesis. Springer Nature,
Cham
Swinscow TDV (1972) Friedrich Welwitsch, 1806–72: a centennial memoir. Biol J Linn Soc
4:269–289
Vaz da Silva B (2015) Evolutionary history of the birds of the Angolan highlands – the missing
piece to understand the biogeography of the Afromontane forests. MSc Thesis. University of
Porto, Porto
Vaz Pinto P (2018) Evolutionary history of the critically endangered giant Sable Antelope
(Hippotragus niger variani): insights into its phylogeography, population genetics, demography
and conservation. PhD Thesis. University of Porto, Porto
Vaz Pinto P (2019) The Giant sable Antelope: Angola’s National Icon. In: Huntley BJ, Russo V,
Walker JF (2004) A certain curve of horn. The hundred-year quest for the giant sable antelope of
Angola. Grove/Atlantic Inc., New York, 514 pp
Weir CR (2019) The Cetaceans (Whales and Dolphins) of Angola. In: Huntley BJ, Russo V,
Werger MJA, van Bruggen AC (eds) (1978) Biogeography and ecology of Southern Africa. The
Junk, Hague, 1444 pp
Zigelski P, Gomes A, Finckh M (2019) Suffrutex dominated ecosystems in Angola. In: Huntley BJ,
Russo V, Lages F, Ferrand N (eds) Biodiversity of Angola. Science & conservation: a modern
synthesis. Springer Nature, Cham
Open Access This chapter is licensed under the terms of the Creative Commons Attribution 4.0
adaptation, distribution and reproduction in any medium or format, as long as you give appropriate
credit to the original author(s) and the source, provide a link to the Creative Commons licence and
indicate if changes were made.
The images or other third party material in this chapter are included in the chapter’s Creative
Commons licence, unless indicated otherwise in a credit line to the material. If material is not
included in the chapter’s Creative Commons licence and your intended use is not permitted by
statutory regulation or exceeds the permitted use, you will need to obtain permission directly from
the copyright holder.
Chapter 2
Angola in Outline: Physiography, Climate
and Patterns of Biodiversity
Brian J. Huntley
Abstract Angola is a large country of 1,246,700 km2 on the southwest coast of

Africa. The key features of the country’s diverse geomorphological, geological,
pedological, climatic and biotic characteristics are presented. These range from the
ultra-desert of the Namib, through arid savannas of the coastal plains to a biologi-
cally diverse transition up the steep western Angolan Escarpment. Congolian rain-
forests are found in Cabinda and along the northern border with the Democratic
Republic of Congo, with outliers penetrating southwards along the Angolan
Escarpment, or up the tributaries of the Congo Basin. Above the escarpment, high
mountains rise to 2620 m above sea level, with isolated remnants of Afromontane
forests and grasslands. Extensive Brachystegia/Julbernardia miombo moist wood-
lands dominate the plateaus and peneplains of the Congo and Zambezi basins, and
dry woodlands of Colophospermum/Acacia occur in the southeast towards the
Cunene River, with Baikiaea/Burkea/Guibourtia woodlands dominating the
Kalahari sands of the endorheic basins of the Cubango and Cuvelai rivers. Rainfall
varies from lower than 20 mm per year in the southwest to over 1600 mm in the
northwest and northeast. At a regional scale, Angola is notable for having represen-
tatives of seven of Africa’s nine biomes, and 15 of the continent’s ecoregions, plac-
ing Angola second only after South Africa for its diversity of African ecoregions.
Keywords Afromontane forest · Biogeography · Biomes · Climate change ·

Congolian forest · Ecoregions · Namib · Kalahari Basin · Zambezian savannas
Introduction
This chapter presents a general outline of the physical geography and biodiversity
characteristics of Angola, as background to the chapters that follow. It draws on the
work of the great Portuguese agro-ecologist Alberto Castanheira Diniz, who
B. J. Huntley (*)
Universidade do Porto, Vairão, Portugal

https://doi.org/10.1007/978-3-030-03083-4_2
16 B. J. Huntley
synthesised the diverse drivers of Angola’s ecological systems and agricultural

potential, based on his many decades of fieldwork in the country (Diniz and Aguiar
1966; Diniz 1973, 1991, 2006). Colonial records of climatic variables (Silveira
1967) are used in the absence of recent time series. The pioneer studies on Angolan
vegetation by Gossweiler and Mendonça (1939) and Barbosa (1970) are fundamen-
tal to any account on Angola biodiversity. Surveys of Angola’s protected areas and
biodiversity ‘hotspots’ (Huntley 1974a, b, 2010, 2015, 2017) provide conservation
context. This outline also draws on the recent regional geographies of Angola by
Mendelsohn and co-workers (Mendelsohn et al. 2013; Mendelsohn and Weber
2013, 2015; Mendelsohn and Mendelsohn 2018). The chapter is also strengthened
by material detailed in the specialist papers that form the core of this volume.
Location and Extent
As a large country of 1,246,700 km2 on the southwest coast of Africa, Angola is

roughly square in outline, lying between 4° 22′ and 18° 02′ south latitude, and 11°
41′ and 24° 05′ east longitude. It is bounded to the west by an arid 1600 km coast-
line along the Atlantic Ocean; to the north by the moist forest and savanna ecosys-
tems of the Republic of Congo and the Democratic Republic of Congo (DRC); to
the east by the moist savanna and woodland ecosystems of the DRC and Zambia;
and by arid woodlands, savannas and desert along its 1200 km southern border with
Namibia.
Geomorphology and Landscape Evolution
The general topography of Angola is illustrated in Fig. 2.1. In summary, coastal

lowlands lying below 200 m altitude and of 10–150 km breadth occupy 5% of the
country’s land surface area, leading to a stepped and mountainous escarpment rising
to 1000 m (23%), and an extensive interior plateau of 1000–1500 m (65%). Seven
percent of the country lies above 1500 m, reaching its highest point at 2620 m on
Mount Moco.
The ecological importance of the major physiographic divisions in Angola was
recognised as early as the 1850s by the pioneer Austrian botanist Friedrich Welwitsch
who categorised the 5000 plant species that he collected in Angola within three
regions: região litoral, região montanhosa, and região alto-plano (Welwitsch 1859).
Besides his remarkable contribution to the founding of Angolan botany, Welwitsch
prepared detailed geological profiles across the landscapes inland of Luanda and
Moçâmedes (Albuquerque and Figueirôa 2018), probably the first such analyses in
western Africa (Fig. 2.2). His understanding of the patterns and relationships of
geology, physiography and vegetation set a strong ecological tradition that has been
followed by successive students of Angola’s biodiversity.
2 Angola in Outline: Physiography, Climate and Patterns of Biodiversity 17
Fig. 2.1 Topography of Angola, indicating provincial boundaries and capitals. The coastal low-
lands, western escarpments, central highlands and plateaus, and the major drainage basins of the
Cuanza, Congo and Zambezi rivers are clearly revealed
A further detailed and indeed classic study of Angola’s geomorphology and local
ecology was that of the German geographer Otto Jessen (1936). Jessen undertook a
series of 11 transects from the coast inland, traversing the escarpment to the interior
plateau from Moçâmedes and thereafter at intervals northwards to Luanda.
Describing, illustrating and mapping selected vegetation communities, geological
exposures, landscapes, landuse and ethnological features of the country, Jessen’s
Angolan work remains unique in its diversity of interest and originality. He recog-
nised five major erosional planation surfaces in western Angola at a time when
geomorphology was evolving as a discipline, and he was recognised by King (1962)
as one of the founders of peneplanation theory. Geomorphological studies in Angola
18 B. J. Huntley
Fig. 2.2 Geological profile from Luanda to Quisonde, scanned from the original manuscript pro-
duced by Friedrich Welwitsch during his expeditions between 1853 and 1860. The lower profile is
a redrafted version of the upper profile adapted from Choffat (1888), and reproduced with permis-
sion from Albuquerque and Figueirôa (2018) and of the Museums of the University of Lisbon
Historical Archives
continued from the 1950s to 1970s by Portuguese researchers, including Marques

(1963), Feio (1964) and Amaral (1969), whose work is summarised by Costa
(2006).
More recent research, in particular that on the evolution and of the major tectonic
and erosional patterns across southern Africa (Cotterill 2010, 2015; Cotterill and De
Wit 2011) and on the biogeography of the freshwater fishes of Angola (Skelton
2019) provide a picture of a very dynamic landscape since the breakup of Gondwana
in the late Cretaceous. These and other authors are providing an improved under-
standing of the processes of uplift, back-tilting, down-warping, deposition, erosion
and river capture on the evolution of the Kalahari Basin. The impacts of sea-level
fluctuations and of the flow of the Congo River on coastal waters and on the ero-
sional forces of the Congo Basin as it impacts on the Zambezi Basin are guiding our
interpretation of the dramatic events shaping the faunal and floral patterns of today.
Cotterill (2015) presents a synthesis of hypotheses on the evolution of the Kalahari
from the late Mesozoic into the early Cenozoic, events which were followed by the
later overlying suite of younger Kalahari sediments – the world’s largest sandsea.
The interplay of geological and paleoclimatic drivers described by Cotterill (2010,
2015), through the pulsing of hot wet and cool dry episodes during the Plio-
Pleistocene, was accompanied by the expansion and contraction of forest and
savanna habitats responding to climatic and fire regimes.
The role of fire in shaping the landscapes of Angola – and particularly of the
dominant miombo moist savanna biome – has become a topic of discussion in recent
years (Zigelski et al. 2019). Maurin et al. (2014) provide evidence based on the
dated phylogenies of 1400 woody species to support the proposal that the ‘under-
ground forests’ (White 1976) that are so prominent across the moist miombo savan-
nas and woodlands of the south-central African plateau, evolved in response to high
fire frequency. They suggest that moist savannas pre-date the emergence of anthro-
pogenic fire and deforestation, becoming a prominent component of tropical vegeta-
tion from the late Miocene (ca. 8 Ma). Maurin et al. (2014) conclude that the
evolution of geoxyles (‘underground trees’) that characterise these moist savannas
define the timing of the transition to fire-maintained savannas occurring in climates
suitable for and previously occupied by forests. The further interpretation of these
key drivers of evolution processes is fundamental to an improved understanding of
the biogeography of Angola.
A major contribution towards an ecological understanding Angola’s contempo-
rary landscapes and natural regions, and their agro-forestry potential, was that of
Castanheira Diniz. Diniz (Diniz and Aguiar 1966, Diniz 1973, 1991) provides a
series of maps illustrating the key features of Angola’s topography, geomorphology,
geology, climate, soils, and phyto-geographic and bio-climatic zones. Diniz’s 11
‘mesological’ units (Fig. 2.3) provide a useful framework for discussions on
Angola’s ecology and biodiversity. Indeed his mesological concept closely corre-
sponds with current perceptions of ecoregions. He also delineated and described 32
agro-ecological zones (Diniz 2006). Although some of his 11 mesological units
need more rigorous and objective definition and delineation, they have become
widely adopted within Angola. Important aspects of these 11 broad units will be
summarised here, integrating these with insights from other sources.
1. Coastal Belt (Faixa litoranea sensu Diniz). This is a mostly continuous plat-
form at 10–200 m above sea level, broken occasionally by broad river valleys.
In contrast to the situation on the east coast of Africa at similar latitudes, the
Angolan coastline is notable for the absence of coral reefs and coastal dune
forests. Long sandbars stretch northwards from rivers such as the Cunene and
Cuanza. Mudflats and mangroves occur at most river mouths from Lobito
northwards, increasing in dimension and diversity towards the Congo. Much of
the coast is uplifted, resulting in sharp sea-cliffs of 10–100 m. In places as nar-
row as 10 km, the coastal belt is mostly of about 40 km width, broadening to
150 km northwards of Sumbe and up the lower Cuanza. The coastal plains are
composed mostly of fossiliferous marine sediments of the Cabinda, Cuanza,
Benguela and Namibe geological basins. The northern coastal platforms are
covered by deep red Pleistocene sands (terras de musseque) of former beaches.
Lying below the sands, and exposed over large areas, are Cretaceous to Miocene
clays, gypsipherous marls, dolomitic limestones and sandstones. Important
beds of Cretaceous fossils occur at Bentiaba and Iembe, the latter including the
sauropod dinosaur Angolatitan adamastor (Mateus et al. 2011, 2019). The
20 B. J. Huntley
Escarpment
Coastal Zone
Zone
Cabinda
M'banza
Congo Congo
Peneplain
Uíge
Lucapa
Cassange
Caxito Basin
Luanda Ndalatando
Saurimo
Coastal Malanje
Zone
Malange
Plateau Congo Peneplain
Sumbe
Upper Cuanza Luena
Cuito Upper
Benguela Escarpment Zambezi
Zone Huambo Massif
Marginal
Mountain
Chain Ancient Plateau
Lubango Menongue
Moçâmedes
Zambezi-Cubango Peneplain
Lower Cunene
Ondjiva
Fig. 2.3 Main geomorphological and landscape units of Angola. (After Diniz 1973)
southernmost segment of the Coastal Belt includes the mobile and mostly
vegetation-less dunes of the Namib Desert.
2. Escarpment Zone (Faixa subplanaltica sensu Diniz; região montanhosa sensu
Welwitsch). A broad transition belt lies between the coastal plains and the inte-
rior plateaus – variable in breadth and gradient. Over much of the zone, the
transition advances up several steep steps of between 400 and 600 m. In the
south, between Moçâmedes and Lubango, the escarpment of the Serra da Chela
is very sharp, rising 1000 m at Tundavala and Bimbe. The geology of the
Escarpment Zone is complex, comprising crystalline rocks of the Precambrian:
granites, gneisses, schists, quartzites and amphibolites. The Escarpment Zone
(also referred to as the Western Angolan Scarp) includes very hilly country,
with mountainous belts in the north, and some major inselbergs in the south, the
most important of which is Serra de Neve, which rises to 2489 m from the sur-
rounding plains and low hills. The Angolan Escarpment has long been recog-
nised for its biogeographic importance (Humbert 1940, Hall 1960a, Huntley
1974a) and has been the centre of interest of many recent studies (Hawkins
1993, Dean 2000, Mills 2010, Cáceres et al. 2015, 2017).
3. Marginal Mountain Chain (Cadeia Marginal de Montanhas sensu Diniz).
Residual mountain lands, mostly at 1800–2200 m, underlain mostly by
Precambrian rocks such as gneiss, granites and migmatites, lie at the western
margin of the extensive interior plateau, and are known as the Benguela,
Huambo and Huíla Highlands. The highest peaks rise to 2420 m on Mount
Namba, 2582 m on Serra Mepo and 2620 m at Mount Moco. The mountains are
of biogeographic importance for their montane grasslands, with some elements
of the Cape flora, and relict patches of Afromontane forests and endemic bird
assemblages (Humbert 1940; Hall 1960b; Hall and Moreau 1962; Huntley and
Matos 1994; Dean 2000; Mills et al. 2011, 2013; Vaz da Silva 2015).
4. Ancient Plateau (Planalto Antigo). This extensive plateau drops eastwards from
below the Marginal Mountain Chain and encompasses the headwaters of the
Cunene, Cubango, Queve and Cutato rivers, comprising rolling landscapes with
wetlands and low ridges with scattered granitic inselbergs. It drops from 1800 m
in the west to 1400 m in central Angola.
5. Lower Cunene (Baixo Cunene). This is a rather artificial unit, leading imper-
ceptibly down from 1400 m on the ‘Ancient Plateau’ to the frontier with
Namibia at 1000 m. The gentle gradient of the eastern half forms the very
clearly defined Cuvelai Basin, which drains as an ephemeral catchment into the
Etosha Pan. West of the Cunene the landscape is more broken, with pockets of
Kalahari sands between low rocky hills.
6. Upper Cuanza (Alto Cuanza). The upper catchments of the Cuanza and its trib-
utary the Luando, at altitudes between 1200 and 1500 m, form a distinct basin
of slow drainage feeding extensive wetlands during the rain season.
7. Malange Plateau (Planalto de Malange). A gently undulating plateau at 1000–
1250 m, dropping abruptly, on its northeastern margin, some several hundred
metres to the Baixa de Cassange and the Cuango drainage. The escarpment
ravines hold important moist forest outliers (such as at Tala Mungongo) that
deserve investigation. To the west, the plateau is drained by the rivers flowing
to the Atlantic, most spectacularly by a tributary of the Cuanza, the Lucala, that
drops over 100 m at the famous Calandula Falls (formerly Duque da Bragança
Falls).
8. Congo Peneplain (Peneplanície do Zaire). This is a vast sandy peneplain,
drained by the northward flowing tributaries of the Cassai/Congo Basin, and
stretching eastwards from the margins of the mountainous northern end of the
Escarpment Zone in Uíge, to the extensive Chanas da Borracha of the Lundas.
These gently dipping plains, mostly at 1100–800 m, are being aggressively dis-
sected by the many northward flowing, parallel tributaries of the Congo Basin.
The Cuango River, draining the Baixa de Cassange, drops to 500 m at the
22 B. J. Huntley
f rontier with the Democratic Republic of Congo. The southern boundary of this
Congo Peneplain is defined imperceptibly by the watershed between the
Zambezi and Congo basins, lying at ca. 1200 m.
9. Cassange Basin (Baixa de Cassange). A wide depression, several hundred
metres below the surrounding plateaus, is demarcated by abrupt escarpments to
the west and the densely dendritic catchment of the Cuango to the northeast.
The geological substrate comprises Triassic Karoo Supergroup sediments of
limestone, sandstone and conglomerates. Within the Basin, several large table-
lands – remnants of the old planation surface – rise above the depression as
extensive plateaus flanked by sheer, 300 m escarpments, exemplified by Serra
Mbango, which awaits biological survey.
10. Zambezi-Cubango Peneplain (Peneplanície do Zambeze-Cubango). This is the
vast peneplain draining deep Kalahari sands, with slow-flowing rivers that
meander across the gently dipping plateau from 1200 m at the watershed with
the Congo Basin to 1000 m at the frontier with Namibia. Within this extensive
peneplain, the Bulozi Floodplain occupies an area in excess of 150,000 km2 in
Angola and Zambia.
11. Upper Zambezi Massif (Maciço do Alto Zambeze). The Calunda Mountains of
eastern Moxico, composed of Precambrian schists and norites, dolorites, sand-
stones and limestones, rise to 1628 m above the Zambezi peneplain which lies
at 1150 m. The mountains form a striking contrast to the almost featureless
landscape that stretches some 800 km eastwards from Huambo to Calunda.
Rivers and Hydrology
Angolan river systems fall into two categories. First, coastal rivers drain the central
and western highlands and flow rapidly westwards where they have penetrated the
steep escarpment to the Atlantic Ocean. Most of these coastal rivers are relatively
short, are highly erosive and carry high sediment loads. Backward erosion by some
of these has produced minor basins, such as the amphitheatres of the upper Queve
and Catumbela. The biogeographic importance of the river captures associated with
these systems, especially the Congo, Cuanza, and Cunene, have been profound, as
described by Skelton (2019). Most of the coastal rivers south of Benguela are
ephemeral.
The second major category of river systems is that of the vast interior plateaus.
Drained by nine large hydrographic basins, seven of which are transnational, Angola
serves as the ‘water tower’ for much of southern and central Africa. Many of these
rivers arise in close proximity on either side of the gently undulating watershed
between the Cuanza, Cassai (Congo), Lungue-Bungo (Zambezi), Cunene, and
endorheic Cubango (Okavango) basins. These rivers drain the vast and deep Kalahari
sands, are slow moving and due to the filtering action of the sands, are crystal clear
and nutrient poor. A separate ephemeral, endorheic system, the Cuvelai Basin,
drains southwards into the Etosha Pan.
The conservation importance of the Angolan river systems is of great signifi-

cance, feeding as they do two wetlands (Okavango and Etosha) of global impor-
tance, and the still under-researched Bulozi Floodplain of Moxico. This is possibly
the largest ephemeral floodplain in Africa – 800 km from north to south and 200 km
from east to west – straddling the Angola/Zambia frontier (Mendelsohn and Weber
2015).
Geology and Soils
The geological history and soil genesis of Angola is complex and interrelated, and
influenced by rainfall, drainage, evaporation and wind. Mateus et al. (2019) provide
a map and stratigraphic profile of the geology of Angola which summarises the
major geological features of the country. The predominance of a broad belt of
Precambrian systems along the western margin of the country, with Cenozoic sys-
tems occupying most of the eastern half, is striking. Over three-quarters of the coun-
try (Fig. 2.4) is covered by two main soil groups arenosols and ferralsols – an
understanding of which provides an essential introduction to Angolan pedology. For
simplicity, soils will be described with reference to their geological substrate.
First, Angola’s main soil groups are the sandy arenosols (solos psamíticos) that
cover more than 53% of the country. These sands are dominant features of three
major landscapes: the dunes of the Namib Desert; the red ‘terras de musseque’ of
the coastal belt northwards from Sumbe; and the vast Kalahari Basin. The great
majority of the arenosols lie to the east of approximately 18° longitude – the aeolian
sands of the Kalahari Basin which cover nearly 50% of Angola and hides nearly all
of the underlying geological formations. The Kalahari Basin, extending across
2500 km from the Cape in the south to the Congo Basin in the north, and up to
1500 km in breadth, is reputedly the largest body of sand in the world. The sands
have been deposited by wind and water over the past 65 million years. Composed of
quartz grains that hold no mineral nutrients, and with very little accumulated organic
matter, they are thus of very low fertility and water-holding capacity. Waters passing
through the vast catchments of the Congo, Cubango and Zambezi basins that drain
the Kalahari are therefore extremely pure.
Second, the higher ground of the western half of Angola (the Ancient Massif) is
dominated by ferralsols (solos ferralíticos) derived from underlying rocks (gneisses,
granites, metamorphosed sediments of the Precambrian Basement Complex; and
schists, limestones and quartzites of the West Congo System). Ferralsols cover
approximately 23% of Angola. The soils are mostly of low water-holding capacity.
Because they are heavily leached in higher rainfall areas, the loss of mineral nutri-
ents and organic matter results in low fertility. They are characteristically reddish
due to the oxidation of their high iron and aluminium content, which also accounts
for the presence in many areas of ferricrete hardpan horizons a metre or two below
the surface, impeding root and water penetration and resulting in the formation of
extensive areas of laterite.
24 B. J. Huntley
Vertisols Podzols Luvisols Ferralsols

Solonetz Phaeozems Lixisols Cambisols
Regosols Nitisols Leptosols Calcisols
Cabinda Gleysols Arenosols
Fluvisols Alisols
M'banza Congo
Acrisols
Uige
Lucapa
Caxito
Luanda
N'dalatando
Malanje
Saurimo
S
Sumbe
Luena
Benguela Cuito
uitto
Huambo
Lubango
Lubang
go Menongue
Moçâmedes
Ondjiva
va
a
Fig. 2.4 Outline of the main soil types of Angola (from Jones et al. 2013), illustrating the pre-
dominance of arenosols in the eastern half of the country, and ferralsols across the western and
central plateaus
These two low-fertility soil groups (arenosols and ferralsols) cover over 76% of
the country, thus despite the adequacy of rainfall over most of Angola, agricultural
production faces the challenges of low soil fertility (Neto et al. 2006; Ucuassapi and
Dias 2006). The natural vegetation types that cover both arenosols and ferralsols –
predominantly miombo woodlands – are well adapted to these soil conditions and
the untransformed landscape gives the appearance of great vitality and luxuriance.
The next soil grouping in terms of landcover, occupying 6% of Angola, are the
shallow regosols (litosolos) of rocky hills and gravel plains, most extensive in the
arid southwest. Other important soil types include luvisols, calcisols and cambisols
(solos calcários, solos calcialíticos), which provide fertile loam soils for crops
(including the ‘coffee forests’ of the Escarpment Zone); alluvial fluvisols (solos
aluvionais) in drainage lines with high organic content and high water retaining
capacity, suitable for crops if not water-logged; gleysol clays (solos hidromórficos),
typically acidic and waterlogged and occasionally very extensive – as on seasonally
flooded plains such as Bulozi Floodplains.
Climate and Weather
The diverse climatic and weather conditions experienced across Angola result from
many atmospheric, oceanic and topographic driving forces.
First, the geographic position of Angola, stretching from near the Equator to
close to the Tropic of Capricorn, across 14 degrees of latitude, accounts for the
overall decrease in solar radiation received and thus annual mean temperatures
experienced from north to south. The latitudinal decrease in mean annual tempera-
ture is illustrated by data from stations in the hot northwest and northeast (Cabinda:
24.7 °C; Dundo: 24.6 °C), compared with stations in the milder southwest and
southeast (Moçâmedes: 20.0 °C; Cuangar: 20.7 °C).
Secondly, both temperature and precipitation are influenced by altitude. The
decrease in mean annual temperature can be illustrated from sites below the Chela
escarpment to the highest weather station in the country: i.e. from Chingoroi: alti-
tude 818 m, mean annual temperature 23.1 °C; Jau: altitude 1700 m, mean annual
temperature 18.0 °C; and finally Humpata-Zootécnica: altitude 2300 m, mean
annual temperature 14.6 °C.
Thirdly, and of greatest importance to the rainfall patterns that determine vegeta-
tion and habitat structure, are the influences of the atmospheric systems which dom-
inate central and southern Africa. Circling the globe near the Equator is a belt of low
pressure where the trade winds of both Northern and Southern Hemispheres con-
verge, creating strong convective activity which generates the dramatic thunder-
storms that characterise the inter-tropics. Known as the Inter-tropical Convergence
Zone (ITCZ), the belt moves southwards over Angola during summer, and then
returns northwards to the Equator as winter approaches. The rainfall season that is
triggered by the ITCZ passes across northern Angola from early summer, reaching
southern Angola in late summer. The climate is strongly seasonal, with hot wet sum-
mers (October to May) and mild to cool dry winters (June to September). Some
stations in northern Angola receive two peaks of rainfall, early summer and late
summer, often with a short drier period in mid-summer.
Moving in tandem with the ITCZ are two high-pressure systems – over the
Atlantic and over southern Africa – the South Atlantic Anticyclone and the Botswana
Anticyclone. In simple terms, these two anticyclones block the southward move-
ment of moist air from the ITCZ during winter (preventing cloud formation) and as
the high-pressure cells move southwards in summer, the conditions required for
cloud formation return. This pulsing of rainfall systems is clearly illustrated in the
series of rainfall maps prepared by Mendelsohn et al. (2013) from weather satellite
imagery (Fig. 2.5).
26 B. J. Huntley
Fig. 2.5 The impact of the southwards and northwards pulsing of the Inter-tropical Convergence
Zone on rainfall seasonality across Angola during 2009/2010. (From Mendelsohn et al. 2013)
During winter and early summer, the Botswana Anticyclone generates strong
winds that blow across Angola from east to west, with impacts on micro-relief over
much of the country. In the southwest, the winds pick up dust from the arid lands
and create hot, choking dust storms that feed the sand dunes of the Namib. The
winds are also notorious in the north, where they desiccate the grasslands of the
Lundas. In the east, the winds and their sand deposits account for dune formation
across the Bulozi Floodplain (Mendelsohn and Weber 2015).
Rainfall and temperature seasonality and other climatic parameters are illus-
trated by the climate diagrams in Fig. 2.6. The distribution of mean annual rainfall
across Angola is summarised in Fig. 2.7.
Fourthly, as noted above, altitude and seasonality determine temperature condi-
tions. However, an anomaly to this general rule occurs in the coastal belt of Angola,
especially in the far south, through the influence of the temperature inversion cre-
ated by the cold, upwelling Benguela Current. The Benguela Current has a stabiliz-
ing effect on the lower atmosphere and prevents the upward movement of moist,
cloud-forming air off the ocean, accounting for the evolution of the Namib Desert.
Its impact also extends as far north as Cabinda, where a narrow belt of arid savanna
woodland and dry forest, of acacias, sterculias and baobabs, flanks the rainforests of
the Maiombe.
Despite the aridity of the coastal zone, the cooling effect of the Benguela Current
results in low stratus cloud and fog (cacimbo) through much of winter, with heavy
dew condensing on vegetation along the coast even during the driest months of win-
ter. The fog belt is most pronounced between Moçâmedes and Benguela, where epi-
phytic lichens reach great abundance in an otherwise desertic environment. The
Benguela Current also results in a gradient of increasing precipitation from south to
north and from west to east. The rainfall gradients are locally accentuated by the
orographic influence of the escarpment and the highland mountain massifs. The
Fig. 2.6 Climate diagrams illustrating rainfall and temperature seasonality and other climatic
parameters. Note weak bimodal rainfall maxima for stations in northern Angola and unimodal
maxima in central and southern Angola
sharp relief of the escarpment creates conditions for orographic rainfall along most of
this zone, supporting the ‘coffee forests’ of Seles, Gabela, Cuanza-Norte and Uíge.
Attempts to synthesise climatic characteristics into simple formulae or graphics
have resulted in a wide range of classification systems. A synthesis of climatic data
provided by the widely used Köppen and Thornthwaite classification systems was
undertaken by Azevedo (1972) to map and quantify, at a national scale, the climatic
regions of Angola, based on the substantial data set available at that time.
Interestingly, despite some of its shortcomings, the Azevedo map provides a closer
fit with general features of Angola’s bio-climatic patterns than a much more recent
map (Peel et al. 2007). The latter map is based on a global synthesis and review of
the Köppen system, and draws on a very limited data set for Angola (5 stations for
temperature; 16 stations for precipitation). The northern region of Angola is typical
of Köppen’s Tropical Wet Savanna (Aw) group, the plateau of the Temperate
Mesothermal (Cw) group, and the southwest and coastal plain the Dry Desert and
Semi-desert (Bsh, Bwh) group.
Mean annual rainfall and mean monthly temperatures for hottest and coldest
months illustrate a few climatic characteristics of the Köppen regions (Table 2.1).
The absence of data on extreme minimum temperatures and of frost occurrence is
regrettable, as these factors, in tandem with fires and herbivory, play significant
roles in the floristic composition and physiognomic structure of Angolan vegetation
(Zigelski et al. 2019).
28 B. J. Huntley
Fig. 2.7 Mean annual rainfall in Angola
Climate Change
Studies on the climate of Angola have been frustrated in recent years by the collapse
of the extensive network of weather stations maintained during the colonial era by
the then Meteorological Services of Angola. The publication by Silveira (1967) of
recordings from 184 stations across all 18 provinces provides an invaluable record
of the country’s climate. According to the Ministry of Environment’s Initial National
Report to the UNFCCC (GoA 2013) the weather recording network collapsed from
225 ‘climatological posts’ in 1974 to zero posts in 2010, while synoptic stations
decreased from 29 in 1974 to 23 operational stations in 2010, 12 being automatic
and 11 conventional. The network has since been strengthened by 22 automatic
Table 2.1 Representative climatic data following the Köppen climate classification system
Köppen Altitude Precipitation Mean of hottest Mean of coldest
symbol Station (m) (mm) month °C month °C
Aw Belize 245 1612 26.7 22.2
Aw Saurimo 1081 1355 23.8 20.3
Bsh Ondgiva 1150 577 26.4 16.7
Bsh Cuangar 1050 596 24.6 15.0
Bsh’ Chitado 1000 405 27.4 19.2
Bsh’ Luanda 44 405 27.0 20.1
Bwh Moçâmedes 44 37 24.2 15.5
Bwh Tômbwa 4 12 24.2 14.5
Bwh’ Benguela 7 184 26.3 18.0
Bwh’ Caraculo 440 123 26.4 17.2
Cwa Menongue 1348 965 23.4 14.5
Cwa Luena 1328 1182 22.7 17.0
Cwb Huambo 1700 1210 20.6 15.7
Cwb Lubango 1760 802 20.7 15.3
Data from Silveira (1967)
stations established by the Southern African Science Service Centre for Climate
Change and Adaptive Land Management (SASSCAL).
The poor national coverage and reliability of climatic data collected over the past
four decades is a challenge for climate change research. However, a recent study
(Carvalho et al. 2017) provides the first analysis and comparison of a set of four
Regional Climate Models (RCMs) with data from 12 meteorological monitoring
stations in Angola. Scenarios of future temperature and precipitation anomaly trends
and the frequency and intensity of droughts are presented for the twenty-first cen-
tury. While there is a difference in the performance of the four RCMs, in particular
for precipitation, consistent results were found for temperature projections, with an
increase of up to 4.9 °C by 2100. The temperature increases are lowest for the north-
ern coastal areas and highest for the southeast. In contrast to temperature rises,
precipitation was projected to fall over the century, with an average of −2% across
the country. Again, the strongest change was projected for the southeast, with
decreases of up to −4%. Due principally to the projected increase in Sea Surface
Temperatures by approximately 3 °C over the Atlantic during the twenty-first cen-
tury, the central coastal region is expected to have a slight increase in
precipitation.
Carvalho et al. (2017) highlight the extreme climate vulnerability of Angola, as
previously noted by other studies (Brooks et al. 2005; Cain and Cain 2015). They
conclude that climate change in Angola will bring stronger and more frequent
droughts through the century, with impacts on water resources, agricultural produc-
tivity and wildfire potential. These factors will no doubt play out in negative ways
on the current trends of land transformation and degradation as described by
Mendelsohn (2019).
30 B. J. Huntley
Biogeography, Biomes and Ecoregions
Overview
Angola’s geographic location, geological history, climate and physiography account

for its rich biological diversity. The comparative paucity of research focused on or
within Angola explains the dependence of descriptions of the country’s biogeogra-
phy on broader regional reviews. While a full synthesis and interpretation of the
evolution of the country’s fauna and flora awaits development (Cotterill 2010,
2015), recent workers have advanced towards consensus on the main patterns, as
discussed in general terms for terrestrial biota in this chapter, for marine systems by
Kirkman and Nsingi (2019) and Weir (2019), and for freshwater fishes by Skelton
(2019) in other chapters of this volume.
In brief, three marine ecoregions (Spalding et al. 2007) are within or overlap with
Angola’s marine environment, namely the Guinea South, Angolan, and Namib
Ecoregions, the first two of which belong to the tropical Gulf of Guinea biogeo-
graphical province whereas the latter is part of the Benguela biogeographical prov-
ince (Kirkman and Nsingi 2019). Most of Angola’s EEZ falls within the Benguela
Current Large Marine Ecosystem, with only Cabinda in the far north being included
in the Guinea Current Large Marine Ecosystem.
The freshwater ecoregions of Africa have been classified and mapped by Thieme
et al. (2005) and the eight ecoregions found within Angola are described in this
volume (Skelton 2019). Skelton (2019) provides an elegant biogeographic model to
explain the patterns and dynamics of freshwater fish faunas of Angola. Neither the
floral nor vegetation patterns reflect the complexities and subtleties embedded in the
ichthyological zoogeography of Angola, given the mobility of terrestrial plant dis-
semination. Here I will confine discussion to the terrestrial biota and ecosystems.
Angola lies between and within two major terrestrial biogeographic regions: the
moist forests and savannas of the Congolian region; and the woodlands, savannas
and floodplains of the Zambezian region. These two major divisions occupy over
97% of Angola. Gallery and escarpment forests of Congolian affinity penetrate
southwards into the Zambezian savannas and woodlands of the Angolan planalto
along deeply incised tributaries of the Congo Basin, and form a broken chain of
forests southwards along the western escarpment. In the south, the extensive
Brachystegia/Julbernardia miombo woodlands that occupy most of central Angola
transition to Baikiaea/Guibourtia/Burkea savannas and woodlands. In the southwest,
the arid Acacia/Commiphora/Colophospermum savannas, dwarf shrublands and
desert of the Karoo-Namib region are found, penetrating northwards as a narrowing
wedge along the coastal lowlands to Cabinda. The smallest of Africa’s centres of
botanical endemism – the Podocarpus Afromontane forests and montane grass-
lands – are represented by extremely restricted, relict patches in the mountains of
the Benguela, Huambo and Huíla highlands.
Early Studies
Beyond general agreement on the above brief outline, botanists and zoologists have
described and debated as many systems of biogeographic classification and of ter-
minologies for Angola and for Africa as there are authors of the papers on the topic
(Werger 1978). The pioneering works of Welwitsch (1859); Gossweiler and
Mendonça (1939) and Barbosa (1970) provided the basis for several subsequent
attempts to integrate the vegetation of Angola within a regional framework (Monteiro
1970; White 1971, 1983; Werger 1978). Zoogeographic classifications (Chapin
1932; Frade 1963; Monard 1937; Hellmich 1957; Crawford-Cabral 1983) are, with
some minor exceptions, compatible with the overall systems of botanists (Werger
1978; Linder et al. 2012), (but see Branch et al. 2019, for comments on lizards). The
Africa-wide synthesis of White (1983) is particularly useful in considering Angola’s
floristic (and in general terms, zoological) patterns and affinities. In broad terms,
and following White’s terminology, Angola includes representation of four ‘regional
centres of endemism’. They comprise the following centres with estimates of the
percentage of their total area in Angola from Huntley (1974a, 2010):
• Guineo-congolian regional centre of endemism - mosaics of forests, thickets, tall
grass savannas – 25.7% (This is Linder et al.’s Congolian Region and includes
their Shaba sub-region);
• Zambezian regional centre of endemism – moist woodlands, savannas, grass-
lands and thickets – 71.6% (The Zambezian Region sensu Linder et al.);
• Karoo-Namib regional centre of endemism – desert, shrublands, arid savannas,
woodlands and thickets – 2.6% (Most of this is placed in Linder et al.’s Southern
African Region as their Southwest Angola sub-region); and
• Afromontane archipelago-like regional centre of endemism – forests, savannas
and grasslands – 0.1%. (This is related to Linder et al.’s Ethiopian Region).
Statistical Regionalisation
Attempts have recently been made to use the massive databases of species distribu-
tion records held by museums and herbaria to bring objectivity and consistency to
the classification of Africa’s floral and faunal regions. A major step towards such
regionalisation is provided by the statistical definition of biogeographical regions of
sub-Saharan Africa by Linder et al. (2012). Using data for 1877 grid cells of one-
degree resolution, the study included data for over a million records of 1103 species
of mammals, 1790 species of birds, 769 species of amphibians, 480 species of rep-
tiles and 5881 species of vascular plants. The databases were analysed using cluster
analysis techniques to define biogeographical units that “comprise grid cells that are
more similar in species composition to each other than to any other grid cells”
(Linder et al. 2012). They proposed seven biogeographical regions for sub-Saharan
32 B. J. Huntley
Africa: Congolian, Zambezian, Southern African, Ethiopian, Somalian, Sudanian

and Saharan. Their analyses demonstrated that patterns of richness and endemism
are positively and significantly correlated among plants, mammals, amphibians,
birds and reptiles and with the overall biogeographical regions revealed by the sum
of the data sets.
The use of modern cluster analysis techniques was taken further, at an Angolan
level, by Rodrigues et al. (2015). Based on a cluster analysis of data for 9880 records
of 140 species of ungulates, rodents and carnivores at a quarter degree resolution,
the study found general congruence with that of Linder et al. (2012) and the earlier
divisions of Angola’s biogeography (Beja et al. 2019). Rodrigues et al. (2015) iden-
tify 18 indicator species for their four main divisions, which agree with the group-
ings based on field surveys undertaken in the 1970s (Huntley 1973) that also
included the enclave of Cabinda, which was not included in the Rodrigues et al.
(2015) analyses.
Both of the above very detailed and objective cluster analyses confirmed the
general patterns of biogeographical regionalisation used for many decades across
Africa, as described at the head of this section, even though terminology and detail
of boundaries and transitions between regions differ from one author to the next.
While objective, it is possible that the cluster analysis approach lacks the subtlety
and flexibility of scale that classical expert systems permit. A particular challenge is
the paucity of geo-referenced data for Angolan taxa, as experienced in a recent
botanical analysis at inter-tropical scale (Droissart et al. 2018). Both cluster analy-
ses and expert systems remain works in progress.
Biomes and Ecoregions
The chorological studies of White (1983) and statistical analyses of Linder et al.
(2012) capture some of the evolutionary history and relationships of Africa’s flora
and fauna, but they do not fully reflect the continent’s diversity of biomes, habitats
and ecosystems – which are based on structural and functional rather than evolu-
tionary relationships. The most comprehensive recent synthesis on African habitats
(Burgess et al. 2004) has been widely adopted as a basis for conservation planning
and is of use for any study of African biomes, ecoregions and habitats (MacKinnon
et al. 2016). At the first level, a global classification and map of the world’s ecore-
gions (Olson et al. 2001) was used to identify the nine biomes of Africa’s three main
biogeographic divisions (Palearctic, Afrotropical and Cape). The biome concept
used was defined as “vegetation types with similar characteristics grouped together
as habitats, and the broadest global habitat categories are called biomes” (Olson
et al. 2001). Of the nine biomes recognised, seven are represented in Angola – the
largest range of biomes represented in any African country. These are:
• Tropical and subtropical moist forests;
• Montane grasslands and shrublands;
• Tropical and subtropical grasslands, savannas, shrublands, and woodlands;

• Tropical and subtropical dry and broadleaf forests;
• Deserts and xeric shrublands;
• Mangroves; and
• Flooded grasslands and savannas.
Within the biomes, Burgess et al. (2004) defined a total of 119 terrestrial ecore-
gions for Africa and its islands. Ecoregions are defined as “large units of land or
water that contain a distinct assemblage of species, habitats and processes, and
whose boundaries attempt to depict the original extent of natural communities
before major land-use change” (Dinerstein et al. 1995). It is impressive to note that
based on the Burgess et al. (2004) assessment, Angola has not only the largest diver-
sity of biomes, but also the second largest representation of ecoregional diversity in
Africa (Table 2.2, Fig. 2.8).
Figure 2.8 (Burgess et al. 2004) provides a useful framework for the understand-
ing of Angola’s biodiversity patterns. Despite its coarse grain, it allows a general
synthesis to be refined as new information becomes available. The relationship
between biomes and ecoregions (sensu Burgess et al. 2004) and the vegetation types
of Barbosa (1970) is summarised in Table 2.3. The very brief notes on key genera
found within the Barbosa vegetation units provide an idea of the floristic composi-
tion that characterises the ecoregion. The photos presented in Fig. 2.9 provide
examples of the main vegetation types and habitats
The Biological Importance of the Angolan Escarpment
While the classification of White (1983) and Linder et al. (2012) are useful at a
continental scale, a more detailed and subtle analysis of the major biomes and habi-
tat groupings is needed at a national scale for both research and conservation
Table 2.2 Representation of biomes and ecoregions in southern African countries

Country Biomes Ecoregions number and total (T) T
Angola 7 8, 32, 42, 43, 49, 50, 51,55, 56, 63, 81, 82, 106, 109, 116 15
Botswana 3 54, 57, 58, 63, 68, 105 6
Congo 3 8, 12, 13, 43,116 5
Republic
D.R. Congo 5 8, 13, 14, 15, 16, 17, 42, 43, 49, 50, 73, 116 12
Mozambique 3 21, 22, 52, 53, 54, 64, 76, 117 8
Namibia 3 51, 55, 58, 67, 105, 106, 107, 108, 109, 110 10
South Africa 5 22, 23, 24, 54, 57, 58, 77, 78, 79, 80, 89, 90, 91, 105, 108, 110, 17
117
Zambia 4 32, 50, 53, 54, 56, 63, 74 7
Zimbabwe 2 51, 53, 54, 57, 58, 76 6
From Burgess et al. (2004)
34 B. J. Huntley
43
Cabinda
M'banza Congo
116
43
Uige
Lucapa 42
Caxito
Luanda
N'dalatando
Malanje
81
Saurimo
S
Sumbe
50
Luena
Benguela
ela Cuito
t
82 Huambo
49
32
56
Lubango
ubango Menongue
Moçâmedes
109
55 63
51
Ondjiva
106
Fig. 2.8 Ecoregions of Angola

8 Atlantic Equatorial Coastal Forest • 32 Zambezian Cryptosepalum Dry Forest • 42 Southern
Congolian Forest-Savanna Mosaic • 43 Western Congolian Forest-Savanna Mosaic • 49 Angolan
Miombo Woodland • 50 Central Zambezian Miombo Woodland • 51 Zambezian Baikiaea
Woodland • 55 Angola Mopane Woodland • 56 Western Zambezian Grassland • 63 Zambezian
Flooded Grasslands • 81 Angolan Scarp Savanna and Woodland • 82 Angolan Montane Forest-
Grassland Mosaic • 106 Kaokoveld Desert • 109 Namib Escarpment Woodlands • 116 Central
African Mangroves. (After Burgess et al. 2004, map used with permission)
Table 2.3 African biomes and ecoregions (as defined by Burgess et al. 2004) and Angolan
vegetation types (Barbosa 1970) with indicative genera
Ecoregion Barbosa n°, name and key
n° Biome Ecoregion genera
8 Tropical and Subtropical Atlantic Equatorial 1,2. Closed Forest
Forest Coastal Forest Gilbertiodendron, Librevillea,
Tetraberlinia
32 Tropical and Subtropical Zambezian 4. Closed Forest
Cryptosepalum
Dry Broadleaf Forest Dry Forest Cryptosepalum, Brachystegia,
Erythrophleum
42 Tropical and Subtropical Southern Congolian 8. Forest-Savanna Mosaic
Grasslands, Savannas, Forest-Savanna Marquesia, Berlinia, Daniella,
Shrublands and Woodlands Mosaic Hymenocardia
43 Tropical and Subtropical Western Congolian 3. Closed Forest
Grasslands, Savannas, Celtis, Albizia, Celtis
Shrublands and Woodlands Forest-Savanna 13. Thicket-Forest Mosaic
Mosaic Annona, Piliostigma,
Andropogon, Hyparrhenia
49 Tropical and Subtropical Angolan Miombo 16, 17, 18. Woodland
Grasslands, Savannas, Woodland Brachystegia, Julbernardia,
Shrublands and Woodlands Guibourtia, Burkea,
Pterocarpus
50 Tropical and Subtropical Central Zambezian 17, 19. Woodland
Grasslands, Savannas, Miombo Woodland Brachystegia, Julbernardia,
Shrublands and Woodlands Cryptosepalum
51 Tropical and Subtropical Zambezian Baikiaea 25. Tree and Shrub Savanna
Grasslands, Savannas, Woodland Baikiaea, Guibourtia,
Shrublands and Woodlands Pterocarpus, Combretum
55 Tropical and Subtropical Angola Mopane 20. Woodland
Grasslands, Savannas, Woodland Colophospermum, Croton,
Shrublands and Woodlands Combretum, Sclerocarya,
Acacia
56 Tropical and Subtropical Western Zambezian 31. Grasslands
Grasslands, Savannas, Grassland Loudetia, Monocymbium,
Shrublands and Woodlands Tristachya, Parinari, Syzygium
63 Flooded Grasslands and Zambezian Flooded 31. Grasslands
Savannas Grasslands Loudetia, Echinochloa, Oryza
81 Montane Grasslands and Angolan Scarp 10, 11, 22, 23. Forest-
Shrublands Savanna and Savanna-Woodland-Thicket
Woodland Mosaic
Adansonia, Acacia, Albizia,
Celtis, Piliostigma
82 Montane Grasslands and Angolan Montane 6, 32. Relict Forest,
Shrublands Forest-Grassland Grasslands
Mosaic Podocarpus, Apodytes,
Pittosporum, Protea, Erica
(continued)
36 B. J. Huntley
Table 2.3 (continued)
Ecoregion Barbosa n°, name and key
n° Biome Ecoregion genera
106 Deserts and Xeric Kaokoveld Desert 28, 29. Desert, Steppes
Shrublands Welwitschia, Zygophyllum,
Stipagrostis, Odyssea
109 Deserts and Xeric Namib Escarpment 27. Steppes
Shrublands Woodlands Acacia Commiphora,
Colophospermum,
Sesamothamnus, Rhigozum
116 Mangroves Central African 14 A. Mangroves
Mangroves Rhizophora, Avicennia,
Raphia, Elaeis
planning in Angola (Revermann and Finckh 2019). What is equally important in

biogeographic analysis is the detection of patterns of endemism and diversity at
dispersed scales – such as the Angolan Escarpment Zone – described by Hall
(1960a) and subsequently recognised by many workers as of great biodiversity and
evolutionary importance (Huntley 1973, 1974a, 2017; Hawkins 1993; Mills 2010;
Clark et al. 2011). Indeed, each taxon-based account in this volume, on plants
(Goyder and Gonçalves 2019), odonata (Kipping et al. 2019), lepidoptera (Mendes
et al. 2019), fishes (Skelton 2019), birds (Dean et al. 2019), amphibians (Baptista
et al. 2019), reptiles (Branch et al. 2019) and mammals (Beja et al. 2019) draws
attention to the importance of the Angolan Escarpment as a centre of endemism and
speciation. Hall (1960a) explained her recognition of the importance of the Angolan
Escarpment as the major speciation hotspot for birds in Angola by it: (i) creating a
barrier between arid-adapted species of the coastal plains and of the miombo wood-
lands of the plateau, (ii) creating a steep ecological gradient, and (iii) functioning as
a refuge for moist forest specialists that were isolated here during the dry periods of
the glacial cycles. Dean et al. (2019) note that 75% of Angola’s endemic birds are
found in this zone.
The Angolan Escarpment and the remote, isolated and fragmentary remnants of
Afro-montane forests of the Angolan Highlands offer ideal testing grounds for bio-
geographic models, as recently explored by Vaz da Silva (2015). The Angolan
Escarpment biogeographic unit awaits clear definition, description and demarca-
tion, but its scientific importance is matched only by the vulnerability of its threat-
ened forest habitats (Cáceres et al. 2015). Linder et al. (2012) similarly recognise
the importance of the Angolan Escarpment, and that of the transition from Congolian
to Zambezian regions along the northern border of Angola (which they place in their
Shaba sub-region). High species replacement values are found across these biologi-
cally rich areas, emphasising the urgency for their protection.
Fig. 2.9 Examples of some of the Ecoregions of Angola with numbering as per map in Fig. 2.8
and summary in Table 2.3. 8. Maiombo Forest, Cabinda; 42. Congolian gallery forest and moist
miombo woodlands and savanna grasslands, Lunda-Norte; 49. Brachystegia/Julbernardia wood-
land Luando Strict Nature Reserve, Malange; 51. Baikiaea/Guibourtia Woodland Mucusso,
Cuando Cubango; 56. Wetlands of the Bulozi Floodplain, Moxico; 81. Angolan Escarpment at
Serra da Chela, Tundavala, Huíla; 82. Remnant patches of Afromontane forest in ravines on Mount
Moco, Huambo; 106 Grasslands of the intermontane plains of central Iona National Park, Namibe.
(Photos: Bulozi – JM Mendelsohn, others by BJ Huntley)
38 B. J. Huntley
Conclusions
This brief outline of the biogeography of Angola demonstrates the country’s unusual
diversity of landscapes, climates and ecoregions, with Angola embracing the high-
est number of biomes represented within any African state.
The many classifications and terminologies applied to Angola’s biogeographic
units over the past century have not yet resulted in a nationally adopted nomencla-
ture for its biomes and habitats. This situation prevails despite the existence of
strong traditions in Angola’s ethnic groups of indigenous taxonomies for habitats,
such as those of the Chokwe of the Lundas, that are as perfect and detailed as mod-
ern systems (Redinha 1961; Huntley 2015). Furthermore, while many vernacular
terms (mato de panda, anharas do alto, floresta cafeeira, muxitos, mulolas, chanas
da borracha, etc.) enjoy wide use, they are imprecise and inadequate for Angola’s
great diversity of biomes and habitats.
The absence of a uniform system of nomenclature limits the use of information
attached to biological collections, which in most cases provide only site locality
data, and more recently, geo-referencing. Several southern African countries have
nationally accepted biome and vegetation maps (e.g. South Africa, Lesotho and
Swaziland – Mucina and Rutherford 2006) with clear descriptors for each biome
and vegetation unit, facilitating communication between researchers and conserva-
tion planners. As Angola re-assesses its biodiversity wealth, and the need to protect
and sustainably utilise these resources, the development of a new map of its vegeta-
tion, ecosystems and biomes becomes a high priority. Equally urgent but similarly
daunting is the study of the evolutionary processes and relationships of the biota of
the Angolan Escarpment and Afromontane forests, and the effective protection of
these fingerprints of the past.
References
Albuquerque S, Figueirôa S (2018) Depicting the invisible: Welwitsch’s map of travellers in

Africa. Earth Sci Hist 37:109–129
Amaral I (1969) ‘Inselberge’ (ou montes-ilhas) e superfícies de aplanação na bacia do Cubal da
Hanha em Angola. Garcia da Orta 17:474–526
Azevedo AL (1972) Caracterizacão Sumária das Condições Ambientais de Angola. Universidade
de Luanda, Luanda, 106 pp
Barbosa LAG (1970) Carta Fitogeográfica da Angola. Instituto de Investigação Científica de
Brooks N, Adger WN, Kelly PM (2005) The determinants of vulnerability and adaptive capacity at
the national level and the implications for adaptation. Glob Environ Chang 15:151–163
Burgess N, Hales JD, Underwood E et al (2004) Terrestrial ecoregions of Africa and Madagascar –
a conservation assessment. Island Press, Washington DC, 499 pp
Cáceres A, Melo M, Barlow J et al (2015) Threatened birds of the Angolan Central Escarpment:
distribution and response to habitat change at Kumbira Forest. Oryx 49:727–734
Cáceres A, Melo M, Barlow J et al (2017) Drivers of bird diversity in an understudied African
centre of endemism: the Angolan Escarpment Forest. Bird Conserv Int 27:256–268
Cain A, Cain A (2015) Climate change and land markets in coastal cities of Angola. In 2015 World
Bank conference on land and poverty. The World Bank, Washington, DC
Carvalho SCP, Santos FD, Pulquério M (2017) Climate change scenarios for Angola: an analysis
of precipitation and temperature projections using four RCMs. Int J Climatol 37:3398–3412
Chapin JP (1932) The birds of the Belgian Congo. Bull Am Mus Nat Hist 65:1–756
Choffat P (1888) Dr. Welwitsch: Quelques notes sur la géologie d’Angola coordonnées et annotées
par Paul Choffat. Separata das Comunicações dos Serviços Geológicos de Portugal 19:1–24
Clark VR, Barker NP, Mucina L (2011) The Great Escarpment of southern Africa: a new frontier
for biodiversity exploration. Biodivers Conserv 20:2543–2561
Costa FL (2006) O conhecimento geomorfológico de Angola. In: Moreira I (ed) Angola:
Agricultura, Recursos Naturais e Desenvolvimento. ISA Press, Lisboa, pp 477–495
Cotterill FPD (2010) The evolutionary history and taxonomy of the Kobus leche species complex
of south-central Africa in the context of Palaeo-drainage dynamics. Unpublished PhD thesis,
University of Stellenbosch
Cotterill FPD (2015) Biogeographical overview of the Lunda region, northeast Angola. In: Huntley
BJ, Francisco P (eds) Avaliação Rápida da Biodiversidade de Região da Lagoa Carumbo,
Lunda-Norte – Angola/Rapid biodiversity assessment of the Carumbo Lagoon Area, Lunda-
Norte – Angola. Ministério do Ambiente, Luanda, pp 77–99
Cotterill F, De Wit M (2011) Geoecodynamics and the Kalahari Epeirogeny: linking its genomic
record, tree of life and palimpsest into a unified narrative of landscape evolution. S Afr J Geol
114:489–514
Crawford-Cabral J (1983) Esboço zoogeográfico de Angola. Unpublished manuscript, Lisbon,
50 pp + 13 maps
Dean WRJ (2000) The birds of Angola: an annotated checklist. BOU Checklist No. 18. British
Ornithologists’ Union. Tring, UK
Dean WRJ, Melo M, Mills MSL (2019) The Avifauna of Angola: richness, endemism and rarity.
Dinerstein E, Olson D, Graham A et al (1995) A conservation assessment of the ecoregions of
Latin America and the Caribbean. World Bank, Washington, DC
Diniz AC, Aguiar FB (1966) Geomorfologia, solos e ruralismo de região central angolana.
Agronomia Angolana 23:11–17
Diniz AC (1973) Características mesológicas de Angola. Missão de Inquéritos Agrícolas de
Angola, Nova Lisboa
Diniz AC (1991) Angola, o meio físico e potencialidades agrárias. Instituto para a Cooperação
Económica, Lisboa, 189 pp
Diniz AC (2006) Características mesológicas de Angola. Instituto Português de Apoio ao
Desenvolvimento, Lisbon, 546 pp
Droissart V, Dauby G, Hardy OJ et al (2018) Beyond trees: biogeographical regionalization of
tropical Africa. J Biogeogr 45:1153–1167
Frade F (1963) Linhas gerais da distribuição geográfica dos vertebrados em Angola. Memórias da
Junta de Investigações do Ultramar 43:241–257
40 B. J. Huntley
Feio M (1964) A evolução da escadaria de aplanações do sudoeste de Angola. Garcia da Orta

12:323–354
GoA (Government of Angola) (2013) Angola initial national communication to the United Nations
framework convention on climate change. Ministry of the Environment, Luanda, 194 pp
Lisboa, 242 pp
Goyder DJ, Gonçalves FMP (2019) The flora of Angola: collectors, richness and endemism. In:
Hall BP (1960a) The faunistic importance of the scarp of Angola. Ibis 102:420–442
Hall BP (1960b) The ecology and taxonomy of some Angolan birds. Bull Br Mus (Nat Hist) Zool
6:367–463
Hall BP, Moreau RE (1962) The rare birds of Africa. Bull Br Mus (Nat Hist) Zool 8:315–381
Angola Scarp Project. In: Proceedings of the VIII Pan-African Ornithological Congress,
pp 279–284. Kigali, Rwanda, 1992. Koninklijk Museum voor Midden-Afrika, Tervuren
Hellmich W (1957) Herpetologische Ergebnisse einer Forschungsreise in Angola.
Veröffentlichungen der Zoologischen Staatssammlung München 5:1–92
Humbert H (1940) Zones et Étages de Végétation dans le Sud-Ouest de l’Angola. Compte-rendu
Sommaire des Séances de la Societé de Biogéographie 17:47–57
Huntley BJ (1973) Distribution and Status of the Larger Mammals of Angola, with particular
reference to Rare and Endangered species: First Progress Report. December 1973. Repartição
Técnica da Fauna, Serviços de Veterinária, Luanda, Mimeograph report, 14 pp
Huntley BJ (1974a) Vegetation and Flora Conservation in Angola. Ecosystem Conservation
Priorities in Angola. Ecologist’s Report 22. Repartição Técnica da Fauna, Serviços de
Veterinária, Luanda, Mimeograph report, 13pp
Huntley BJ (1974b) Ecosystem Conservation Priorities in Angola. Ecologist’s Report 26.
Repartição Técnica da Fauna, Serviços de Veterinária, Luanda, Mimeograph report
Huntley BJ (2010) Estratégia de Expansão da Rede de Áreas Protegidas da Angola/Proposals for
an Angolan Protected Area Expansion Strategy (APAES). Unpublished report to the Ministry
of Environment, Luanda, 28 pp + map
Huntley BJ (2015) Biophysical profile of Lunda-Norte. In: Huntley BJ, Francisco P (eds) Avaliação
Rápida da Biodiversidade de Região da Lagoa Carumbo, Lunda-Norte – Angola / Rapid
Biodiversity Assessment of the Carumbo Lagoon Area, Lunda-Norte – Angola. Ministério do
Ambiente, Luanda, pp 31–75
Huntley BJ (2017) Wildlife at war in Angola: the rise and fall of an African Eden. Protea Book
House, Pretoria, 432 pp
Huntley BJ, Matos EM (1994) Botanical diversity and its conservation in Angola. Strelitzia
1:53–74
Jessen O (1936) Reisen und Forschungen in Angola. Dietrich Reimer Verlag, Berlin
Jones A, Breuning-Madsen H, Brossard M et al (2013) Soil atlas of Africa. Publications Office of
the European Union, Brussels
King LC (1962) Morphology of the Earth. Oliver & Boyd, London, 699 pp
Linder HP, de Klerk HM, Born J et al (2012) The partitioning of Africa: statistically defined bio-
geographical regions in sub-Saharan Africa. J Biogeogr 39:1189–1925
MacKinnon J, Aveling C, Olivier R et al (2016) Inputs for an EU strategic approach to wildlife
conservation in Africa – regional analysis. European Commission, Directorate-General for
International Cooperation and Development, Brussels
Marques MM (1963) Notas sobre a geomorfologia da Angola 1. Significado morfológico de algu-
mas ‘anharas do alto’. Garcia da Orta 11:541–560
Maurin O, Davies TJ, Burrows JE et al (2014) Savanna fire and the origins of the ‘underground
forests’ of Africa. New Phytol 204(1):201–214
Mateus O, Callapez P, Polcyn M et al (2019) Biodiversity in Angola through time: a paleonto-
logical perspective. In: Huntley BJ, Russo V, Lages F, Ferrand N (eds) Biodiversity of Angola.
and the first record from Angola. Ann Braz Acad Sci 83(1):221–233
Cham
Mendelsohn JM, Mendelsohn S (2018) Sudoeste de Angola: um Retrato da Terra e da Vida. South
West Angola: a portrait of land and life. Raison, Windhoek
Mendelsohn J, Weber B (2013) An atlas and profile of Huambo: its environment and people.
Development Workshop, Luanda, 80 pp
Mendelsohn J, Weber B (2015) Moxico: an atlas and profile of Moxico, Angola. Raison, Windhoek,
44 pp
Mendelsohn J, Jarvis A, Robertson T (2013) A profile and atlas of the Cuvelai-Etosha Basin.
Raison & Gondwana Collection, Windhoek, 170 pp
tion: A modern synthesis. Springer Nature, Cham
Mills MSL, Melo M, Vaz A (2013) The Namba mountains: new hope for Afromontane forest birds
in Angola. Bird Conserv Int 23:159–167
Mills MSL, Olmos F, Melo M et al (2011) Mount Moco: its importance to the conservation of
Swierstra’s Francolin Pternistis swierstrai and the Afromontane avifauna of Angola. Bird
Conserv Int 21:119–133
Monard A (1937) Contribution à l’herpétologie d’Angola. Arquivos do Museu Bocage 8:19–154
Monteiro RFR (1970) Estudo da Flora e da Vegetacão das Florestas Abertas do Planalto do Bié.
Instituto de Investigacão Científica de Angola, Luanda, 352 pp
Mucina L, Rutherford MC (2006) The vegetation of South Africa, Lesotho and Swaziland.
Strelitzia 19:1–807
Neto AG, Ricardo RP, Madeira M (2006) O alumínio nos solos de Angola. In: Moreira I (ed)
Angola: Agricultura, Recursos Naturais e Desenvolvimento. ISA Press, Lisboa, pp 121–143
Olson DM, Dinerstein E, Wikramanayake ED et al (2001) Terrestrial ecoregions of the World: a
new map of life on Earth. Bioscience 51:933–938
Peel MC, Finlayson BL, McMahon TA (2007) Updated world map of the Köppen-Geiger climate
classification. Hydrol Earth Syst Sci 11:1633–1644
Redinha J (1961) Nomenclaturas nativas para as formações botânicas do nordeste de Angola.
Revermann R, Finckh M (2019) Vegetation survey, classification and mapping in Angola. In:
Rodrigues P, Figueira R, Vaz Pinto P et al (2015) A biogeographical regionalization of Angolan
mammals. Mammal Rev 45:103–116
Silveira MM (1967) Climas de Angola. Serviço Meteorólogico de Angola, Luanda, 44 pp
42 B. J. Huntley
Skelton PH (2019) The Freshwater Fishes of Angola. In: Huntley BJ, Russo V, Lages F, Ferrand
Cham
Spalding MD, Fox HE, Allen GR et al (2007) Marine ecoregions of the world: a bioregionalization
of coastal and shelf areas. Bioscience 57:573–583
Thieme ML, Abell R, Stiassny ML et al (eds) (2005) Freshwater ecoregions of Africa and
Madagascar: a conservation assessment. Island Press, Washington DC
Ucuassapi AP, Dias JCS (2006) Acerca da fertilidade dos solos de Angola. In: Moreira I (ed)
piece to understand the biogeography of the Afromontane forests. MSc Thesis, University of
Porto, Porto
Weir CR (2019) The whales and dolphins of Angola. In: Huntley BJ, Russo V, Lages F, Ferrand
Cham
Welwitsch F (1859) Apontamentos phyto-geographicos sobre a Flora da Provincia de Angola na
Africa Equinocial servindo de relatório preliminar acerca da exploração botanica da mesma
provincia. Annaes do Conselho Ultramarino (Ser. 1):527–593
Werger MJA (1978) Biogeographical division of southern Africa. In: Werger MJA, van Bruggen
AC (eds) Biogeography and ecology of Southern Africa. Junk, The Hague, pp 145–170
White F (1971) The taxonomic and ecological basis of chorology. Mitteilungen Botanischen
Staatsammlung München 10:91–112
White F (1976) The underground forests of Africa: a preliminary review. The Gardens’ Bull
Singapore 24:57–71
White F (1983) The vegetation of Africa – a descriptive memoir to accompany the UNESCO/
AETFAT/UNSO vegetation map of Africa. UNESCO, Paris, 356 pp
Chapter 3
Marine Biodiversity of Angola:
Biogeography and Conservation
Stephen P. Kirkman and Kumbi Kilongo Nsingi
Abstract Some major physical and oceanographic features of the Angolan marine
system include a narrow continental shelf, the warm, southward flowing Angola
Current, the plume of the Congo River in the north and the Angola-Benguela Front
in the south. Depth, substrate types and latitude have been shown to account for
species differences in demersal faunal assemblages including fish, crustaceans, and
cephalopods. The extremely narrow shelf between Tômbwa (15°48′S) and Benguela
(12°33′S) may serve as a barrier for the spreading of shelf-occurring species
between the far south, which is influenced by the Angola-Benguela Front, and the
equatorial waters of the central and northern areas. A similar pattern is evident for
coastal and shallow-water species, including fishes, intertidal invertebrates and
seaweeds, with species that have temperate affinities found in the far south and
tropical species further to the north. In general the fauna and flora of the littoral zone
appears to be consistent with a pattern of relatively low diversity of the shore and
near-shore areas, that is characteristic of West Africa, but paucity of data for Angola
may make such comparisons of diversity with other areas inappropriate at this stage.
The Congo River delta and many features that are interspersed along the coast such
as estuaries and associated floodplains, wetlands, lagoons, salt marshes and
mangroves, support a rich suite of species, many of which are rare, endemic,
migratory, and/or threatened, and provide important ecosystem services. While the
ecological value of many areas or features is recognised, lack of any legal protection
in the form of marine protected areas (MPAs) has been identified as one of the main
challenges facing conservation and sustainable use of Angola’s marine and coastal
biodiversity and habitats, in the face of multiple threats. A current process to identify
and describe ecologically or biologically significant marine areas (EBSAs) could
provide a foundation for designating some MPAs in future.
S. P. Kirkman (*)
Department of Environmental Affairs, Oceans and Coastal Research,
Cape Town, South Africa
e-mail: skirkman@environment.gov.za
K. K. Nsingi
Benguela Current Convention, Swakopmund, Namibia
e-mail: kkilongo@gmail.com

https://doi.org/10.1007/978-3-030-03083-4_3
44 S. P. Kirkman and K. K. Nsingi
Keywords Benguela current · Ecologically or biologically significant marine areas

· Important bird areas · Fish · Marine protected areas · Marine spatial planning ·
Seaweed · Systematic conservation planning · West Africa
Physical and Oceanographic Context
The coastline of Angola, which is approximately 1650 km long, consists of sandy

and rocky stretches of coastline, punctuated by numerous coastal features such as
estuaries, mangroves, coastal lakes, wetlands and tidal flats (Harris et al. 2013).
Between Rio Bero (north of Moçâmedes) in Namibe Province and to the north of
Rio Coporolo in Benguela Province are rocky shores; the rest of the coastline is
predominantly sandy although there are some scattered rocky shores further north
of Lobito (Harris et al. 2013). The continental shelf, which extends to about 200 m
depth, is relatively narrow especially near the south where it is as little as 6 km wide
and very steep in parts of Namibe and Benguela, but it widens further north to
33 km width near the mouth of the Congo River, and in the south it widens a little
between Tômbwa and Cunene (Figure 1, Bianchi 1992). The neritic zone (i.e. waters
overlying the continental shelf) covers about one third of Angola’s Exclusive
Economic Zone (EEZ), which also includes extensive bathyal and abyssal zones,
with depths up to 4000 m in the latter (Nsiangango et al. 2007).
Hutchings et al. (2009) describe the marine system of Angola’s continental shelf
area as a subtropical transition zone between the Equatorial Atlantic to the north and
the Benguela’s wind-driven upwelling system to the south. The conspicuous,
dynamic but relatively shallow Angola-Benguela Front at 17°S in the south of
Angola forms the boundary with the upwelling system, and the boundary to the
north is near the plume of the Congo River. Seasons are well-defined and there is
intermediate productivity; moderate to weak upwelling occurs year-round in the
south and all along the coast in winter with strengthening of southeast trade winds.
The major oceanographic feature of the system is the warm (>24 °C) Angola
Current, which flows southward along the shelf and slope as an extension of the
South Equatorial Counter-current, extending down to 200 m depth and with a mean
flow at 50 m depth of 5–8 cm s−1 (Kopte et al. 2017). During winter and spring the
Angola Current tends to retreat to the northwest and is replaced by slightly cooler
waters; this is linked to the intensity of wind-driven upwelling off the Namibian
coast (Meeuwis and Lutjeharms 1990; O’Toole 1980).
Other important drivers of the system are Kelvin waves propagating from the
Equatorial Atlantic and the South Equatorial Counter Current (Florenchie et al.
2004; Shillington et al. 2006), as well as southward flow of brackish water with high
nutrient loads from the Congo River outflow and solar heating (Veitch et al. 2010),
Both result in stratification of the water column (Kirkman et al. 2016), with thermo-
cline depth ranging from 10 m in the north down to about 50 m off central Angola
(Bianchi 1992). Another feature of the Angolan system is the cold water Angola
Dome, found offshore of the Angola Current. This is a cyclonic eddy that causes
3 Marine Biodiversity of Angola: Biogeography and Conservation 45
doming of the thermocline, centred near 10°S and 9°E (Lass et al. 2000). The
Angola Dome has lower salinity and concentrations of oxygen than surrounding
waters, but it does not exist in winter and its width and extension depend on the
intensity and horizontal shear of southeasterly trade winds (Signorini et al. 1999).
Phytoplankton production associated with the Angola Dome strongly influences the
shelf ecosystem throughout northern Angola (Monteiro et al. 2008).
Biodiversity and Biogeography
There is a high diversity of demersal species in Angola relative to the temperate

Benguela Ecosystem to the south, with species richness greatest at about 100 m
depth according to research surveys (Kirkman et al. 2013). Demersal fish stocks are
exploited by a multispecies bottom trawl fishery extending from southern to northern
Angola, that exploits over 30 species of fish belonging to the families Sparidae
(seabreams), Scianidae (croakers), Serranidae (groupers), Haemulidae (grunts) and
Merlucidae (hakes). Some of the most commercially important species include
Benguela Hake Merluccius polli and demersal sparid fish such as Dentex spp.
(Kirkman et al. 2016); there is also bottom fishing for crustaceans, most importantly
deep-sea crab Chaceon maritae and shrimps Aristeus varidens and Parapenaeus
longirostris (Japp et al. 2011; Kirkman et al. 2016). The most important fish targeted
by small pelagic fisheries include Kunene Horse Mackerel Trachurus trecae and
Sardinella species, with most large pelagic fishing (tuna spp.) taking place in the
south (Japp et al. 2011, Kirkman et al. 2016). Several of the above stocks are targeted
both by industrial and artisanal fisheries (Duarte et al. 2005; Japp et al. 2011). The
Angolan fisheries are described by Hutchings et al. (2009) as being of moderate
intensity with stocks generally declining. There is also a rapidly growing local and
foreign recreational shore-fishery sector in southern Angola targeting mainly
Leerfish Lichia amia, West Coast Dusky Kob Argyrosomus coronus and Shad
Pomatomus saltatrix (Potts et al. 2009).
Bianchi (1992) and later Nsiangango et al. (2007) studied the structure of demer-
sal assemblages of the continental shelf and upper Angolan slope, including fish,
crustaceans and cephalopods, based on trawl surveys. It was shown that thermal,
depth-dependent stratification explained the main faunal groupings, with certain
species generally restricted to waters shallower than where the lower limit of the
thermocline meets the shelf, and others usually occurring in deeper waters than this.
Species such as Shallow-water Croaker Pteroscion peli, Red Pandora Pagellus bel-
lotti, Lesser African Threadfin Galeiodes decadactylus and Grunt Pomadasys inci-
sus dominated in the shallower demersal water (coast to 100 m), with some sea
breams in low densities, whereas deeper waters of the shelf and upper slope were
dominated by such species as Splitfin Synagrops microlepis, Atlantic Green-eye
Chlorophthalmus atlanticus, Angolan Dentex D. angolensis and M. polli. Within
the different depth strata, substrate type and latitudinal gradients were the main
factors affecting the composition of species assemblages, and a major latitudinal
shift both in shallow- and deep-water assemblages was shown to occur in southern
Angola between Tômbwa and Cunene where the shelf widens and where Large-
eyed Dentex D. macrophthalmus dominated the catches. Bianchi (1992) related the
shift to the southern limit of warmer equatorial waters, the presence of the Angola-
Benguela Front where there is year-round upwelling and cooler waters, and the
extremely narrow shelf to the north of Tômbwa (up to Benguela), which may serve
as a barrier to the spreading of northern species to the south and vice versa.
While deep-water coral reefs have been documented for Angola’s continental
slope (Le Guilloux et al. 2009), shallow-water coral reefs are absent and in general
the fauna and flora of littoral zone seems to be consistent with the pattern of
relatively low diversity of the shore and near-shore areas of West Africa (John and
Lawson 1991). Factors that could account for this include the lack of hard substrata
(most of the shoreline being sandy), upwelling of cooler water in areas, high
turbidity and sediment input from a massive river such as the Congo, or loss of
species associated with reductions in sea-temperatures that considerably reduced
the tropical zone during Pleistocene glaciations (van den Hoek 1975; John and
Lawson 1991). However, while recent studies (e.g. Hutchings et al. 2007; Anderson
et al. 2012) have added to the existing species lists (e.g. Lawson et al. 1975; Penrith
1978 and others) for coastal fishes, sandy beach macrofauna, rocky shore
invertebrates and seaweeds, at this stage the paucity of information in Angola may
not make comparison with other areas appropriate. In general the data that exist
both for coastal and estuarine fishes (Whitfield 2005; Hutchings et al. 2007) but also
offshore fish species (Kirkman et al. 2013; Yemane et al. 2015) of Angola show
decreasing species richness from north to south, seemingly supporting the
established trend of decreasing diversity with latitude as one moves polewards from
tropical regions (e.g. Rex et al. 2000; Willig et al. 2003).
Based on the latitudinal distributions of intertidal fauna of rocky shores (Kensley
and Penrith 1980), the southern limit of tropical biota has previously been reported
to be around the border of Angola and Namibia. Lawson (1978) on the other hand,
based on analyses of seaweed flora, considered Angola to be intermediate in nature
between tropical and temperate. Results of surveys of intertidal invertebrates and
seaweeds by Hutchings et al. (2007) however, showed that although there was a
marked discontinuity between the biota of Angola and that of northern Namibia,
which supports a cool-temperate intertidal flora up to nearby the Cunene River (Rull
Lluch 2002), a number of taxa found in the south of Angola had temperate affinities.
This led the authors to suggest that the inshore biota of the south of Angola may be
intermediate in nature, and that of the north truly tropical. This is confirmed by
Anderson et al. (2012) who conclude that the overall affinities of the Angolan
seaweed flora is Tropical West African, but with a well-developed temperate element
in southern Angola (from about 13°S) comprising mainly cooler-water species.
Broadly, this supports the division of the Angolan coast into at least two sub-areas,
with the more temperate south influenced by the cooler waters of the Angola-
Benguela Front. This is similar to the division between demersal assemblages of
north and south (Bianchi 1992) and also congruent with a break in the pelagic
ecosystem of the inshore as determined from classification of key oceanographic
variables and depth (Lagabrielle 2011). It also ties in with global mapping classifi-
Fig. 3.1 Delineations of marine ecoregions (Spalding et al. 2007) and large marine ecosystems
(Sherman 2014) that coincide with Angola. The ecoregions extend from the coast to the shelf edge.
Also shown are recognised marine and coastal biodiversity areas and the approximate situations of
important oceanographic processes
cation of coastal and shelf areas based on species distributions and levels of ende-
mism of benthic and pelagic biota (Spalding et al. 2007; Briggs and Bowen 2012),
that puts the divide between the temperate Benguela Province and the tropical Gulf
of Guinea Province, near Moçâmedes (Fig. 3.1). Spalding et al. (2007) situate the
majority of the Angolan EEZ in the Angolan ecoregion of the Gulf of Guinea
Province, but include the area north of 6°30′S in the more tropical Gulf of Guinea
south ecoregion. This is slightly incongruent with the mapping of large marine eco-
systems (LMEs) of the world (which is expert- rather than data-derived), whereby
most of Angola is included in the Benguela Current LME bounded to the north by
the Angola Front (ca. 5°S), and only Cabinda in the far north included in the Guinea
Current LME (Sherman 2014).
arine and Coastal Biodiversity Hotspots, Threats

M
and the Need for Protection
Whereas the Angolan coastal and shallow habitats are considered to be relatively
low in biodiversity, coastal features such as the Congo River Delta, estuaries such as
the Cuanza, Catumbela, Longa and Cunene, and associated floodplains, wetlands,
lagoons, salt marshes and (north of Lobito) mangroves support a rich suite of
species, often in high abundance (Hughes and Hughes 1992; van Niekerk et al.
2008; Harris et al. 2013). This includes several rare, endemic, migratory, and/or
threatened fauna such as the African Manatee Trichechus senegalensis, turtle species
and diverse waterbird species. Recognised ecosystem services of such features
include (amongst others) providing habitat for important food-fish and crustacean
species and their critical life stages (e.g. performing important nursery functions for
many marine fishes) or providing plant species that are useful for medicinal,
subsistence or construction purposes (Hughes and Hughes 1992; van Niekerk et al.
2008). While Angola is not currently a contracting party of the Ramsar Convention,
some coastal wetland sites have been identified as being potential Ramsar sites,
including Quiçama National Park between the Cuanza and Longa Rivers (Fig. 3.1),
which is also a confirmed Important Bird and Biodiversity Area (IBA). Other
confirmed coastal IBAs in Angola include Mussulo just south of Luanda and the
Iona National Park in the south between the Cunene and Curoca Rivers. These IBAs
are important for numerous waterbirds and are frequented by wintering seabird
species that breed further south on the sub-continent such as Cape gannet Morus
capensis (IUCN Red List – Endangered) and Damara tern Sternula balaenarum
(Vulnerable) (Birdlife International 2002), the latter of which is known to also breed
in the Iona National Park (Simmons 2010).
While the ecological value of these and other areas is recognised, the lack of
formal protection of key biodiversity areas or features in Angola’s marine and
coastal environments has been noted as a concern (e.g. Tarr et al. 2007). As part of
a regional systematic conservation planning (SCP) project involving all three
member states of the Benguela Current Convention (BCC; a legally constituted
collaborative mechanism representing Angola, Namibia and South Africa), Holness
et al. (2014) showed that Angola is particularly poorly off in terms of spatial
protection of its marine systems, with 102 out of 133 identified ecosystem types
having no protection at all. Whereas some legislative protection of coastal ecosystem
types in the Cuanza, Cunene and Tômbwa areas may be afforded by terrestrial
national parks (Quiçama and Iona) or reserves (Namibe Partial Reserve), this may
have value for conservation of the adjacent marine areas if effective management of
these areas is achieved through the provision of increased human and financial
resources. Holness et al. (2014) therefore opined that a programme of rapid
expansion of protected areas for the Angolan marine systems is urgently required,
and the ultimate product of their study was the prioritisation of sites for protection
(ideally within a MPA network).
The current lack of marine protected areas (MPAs) was described by Tarr et al.
(2007) as amongst the main challenges facing conservation and sustainable use of
Angola’s marine and coastal biodiversity and habitats, in light of multiple threats to
the ecosystem that are likely to worsen over time. These threats include (but are not
limited to) rapid, unplanned coastal urbanisation causing habitat destruction and a
severe problem with waste management along the coast, particularly in the area of
Luanda; escalation in over-exploitation of living marine resources related to rapid
urbanisation and human migration to the coastal nodes, especially since the end of
the civil war; industrial pollution caused e.g. by deposition of industrial wastes in
catchment areas or cleaning of ships; offshore oil exploitation in the north, with
potential for oil spills; loss of mangroves, which includes threats from pollution and
wood collection for firewood and construction; rapid growth of the tourism industry;
and impacts of climate change (Tarr et al. 2007; Heileman and O’Toole 2009).
With such threats in mind, Angola, like the other two member states of the BCC,
has committed to implementing ecosystem-based management (EBM) of the marine
environment to address responsible use of its ocean and its resources and put in
practice the principles of sustainable development (BCC 2014). EBM is an
integrative approach to management that takes into account all interactions in the
ecosystem (including those involving human activities) and their cumulative
impacts in space and time (Long et al. 2015). To be able to assist EBM with regard
to the allocation and siting of ocean uses or protection measures, there is an initiative
to implement marine spatial planning (MSP) in Angola and the other countries of
the region (Kirkman et al. 2016). A pilot area for an experimental MSP project,
covering an area of approximately 107,000 km2 between south of Palmerinhas and
the Tapado River mouth (GNC-OEM 2018), has recently been identified. A key
element of the process is to identify and describe a network of Ecologically or
Biologically Significant Marine Areas (EBSAs) - geographically or
oceanographically discrete areas that have been identified as important for the
services that they provide and for the healthy functioning of oceans (Dunstan et al.
2016) and to include these in marine spatial plans.
Currently, only two Angolan EBSAs have been described and subsequently
endorsed by CBD (CBD 2014), namely the Ramiros-Palmerinhas Coastal Area
which partly adjoins the Mussulo Peninsula south of Luanda, and the Cunene-
Tigres EBSA which overlaps with northern Namibia and is adjacent to the Iona
National Park on the Angola side (Fig. 3.1). The former includes estuaries with
mangroves and salt marshes and has special importance for bird aggregations and
breeding turtles. The latter includes the Cunene estuary and associated wetland as
well as the Baía dos Tigres Island-Bay complex to the north of it, and has special
importance for migratory birds and in terms of its nursery function for many marine
species. Both of these areas have undergone thorough assessment processes with a
view to expanding their areas in order to include other relevant features such as
estuaries, sensitive coastline, canyons and seamounts.
Currently Angola is in the process of describing new potential EBSAs, in coastal
and offshore areas, as part of a collaborative regional project with Namibia and
South Africa, coordinated by the BCC (http://www.benguelacc.org). Currently, five
new areas have been proposed as EBSAs which include coastal and offshore areas
in the provinces of Cabinda, Zaire, Luanda, Cuanza-Sul and Namibe. Although
EBSA status itself does not carry any conservation or protection interventions, legal
protection is among the management measures that can be applied to secure the
persistence of these special features and their ecosystem services. Therefore the
process of expanding the EBSA network could provide a foundation for initiating a
network of MPAs in Angola. In this regard, there is a recent project proposal for the
establishment of the first MPA in Angola in the offshore area adjacent to the Iona
National Park.
References
Anderson RJ, Bolton JJ, Smit AJ et al (2012) The seaweeds of Angola: the transition between trop-
ical and temperate marine floras on the west coast of southern Africa. Afr J Mar Sci 34:1–13
BCC (Benguela Current Commission) (2014) Strategic action programme 2015–19. Swakopmund,
Namibia, 36 pp. http://benguelacc.org/index.php/en/publications
Bianchi G (1992) Demersal assemblages of the continental shelf and upper slope of Angola. Mar
Ecol Prog Ser 81:101–120
BirdLife International (2002) Important bird areas and potential Ramsar sites in Africa. BirdLife
International, Cambridge, MA, 136 pp + appendices
Briggs JC, Bowen BW (2012) A realignment of marine biogeographic provinces with particular
reference to fish distributions. J Biogeogr 39:12–30
CBD (Convention on Biological Diversity) (2014) Decision adopted by the Conference of the
Parties to the Convention on Biological Diversity. XII/22. Marine and Coastal Biodiversity:
Ecologically or Biologically Significant Marine Areas (EBSAs). Twelfth meeting of the
Conference for the Parties, 6–17 October 2014, Pyeongchang, Republic of Korea. UNEP/CBD/
COP/DEC/XII/22
Duarte A, Fielding P, Sowman M, et al (2005) Overview and analysis of socio-economic and
fisheries information to promote the management of artisanal fisheries in the Benguela Current
Large Marine Ecosystem (BCLME) region (Angola). Unpublished Final Report. Rep. No.
LMRAFSE0301B. Cape Town Environmental Evaluation Unit, University of Cape Town,
Cape Town
Dunstan PK, Bax NJ, Dambacher JM et al (2016) Using ecologically or biologically significant
marine areas (EBSAs) to implement marine spatial planning. Ocean Coast Manag 121:116–127
Florenchie P, Reason CJC, Lutjeharms JRE et al (2004) Evolution of interannual warm and cold
events in the Southeast Atlantic Ocean. J Clim 17:2318–2334
Grupo Nacional de Coordenação para o Ordenamento do Espaço Marinho (GNC-OEM) (2018).
Relatório Preliminar sobre o Ordenamento do Espaço Marinho em Angola: Área Experimental
Palmeirinhas – Tapado. Unpublished report
Harris L, Holness S, Nel R et al (2013) Intertidal habitat composition and regional-scale shoreline
morphology along the Benguela coast. J Coast Conserv 17:143–154
Heileman S, O’Toole MJ (2009) I West and Central Africa: I-1 Benguela current LME. In: Sherman
K, Hempel G (eds) The UNEP large marine ecosystems report: a perspective on changing con-
ditions in LMEs of the World’s regional seas. UNEP Regional Seas Report and Studies No.
182. United Nations Environment Programme, Nairobi, pp 103–115
Holness S, Kirkman S, Samaai T, et al (2014) Spatial biodiversity assessment and spatial manage-
ment, including marine protected areas. Final report for the Benguela Current Commission
project BEH 09-01, 105 pp + annexes
Hughes RH, Hughes JS (1992) A directory of African wetlands. IUCN/UNEP/WCMC, Gland/
Cambridge/Nairobi/Cambridge, xxiv + 820 pp, 48 maps
Hutchings K, Clark B, Steffani, Anderson R (2007) Identification of communities, biotopes and
species in the offshore areas and along the shoreline and in the shallow subtidal areas in the
BCLME region. Section B. Angola field trip report. Final report for Benguela Current Large
Marine Ecosystem Programme project BEHP/BAC/03/03
Hutchings L, van der Lingen CD, Shannon LJ et al (2009) The Benguela current: an ecosystem of
four components. Prog Oceanogr 83:15–32
Japp DW, Purves MG, Wilkinson S (eds) (2011) State of stocks review. Report No. 2 (Updated by
C Kirchner). Benguela Current Large Marine Ecosystem State of Stocks Report 2011, 105 pp
John DM, Lawson GW (1991) Littoral ecosystems of tropical western Africa. In: Mathieson AC,
Nienhuis PH (eds) Ecosystems of the world, vol 24. London. New York, Tokyo, pp 297–321
Kensley B, Penrith ML (1980) The constitution of the fauna of rocky intertidal shores of south
West Africa. Part III. The north coast from False Cape Frio to the Kunene River. Cimbebasia
(Series A) 5:201–214
Kirkman SP, Blamey L, Lamont T et al (2016) Spatial characterisation of the Benguela ecosystem
for ecosystem based management. Afr J Mar Sci 38:7–22
Kirkman SP, Yemane D, Kathena J et al (2013) Identifying and characterizing of demersal bio-
diversity hotspots in the BCLME: relevance in the light of global changes. ICES J Mar Sci
70:943–954
Kopte R, Brandt P, Dengler M et al (2017) The Angola current: flow and hydrographic character-
istics as observed at 11°S. J Geophys Res Oceans 122:1177–1189
Lagabrielle E (2011) A pelagic bioregionalisation of the Benguela current system. Appendix 4
In: Holness S, Kirkman S, Samaai T, et al (2014) Spatial Biodiversity Assessment and Spatial
Management, including Marine Protected Areas. Final report for the Benguela Current
Commission project BEH 09-01, 105 pp + annexes
Lass HU, Schmidt M, Mohrholz V et al (2000) Hydrographic and current measurements in the area
of the Angola-Benguela front. J Phys Oceanogr 30:2589–2609
Lawson GW (1978) The distribution of seaweed floras in the tropical and subtropical Atlantic
Ocean: a quantitative approach. Bot J Linn Soc 76(3):177–193
Lawson GW, John DM, Price JH (1975) The marine algal flora of Angola: its distribution and
affinities. Bot J Linn Soc 70(4):307–324
Le Guilloux E, Olu K, Bourillet JF et al (2009) First observations of deep-sea coral reefs along the
Angola margin. Deep Sea Research Part II: Tropical Studies in Oceanography 56:2394–2403
Long RD, Charles A, Stephenson RL (2015) Key principles of marine ecosystem-based manage-
ment. Mar Policy 57:53–60
Meeuwis JM, Lutjeharms JRE (1990) Surface thermal characteristics of the Angola-Benguela
front. S Afr J Mar Sci 9:261–279
Monteiro PMS, van der Plas AK, Mélice J-L et al (2008) Interannual hypoxia variability in a
coastal upwelling system: ocean-shelf exchange, climate and ecosystem-state implications.
Deep-Sea Res I 55:435–450
Nsiangango S, Shine K, Clark B (2007) Identification of communities, biotopes and species in
the offshore areas and along the shoreline and in the shallow subtidal areas in the BCLME
region. Section C3. Biogeographic patterns and assemblages of demersal fishes on the coast
of Angola. Final report for Benguela Current Large Marine Ecosystem Programme project
BEHP/BAC/03/03
O’Toole MJ (1980) Seasonal distribution of temperature and salinity in the surface waters off south
West Africa, 1972–1974. Investig Rep S Afr Sea Fish Inst 121:1–25
Penrith MJ (1978) An annotated check-list of the inshore fishes of southern Angola. Cimbebasia
(Series A) 4:179–190
Potts WM, Childs AR, Sauer WHH et al (2009) Characteristics and economic contribution of a
developing recreational fishery in southern Angola. Fish Manag Ecol 16:14–20
Rex MA, Stuart CT, Coyne G (2000) Latitudinal gradients of species richness in the deep-sea
benthos of the North Atlantic. Proc Natl Acad Sci USA 97:4082–4085
Rull Lluch JR (2002) Marine benthic algae of Namibia. Sci Mar 66. (suppl. 3:5–256
Sherman K (2014) Toward ecosystem-based management (EBM) of the world’s large marine eco-
systems during climate change. Environ Dev 11:43–66
Shillington FA, Reason CJC, Duncombe Rae CM et al (2006) Large scale physical variability
of the Benguela Current Large Marine Ecosystem (BCLME). In: Shannon V, Hempel G,
Malanotte-Rizzoli P et al (eds) Benguela: predicting a large marine ecosystem, vol 14. Elsevier,
Amsterdam, pp 49–70
Signorini SR, Murtuguddo RG, McClain CR et al (1999) Biological and physical signatures in the
tropical and subtropical Atlantic. J Geophys Res 104:18367–18382
Simmons RE (2010) First breeding records for Damara Terns and density of other shorebirds along
Angola’s Namib Desert coast. Ostrich 81:19–23
Tarr P, Krugmann H, Russo V, Tarr J, et al (2007) Analysis of threats and challenges to marine bio-
diversity and marine habitats in Namibia and Angola. Final report for Benguela Current Large
Marine Ecosystem Programme project BEHP/BTA/04/01. 132 pp + annexes
Van den Hoek C (1975) Phytogeographic provinces along the coast of the northern Atlantic Ocean.
Phycologia 14:317–330
Van Niekerk L, Neto DS, Boyd AJ, et al (2008) Baseline surveying of species and biodiversity in
estuarine habitats. BCLME project BEHP/BAC/03/04. 118 pp + appendices
Veitch JA, Penven P, Shillington F (2010) Modeling equilibrium dynamics of the Benguela Current
System. J Phys Oceanogr 40:1942–1964
Whitfield AK (2005) Preliminary documentation and assessment of fish diversity in sub-Saharan
African estuaries. Afr J Mar Sci 27(1):307–324
Willig MR, Kaufman DM, Stevens RD (2003) Latitudinal gradients of biodiversity: pattern, pro-
cess, scale, and synthesis. Annu Rev Ecol Evol Syst 34:273–309
Yemane D, Mafwila SK, Kathena J et al (2015) Spatio-temporal trends in diversity of demersal fish
species in the Benguela Current Large Marine Ecosystem (BCLME) region. Fish Oceanogr 24.
(Suppl. 1:102–121
Chapter 4
The Fossil Record of Biodiversity
in Angola Through Time: A Paleontological
Perspective
Octávio Mateus, Pedro M. Callapez, Michael J. Polcyn, Anne S. Schulp,

António Olímpio Gonçalves, and Louis L. Jacobs
Abstract This chapter provides an overview of the alpha paleobiodiversity of

Angola based on the available fossil record that is limited to the sedimentary rocks,
ranging in age from Precambrian to the present. The geological period with the
highest paleobiodiversity in the Angolan fossil record is the Cretaceous, with more
than 80% of the total known fossil taxa, especially marine molluscs, including
ammonites as a majority among them. The vertebrates represent about 15% of the
known fauna and about one tenth of them are species firstly described based on
specimens from Angola.
O. Mateus (*)
GeoBioTec, Faculdade de Ciências e Tecnologia, Universidade Nova de Lisboa,
Lisbon, Portugal
Museu da Lourinhã, Rua João Luis de Moura, Lourinhã, Portugal
e-mail: omateus@fct.unl.pt
P. M. Callapez
CITEUC; Departamento de Ciências da Terra, Faculdade de Ciências e Tecnologia,
Universidade de Coimbra, Coimbra, Portugal
e-mail: callapez@dct.uc.pt; jacobs@smu.edu
M. J. Polcyn · L. L. Jacobs
Roy M. Huffington Department of Earth Sciences, Southern Methodist University,
Dallas, TX, USA
e-mail: mpolcyn@smu.edu
A. S. Schulp
Naturalis Biodiversity Center, Leiden, The Netherlands
Faculty of Earth and Life Sciences, VU University Amsterdam, Amsterdam, The Netherlands
e-mail: anne.schulp@naturalis.nl
A. O. Gonçalves
Departamento de Geologia, Faculdade de Ciências, Universidade Agostinho Neto,
Luanda, Angola
e-mail: antonio.goncalves@geologia-uan.com

https://doi.org/10.1007/978-3-030-03083-4_4
54 O. Mateus et al.
Keywords Ammonites · Benguela Basin · Cenozoic · Cretaceous · Cuanza Basin ·

Dinosaur · Invertebrates · Mammals · Mollusca · Mosasaur · Namibe Basin ·
Paleobiodiversity · Pleistocene · Vertebrate · Plesiosaur · Turtle
Studies of Paleobiodiversity
The study of paleobiodiversity, i.e., the development of biodiversity through geo-

logical time, is challenging at multiple levels. In addition to the issues and biases
affecting the study of the diversity of modern life, understanding paleobiodiversity
faces extra challenges, mostly because of the dependency on the fossil record.
Glimpses of entire ecosystems and clades may never reach the paleontologist’s eyes
if appropriate rocks of that exact time and space were not formed, or if formed, did
not preserve fossils, or are eroded away or otherwise inaccessible (see Jackson and
Johnson 2001; Crampton et al. 2003).
The study of life’s diversity in the past is filtered by the remains that can leave
traces and fossilize, remains that actually fossilized, fossils existing today, fossils
accessible today, fossils collected (number of fossils accessible to scientists), and
species recognised (Fig. 4.1). Moreover, the definition and discrimination of species
in the fossil record can be problematic.
Angola has no known fossiliferous rocks from the Paleozoic (541–251 Ma – mil-
lions of years ago) nor the Jurassic (199–145 Ma) leaving only windows to life in
the territory during the Triassic (251–199 Ma), Cretaceous (145–66 Ma), and
Cenozoic (66 Ma -Present) (Fig. 4.2). The chances of finding Paleozoic or Jurassic
fossils from Angola are essentially zero. Thus, within the last 550 Ma, the known
1. Life diversity
2. Life remains that can leave traces and fossilize
3. Remains that actually fossilized
4. Fossils existing today and rock outcrops
5. Fossil accessibility
6. Fossil collection
7. Species
recognition
Fig. 4.1 The seven layers of filters of uneven preservation in the fossil record that obscure accu-
rate reconstruction of paleobiodiversity
4 The Fossil Record of Biodiversity in Angola Through Time: A Paleontological… 55
Fig. 4.2 The geological record of Angola, in a stratigraphic log (left) and geological map (right),
leaves more than 350 Ma of blank geological record. Map extracted after Africa Geological Map
1:30.000.000 by U.S. Geological Survey, 2002 available at www.uni-koeln.de/sfb389/e/e1
rocks of Angola represent less than 196 million years of geologic time, leaving more
than 354 million years (64% of the time) with no known fossil record.
Despite the incompleteness of the fossil record and consequent limitations to the
study of the paleobiodiversity, cooperative research and modern databases can,
however, improve approximation of the estimated number of species in the fossil
record. The PaleoBiology Database (paleobiodb.org) is, by far, the most compre-
hensive database of fossils, which, together with the scientific literature and our own
research, contributed to the Supplementary Material and its summary in Table 4.1
that compiles the list of the fossil taxa in Angola, with updated taxonomy, geologi-
cal age, locality, and references. The fossil record can be used as a lower limit for
the alpha paleobiodiversity for specific times and locations in Angola, although it is
likely an underestimate of true paleodiversity in the vast majority of cases. The total
of all fossil species is a gross underestimation of paleodiversity for the full extent of
the time involved, exacerbated by missing intervals of fossiliferous rocks. However,
the pattern through time can inform an understanding of general trends.
Fossil collecting in Angola has been conducted since the nineteenth century with
Friederich Welwitsch, José de Anchieta (1885), Freire de Andrade, Augusto Eduardo
Neuparth and others (Brandão 2008, 2010; Silva and Geirinhas 2010; Callapez et al.
2011; Masse and Laurent 2016). Numerous paleontologists have contributed to the
56
Table 4.1 Summary of the known fossil record of Angola

Period
COUNT (Genus/sp, Family Indet) Triassic Cretaceous Paleogene Neogene Quaternary Triassic Grand Total
Bacteria Cyanobacteria 1 1 0%
Bacterial total 1 1 0%
Plantae Chlorophyta 1 1 0%
Cycadophyta 1 1 0%
Ginkgophyta 1 1 0%
Pinophyta 1 1 1 2 0%
Pteridosperma-tophyta 4 4 4 0%
Rhodophyceae 2 2 0%
Plantae total 5 6 5 9 1%
Protista and Invertebrates Foraminifera 207 207 16%
Anthozoa 1 1 2 2 6 0%
Brachiopoda 1 1 0%
Mollusca 677 27 22 72 798 61%
Echinodermata 61 1 4 1 67 5%
Arthropoda 13 3 1 1 2 13 20 2%
Protista and invertebrates total 13 950 30 29 77 13 1102 84%
Vertebrata Chondrichthyes 2 63 1 18 2 84 6%
Actinopterygii 6 10 5 6 21 2%
Sarcopterygii 3 3 3 0%
Reptilia 21 5 26 2%
Mammalia 12 1 54 67 5%
Vertebrata total 11 94 18 24 54 11 201 15%
Grand Total 29 1052 48 53 131 29 1313 100%
2% 80% 4% 4% 10% 2% 100%
O. Mateus et al.
understanding the paleobiodiversity of Angola, since the first explorations and stud-
ies: Fernando Mouta, Paul Choffat, Carlos Teixeira, Carlos Freire de Andrade,
Edgard Casier, A Jamotte, M Leriche, Heitor de Carvalho, E Dartevelle, Miguel
Telles Antunes, Alexandre Borges, Philippe Brebion, Gaspar Soares de Carvalho,
Louis Dollo, Henri Douvillé, Otto Haas, Manuel Mascarenhas Neto, Arménio
Tavares da Rocha, Gumerzindo Henriques da Silva, LF Spath, António Ferreira
Soares, Maurice Collignon, among many others (see bibliography compiled by
Nunes 1991). In vertebrate paleontology the work of Miguel Telles Antunes and
co-authors is noteworthy. Today, various researchers work on the paleontology of
Angola, among them is the team of the Projecto PaleoAngola (paleolabs.org/paleo-
angola), with regular yearly scientific expeditions since 2005 (Jacobs et al. 2006,
2016).
A Brief Geological History and Context of Angola
The most significant geological event governing the paleogeography of Angola is

the opening of the South Atlantic Ocean, in which Africa and South America rifted
apart beginning in the Early Cretaceous Epoch about 134 million years ago and the
subsequent drifting apart of these continents as the South Atlantic grew (Guiraud
and Maurin 1991; Buta-Neto et al. 2006; Quirk et al. 2013; Pérez-Díaz and Eagles
2017). After about 120 million years ago, marine deposition along the coast began
to preserve fossils. Africa’s place in Gondwana prior to this time resulted in the lack
of a marine record for the entire Paleozoic Era and the consequent lack of a fossil
record for that time.
As the South Atlantic opened it was colonised by species moving in from the
southern ocean and, as a connection between the North Atlantic and South Atlantic
oceans developed, from the north. Sea turtles (Angolachelys mbaxi) and mosasaurs
(Angolasaurus bocagei and Tylosaurus iembeensis), with relatives to the north, first
occur at about 88 million years ago. Along with them were plesiosaurs, probably
with southern affinities. Washed into the sea and found with marine creatures, is the
sauropod dinosaur Angolatitan adamastor, which is probably a remnant of a more
broadly distributed Gondwanan dinosaur assemblage. At that time, coastal Angola
lay approximately 10–12 degrees further south than today.
Geologic uplift along the coast resulted in the erosion of rock and loss of the fos-
sil record from effected strata. Permian and Jurassic uplift eliminated those intervals
from the terrestrial fossil record. Early and Late Cretaceous uplift also occurred,
prior to burial of the remaining Cretaceous with up to 1.5–2 km of sediments, since
removed by erosion accompanying mid-Cenozoic uplift beginning around 30 Ma
and 20 Ma (Green and Machado 2015). Uplift from as young as 45,000 years ago
also has been recorded along the Angolan coast (Walker et al. 2016), resulting in a
large diversity of fossiliferous raised-beach deposits, frequently associated with
pre-historic shell-middens with Paleolithic industries.
58 O. Mateus et al.
The Precambrian – The First Fossils
Precambrian Era (geologic time since the formation of Earth and prior to 541 Ma)
rocks worldwide are mostly devoid of fossils as life was unicellular for most time
and only macroscopic in the last stages. The shale-limestone series of the Bembe
System in Angola includes dolomitic limestones, mostly devoid of fossils, but con-
taining levels with concentrations of coalescent stromatolites (structures due to cya-
nophilic activity) attributable to the genera Collenia and Conophyton in Mavoio,
Alto Zambeze, and Humpata (Vasconcelos 1951; Antunes 1970; Duarte et al. 2014).
In Angola, no fossils are known from the Paleozoic (Cambrian through Permian
periods), which represents a time gap of more than 290 million years (Fig. 4.2).
The Triassic – Inner Basins
The Triassic is a geological period stretching from about 250 to 200 Ma. The begin-
ning and end of this interval are both marked by mass extinction events, the older
Permian-Triassic extinction event, concomitant with the Siberian Magmatic event,
marking the initiation of the Triassic Period. The vast supercontinent Pangea existed
until the Triassic, after which it gradually began to break-up apart, separating the
two masses of land, Laurasia to the north and Gondwana to the south. The global
climate during the Triassic Period was warm and dry, with deserts covering much of
the interior of Pangea. The end of the Triassic was marked by another major mass
extinction related with the Central Atlantic Magmatic Province and the early open-
ing of the North Atlantic. Therapsids (a large group containing mammals and their
extinct relatives) and archosaurs (dinosaurs, birds, crocodiles and their relatives)
were the major terrestrial vertebrates during this time. The dinosaurs first appeared
in the Triassic. The first true mammals, which are derived therapsids, evolved dur-
ing this period, as well as the first flying vertebrates, the pterosaurs.
Triassic outcrops of Angola are restricted to the Baixa de Cassange (Cassange
Depression) in Malanje and Lunda-Norte in rocks referred to the Karoo Supergroup
geological unit.
Plants and Invertebrates
The Triassic paleoflora of Angola includes the extinct genera Glossopteris,

Sphenopteris, and Noeggerathiopsis (Teixeira 1948a, 1961). A few fossils indicate
a freshwater environment and a Triassic age. These include the coleopteran
Coptoclavidae insect Coptoclavia africana and 12 conchostracan crustaceans:
Estheriella moutai Leriche 1932, E. cassambensis Teixeira 1958, Estheria anchietai
Teixeira 1947, E. (Echinestheria) marimbensis Marliére 1950, E. (Euestheria) man-
galiensis Jones 1862, Palaeolimnadiopsis reali Teixeira 1958, Palaeolimnadia
(Palaeolimnadia) wianamattensis (Mitchell), P. (Grandilimnadia) oesterleni Tasch

1987, P. (G.) africania Tasch 1987, Gabonestheria gabonensis (Marliére 1950),
Cornia angolata Tasch 1987, and Estheriina (Nudusia) cf. rewanensis Tasch 1979.
Vertebrates
The only Triassic vertebrates known from Angola are fishes, including the
Elasmobranchii Lissodus cassangensis, the paleoniscoids Perleidus lutoensis
Teixeira 1947, Palaeonisciformes canobiid Marquesia moutai, Halecostomi
Angolaichthys lerichei Teixeira, the ray-finned Teffichthys lehmani and T. lutoensis,
and the Sarcopterygii lungfish Microceratodus angolensis.
This faunal assemblage indicates a freshwater environment with insects and a
nearly exclusively endemic fauna. Based on the fishes, a Lower Triassic age (252–
247 Ma) is indicated for Lunda and Baixa de Cassange rocks (Murray 2000; Antunes
et al. 1990). No tetrapods have been collected so far.
The Early Cretaceous – The Opening of South Atlantic
Most of the fossils and outcrops of Cretaceous age in Angola are in Mesozoic-
Cenozoic basins of coastal Angola: the Cabinda, Zaire, Cuanza, Benguela, and
Namibe sedimentary basins (Antunes 1964; Séranne and Anka 2005; Guiraud et al.
2010), bordered by basement rocks. Almost all formations are mostly marine except
for the ichnofauna from the Catoca mine (Marzola et al. 2014; Mateus et al. 2017),
in Lunda-Sul. The oldest fossiliferous Cretaceous formations seen in outcrop seem
to be Barremian to Aptian lacustrine deposits that contain unidentified gastropods
(Ceraldi and Green 2016).
Plants, Protists and Invertebrates
Cretaceous plant remains in Angola seem to be rare (see Supplementary Material),

but several examples of unstudied field contexts with transitional facies, namely
above the Cuvo units, might come to reveal new fossil sites. The same situation is
likely for palynomorphs and dinoflagellates of Early Cretaceous and more recent
ages. Plants are known from pollen taxa such as Classopolis sp. and Eucommiidites
sp. Pachypteris montenegroi Teixeira 1948 is a Ginkgophyta Umkomasiaceae from
Early Albian lagoonal sediments of the Cuanza Basin (Teixeira 1948b; Antunes
1964; Neto 1970; Nunes 1991) and first found in Angola. One species of Chlorophyta,
one Cycadophyta, one Rhodophyceae, one Ginkgophyta, and two Pinophyta have
been reported from Cabinda, Benguela and Cuanza Basin (Antunes 1964; Neto
1970; Nunes 1991; Araújo and Guimarães 1992; Tavares 2006).
60 O. Mateus et al.
More than 200 foraminifera taxa were identified in the Cretaceous of Angola,
and many indicate an Albian age, such as Globotruncana ventricosa (Rocha 1984;
Antunes and Cappetta 2002; Antunes 1964; Jacobs et al. 2006).
Crustaceans are known from the Decapoda Parapirimela angolensis Van Straelen
1937 from the Albian of Iela beach, Benguela Basin (Ferreira 1957; Van Straelen
1937; Antunes 1964) and the ostracods Chloridella angolia and Petrobrasia tenui-
striata longinsuela from Quiçama, Cuanza Basin, (Berry 1939; Antunes 1964) and
Cabinda (Araújo and Guimarães 1992), respectively.
The Early Cretaceous of Angola yielded fossils of seven species of Scleractinean
corals, one of brachiopod and eight crustaceans.
Of the more than known 600 species of molluscs from the Cretaceous of Angola,
the vast majority are ammonites, many are unique to Angola and received related
specific epithets such as the Anisoceras teixeirai, Durnovarites autunesi Collignon
1978, Durnovarites netoi, Elobiceras lobitoense Spath 1922, Hamitoides angolanus
(Tavares 2006), followed by bivalves such as Neithea angoliensis Newton 1917, and
gastropods such as “Cerithium” monteroi Choffat. By far, the ammonites are the
most relevant portion of the Cretaceous biodiversity of Angola (Tavares et al. 2007;
Haas 1942, 1943) and also age-indicators for geologists and paleontologists. Haas
(1942) described and reported many ammonites from the Albian, some as new spe-
cies including Hysteroceras falcicostatum Haas 1942 and H. intermedium Haas
1942. See the Supplementary Material for the complete list.
Echinoderms, mostly echinoids, are remarkably common in the Early Cretaceous
of Angola, with about 50 taxa known, but that number depends on the validity and
synonymisation of the taxa addressed (see the Supplementary Material for the com-
plete list). A few echinoderms received names after Angolan toponyms and research-
ers such as the Douvillaster benguellensis Loriol 1888 and D. carvalhoi Loriol 1888
and Epiaster catumbelensis Loriol 1888, and Holaster domboensis Loriol 1888
from the Lower Cretaceous of Dombe Grande, Catumbela and Praia da Hanha
(Loriol 1888; Ferré and Granier 2001; Tavares 2006; Tavares et al. 2007).
Vertebrates
In the Catoca Diamond Mine, in Lunda-Sul Province, mammaliamorph, crocodylo-

morph, and sauropod tracks were discovered in Early Cretaceous crater lake sedi-
ments. One sauropod track has skin impressions preserved. These are the only fossil
vertebrate tracks known in Angola. The most surprising feature is the unexpectedly
large size of the mammaliamorph footprints considering the age (Mateus et al.
2017). The Catoca Diamond Mine has an eruption age of around 118 Ma (Aptian;
Robles-Cruz et al. 2012). A fragment of a caudal vertebra of a sauropod dinosaur
was recovered from Tzimbio, in northern Namibe, from strata that are likely Albian
in age.
The Late Cretaceous – Marine Reptiles Flourish
The Late Cretaceous is the time of the Cretaceous Period between 100.5 Ma and 66
million years ago. It is subdivided into the Cenomanian, Turonian, Coniacian,
Santonian, Campanian and Maastrichtian ages, from the oldest to the most recent.
The climate was warmer than present, although with a cooling trend throughout the
period. In the oceans, where the sea-level was much higher than today, mosasaurs (a
group of marine lizards) suddenly appeared and underwent a spectacular evolution-
ary radiation (Polcyn et al. 2014). Modern grade sharks also appeared and plesio-
saurs diversified. These predators fed on the numerous teleost fishes, which in turn
evolved into new advanced and modern forms (Neoteleostei). Ichthyosaurs and
pliosaurs (a group of short-necked plesiosaurs), on the other hand, became extinct
during the Cenomanian-Turonian anoxic event (Schlanger et al. 1987) and are not
known from Angola. The end of Cretaceous is marked by the mass extinction of
some three-quarters of plant and animal species on Earth, known as the Cretaceous-
Paleogene (K/Pg) event (Archibald et al. 2010).
Protists and Invertebrates
The Late Cretaceous Foraminifera of Angola were studied by various researchers

including Ferreira and Rocha (1957), Lapão and Simões (1972), Rocha (1984) and
Blake et al. (1996) who list more than 180 taxa. Foraminifera are known from 15
taxa of Granuloreticulosea, such as the Gavelinellidae Anomalia berthelini from the
Aptian to Cenomanian of Cabinda, Cuanza and Benguela Basins (Araújo and
Guimarães 1992; Rocha 1984).
Molluscs of the Late Cretaceous count more than 240 known taxa, mostly ammo-
nites (see Supplementary Material for the full list). Some were named after Angolan
toponyms or after paleontologists that worked in Angola (Borges 1946; Carvalho
1961; Haas 1943; Howarth 1965; Cooper 1972, 1982; Cooper and Kennedy 1979):
Acera choffati Rennie 1945, Axonoceras angolanum Haas 1943, Didymoceras cf.
angolaense Haughton 1924 (Howarth 1965), Eutrephoceras egitoense Miller and
Carpenter 1956, Kitchinites angolaensis Howarth 1965, Libycoceras dandense
Howarth 1965, Lucina egitoensis Rennie 1945, L. angolensis Rennie 1929,
Mammites mocamedensis Howarth 1966, Nostoceras mariatheresianum Haas 1943,
Oiophyllites angolaensis Spath 1953, Prionocyclus carvalhoi Howarth 1966,
Protacanthoceras angolaense Spath 1931, Prohysteroceras hanhaense (Haas
1942), P. angolaense (Boule et al. 1907), Protocardia moutai Rennie 1945,
Pseudocalycoceras angolaense (Spath 1931), Pseudomelania salenasensis Rennie,
Pterotrigonia borgesi Rennie 1945, Mortoniceras (Angolaites) stolikzcai (Spath
1922), Mortoniceras (?) rochai Collignon 1978, M. (Deiradoceras) reali Collignon
1978, Collignoniceras (Selwynoceras) reali Collignon 1978, and Solenoceras
62 O. Mateus et al.
bembense Haas 1943. The ammonites are found in almost all Upper Cretaceous sec-
tions, including those of the Quissonde Formation from the beaches of Quimbala,
Chamure, Cabeça da Baleia, Egito in Benguela Basin, Teba, Bembe in Cuanza
Basin, Bentiaba and Salinas in Namibe, and Iembe in Bengo Province (Segundo
et al. 2014).
Rennie (1945) describes ten species of gastropods and bivalves from Cabeça da
Baleia, Egito-Praia: Trigonia (Scabrotrigonia) borgesi, Lucina egitoensis,
Protocardia moutai, Pseudomelania egitoensis, Confusiscala angolensis, Acirsa
(Plesioacirsa?) egitoensis, Dicroloma (Perissoptera) o’donnelli, Paleopsephaea
o’donnelli, Acera choffati, and Ringicula moutai (Lapão and Pereira 1971).
More than ten echinoderm taxa are known from the Late Cretaceous of Angola,
mostly echinoids, some received Angola-related names, such as the Toxasteridae
Epiaster angolensis Haughton 1924, collected 150 m below the Itombe Formation
at Zenza do Itombe, and E. carvalhoi Dartevelle 1953 (Haughton 1924; Kier and
Lawson 1978; Néraudeau and Mathey 2000), Leiostomaster angolanus Greyling
and Cooper 1995, Palaeostomatidae and Tholaster carvalhoi Greyling and Cooper
1995, and Holasteridae from the Middle Campanian of Egito Praia, near Quimbala.
Vertebrates
In Angola, a peak of vertebrate paleobiodiversity is observed in the Late Cretaceous,

with more than 100 taxa recognised (Antunes 1964; Antunes and Cappetta 2002;
Jacobs et al. 2006, 2009, 2016). This peak is in comparison with other time intervals
and is likely an artifact of the inadequacies of the fossil record rather than a state-
ment of biological reality. Most known Late Cretaceous sedimentary rocks from
Angola are marine, thus the fossil assemblage also reflects this ecosystem. Non-
marine organisms are occasionally drifted into a marine setting, including the sau-
ropod dinosaur Angolatitan adamastor Mateus et al. (2011) from Iembe, in Bengo
Province, originally considered Turonian in age but following the discovery of the
ammonite Protexanites sp. at Iembe, it is now considered Coniacian. Pterosaurs are
known from the Maastrichtian marine sediments of Bentiaba, in Namibe Province.
Angolatitan adamastor is the first dinosaur found in Angola. Its generic name means
‘Titan of Angola’ and the specific name refers to Adamastor made famous by Luís
de Camões in the Lusíadas. It was 13 m long and lived in an arid environment
(Mateus et al. 2011). A number of bones of large flying reptiles, pterosaurs, col-
lected in the marine sediments of the Maastrichtian of Bentiaba are the only know
pterosaur remains in sub-saharian Africa for this stage.
The fossil record of chondrichthyans shows a peak in Late Cretaceous deposits
with 64 species known, including taxa only known in Angola or with Angolan top-
onyms such as Angolabatis angolensis, A. benguelaensis, Chlamydoselachus graci-
lis, Cretascymnus quimbalaensis, and Echinorhinus lapaoi named by Antunes and
Cappetta (2002). The diversity of chondrichthyans includes representatives of most
of the Cretaceous lineages: Hexanchiformes, Squaliformes, Rajiformes, Orectolo-
biformes, Odontaspidida, Lamniformes, Carcharhiniformes, and Squaliformes
(Antunes and Cappetta 2002).
In contrast to the diversity of chondrichthyans, the Late Cretaceous bony ray-

finned fish from Angola are represented by ten taxa only, most likely due to the
difficulty of identification of isolated remains, in comparison with the abundance of
shark and ray teeth. The teleost genus Enchodus is known from various species
(Enchodus bursauxi, E. crenulatus, E. elegans, E. faujasi and E. libycus). Other
teleosts include Eodiaphyodus lerichei, Pseudoegertonia bebianoi, and Stephanodus
libycus from the Late Campanian and Maastrichtian in Benguela and Namibe prov-
inces (Antunes and Cappetta 2002) (Fig. 4.3).
Two Upper Cretaceous fossil localities from Angola are remarkably rich in large
reptiles: Iembe in Bengo Province (Fig. 4.4) of Coniacian age (~88 Ma), and
Bentiaba in Namibe (Maastrichtian). The Coniacian provided the bizarre duropha-
gous cryptodiran turtle Angolachelys mbaxi Mateus et al. (2009) that justified the
erection of its own clade Angolachelonia, and the sauropod dinosaur Angolatitan
adamastor. Indeterminate plesiosaur remains have also been recovered from this
region. Mosasaurs are represented by Angolasaurus bocagei (Fig. 4.3), Tylosaurus
iembeensis Antunes (1964) and an indeterminate halisaurine.
The richest vertebrate locality in Angola is near Bentiaba in Namibe province
(Strganac et al. 2014, 2015a, b). The reptile list includes the cheloniid turtles
Euclastes sp., Protostega sp. and Toxochelys sp. Terrestrial reptiles include isolated
remains that allow undetermined large pterosaurs, an undetermined sauropod based
on a metapodial, and a possible hadrosaur based on an isolated phalanx (Mateus
et al. 2012). The squamate mosasaurs are, by far, the most abundant and speciose
group of tetrapods, and include Globidens phosphaticus, Prognathodon kianda,
Halisaurus sp., Mosasaurus sp., Phosphorosaurus sp., and ‘Platecarpus’ ptychodon
(Polcyn et al. 2007, 2010, 2014; Schulp et al. 2006, 2008, 2013). The plesiosaur
Fig. 4.3 The mosasaur Angolasaurus bocagei skull and anterior postcrania (Museum of Geology,
Universidade Agostinho Neto). (Photo: Hillsman Jackson, Southern Methodist University)
64 O. Mateus et al.
Fig. 4.4 Reconstruction of the fauna during the Late Cretaceous, based on the Coniacian fauna of
Iembe. Illustration using acrylic painting with brush by Fabio Pastori
Elasmosauridae are also abundant at Bentiaba where two taxa are known:
Aristonectinae indet. and Cardiocorax mukulu (Araújo et al. 2015a, b).
The Paleogene – Mammals Take Over
The Paleogene Period begins at the end of the Cretaceous (66 Ma) and lasted until
the Neogene Period 23.03 Ma. The Paleogene is the interval of Earth’s history in
which mammals diversified and flourished after the K/Pg mass extinction, when
most large reptiles, and belemnites and ammonites went extinct. There is a dearth of
terrestrial vertebrates, especially mammals, from the Paleogene of Angola. During
the global Paleocene-Eocene Thermal Maximum (PETM), 55.8 million years ago,
there occurred a sudden change of the climate that marked the end of the Paleocene
and the beginning of the Eocene, one of the most significant periods of climate
change in the Cenozoic era (Zachos et al. 2005). The best known Paleogene geo-
logic section in Angola is that of Lândana in Cabinda Province. A recent study of
the Paleocene and Eocene biota and strata of Lândana indicated that the PETM is
missing from that section, either because the event was too short to be recorded at
the sampling interval used or that it falls within one of the several stratigraphic gaps
documented in the Lândana section (Solé et al. 2018).
Protista and Invertebrates
Among the protists, foraminifera stand out as the most important and well-known
taxonomic group in the Angolan marine series of Paleogene age, due to their impor-
tance for biostratigraphic correlations in offshore oil drilling and their equivalence
with landward outcrops. Important works include those of Rocha (1973) and Kender
et al. (2008), which include many Eocene and Oligocene characteristic taxa such as:
Cyclammina cf. compressa, Nonion centrosulcatum, Cassidulina subglobosa,
Globigerina ampliapertura, Bolivina cf. pygmaea, Bulimina alsatica, B. kackso-
nensis and B. nkomi.
In the Paleogene of Angola, molluscs remain the most speciose clade of inverte-
brates, comprising at least three nautiloids, 14 bivalves, and 21 gastropods (see
Supplementary Material). The bivalves are mainly known from the Eocene
(Lutetian) Quimbriz Formation along the Luculo River (Tavares et al. 2007) and
include Leda africae, Noetia veatchi, ‘Cardium’ luculensis, ‘C.’ sandigii, Crassatella
schoonoverae, Lucina cf. landanensis, Macrocallista palmerae, Metis olssoni, Pitar
quimbrizensis, P. quipayensis, Protonoetia nigeriensis, Raetomya schweinfurthi,
Venericardia angolae, and V. heroyi (Tavares et al. 2007). The three nautiloids are
Cimomia landanensis, Deltoidonautilus caheni, Hercoglossa diderrichi from the
Danian of Cabinda Basin, Landana (Soares 1965), and the gastropods are also
mainly known from the Eocene, Lutetian of Luculo River (Tavares et al. 2007):
Ficula roscheni, Fulguroficus harrisi, Pleurotoma angolae, P. rebeccae, Polinices
(Neverita) angolae, Ringicula hughesae, Sinum dusenberryi, Surcula cf. ingens,
and Turricula (Knefastia) angolensis.
Other groups such as Anthozoa, Arthropoda and Echinodermata, exist but are
low in numbers (Dartevelle 1953).
Vertebrates
Paleogene marine sediments have been studied mostly in the Benguela Basin and in
Cabinda. Adnet et al. (2009) recognized a new species of Eocene lamniform shark,
Xiphodolamia serrata, from Benguela. Sharks and bony fishes from Cabinda have
been listed by Solé et al. (2018), and Taverne (2016) has provided new information
about osteoglossid fishes from Lândana. The tetrapods from the Paleogene of
Angola are mostly from the fossil sites of Lândana (Solé et al. 2018) and Malembo
in Cabinda. The section begins with Lower Paleocene strata at Lândana. Reptiles
comprise the dyrosaurid crocodylomorph Congosaurus bequaerti, and indetermi-
nate crocodilians (Jouve and Schwarz 2004; Schwarz 2003; Schwarz et al. 2006),
the turtles Taphrosphys congolensis, a toxochelyid, and Cabindachelys landanensis
(Myers et al. 2017). Along the Chiloango River, Cabinda, a vertebra of the snake
Palaeophis was reported by Antunes (1964).
The youngest portion of the Cabinda section is at Malembo Point, south of
Lândana, which was originally considered Miocene. The Malembo mammal fauna
is comparable to the Early Oligocene fauna of the Fayum, Egypt because of the
66 O. Mateus et al.
presence of the embrithopod Arsinoitherium, hyracoids such as Geniohyus aff.

Mirus and Bunohyrax aff. Fajumensis, the proboscidean cf. Phiomia or
Hemimastodon, the sirenian Halitherium, and a reported anthropoid canine (Hooijer
1963; Pickford 1986; Jacobs et al. 2016). Recent findings by Projecto PaleoAngola
in Malembo include a ptolemaiidan molar more similar to Kelba from the Miocene
of Songhor, Kenya (19.5 Ma) than to Fayum Ptolemaia, and an isolated premolar
tooth of a large primate comparable in size to that of a female gorilla, likely an
undescribed taxon (Jacobs et al. 2016), and not representing any of the numerous
Fayum primate taxa. In addition, Arsinoitherium is now known from the latest
Oligocene of Kenya. The presence of a Kelba-like ptolemaiadan, a unique primate,
and Arsinoitherium may indicate a latest Oligocene or even earliest Miocene age for
Malembo Point. The assemblage certainly has differences from the Fatum fauna and
may indicate the presence of a lowland West Africa faunal province near the
Paleogene-Neogene boundary in age and distinct from other regions such as the
East African Rift Valley or the Fayum.
The Neogene – The Founding of Modern Biodiversity
The Neogene began about 23 Ma and extends until the Pleistocene (1.8 Ma). It is
divided into Miocene (23 Ma to 5.3 Ma) and Pliocene (5.3 Ma to 2.6 Ma), from the
oldest to the more recent. This period saw the expansion of the large mammals, and
the appearance of hominids. In the Miocene the climate warms again and grasslands
and savannas spread. In the Pliocene, the Earth had become similar to the one we
know today.
Protists and Invertebrates
The Neogene foraminifera of the Angolan coastal basins, including those from the
Miocene Quifandongo series of the Cuanza Basin, are known from a diversity of
planktonic and benthic taxa widely used in oil industry offshore correlations or as
palaeoenvironmental indicators. Rocha (1957), Graham et al. (1965), Mcmillan and
Fourie (1999) and Kender et al. (2009), among others describe the essentials of
these West African foraminiferal assemblages, which include planktonic taxa such
as Globigerina praebulloides, Globigerinella obesa, Globigerinoides bisphericus,
G. immaturus, G. trilobus, Globorotalia peripheroronda and Orbulina bilobata.
Invertebrates are surprisingly poorly known and comprise, at least, the nautiloid
mollusc Aturia luculoensis, the decapod crustacean Callianassa floridana from the
Miocene Burdigalian of Cabinda Basin (Newton 1917), several taxa of Miocene
echinoid echinoderms such as Clypeaster borgesi, Echinolampas antunesi, Rotula
deciesdigitata, Rotuloidea vieirai, Amphiope neuparthi, and Plagiobrissus sp.
(Loriol 1905; Dartevelle 1953; Gonçalves 1971; Kroh 2010; Silva and Pereira 2014;
Pereira and Stara 2018), and two species of anthozoan corals Flabellum extensum
and Stylophora raristella (Chevalier 1970). Nevertheless, bivalve and gastropod

molluscs are undoubtedly the most diverse and abundant invertebrate taxa of the
marine Neogene of Luanda, Benguela and Namibe, with several rich fossil sites,
some of them presently in course of study. The molluscan faunas of these coastal
basin areas, including new species such as Pereiraea africana, Clavatula loandensis
or Chlamys silvai Antunes (1964), were the focus of Douvillé (1933), Keller (1934),
Dartevelle (1952, 1953), Dartevelle and Roger (1954), Soares (1961, 1962), Silva
(1962), Silva and Soares (1962), Antunes (1964), and more recently Antunes
(1984), Lozouet and Gourgues (1995), among others.
Vertebrates
Mammals are known from Benguela and the Cuanza provinces where Projecto
PaleoAngola collected skulls of fossil baleen whales. An odontocete has also been
found from Barra da Cuanza.
In Angola the most abundant group of Neogene vertebrates are the
Elasmobranchii chondrichthyans (18 taxa; see Supplementary Material). The
following taxa are from the Pliocene of Farol das Lagostas (Cuanza Basin):
Aetobatus, Carcharhinus egertoni, Carcharhinus priscus, Carcharias taurus,
Carcharocles megalodon, Carcharodon carcharias, Galeocerdo cuvier,
Hemipristis serra, Isurus benedeni, Isurus oxyrinchus, Mitsukurina, Myliobatis,
Negaprion brevirostris, Paragaleus, Pristis, Pteromylaeus bovina, Rhinoptera
brasiliensis, and Sphyrna zygaena (Antunes 1964). Five bony fishes are known,
the actinopterygians Cybium, Sparus, Sphyraena barracuda, Tachysurus, and
Tetrodon.
The Quaternary – The Dominance of Humans
The Quaternary (2.6 Ma to present, including the Pleistocene and Holocene) is the
third geological period of the Cenozoic era and the most recent in the geological
time scale. This period is characterised by the return of glaciations at higher eleva-
tions and latitudes, the dominant role of the genus Homo in all terrestrial habitats
and the extinction of much of the megafauna.
Invertebrates
In Angola, the Quaternary biodiversity is again marked by the high number of mol-
lusc taxa (73 or more), of which 29 are bivalves such as Arcopsis afra, Barbatia
complanata, Cardium indicum, Chama crenulata, Glycymeris concentrica, Lutraria
senegalensis, Noetiella congoensis, Ungulina cuneata from the Middle Pleistocene
of Pipas (Namibe Basin) and 44 gastropods mainly known from Namibe Basin,
such as Cantharus viverratus, Columbella adansoni, Conus babaensis, Siphonaria
68 O. Mateus et al.
capensis, and Terebra senegalensis (Miller and Carpenter 1956; Sessa et al. 2013).
Other invertebrates such as corals, arthropods and echinoderms are known but
reduced to a handful of known taxa, such as Cladangia carvalhoi from the
Pleistocene of Salinas de Bero, Saco, Namibe (Wood 1973). In most situations they
occur in a variety of raised-beach and lagoonal deposits related to coastal uplift and
major sea-level changes (Carvalho 1961). The post-glacial Holocene is marked by
the accretion of sand-spits and deltaic facies with rich coquinas, including the
bivalve Senilia senilis as a typical species (Dinis et al. 2016). See Supplementary
Material for the updated list.
Vertebrates
A remarkable fossilized jaw (dentary bone) of Blue Whale Balaenoptera musculus

present in the Museu Nacional de História Natural in Luanda measuring nearly
seven meters in length is not only the largest known fossilized bone but also one of
the largest whales, thus animals, ever recorded. Large land mammals, including
Bubalus, Syncerus cf. nanus Boddaert; Phacochoerus sp., Equus, Hippotigris cf.
zebra have been reported from the site called Cemitério dos Ossos, north of Luanda
(Antunes 1961).
The caves of Humpata, in Huíla Province, in southern Angola, are formed in
Chela Dolomite that hosts fossiliferous caves and fissures (Amaral 1973; Antunes
1965; Arambourg and Mouta 1952; França 1964; Mouta 1950). Pickford et al.
(1990, 1992, 1994) listed taxa of mammals from the Humpata Caves. These include
the insectivore Crocidura, a Macroscelididae, the chiropteran Rhinolophus,
Miniopterus, Nycteris, 19 genera of rodents (Uranomys, Acomys, Dasymys,
Aethomys, Thallomys, Zelotomys, Mus, Pelomys, Malacomys, Praomys,
Grammomys, Dendromus, Steatomys, Petromyscus, Tatera, Otomys, Cryptomys,
Graphiurus, and Hystrix), the lagomorph Serengetilagus, Mustelidae, Viverridae,
Canidae, and the Hyaenidae cf. Chasmoporthetes, the Hyracoidea Gigantohyrax
and Procavia, Rhinocerotidae, Equidae, Suidae Metridiochoerus andrewsi and the
Bovidae Hippotragini and Connochaetes.
The most thoroughly studied of the Humpata fossils are those of the extinct
baboon. Cercopithecid primates from Humpata caves include Soromandrillus qua-
dratirostris, cf. Theropithecus sp., and Cercopithecoides sp. (dated as ca. 2.0–
3.0 Ma) (Minkoff 1972; Jablonski 1994; Jablonski and Frost 2010; Gilbert 2013).
Pleistocene deposits in Namibe provided remains of fossilized ostrich Struthio
eggshells, artiodactyl bones and numerous human artifacts - including Acheulean
hand axes (amygdaloid bifaces) suggesting the presence of early humans, such as
Homo ergaster or H. erectus, whereas bones of early humans are not known in
Angola.
Final Remarks on the Fossil Record and Paleobiodiversity
Measuring paleobiodiversity is challenging due to the sparse and limited availabil-

ity of data, compared with modern extant faunas. The paleobiodiversity of Angola
is mostly known from Cretaceous and Cenozoic fossils that comprise 90% or more
of all known fossil records of Angolan taxa (see Table 4.1 and Supplementary
Material). The vast preponderance of the fossil taxa is marine which is consistent
with the geological settings and paleogeography, related to the opening of the South
Atlantic and repeated inundation of the Angolan continental margin.
For this study, we compiled a list of taxa using species, genus or the lowest
known taxonomical clade reported in the scientific literature for Angola (see
Supplementary Material). Of the resulting list of more than 1300 fossil taxa, many
may require systematic revision and the final number will depend on the validity of
the taxonomy.
By far the most speciose group are the molluscs (about 61% of taxa, more than
half being Cretaceous ammonites) and Cretaceous foraminiferans (16%), followed
by vertebrates with about 15% of taxa. Chondrichthyes and mammals represent 6%
and 5% of taxa, respectively.
About 10% of vertebrate taxa listed are unique or were first recognised in Angola,
most of them receiving species names after localities in Angola or of geologists that
worked in the country. According to current knowledge, at least 67 taxa (6.1%) of
invertebrates are endemic or were first mentioned from Angola.
References
Adnet S, Hosseinzadeh R, Antunes MT et al (2009) Review of the enigmatic Eocene shark genus
Xiphodolamia (Chondrichthyes, Lamniformes) and description of a new species recovered
from Angola, Iran and Jordan. J Afr Earth Sci 55(3–4):197–204
Amaral L (1973) Nota sobre o “karst” ou carso do Planalto do Humpata (Huila), no Sudoeste de
Angola. Garcia de Orta 1:29–36
Anchieta J (1885) Traços geológicos da África Occidental Portuguesa. Tipografia Progresso,
Benguela, 15 pp
Antunes MT (1961) A jazida de vertebrados fósseis do Farol das Lagostas: II Paleontologia.
Boletim dos Serviços de Geologia e Minas de Angola 3:1–18
Antunes MT (1964) O Neocretácico e o Cenozóico do litoral de Angola. Junta de Investigações
Ultramar, Lisboa, 255 pp
Antunes MT (1965) Sur la faune de vertébrés du Pléistocène de Leba, Humpata (Angola). Actes du
Ve Congrès Panafricain de Préhistoire et de l’Étude du Quaternaire. Tenerife, 127–128
Antunes MT (1970) Paleontologia de Angola. In: Curso de Geologia do Ultramar. Junta de
Investigações do Ultramar, Lisboa, pp 126–143
Antunes MT (1984) Étude d’une faune gastéropodes miocène récoltés par M. M Feio dans le Sud
de l’Angola Comunicações dos Serviços Geológicos de Portugal 70(1):126–128
Antunes MT, Cappetta H (2002) Sélaciens du Crétacé (Albien-Maastrichtien) d’Angola.
Palaeontographica, Abteilung A 264 (5–6):85–146
Antunes MT, Maisey JG, Marques MM, et al (1990) Triassic fishes from the Cassange depression
(R.P. de Angola). Ciências da Terra (UNL), special number 1:1–64
70 O. Mateus et al.
Arambourg C, Mouta F (1952) Les grottes et fentes à ossements du sud de l’Angola. Actes du
IIème Congrès Panafricain de Préhistoire d’Alger 12:301–301
Araújo AG, Guimarães F (1992) Geologia de Angola, Notícia explicativa da Carta Geológica à
escala 1: 1 000 000. Serviço Geológico de Angola, Luanda, 140 pp
Araújo R, Polcyn MJ, Lindgren J et al (2015a) New aristonectine elasmosaurid plesiosaur speci-
mens from the Early Maastrichtian of Angola and comments on paedomorphism in plesiosaurs.
Neth J Geosci 94(1):93–108
Araújo R, Polcyn MJ, Schulp AS et al (2015b) A new elasmosaurid from the early Maastrichtian
of Angola and the implications of girdle morphology on swimming style in plesiosaurs. Neth
J Geosci 94(1):109–120
Archibald JD, Clemens WA, Padian K et al (2010) Cretaceous extinctions: multiple causes.
Science 328(5981):973–973
Berry CT (1939) A summary of the fossil Crustacea of the Order Stomatopoda, and a description
of a new species from Angola. Am Midl Nat 21(2):461–471
Blake DB, Breton G, Gofas S (1996) A new genus and species of Asteriidae (Asteroidea;
Echinodermata) from the Upper Cretaceous (Coniacian) of Angola, Africa. Paläontol Z
70(1–2):181–187
Borges A (1946) A costa de Angola da Baía da Lucira à Foz do Bentiaba (entre Benguela e
Mossâmedes). Boletim da Sociedade Geológica de Portugal 5(3):141–150
Boule M, Lemoine P, Thevenin A (1907) Paléontologie de Madagascar. III Céphalopodes crétacés
des environs de Diègo-Suarez. Annales de Paléontologie 2:1–56
Brandão JM (2008) “Missão Geológica de Angola”: contextos e emergência. Memórias e Notícias,
new series 3:285–292
Brandão JM (2010) O “Museu de Geologia Colonial” das Comissões Geológicas de Portugal:
contexto e memória. Revista Brasileira de História da Ciência 3(2):184–199
Buta-Neto A, Tavares TS, Quesne D et al (2006) Synthèse préliminaire des travaux menés sur le
bassin de Benguela (Sud Angola): implications sédimentologiques et structurales. Áfr Geosci
Rev 13(3):239–250
Callapez PM, Gomes CR, Serrano Pinto M et al (2011) O contributo do Museu e Laboratório
Mineralógico e Geológico da Universidade de Coimbra para os estudos de Paleontologia
Africana. In: Neves LF, Pereira AC, Gomes CR, Pereira LCG, Tavares AO (eds) Modelação de
Sistemas Geológicos. Homenagem ao Professor Doutor Manuel Maria Godinho. Laboratório
de Radioactividade Natural da Universidade de Coimbra, Coimbra, pp 159–174
Carvalho GS (1961) Geologia do deserto de Moc̦âmedes, (Angola): Uma contribuição para
o conhecimento dos problemas da orla sedimentar de Moc̦âmedes. Memórias da Junta de
Investigações do Ultramar 26:1–227
Ceraldi TS, Green D (2016) Evolution of the South Atlantic lacustrine deposits in response to
Early Cretaceous rifting, subsidence and lake hydrology. In: Ceraldi S, Hodgkinson RA, Backe
G (eds) Petroleum geoscience of the West Africa Margin. Geological Society, London, Special
Publications, 438, doi: https://doi.org/10.1144/SP438.10
Chevalier JP (1970) Les Madreporaires du Neogene et du Quaternaire de l’Angola [Neogene
and Quaternary corals from Angola]. Annalen Koninklijk Museum voor Midden-Afrika 8:
Geologische Wetenschappen 68:13–33
Collignon M (1978) Ammonites du Crétacé Moyen-Supérieur de l’Angola. 2° Centenário
Academia das Ciências. Estudos de Geologia e Paleontologia e de Micologia, Academia das
Ciências, Lisboa, pp 1–75
Cooper MR (1972) The Cretaceous stratigraphy of San Nicolau and Salinas, Angola. Ann S Af
Mus 6(8):245–251
Cooper MR (1982) Lower Cretaceous (Middle Albian) ammonites from Dombe Grande, Angola.
Ann S Af Mus 89:265–314
Cooper MR, Kennedy WJ (1979) Upper most Albian (Stoliczkaia dispar zone) ammonites from
the Angolan littoral. Ann S Af Mus 77:175–308
Crampton JS, Beu AG, Cooper RA et al (2003) Estimating the rock volume bias in paleobiodiver-
sity studies. Science 301(5631):358–360
Dartevelle E (1952) Echinides fossiles du Congo et de l’Angola. Partie 1: Introduction historique
et stratigraphique. Annales du Musée Royal du Congo Belge, série 8, Sciences Géologiques
12:1–70
Dartevelle E (1953) Echinides fossiles du Congo et de l’Angola. Partie 2: description systématique
des échinides fossiles du Congo et de l’Angola. Annales du Muséum Royal du Congo Belge,
série 8, Sciences Géologiques 13:1–240
Dartevelle E, Roger J (1954) Contribution à la connaissance de la faune du Miocène de l’Angola.
Comunicações dos Serviços Gelógicos de Portugal 35:227–312
Dinis P, Huvi J, Cascalho J, Garzanti E, Vermeesch P, Callapez P (2016) Sand-spits systems from
Benguela region (SW Angola). An analysis of sediment sources and dispersal from textural and
compositional data. J Afr Stud 117:181–192
Douvillé H (1933) Le Tertiaire de Loanda. Boletim do Museo e Laboratorio Mineralógico e
Geológico da Universidade de Lisboa, 1:63–118
Duarte LV, Callapez PM, Kalukembe A, et al (2014) Do Proterozoico da Serra da Leba (Planalto
da Humpata) ao Cretácico da Bacia de Benguela (Angola). A geologia de lugares com elevado
valor paisagístico. Comunicações Geológicas 101(Especial III):1255–1259
Ferré B, Granier B (2001) Albian roveacrinids from the southern Congo Basin off Angola. J S Am
Earth Sci 14:219–235
Ferreira JM, Rocha AT (1957) Foraminíferos do Senoniano de Catumbela (Angola). Garcia de
Orta 5(3):517–545
Ferreira OV (1957) Acerca de Parapirimela angolensis Van Straelen nas Camadas de Iela, Angola.
Comunicações dos Serviços Geológicos de Portugal 38:465–468
França JC (1964) Nota preliminar sobre uma gruta pré-histórica do Planalto da Humpata (Angola).
Junta de Investigações do Ultramar 2(50):59–67
Gilbert CC (2013) Cladistic analysis of extant and fossil African papionins using craniodental data.
J Hum Evol 64(5):399–433
Gonçalves F (1971) Echinolampas antunesi, nov. sp. Cassidulidae, échinide nouveau du Miocène
de la région de Luanda, Angola. Revista da Faculdade de Ciências, C – Ciências Naturais
16(2):307–310
Graham JJ, Klasz I, Rerat D (1965) Quelques importants foraminifères du Tertiaire du Gabon
(Afrique Equatoriale). Revue de Micropalèontologie 8:71–84
Green PF, Machado V (2015) Pre-rift and synrift exhumation, post-rift subsidence and exhuma-
tion of the onshore Namib1e margin of Angola revealed from apatite fission track analysis. In:
Sabato Ceraldi T, Hodgkinson RA, Backe G (eds) Petroleum geoscience of the West Africa
Margin. Geological Society, London, Special Publications 438, pp 99–118
Greyling MR, Cooper MR (1995) Two new irregular echinoids from the Upper Cretaceous (mid-
Campanian) of Angola. Durban Museum Novitates 20(1):63–71
Guiraud R, Maurin JC (1991) Le rifting en Afrique au Crétacé Inférieur: Synthèse structural, mise
en évidence de deux phases dans la genèse des bassins, relations avec les ouvertures océaniques
péri-africaines. Bulletin de la Société Géologique de France 165(5):811–823
Guiraud M, Buta-Neto A, Quesne D (2010) Segmentation and differential post-rift uplift at the
Angola margin as recorded by the transform – rifted Benguela and oblique-to-orthogonal-rifted
Kwanza basins. Mar Pet Geol 27:1040–1068
Haas O (1942) The vernay collection of cretaceous (Albian) ammonites from Angola. Bull Am
Mus Nat Hist 81(1):1–224
Haas O (1943) Some abnormally coiled ammonites from the Upper Cretaceous of Angola. Am
Mus Novit 1222:1–17
Haughton SH (1924) Notes sur quelques fossiles crétacés de l’Angola (Céphalopodes et Échinides).
Comunicações dos Serviços Geológicos de Portugal 15:79–106
72 O. Mateus et al.
Hooijer DA (1963) Miocene mammals of Congo. Annales du Museum Royal d’Afrique Centrale,
Series 8 Sciences Géologiques 46:1–77
Howarth MK (1965) Cretaceous ammonites and nautiloids from Angola. Bull Br Mus Nat Hist
Geol 10:335–412
Howarth MK (1966) A mid-Turonian ammonite fauna from the Moçâmedes desert, Angola. Garcia
de Orta 14(2):217–228
Jablonski NG (1994) New fossil cercopithecid remains from the Humpata Plateau, southern
Angola. Am J Phys Anthropol 94(4):435–464
Jablonski NG, Frost S (2010) Cercopithecoidea. In: Werdelin L, Sanders WJ (eds) Cenozoic
Mammals of Africa. University of California Press, Berkeley, pp 393–428
Jackson JB, Johnson KG (2001) Measuring past biodiversity. Science 293(5539):2401–2404
Jacobs LL, Mateus O, Polcyn MJ et al (2006) The occurrence and geological setting of Cretaceous
dinosaurs, mosasaurs, plesiosaurs, and turtles from Angola. J Paleontol Soc Korea 22:91–110
Jacobs LL, Mateus O, Polcyn MJ et al (2009) Cretaceous paleogeography, paleoclimatology, and
amniote biogeography of the low and mid-latitude South Atlantic Ocean. Bull Geol Soc Fr
180(4):333–341
Jacobs LL, Polcyn MJ, Mateus O et al (2016) Post-Gondwana Africa and the vertebrate history of
the Angolan Atlantic Coast. Mem Mus Vic 74:343–362
Jones TR (1862) A Monograph of the fossil Estheriae. Palaeontol Soc 14:1–134
Jouve S, Schwarz D (2004) Congosaurus bequaerti, a Paleocene dyrosaurid (Crocodyliformes;
Mesoeucrocodylia) from Landana (Angola). Bulletin de l’Institut Royal des Sciences Naturelles
de Belgique, Sciences de la Terre 74:129–146
Keller A (1934) Contribution a la géologie de l’Angola. Le Tertiaire dc Luanda. Description
des especes, Mollusque. Lamellibranches. Boletim do Museo e Lab. Mineral. e Geol. do
Universidade de Lisboa, la Ser, 3, pp 219–250
Kender S., Kaminski MA, Jones, BW (2008) Oligocene deep-water agglutinated foraminifera
from the Congo Fan, offshore Angola: Palaeoenvironments and assemblage distributions. In:
Kaminski MA, Coccioni R (eds) Proceedings of the seventh international workshop on aggluti-
nated foraminifera. Grzybowski Foundation Special Publication 13, London, pp 107–156
Kender S, Kaminski MA, Jones BW (2009) Early to middle Miocene foraminifera from the deep-
sea Congo Fan, offshore Angola. Micropaleontology 54(6):477–568
Kier PM, Lawson M (1978) Index of living and fossil echinoids, 1924-1970. Smithson Contrib
Paleobiol 34:1–182
Kroh A (2010) Index of Living and Fossil Echinoids 1971-2008. Annalen des Naturhistorischen
Museums in Wien 112:195–470
Lapão LGP, Pereira ES (1971) Notícia explicativa Carta Geológica de Angola, escala 1:100 000,
folha n° 206, Egito Praia. Direcção Provincial dos Serviços de Geologia e Minas, Luanda,
42 pp
Lapão LGP, Simões MC (1972) Notícia Explicativa da Carta Geológica de Angola, escala 1:100
000, Folha n°184, Novo Redondo. Direcção Provincial dos Serviços de Geologia e Minas,
Luanda, 54 pp
Leriche M (1932) Sur les premiers fossiles découverts au nord de l’Angola, dans le prolongement
des couches du Lubilash et des couches du Lualaba. Association Française pour l’Avancement
des Sciences, Compte Rendu 56éme session: 1–6
Loriol P (1888) Matériaux pour l’étude stratigraphique et paléontologique de la province d’Angola.
Description des Echinides. Mémoires de la Société de Physique et d’histoire naturelle de
Genève 30(2):97–114
Loriol P (1905) Notes pour servir à l’étude des échinodermes. Fasc. 2 (3). Georg Editeur, Bâle/
Genève, pp 119–146
Lozouet P, Gourgues D (1995) Senilia (Bivalvia: Arcidae) et Anazola (Gastropoda: Olividae) dans
le Miocène d’Angola et de France, témoins d’une paléo-province Ouest-Africaine. Haliotis,
24, pp 101–108
Marliére R (1950) Ostracodes and Phyllopodes au Système du Karroo au Congo Belge et les
régions avoisinantes. Annales du Muséum Royal du Congo Belge, Sciences Géologiques, Série
8(6):1–43
Marzola M, Mateus O, Schulp A, et al (2014) Early Cretaceous tracks of a large mammalia-
morph, a crocodylomorph, and dinosaurs from an Angolan diamond mine. J Vertebr Paleontol,
Program and Abstracts, 181
Masse P, Laurent O (2016) Geological exploration of Angola from Sumbe to Namibe: a review
at the frontier between Geology, natural resources and the history of Geology. C R Geosci
348(1):80–88
Mateus O, Jacobs LL, Polcyn MJ et al (2009) The oldest African eucryptodiran turtle from the
Cretaceous of Angola. Acta Paleontol Pol 54:581–588
and the first record from Angola. Anais da Academia Brasileira de Ciências 83(1):221–233
Mateus O, Polcyn MJ, Jacobs LL, et al (2012) Cretaceous amniotes from Angola: dinosaurs, ptero-
saurs, mosasaurs, plesiosaurs, and turtles. V Jornadas Internacionales sobre Paleontología de
Dinosaurios y su Entorno, Salas de los Infantes, Burgos, pp 75–105
Mateus O, Marzola M, Schulp AS et al (2017) Angolan ichnosite in a diamond mine shows the
presence of a large terrestrial mammaliamorph, a crocodylomorph, and sauropod dinosaurs in
the Early Cretaceous of Africa. Palaeogeogr Palaeoclimatol Palaeoecol 471:220–232
Mcmillan IK, Fourie A (1999) Kwanza Basin coastal stratigraphy with atlas of Albian to Holocene
Foraminifera species. De Beers Marine (Pty) Limited, Luanda, 167 pp
Miller AK, Carpenter LB (1956) Cretaceous and Tertiary Nautiloids from Angola. Estudos,
Ensaios e Documentos da Junta de Investigações do Ultramar 21:1–48
Minkoff EC (1972) A fossil baboon from Angola, with a note on Australopithecus. J Paleontol
46(6):836–844
Mouta F (1950) Sur la présence du Quaternaire ancien dans les hauts plateaux du Sud de l’Angola
(Humpata, Leba). Compte Rendu sommaire des Scéances de la Société géologique de France
14:261–262
Murray AM (2000) The palaeozoic, mesozoic and early cenozoic fishes of Africa. Fish Fish
1(2):111–145
Myers TS, Polcyn MJ, Mateus O et al (2017) A new durophagous stem cheloniid turtle from the
lower Paleocene of Cabinda, Angola. Pap Palaeontol 2017:1–16
Néraudeau D, Mathey B (2000) Biogeography and diversity of South Atlantic Cretaceous echinoids:
implications for circulation patterns. Palaeogeogr Palaeoclimatol Palaeoecol 156(1–2):71–88
Neto MGM (1970) O sedimentar costeiro de Angola. Algumas notas sobre o estado actual do seu
conhecimento. In: Curso de Geologia do Ultramar, vol 2. Publicações da Junta de Investigações
do Ultramar, Lisboa, pp 193–232
Newton RB (1917) On some Cretaceous Brachiopoda and Mollusca from Angola, Portuguese
West Africa. Earth and environmental science. Trans R Soc Edinb 51(3):561–580
Nunes AF (1991) A Investigação Geológico-Mineira em Angola. Ministérios dos Negócios
Estrangeiros, Ministério das Finanças, Instituto para a Cooperação Económica, Lisboa, 387 pp
Pereira P, Stara P (2018) Redefinition of Amphiope neuparthi de Loriol, 1905 (Echinoidea,
Astriclypeidae) from the early-middle Miocene of Angola. Comunicações Geológicas, 104(1):
in press
Pérez-Díaz L, Eagles G (2017) South Atlantic paleobathymetry since early Cretaceous. Sci Rep
7:11819. https://doi.org/10.1038/s41598-017-11959-7
Pickford M (1986) Première découverte d’une faune mammalienne terrestre paléogène d’Afrique
sub-saharienne. Comptes rendus de l’Académie des Sciences Paris Série II 19:1205–1210
Pickford M, Fernandez T, Aço S (1990) Nouvelles découvertes de remplissages de fissures à pri-
mates dans le ‘Planalto da Humpata’, Huilà, Sud de l’Angola. Comptes Rendus de l’Académie
des Sciences Paris, Série II 310:843–848
74 O. Mateus et al.
Pickford M, Mein P, Senut B (1992) Primate-bearing Plio-Pleistocene cave deposits of Humpata,

southern Angola. Hum Evol 7:17–33
Pickford M, Mein P, Senut B (1994) Fossiliferous Neogene karst fillings in Angola, Botswana and
Namibia. S Afr J Sci 90:227–230
Polcyn M, Jacobs L, Schulp A, et al (2007) Halisaurus (Squamata: Mosasauridae) from the
Maastrichtian of Angola. J Vertebr Paleontol 27(Suppl. to 3):130A
Polcyn MJ, Jacobs LL, Schulp AS et al (2010) The North African Mosasaur Globidens phosphati-
cus from the Maastrichtian of Angola. Hist Biol 22(1–3):175–185
Polcyn MJ, Jacobs LL, Araújo R et al (2014) Physical drivers of mosasaur evolution. Palaeogeogr
Palaeoclimatol Palaeoecol 400:17–27
Quirk DG, Hertle M, Jeppesen JW, et al (2013) Rifting, subsidence and continental break-up above
a mantle plume in the central South Atlantic. In: Mohriak WU, Danforth A, Post PJ, et al (eds)
Conjugate Divergent Margins. Geological Society, London, Special Publication, 369: 185–214
Rennie JVL (1929) Cretaceous fossils from Angola (Lamellibranchia and Gastropoda). Ann S Afr
Mus 28:1–54
Rennie JVL (1945) Lamelibrânquios e gastrópodes do Cretácico Superior de Angola (vol. 1). Junta
das Missões Geográficas e de Investigações Coloniais 1:1–141
Robles-Cruz SE, Escayola M, Jackson S et al (2012) U-Pb SHRIMP geochronology of zircon
from the Catoca kimberlite, Angola: Implications for diamond exploration. Chem Geol 310-
311:137–147. https://doi.org/10.1016/j.chemgeo.2012.04.001
Rocha AT (1957) Contribuição para o estudo dos foraminíferos do Terciário de Luanda. Garcia de
Orta 5(2):297–312
Rocha AT (1973) Contribution à l’étude des foraminifères paléogènes du bassin du Cuanza
(Angola). Memórias e Trabalhos do Instituto de Investigação Científica de Angola 12:1–309
Rocha AT (1984) Notas micropaleontológicas sobre as formações sedimentares da orla meso-
cenozóica de Angola - V. O Maestrichtiano inferior da mancha de Cabeça da Baleia (a norte de
Egito-Praia). Garcia de Orta, Série Geológica 7(1–2):97–108
Schlanger SO, Arthur MA, Jenkyns HC et al (1987) The Cenomanian-Turonian Oceanic Anoxic
Event, I. Stratigraphy and distribution of organic carbon-rich beds and the marine δ13C excur-
sion. Geol Soc Lond, Spec Publ 26(1):371–399
Schulp AS, Polcyn MJ, Mateus O et al (2006) New mosasaur material from the Maastrichtian
of Angola, with notes on the phylogeny, distribution and palaeoecology of the genus
Prognathodon. On Maastricht Mosasaurs. Publicaties van het Natuurhistorisch Genootschap
in Limburg 45(1):57–67
Schulp AS, Polcyn MJ, Mateus O et al (2008) A new species of Prognathodon (Squamata,
Mosasauridae) from the Maastrichtian of Angola, and the affinities of the mosasaur genus
Liodon. In: Proceedings of the Second Mosasaur Meeting, vol 3. Fort Hays State University
Hays, Kansas, pp 1–12
Schulp AS, Polcyn MJ, Mateus O et al (2013) Two rare mosasaurs from the Maastrichtian of
Angola and the Netherlands. Neth J Geosci 92(1):3–10
Schwarz D (2003) A procoelous crocodilian vertebra from the lower Tertiary of Central Africa
(Cabinda enclave, Angola). Neues Jahrbuch für Geologie und Paläontologie, Monatshefte
2003:376–384
Schwarz D, Frey E, Martin T (2006) The postcranial skeleton of the Hyposaurinae (Dyrosauridae;
Crocodyliformes). Palaeontology 49(4):695–718
Segundo J, Duarte LV, Callapez PM (2014) Lithostratigraphy of the Quissonde Formation marl-
limestone succession (Albian) of the Ponta do Jomba-Praia do Binge sector (Benguela Basin,
Angola). Comunicações Geológicas 101(Especial III):567–571
Séranne M, Anka Z (2005) South Atlantic continental margins of Africa: a comparison of the tec-
tonic vs climate interplay on the evolution of equatorial west Africa and SW Africa margins.
J Afr Earth Sci 43(1–3):283–300
Sessa JA, Callapez PM, Dinis PA et al (2013) Paleoenvironmental and paleobiogeographical
implications of a Middle Pleistocene mollusc assemblage from the marine terraces of Baía das
Pipas, southwest Angola. J Paleontol 87(6):1016–1079
Silva GH (1962) Fósseis do Miocénico de Luanda (Angola). Associação Portuguesa para o
Progresso das Ciências. Actas do XXVI Congresso Luso-Espanhol (Porto, 22–26 June, 1962),
sections II and IV, 3 pp
Silva R, Geirinhas F (2010) Colecções geológicas das antigas Províncias Ultramarinas Portuguesas
arquivadas na Litoteca do LNEG. e-Terra 15(4):1–4
Silva R, Pereira P (2014) Redescoberta dos equinodermes fósseis das coleções históricas ultrama-
rinas do LNEG. Comunicações Geológicas 101(Especial III):1379–1382
Silva GH, Soares AF (1962) Contribuição para o conhecimento da fauna miocénica de S. Pedro da
Barra e do Farol das Lagostas (Luanda, Angola). Garcia de Orta 9:721–736
Soares AF (1961) Nouvelle espèce de Chlamys du Miocène de la région de Luanda (Angola).
Memórias e Notícias 51:1–6
Soares AF (1962) Nota sobre alguns lamelibrânquios e gastrópodes do Miocénico de Luanda
(Angola). Memórias e Notícias 53:31.35
Soares AF (1965) Contribuição para o estudo dos lamelibrânquios Cretácicos da região de
Moçâmedes. Serviços de Geologia e Mina de Angola Boletim 11:137–168
Solé F, Noiret C, Desmares D et al (2018) Reassessment of historical sections from the Paleogene
marine margin of the Congo Basin reveals an almost complete absence of Danian deposits.
Geosci Front. https://doi.org/10.1016/j.gsf.2018.06.002
Spath LF (1922) On Cretaceous ammonites from Angola, collected by Prof. J.W. Gregory, D.Sc.,
F.R.S. Trans R Soc Edinb Earth Sci 53:91–160
Spath LF (1931) On cretaceous Ammonoidea from Angola, collected by Pr. J.W. Gregory. Trans
Geol Soc Edinb. 53 (1):91–160
Spath LF (1953) The Upper Cretaceous cephalopod fauna of Grahamland. Sci Rep Falkland
Islands Depend Surv 3:1–60
Strganac C, Salminen J, Jacobs LL et al (2014) Carbon isotope stratigraphy, magnetostratigraphy,
and 40Ar/39Ar age of the Cretaceous South Atlantic coast, Namibe Basin, Angola. J Afr Earth
Sci 99:452–462
Strganac C, Jacobs LL, Polcyn M et al (2015a) Stable oxygen isotope chemostratigraphy and
paleotemperature regime of mosasaurs at Bentiaba, Angola. Neth J Geosci 94(1):137–143
Strganac C, Jacobs LL, Polcyn MJ et al (2015b) Geological setting and paleoecology of the
Upper Cretaceous Bench 19 marine vertebrate bonebed at Bentiaba, Angola. Neth J Geosci
94(1):121–136
Tasch P (1979) Permian and Triassic Conchostraca from the Bowen Basin (with a note on a
Carboniferous leaiid from the Drummond Basin), Queensland. Aust Bureau Mineral Resour
Geolo Geophys Bull 185:33–44
Tasch P (1987) Fossil Conchostraca of the Southern Hemisphere and continental drift: Paleontology,
biostratigraphy, and dispersal. Memoir of the Geological Society of America 165:1–290
Tavares T (2006) Ammonites et échinides de l’Albien du bassin de Benguela (Angola).
Systématique, biostratigraphie, paléogéographie et paléoenvironnement. Unpublished PhD
Thesis, Université de Bourgogne, Dijon, 389 pp
Tavares T, Meister C, Duarte-Morais ML et al (2007) Albian ammonites of the Benguela Basin
(Angola): a biostratigraphic framework. S Afr J Geol 110(1):137–156
Taverne L (2016) New data on the osteoglossid fishes (Teleostei, Osteoglossiformes) from
the marine Danian (Paleocene) of Landana (Cabinda Enclave, Angola). Geo-Eco-Trop
40(4):297–304
Teixeira C (1947) Contribuição para o conhecimento geológico do Karroo da África Portuguesa.
I-Sobre a flora fóssil do Karroo da região de Téte. Anais da Junta de Investigações do Ultramar,
Estudos de Geologia e Paleontologia 2(2):9–28
Teixeira C (1948a) Fósseis vegetais do Karroo de Angola. Boletim da Sociedade Geológica de
Portugal 7(1–2):73–77
76 O. Mateus et al.
Teixeira C (1948b) Vegetais fósseis do grés do Quilungo. Anais da Junta de Investigações Coloniais
2:85–92
Teixeira C (1958) Note paléontologique sur le Karroo de la Lunda, Angola. Boletim da Sociedade
Geológica de Portugal 12(3):83–92
Teixeira C (1961) Paleontological notes on the Karroo of the Lunda (Angola). Garcia de Orta
9(2):307–311
Van Straelen V (1937) Parapirimela angolensis. Brachyure nouveau du Miocène de l’Angola.
Bulletin du Musée Royal d’Histoire Naturelle de Belgique 8(5):1–4
Vasconcelos P (1951) Sur la découverte d’algues fossiles dans les terrains anciens de l’Angola. Int
Geol Congr XVIIIth Sess 14:288–293
Walker RT, Telfer M, Kahle RL et al (2016) Rapid mantle-driven uplift along the Angolan margin
in the late Quaternary. Nat Geosci 9(12):909–914. https://doi.org/10.1038/NGEO2835
Wood RC (1973) Fossil marine turtle remains from the Paleocene of the Congo. Annales du Musée
Royal d’Afrique Centrale, Sciences Geologiques 75:1–28
Zachos JC, Röhl U, Schellenberg SA et al (2005) Rapid acidification of the ocean during the
Paleocene-Eocene thermal maximum. Science 308(5728):1611–1615
Part II
Flora, Vegetation and Landscape Change
Chapter 5
The Flora of Angola: Collectors, Richness
and Endemism
David J. Goyder and Francisco Maiato P. Gonçalves
Abstract Angola is botanically rich and floristically diverse, but is still very

unevenly explored with very few collections from the eastern half of the country.
We present an overview of historical and current botanical activity in Angola, and
point to some areas of future research. Approximately 6850 species are native to
Angola and the level of endemism is around 14.8%. An additional 230 naturalised
species have been recorded, four of which are regarded as highly invasive. We draw
attention to the paucity of IUCN Red List assessments of extinction risk for Angolan
vascular plants and note that the endemic aquatic genus Angolaea (Podostemaceae),
not currently assessed, is at high risk of extinction as a result of dams built on the
Cuanza river for hydro-electric power generation. Recent initiatives to document
areas of high conservation concern have added many new country and provincial
records and are starting to fill geographic gaps in collections coverage.
Keywords Botanical collectors · Botanical diversity · Botanical history ·

Gossweiler · Invasive species · Welwitsch
D. J. Goyder (*)
Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, UK
e-mail: D.Goyder@kew.org
F. M. P. Gonçalves
Instituto Superior de Ciências da Educaҫão da Huíla, Lubango, Angola
University of Hamburg, Institute for Plant Science and Microbiology, Hamburg, Germany
e-mail: francisco.maiato@gmail.com

https://doi.org/10.1007/978-3-030-03083-4_5
80 D. J. Goyder and F. M. P. Gonçalves
History of Botanical Exploration in Angola
It appears that the earliest extant botanical collections from Angola date from either
1669 (Exell 1939; Martins 1994) or more probably 1696 (Dandy 1958; Exell 1962;
Mendonça 1962; Figueiredo et al. 2008), and were made by Mason in the Luanda
region, and by John Kirckwood in Cabinda. These reached Hans Sloane whose
plant and insect collections formed the core of the British Museum (now the Natural
History Museum), London, via James Petiver who encouraged surgeons on English
ships to send him natural history collections from their overseas travels. Other Pre-
Linnean collections from Angola in the Sloane Herbarium were made by Gladman
and William Browne (Fig. 5.1). The earliest known Portuguese collector was the
naturalist Joaquim José da Silva, who collected along the Angolan coast and the
western escarpment between 1783 and 1804. This material was taken from Lisbon
to Paris, where it now resides, in 1808 during the Napoleonic Peninsula War
(Mendonça 1962; Figueiredo et al. 2008).
Mendonça (1962) presents a historical account of plant collectors in Angola, giv-
ing helpful insights to the itineraries of a number of early expeditions. A more com-
plete list of collectors is given by Figueiredo et al. (2008), which volume also
includes a useful listing of references relevant to the study of the flora of Angola.
Fig. 5.1 One of the

earliest herbarium
specimens collected in
Angola, in 1706 or 1707,
by W Browne and now
housed in the Sloane
Herbarium at the Natural
History Museum in
London. The New World
starch crop – cassava
Manihot esculenta
(Euphorbiaceae)
5 The Flora of Angola: Collectors, Richness and Endemism 81
Most eighteenth and early nineteenth century explorers visited only coastal
regions of Angola, but by the 1850s, botanists and explorers were starting to
document plants from more elevated parts of the interior. Friedrich Welwitsch, who
spent 6 years in Angola, amassed over 8000 collections of plants representing
around 5000 species, of which around 1000 were new to science (Albuquerque
2008; Albuquerque et al. 2009; Albuquerque and Figueirȏa 2018). He spent his first
year in Angola in the coastal zone between the mouth of the Rio Sembo (‘Quizembo’)
just north of Ambriz, and the mouth of the Cuanza. In September 1854 he embarked
on a three-year excursion, initially following the Bengo River and reaching Golungo
Alto (Cuanza-Norte). He based himself eventually at Sange from where he made
excursions to Ndalatando (‘Cazengo’) and the banks of the Luinha. In October 1856
he arrived at Pungo Andongo (Malange) where he was based for the next eight
months, making collections from Pedras Negras, Pedras de Guinga and localities
along the Cuanza River – the furthest point he reached upstream was Quissonde,
south of Malange. After an extended period back in Luanda, he headed south via
Benguela to Namibe (‘Little Fish Bay’) in June 1859, gradually extending his
journeys along the coast to Cabo Negro, the port of Pinda (probably Tômbua) and
Baía dos Tigres. In October 1859 he headed inland from Namibe, following the Rio
Giraul (‘Maiombo river’) to Bumbo on the slopes of the Serra da Chela. He was
based at Lopollo on the Huíla plateau until 1860. In 1866, José Anchieta moved to
Angola and was based at Caconda on the Huíla plateau. And by the 1880s,
missionaries such as José Maria Antunes and Eugène Dekindt, and collectors such
as Francisco Newton and Henry Johnston were also making significant collections
from this region.
Three nineteenth century German expeditions to the Congo travelled through
Angola – Pechuël-Lösche’s 1873 Loango Expedition with Paul Güssfeldt and
Hermann Soyaux started from Cabinda; Pogge, Buchner and Wissmann’s Cassai
Expedition made collections from Malange and the Lundas (Mona Quimbundo,
Saurimo, Cuango River) in 1876; while Teucsz and Mechow’s Cuango Expedition
made collections from Dondo (Cuanza-Norte), Pungo Andongo and Malange
(Malange), and the Cuango river (Uíge) in 1879–1881. A fourth German expedition,
the Kunene-Sambesi Expedition, left Namibe on 11 August 1899 and travelled east,
through present-day Cunene and Cuando Cubango provinces, reaching the Cuando
River in March 1900 before returning to Namibe in June of that year. Over 1000
collections were made on this expedition by the botanist Hugo Baum (Warburg
1903; Figueiredo et al. 2009b).
The first half of the twentieth century was dominated by the efforts of Kew-
trained Swiss botanist John Gossweiler who in the course of 50 years’ work collected
in all of Angola’s provinces, and amassed over 14,000 collections. His final 2 years’
collections, in 1946 and 1948, were from the remote northeast of the country, and
formed the basis of Cavaco’s Flora of Lunda (Cavaco 1959). Other significant
colonial era collectors included Portuguese and British participants of the Missões
Botânicas such as Luiz Carrisso, Francisco Mendonça, Arthur Exell and Francisco
de Sousa (as well as John Gossweiler), whose work formed the basis of early parts
of the Conspectus Florae Angolensis, and the first vegetation map of Angola
2015
2000
1990
10°N 1980
1970
1960
0°
1950
1940
1930
10°S
1920
1910
20°S 1900
1890
1780
10°W 0° 10°E 20°E 30°E 40°E
Fig. 5.2 Time lapse of botanical collecting history across tropical Africa. The map represents the
date of the first botanical collection made within each 0.5° sampling unit. Dashed lines represent
the limits for tropical Africa as defined by Sosef et al. (Used with permission from Sosef et al.
2017: http://rainbio.cesab.org)
(Gossweiler and Mendonça 1939). There are too many other collectors from 1950–
1975 to list (see Figueiredo et al. 2008), but two specialist collections are here
noted – Hans Hess’s aquatic and wetland plants from many of the rivers of western
Angola in 1950–1952 are now housed principally in Zurich, and Larry Leach and
IC Cannell who travelled up the arid and semi-arid coastal plain between 1967 and
1973, focussed mostly on the succulent flora. After Angolan independence in 1975
and the commencement of the long-running civil war, collection of plants essen-
tially ground to a halt until the end of the twentieth century. Several recent collect-
ing programmes will be described in a later section of this paper. Despite Gossweiler
and his successor Brito Teixeira’s efforts to survey little known regions of Angola,
plant collection coverage and intensity is skewed heavily to the western half of the
country, and large parts of Moxico, Cuando Cubango, the Lundas and Uíge are still
devoid of collections (Sosef et al. 2017: Fig. 5.2; http://rainbio.cesab.org).
Floristic Diversity and Endemism
Under the leadership of Estrela Figueiredo and Gideon Smith, thirty-two authors
from around the world compiled the first checklist of vascular plants for Angola
(Figueiredo and Smith 2008; Smith and Figueiredo 2017). A total of 6735 native
species were recorded with an additional 226 non-native species. The exotic flora of
Angola was documented by Gossweiler (1948, 1949, 1950). Four of these alien spe-
cies pose particular threats as they are highly invasive in Angola (Rejmánek et al.
2017). Forty-four additional species have been described or entered onto the
International Plant Names Index since publication of Figueiredo and Smith (2008),
and inventories in Lunda-Norte (see below) and elsewhere added a further 70 or so
species to the Angolan list. So the current estimate of the vascular plants native to
Angola is around 6850 species. Current accepted nomenclature for plants can be
checked on the African Plants Database (2018), and local plant names in Gossweiler
(1953) and Figueiredo and Smith (2012).
Figueiredo et al. (2009a) reported that 997 species (14.8%) are endemic to the
country. This percentage is considerably lower than the estimate of 27.3% by Exell
and Gonçalves (1973) based on a limited sample of the flora, or studies of individual
families of plants where 19% of both Rubiaceae (Figueiredo 2008) and legume
species (Soares et al. 2009) were recorded as endemics. Several genera are endemic
to Angola, including Calanda K.Schum. and Ganguelia Robbr. (Rubiaceae);
Carrissoa Baker f. (Leguminosae); and Angolaea Wedd. (Podostemaceae) – the
latter now possibly extinct as it was described from the Cambambe rapids on the
now heavily dammed Cuanza River.
Legumes (934 spp.), grasses (526 spp.), Compositae (463 spp.) and Rubiaceae
(444 spp.) are the most diverse families in the flora, and Crotalaria L. and Euphorbia
L. each have more than 40 Angolan endemic species.
Two of the six tropical African centres of endemism identified by Linder (2001)
fall partially or entirely within Angola. A recent analysis of RAINBIO data
(Droissart et al. 2018) identifies the western Angolan highlands as a distinct floristic
bioregion, although the limited data preclude statements on the remainder of the
country. The Huíla plateau consistently stands out as being rich in endemic species
(Exell and Gonçalves 1973, Brenan 1978: 472, Linder 2001) and Soares et al.
(2009) record 83 endemic legumes from the province. For Rubiaceae, Cabinda has
the highest level of diversity with 175 species, but Huíla possesses the most
endemics (Figueiredo 2008). Figueiredo (2008) also demonstrates that for
Rubiaceae, Huíla is the most intensively collected province. However, our experience
is that many of these collections have not necessarily been well studied. Clark et al.
(2011) state that the western highlands of Angola comprise the least well-
documented stretch of the Great Escarpment of southern Africa.
The western margin of the Huíla Plateau reaches its highest elevation along the
Lubango Escarpment of the Serra da Chela and runs in a southwesterly direction
from near Tundavala c. 15 km NW of Lubango to Bimbe c. 20 km NW of Humpata.
It reaches a height of just over 2200 m and Goyder et al. (in prep.) estimate around
200 species are endemic to this area. However, as other mountains further to the
north are surveyed botanically, some of these supposed local endemics may prove
to be more widely distributed than currently thought.
Linder’s (2001) second area of high species diversity and endemism, the
Zambezi-Congo watershed, encompasses eastern Angola, northern Zambia and the
Katanga region of the DR Congo. This area has not been well documented in
Angola.
Biogeography, Regional Centres of Endemism and Vegetation
With its extremes of landform, climate and rainfall, Angola is host to six of White’s
(1983) phytochoria, or regional centres of endemism.
Outliers of the Guineo-Congolian forests in Cabinda, Uíge and Cuanza-Norte
are progressively smaller in area to the south, ending in the isolated coffee forests of
Gabela and Cumbira in Cuanza-Sul. The northward-draining tributaries of the
Cuango and Cassai rivers in Uíge and Lunda-Norte have fingers of pure Congolian
forest along them. However, much of northern Angola forms a transition zone
between Guineo-Congolian vegetation and Zambezian – the latter covers the rest of
the country with the exception of the fragmented Afromontane centre of endemism
at higher elevations, and the more arid Karoo-Namib and Kalahari-Highveld zones
in the southwest.
Geologically, the eastern half of Angola is notable for its deep deposits of
Kalahari sand, while to the west crystalline rocks predominate. Marine sediments
and recent sands cover the coastal plain (Huntley and Matos 1994; Huntley 2019).
The coastal plain is arid in the south due to the cold, upwelling Benguela current,
and semi-arid further to the north. Most of the rainfall occurs on the escarpment and
the plateau, again with a steady increase to the north. Central Angolan headwaters
of major river systems drain into the Okavango (Cuito and Cubango), the Indian
Ocean (Cuando, Lungué Bungo and Zambezi) and the Atlantic (Cassai, Cuango,
Cuanza and Cunene).
The standard work for vegetation is Barbosa’s (1970) Carta Fitogeográfica de
Angola which recognises 32 vegetation types ranging from desert to moist evergreen
and swamp forests. Huntley and Matos (1994) present a concise summary. Barbosa’s
vegetation map built on the painstaking pioneering work of Gossweiler and
Mendonça (1939) – a major contribution that reached a wider audience through the
extended English summary by Airy-Shaw (1947).
Angola has a diverse seaweed flora and 169 species have been recorded (Lawson
et al. 1975; Anderson et al. 2012). Biogeographically, Angola’s marine algae group
with those of tropical West Africa, but with a well-developed southern element from
around 13°S comprising mainly cooler-water species from the Benguela Marine
Province of Namibia and western South Africa.
Recent Botanical Survey Initiatives
In 1968, Angola had only three National Parks (Quiçama, Cameia and Iona) and
two Nature reserves (Mupa and Luando), plus a number of forest and game reserves
(Teixeira 1968a). Between 1971 and 1975 a programme of field surveys was
undertaken to identify areas of high importance for biodiversity conservation
(Huntley 1973, 1974; Huntley and Matos 1994). These were supplemented by
fieldwork in Huíla, Namibe, Cuanza-Sul and Huambo (Huntley 2009; Mills et al.
2011), and synthesised into an ‘Angolan Protected Area Expansion Strategy –

APAES’ (Huntley 2010). The APAES report was submitted to the Angolan Ministry
of Environment in 2010, and formed the basis for the proposals approved by the
Angolan Conselho do Ministros on 28th April 2011 (GoA 2011).
Much of the recent botanical activity in Angola has focused on the eleven areas
highlighted in this conservation planning document. The areas proposed for
protection were: Maiombe (Cabinda), Serra do Pingano (Uíge), Lagoa Carumbo
(Lunda-Norte), Serra Mbango (Malange), Gabela and Cumbira Forests (Cuanza-
Sul), Morro Namba (Cuanza-Sul), Morro Moco (Huambo) Serra da Neve (Namibe),
Serra da Chela (Huíla) and Luiana (Cuando Cubango). A listing of post-Independence
botanical collectors in Angola is given in Appendix, following the format used for
earlier collectors used by Figueiredo et al. (2008).
A collaborative Rapid Biodiversity Assessment and training expedition to the
Huíla Plateau and to Iona National Park, with 30 scientific participants from 10
countries and with 15 Angolan students, was convened in 2009. Over 2700 botanical
collections were made and deposited in the National Herbarium, Pretoria with
duplicates deposited in the ISCED-Huíla Herbarium in Lubango (Huntley 2009).
In northern Angola botanical surveys have been initiated in the moist coffee for-
ests of Serra do Pingano, and more widely in Uíge Province, by a team from Dresden
in cooperation with the Universidade Kimpa Vita (Lautenschläger and Neinhuis
2014; Neinhuis and Lautenschläger 2014). These have resulted in a revised list of
bryophytes for Angola (Müller 2014; Müller et al. 2018), the description of new
species of vascular plant (Abrahamczyk et al. 2016), and ethnobotanical assess-
ments (Göhre et al. 2016; Mawunu et al. 2016; Heinze et al. 2017; Lautenschläger
et al. 2018). In total, about 820 species were identified; several of these are new
records for Angola.
Lagoa Carumbo and the Luxico, Luele and Lovua valleys were surveyed by a
team from Kew, the Ministry of the Environment and Agostinho Neto University,
Luanda in 2011, and again in 2013, trebling the known flora of Lunda-Norte as
compared to Cavaco (1959) – the combined report documents 752 taxa including 72
additions to the flora of Angola, and 22 potential new species (Darbyshire et al.
2014; Cheek et al. 2015). This part of Lunda-Norte has Congolian swamp forest in
the river valleys, moist miombo woodland on the slopes, and Zambezian savanna
grasslands on the plateau.
The isolated patch of Guineo-Congolian forest at Cumbira was the subject of a
rapid botanical assessment with more than a hundred botanical specimens collected,
including new Guineo-Congolian records for Angola and species potentially new to
science (Gonçalves and Goyder 2016).
Plants collected from Mount Namba are currently being studied by the Kew/
Lubango team – this work may inform studies on the Lubango Escarpment further
to the south. Both share a mosaic of Afromontane forest, grassland and miombo
woodland habitats, although most of Lubango’s woody vegetation is now heavily
degraded. Comparisons with the much better preserved vegetation on Mount Namba
might inform habitat restoration initiatives in the area.
Serra da Neve and Serra da Chela were visited briefly in 2013 as part of a wider
floristic survey of the Angolan Escarpment led by a team from Rhodes University in
South Africa, ISCED-Huíla in Lubango, and Kew. One or two new species have
been published from these collections (Hind and Goyder 2014), but wider analysis
of the flora is still on-going. Through the German-funded Southern African Science
Service Centre for Climate Change and Adaptive Land Management (SASSCAL)
project, researchers at the Lubango Herbarium are working on vegetation
classification of the woodlands of Huíla Province, towards a new vegetation map for
the region (Chisingui et al. 2018). A checklist of the Huíla flora is one of the expected
early outputs.
In addition to the Protected Areas Expansion Strategy sites mentioned above,
three cross-border initiatives have focused on the catchment of the Okavango system
in Angola, Namibia and Botswana in recent years. Botswana’s flagship wetland
ecosystem – the Okavango Delta – is dependent entirely on the two main Angolan
tributaries (Cuito and Cubango) for its hydrology. The Southern Africa Regional
Environmental Program (SAREP) and OKACOM organised fieldwork in Cuando
Cubango in 2013 with botanists from Kew and the University of Botswana. About
350 collections were made from the southeast corner of Angola, as far east as the
Cuando river, thus contributing to the documentation of the Luiana proposed
protected area. The Future Okavango (TFO) project led by a research team from
Hamburg focused on two research sites in Angola (Cusseque, Bié Province;
Caiundo, Cuando Cubango Province) both in the more westerly Cubango catchment,
one in Namibia (Mashare), and Seronga in Botswana. This project contributed
significantly to a better understanding of Angolan miombo and Baikiaea-Burkea
woodlands in terms of recovery following disturbance caused by shifting cultivation
(Wallenfang et al. 2015, Gonçalves et al. 2018, Gonçalves et al. 2017). A checklist
of woody species and geoxylic suffrutices in the grasslands of south-central Angola
was provided, documenting potential new species and new records for the country
(Gonçalves et al. 2016; Revermann et al. 2017, 2018). Further vegetation and
ecological studies are published in Oldeland et al. (2013).
The easterly Cuito and Cuanavale catchment has been the focus of the National
Geographic Okavango Wilderness Project from 2015 onwards. Surveys in the upper
Cubango were initiated in 2017. To date, over 1300 plant collections have been
made by a Kew, South African and Angolan team, who have recorded 417 species
of vascular plant from the high-rainfall upper Cuito and Cuanavale drainage system,
and 176 from the lower rainfall zones further south (e.g., Fig. 5.3). Over 100 new
provincial records were reported for Moxico, with a further 24 for Cuando Cubango,
underlining how poorly documented and understood this vast and sparsely inhab-
ited part of Angola is, even now (Goyder et al. 2018). Baseline botanical collection
data such as these feed into wider biodiversity assessments of the area and provide
vital evidence in building a case to protect the headwaters of not only the Okavango
system, but other major river systems originating in central Angola (NGOWP 2018).
Fig. 5.3 Some plants collected during recent fieldwork in central and eastern Angola as part of the
National Geographic Okavango Wilderness Project. Top to bottom, left to right: Protea poggei
subsp. haemantha (Proteaceae); Clerodendrum baumii (Lamiaceae); Erythrina baumii
(Leguminosae); Monotes gossweileri (Dipterocarpaceae); Gloriosa sessiliflora (Colchicaceae);
Raphionacme michelii (Apocynaceae). All photos: David Goyder
Future Botanical Work
Almost every botanical survey made in recent years in Angola has revealed unde-
scribed species and new country or provincial records. Eastern and northern prov-
inces are in most need of collecting programmes and botanical documentation.
Most national parks lack basic botanical inventories. To give one example, Teixeira’s
(1968b) work on plant diversity in Bicuar National Park (Huíla Province) resulted
in the recognition of six vegetation types in the park. But recent SASSCAL-funded
surveys revealed species unaccounted for by Figueiredo and Smith (2008),
underlining the need for more botanical surveys in both existing and newly proposed
areas of conservation concern.
Analysis of the collections from recent surveys is starting to reveal little-
documented areas of endemism. The Lubango Escarpment is one obvious focus, but
so too is the highly leached high-rainfall Kalahari sand system of Moxico Province
and adjacent area that has its own peculiar and little-understood flora.
Only 399 species of vascular plant in Angola have been formally assessed for
extinction risk through the IUCN Red List system (IUCN 2018), and a mere 36 of
these appear in threatened categories. None of the genera listed in an earlier section of
this paper as Angolan endemics have been assessed. Much work is needed in this area.
Four Angolan institutions are listed in Index Herbariorum (Thiers, continually
updated), LUAI (ex-Centro Nacional de Investigação Cientifica (CNIC), Luanda),
LUA (Instituto de Investigação Agronómica, Huambo), LUBA (Instituto Superior
de Ciencias da Educação, Lubango), and DIA (Museu do Dundo). While the Dundo
Museum has been refurbished and reopened to the public in 2012, it appears that the
herbarium collections formerly housed there no longer exist. The LUA herbarium
contains 40,000 collections. It was evacuated to Luanda in 1995, and has now
returned to Huambo, but is in poor condition and funds are needed to employ well-
trained young staff to conserve, rehabilitate and work on this important collection.
LUAI contains 35,000 collections and LUBA around 50,000. There are ongoing
digitisation programmes at both institutions that will make these collections more
widely accessible.
Outside of Angola, Portuguese institutes in Coimbra (COI) and Lisbon (LISC,
LISU) hold the largest collections of Angolan plants, an estimated 90,000 collections
(Figueiredo and César 2008). 8700 of Gossweiler’s Angolan collections are housed
at COI and these are available online. The collections at LISC are also available
digitally, and are now being incorporated into the Lisbon University herbarium
LISU. Most other herbaria with significant Angolan holdings have only digitised
their type collections, although mass digitisation of entire national collections has
made material in the Paris Natural History Museum (P) and Leiden’s Naturalis (L,
WAG, U) accessible. In the UK, the Natural History Museum (BM) and Royal
Botanic Gardens, Kew (K) in London – both of which contain significant Angolan
holdings, and Royal Botanic Gardens, Edinburgh (E) have plans to follow suit. In
Germany, the collection of Technische Universität Dresden (DR) comprises 2400
specimens, kept separately from the main herbarium. The Future Okavango project
has augmented Hamburg’s (HBG) Angolan collections by around 2000 numbers.
Once these combined resources are available online, georeferencing the Angolan
material should be a priority. Such collections data could then be used in a variety
of projects or programmes. Georeferenced specimen data underpins IUCN
conservation assessments, for example, and these in turn inform Important Plant
Area designations (Darbyshire et al. 2017) and other forms of conservation planning.
Acknowledgements We are delighted to acknowledge the support of the former Minister of

Environment, Dr. Fátima Jardim, and the present Minister, Dr. Paula Francisco, at the Ministério
do Ambiente, Luanda, in our attempts to provide the botanical evidence for the conservation of
Angola’s unique flora. We are grateful to Thea Lautenschläger for providing biographical and
other information relating to projects in Uíge Province.
Appendix
Post-Independence collectors in Angola. Entries follow a format developed from

Figueiredo et al. (2008).
Surname, first names (birth–death); C: period when collecting in Angola; H: her-
baria [abbreviations after Thiers, continuously updated; FC-UAN = Faculdade
de Ciências, Universidade Agostinho Neto, Luanda; INBAC = Instituto Nacional
da Biodiversidade e Áreas de Conservação of the Ministério do Ambiente,
Luanda]; L: provinces abbreviated after Figueiredo and Smith 2008: principal
localities; B: biographical information.
Alcochete, António (1963–)
C: 1991; H: K; L: CU HI NA; B: Angolan botanist, collected with Gerrard, Matos
and Newman.
Baragwanath, S.
C: 1994. H: PRE.
Barker, Nigel P.(1962-)
C: 2013, 2015, 2017; H: GRA, INBAC, K, LUBA, PRE; L: CC HI NA: Lubango
Escarpment, Mt. Tchivira, Serra da Neve, Mundondo Plateau, Okavango, Cuito
and Longa Rivers; B: South African Professor of Plant Science at University of
Pretoria, formerly at Rhodes University.
Bester, Stoffel Petrus (Pieter) (1969–)
C: 2009, 2015; H: GRA, INBAC, K, LUBA, PRE; L: CC CU HI NA: Iona, Lubango
Escarpment, Bicuar, Okavango, Cuito and Longa Rivers; B: South African
botanist based at PRE.
Bruyns, Peter Vincent (1957–)
C: 2006, 2007; H: BOL, E, K, NBG, PRE; L: BE HI NA: Lubango Escarpment and
coastal plain; B: South African mathematician and botanist with particular
interest in succulent plants.
Cardoso, João Francisco (1974–)

C: 2005, 2006; H: LISC, LUAI; L: HI NA: Serra da Leba, Virei, Caraculo, Cainde;
B: Agronomist with Agostinho Neto University.
Cheek, Martin Roy (1960–)
C: 2012; H: K; L: CA; B: British botanist at Royal Botanic Gardens Kew, specialist
on West African flora.
Clark, Vincent Ralph (1977–)
C: 2013; H: GRA, K, LUBA, PRE; L: HI NA: Lubango Escarpment, Mt. Tchivira,
Serra da Neve; B: South African botanist.
Cooper, C.E.
C: 1997; H: PRE.
Crawford, Frances Mary (1981–)
C: 2009, 2011; H: INBAC, K, PRE; L: HI LN NA: Lucapa, Lagoa Carumbo, Iona,
Lubango Escarpment; B: British botanist, Curator of WIND herbarium, formerly
at Royal Botanic Gardens, Kew; collected with Darbyshire and Goyder in LN.
Daniel, José Maria (1943–2015)
C: 1964–2008; H: LUBA, LUA, LUAI, L: Collected in all Angolan Provinces; B:
Angolan botanist at Lubango Herbarium until his retirement; collected with
Huntley, Matos and Gonçalves.
Darbyshire, Iain Andrew (1976–)
C: 2011, 2013; H: INBAC, K, LISC; L: LN: Lucapa, Lagoa Carumbo; B: British
botanist at Royal Botanic Gardens, Kew; collected with Crawford, Gomes,
Goyder & Kodo.
Dexter, Kyle Graham (1980–)
C: 2017–; H: E, COLO, LUBA, WIND; L: CU HI NA; B: Senior Lecturer at
University of Edinburgh and Associate Researcher at Royal Botanic Garden
Edinburgh.
Ditsch, Barbara (1961–)
C: 2013, 2015 H: DR, LUA; L: UI: Serra do Pingano, Municipality of Uíge,
Kimbele, Damba, Mucaba; B: German botanist at Dresden Botanic Garden.
Finckh, Manfred (1963–)
C: 2011–; H: HGB, LUBA, WIND; BI CC HA HI MO: Chitembo (Cusseque),
Caiundo, Cachingues, Savate, Cuangar, Bicuar National Park, Cameia National
Park, Tundavala Observatory under TFO and SASSCAL Projects; B: Ecologist
at University of Hamburg, Germany.
Francisco, Domingos Mumbundu (1974–)
C: 2008–; H: LISC, LUAI, LUBA; L: CA CC LA MA NA ZA: Barra do Cuanza,
Iona, Cangandala, Quiçama National Parks; B: Angolan botanist at Universidade
Agostinho Neto, Centro de Botânica, LUAI Herbarium.
Frisby, Arnold.
C: 2016, 2017; H: INBAC, K, LUBA, PRE; L: BI CC: Cubango and Cuito Rivers;
B: South African botanist at University of Pretoria.
Gerrard, Jacqueline
C: 1991; H: K; L: CU HI NA.
Godinho, Elizeth
C: 2013; H: INBAC, K, LISC; L: LN: Lagoa Carumbo; B: Angolan botanist at
INBAC; collected with Darbyshire, Goyder and Kodo.
Göhre, Anne (1990–).
C: 2014–2016; H: B, BR, BONN, P; L: UI: Municipality of Uíge, Kimbele, Damba,
Mucaba; B: German botanist at Dresden Botanic Garden.
Gomes, Amândio Luís (1971–).
C: 2010–; H: FC-UAN, INBAC, K, LISC, LUAI, LUBA; L: BE BI BO CC CN CS
HA LN ZA: Lucapa, Lagoa Carumbo, Chitembo (Cusseque), Tundavala
Observatory under TFO and SASSCAL Projects; B: Angolan botanist at
Universidade Agostinho Neto, Luanda; collected with Crawford, Darbyshire and
Goyder in LN.
Gonçalves, Francisco Maiato Pedro (1982–).
C: 2008–; H: HBG, INBAC, K, LUBA; L: BI CC CU CS HA HI LA NA MO:
Chitembo (Cusseque), Cumbira forest, Mt. Namba, Lubango Escarpment,
Okavango headwaters, Huíla Province SASSCAL Project; B: Angolan botanist
at Lubango Herbarium, ISCED Huíla, Lubango.
Goyder, David John (1959–)
C: 2011–; H: GRA, INBAC, K, LUBA, PRE; L: BI CC CS HI LN MO NA:
Cumbira, Mt. Namba, Serra da Neve, Lubango Escarpment, Mt. Tchivira,
Okavango headwaters, Lucapa, Lagoa Carumbo; B: British botanist at Royal
Botanic Gardens, Kew; collected with Crawford, Darbyshire, Godinho, Gomes
and Kodo in LN, with Barker and Clark on the western escarpment, with
Gonçalves in CS and Okavango headwaters, with Barker, Bester, Frisby and
Janks in CC.
Harris, Timothy (1982–)
C: 2013; H: K, LUAI, PSUB, WIND; L: CC: Okavango, Cuito and Cuando Rivers;
B: British botanist; collected with Murray-Hudson.
Heinze, Christin (1993–).
C: 2014–2017; H: DR, LUA; L: CN: all municipalities; B: German botanist at
Technische Universität Dresden.
Janks, Matthew.
C: 2015; H: GRA, INBAC, LUBA, PRE; L: CC: Okavango, Cuito and Longa
Rivers; B: South African botanist; collected with Barker, Bester & Goyder.
Jürgens, Norbert (1953–)

C: 2008–; H: HGB, WIND, LUBA; L: CU HI NA; B: Professor at Institute for
Plant Science and Microbiology, University of Hamburg, Germany.
Kodo, Felipe
C: 2013; H: INBAC, K, LISC; L: LN: Lagoa Carumbo; B: Angolan botanist at
INBAC; collected with Darbyshire, Godinho and Goyder.
Lautenschläger, Thea (1980–)
C: 2012–2018; H: DR, LUA; L: UI: Municipality of Uíge, Mucaba, Maquela do
Zombo, Quitexe, Milunga, Sanza Pombo, Kimbele, Ambuila, Songo, Bungo,
Bembe, Puri, Negage, Altocauale, Damba; B: German botanist at Technische
Universität Dresden.
Luís, José Camôngua (1984–)
C: 2015–. H: K, LUBA; L: CS HI: Lubango Escarpment, Mt. Namba; B: Angolan
botanist.
Maiato, Francisco
See Gonçalves, Francisco Maiato Pedro.
Manning, Stephen D.
C: 1986–1998.
Matos, Elizabeth (Liz), Merle (1938–)
C: 1975–; B: British botanist, founder and director of Angola’s National Plant
Genetic Resources Centre, Agostinho Neto University, Luanda. Retired in 2008.
Mawunu, Monizi (1973–)
C: 2013–2018; H: DR, LUA; L: UI, whole province; B: Angolan botanist at
Universidade Kimpa Vita.
Müller, Frank (1966–)
C: 2015; H: DR, LUA; L: UI: Municipality of Uíge, Songo, Mucaba; B: German
botanist at Technische Universität Dresden.
Murray-Hudson, Frances
C: 2013; H: K, LUAI, PSUB, WIND; L: CC: Okavango, Cuito and Cuando Rivers;
B: Volunteer at Peter Smith University of Botswana Herbarium (PSUB);
collected with Harris.
Neinhuis, Christoph (1962–)
C: 2012–2018; H: DR, LUA; L: UI: Municipality of Uíge, Mucaba, Maquela do
Zombo, Quitexe, Milunga, Sanza Pombo; B: German botanist at Technische
Universität Dresden, director of the Botanical Garden TU Dresden.
Newman, Mark Fleming (1959–)
C: 1991; H: K; L: CU HI NA; B: British botanist at Royal Botanic Garden
Edinburgh. In 1991, at the Seed Bank, Royal Botanic Gardens, Kew; collected
with Alcochete, Gerrard and Matos, mainly for seeds, with herbarium voucher
specimens for identification.
Rejmánek, Marcel (Marek/Marc) (1946–)

C: 2014; H: LUBA, STE; L: BE BO CN CS HA HI MA NA UI; B: Czech botanist
based at University of California, Davis, working on biological invasions.
Conducted a rapid inventory of invasive plants in Angola in 2014 with Huntley,
Roux and Richardson.
Revermann, Rasmus (1979–)
C: 2011–; H: HGB, WIND, LUBA; L: BI CC HA HI: Chitembo (Cusseque),
Caiundo, Cachingues, Savate, Cuangar under TFO and SASSCAL Projects; B:
Ecologist at University of Hamburg, Germany.
Roux, Jacobus Petrus (Koos) (1954–2013)
C: 2001; H: PRE; B: South African Pteridophyte specialist.
Tripp, Erin Anne (1979–)
C: 2017–; H: COLO, E, LUBA, WIND; L: CU HI NA; B: Researcher at Colorado
Herbarium, University of Colorado.
References
Abrahamczyk S, Janssens S, Xixima L et al (2016) Impatiens pinganoensis (Balsaminaceae), a

new species from Angola. Phytotaxa 261:240–250
African Plant Database version 3.4.0 (2018) Conservatoire et Jardin Botaniques de la Ville de
Genève and South African National Biodiversity Institute, Pretoria. http://www.ville-ge.ch/
musinfo/bd/cjb/africa
Airy-Shaw H (1947) The vegetation of Angola. J Ecol 35:23–48
Albuquerque S (2008) Friedrich Welwitsch. Figueiredo E, Smith GF Plants of Angola/Plantas de
Angola. Strelitzia 22: 2–3
Albuquerque S, Figueirȏa S (2018) Depicting the invisible: Welwitsch’s map of travellers in
Africa. Earth Sci Hist 37(1):109–129
Albuquerque S, Brummitt RK, Figueiredo E (2009) Typification of names based on the Angolan
collections of Friedrich Welwitsch. Taxon 58:641–646
Anderson RJ, Bolton JJ, Smit AJ et al (2012) The seaweeds of Angola: the transition between trop-
ical and temperate marine floras on the west coast of southern Africa. Afr J Mar Sci 34:1–13
Barbosa LAG (1970) Carta Fitogeográfica de Angola. Instituto de Investigação Científica de
Angola, Luanda
Brenan JPM (1978) Some aspects of the phytogeography of tropical Africa. Ann Mo Bot Gard
65(2):437–478
Cavaco A (1959) Contribution à l'Étude de la Flore de la Lunda d'Après les Récoltes de Gossweiler
(1946–1948). Publicações Culturais da Companhia de Diamantes de Angola 42, 230 pp
Cheek M, Lopez Poveda L, Darbyshire I (2015) Ledermanniella lunda sp. nov. (Podostemaceae)
of Lunda-Norte, Angola. Kew Bull 70:10
Chisingui AV, Gonçalves FMP, Tchamba JJ et al (2018) Vegetation survey of the woodlands of
Huíla Province. Biodivers Ecol 6:426–437
for biodiversity exploration. Biodivers Conserv 20:2543–2561
Dandy JE (1958) The Sloane Herbarium. An annotated list of the horti sicci composing it; with
biographical accounts of the principal contributors. British Museum (Natural History), London
Darbyshire I, Goyder D, Crawford F, et al (2014) Update to the Report on the Rapid Botanical
Survey of the Lagoa Carumbo Region, Lunda-Norte Prov., Angola for the Angolan Ministry
of the Environment, following further field studies in 2013, incl. Appendix 2: checklist to the
flowering plants, gymnosperms and pteridophytes of Lunda-Norte Prov, Angola. Ministério do
Ambiente, Luanda
Darbyshire I, Anderson S, Asatryan A et al (2017) Important Plant Areas: revised selection criteria
for a global approach to plant conservation. Biodivers Conserv 26:1767–1800
Droissart V, Dauby G, Hardy OJ et al (2018) Beyond trees: biogeographical regionalization of
tropical Africa. J Biogeogr 2018:1–15
Exell AW (1939) Notes on the flora of Angola. IV. 1. Collections from Angola in the Sloane
Herbarium. J Bot 77:146–147
Exell AW (1962) Pre-Linnean collections in the Sloane Herbarium from Africa south of the Sahara.
In: Fernandes A (ed) Comptes Rendus de la IVe Réunion Plénière de l'Association pour l'Étude
Taxonomique de la Flore d'Afrique Tropicale (Lisbonne et Coïmbre, 16–23 Septembre, 1960).
Junta de Investigações do Ultramar, Lisbon, pp 47–49
Exell AW, Gonçalves ML (1973) A statistical analysis of a sample of the flora of Angola. Garcia
de Orta, Série de Botânica 1:105–128
Figueiredo E (2008) The Rubiaceae of Angola. Bot J Linn Soc 156:537–638
Figueiredo E, César J (2008) Herbaria with collections from Angola/Herbários com colecções de
Angola. Strelitzia 22:11–12
Figueiredo E, Smith GF (eds) (2008) Plants of Angola/Plantas de Angola. Strelitzia 22:1–279
Figueiredo E, Smith GF (2012) Common names of Angolan plants. Inhlaba Books, Pretoria
Figueiredo E, Matos S, Cardoso JF et al (2008) List of collectors/Lista de colectores. Strelitzia
22:4–11
Figueiredo E, Smith GF, César J (2009a) The flora of Angola: first record of diversity and ende-
mism. Taxon 58:233–236
Figueiredo E, Soares M, Siebert G et al (2009b) The botany of the Cunene-Zambezi Expedition
with notes on Hugo Baum (1867-1950). Bothalia 39:185–211
de Angola. Direcção Nacional da Biodiversidade, Ministério do Ambiente, Luanda, 35 pp
Göhre A, Toto-Nienguesse AB, Futuro M et al (2016) Plants from disturbed savannah vegetation
and their usage by Bakongo tribes in Uíge, Northern Angola. J Ethnobiol Ethnomed 12:42
Gonçalves FMP, Goyder DJ (2016) A brief botanical survey into Kumbira forest, an isolated patch
of Guineo-Congolian biome. PhytoKeys 65:1–14
Gonçalves FMP, Tchamba JJ, Goyder DJ (2016) Schistostephium crataegifolium (Compositae:
Anthemideae), a new generic record for Angola. Bothalia 46:a2029
Gonçalves FMP, Revermann R, Gomes AL, et al (2017) Tree species diversity and composition of
Miombo woodlands in south-central Angola, a chronosequence of forest recovery after shifting
cultivation. Int J For Res 2017(Article ID 6202093), 13 pp
Gonçalves FMP, Revermann R, Cachissapa MJ, et al (2018) Species diversity, population structure
and regeneration of woody species in fallows and mature stands of tropical woodlands of SE
Angola. J Forest Res. Published online 13 January 2018
Gossweiler J (1948) Flora exótica de Angola. Nomes vulgares e origem das plantas cultivadas ou
sub-espontâneas. Agronomia Angolana 1:121–198
Gossweiler J (1953) Nomes indígenas das plantas de Angola. Agronomia Angolana 7:1–587
Lisboa, 242 pp
Goyder DJ, Barker N, Bester SP et al (2018) The Cuito catchment of the Okavango system: a vas-
cular plant checklist for the Angolan headwaters. PhytoKeys 113:1–31. https://doi.org/10.3897/
phytokeys.113.30439
Heinze C, Ditsch B, Congo MF et al (2017) First Ethnobotanical Analysis of Useful Plants in
Cuanza Norte North Angola. Res Rev J Bot Sci 6:44
Hind DJN, Goyder DJ (2014) Stomatanthes tundavalaensis (Compositae: Eupatorieae:
Eupatoriinae), a new species from Huíla Province, Angola, and a synopsis of the African spe-
cies of Stomatanthes. Kew Bull 69(9545):1–9
Huntley BJ (1973) Proposals for the creation of a strict nature reserve in the Maiombe forest
of Cabinda. Report 16. Repartição Técnica da Fauna, Serviços de Veterinária, Luanda,
Mimeograph report, 10 pp
Huntley BJ (1974) Ecosystem conservation priorities in Angola. Report 28. Repartição Técnica da
Fauna, Serviços de Veterinária, Luanda, Mimeograph report, 22 pp
ect. Report on Pilot Project. Unpublished Report to Ministry of Environment, Luanda, 97 pp,
27 figures
Huntley BJ (2010) Estratégia de Expansão de Rede da Áreas Protegidas da Angola/Proposals for
an Angolan Protected Area Expansion Strategy (APAES). Unpublished Report to the Ministry
of Environment, Luanda, 28 pp, map
Huntley BJ (2019) Angola in Outline: Physiography, Climate and Patterns of Biodiversity. In:
1:53–74
IUCN (2018) The IUCN Red List of Threatened Species. Ver. 2017-3. http://www.iucnredlist.org.
Downloaded on 27 March 2018
Lautenschläger T, Neinhuis C (eds) (2014) Riquezas Naturais de Uíge – uma Breve Introdução
Sobre o Estado Atual, a Utilização, a Ameaça e a Preservação da Biodiversidade. Technische
Universität Dresden, Dresden
Lautenschläger T, Monizi M, Pedro M et al (2018) First large-scale ethnobotanical survey in the
province Uíge, northern Angola. J Ethnobiol Ethnomed 14:51
Lawson GW, John DM, Price JH (1975) The marine algal flora of Angola: its distribution and
affinities. Bot J Linn Soc 70:307–324
Linder HP (2001) Plant diversity and endemism in sub-Saharan tropical Africa. J Biogeogr
28:169–182
Martins ES (1994) John Gossweiler. Contribuição da sua obra para o conhecimento da flora ango-
lana. Garcia de Orta, Série de Botânica 12:39–68
Mawunu M, Bongo K, Eduardo A et al (2016) Contribution à la connaissance des produits fores-
tiers non ligneux de la Municipalité d’Ambuila (Uíge, Angola): Les plantes sauvages comes-
tibles [Contribution to the knowledge of no-timber forest products of Ambuila Municipality
(Uíge, Angola): The wild edible plants]. Int J Innov Sci Res 26:190–204
Mendonça FA (1962) Botanical collectors in Angola. In: Fernandes A (ed) Comptes Rendus de
la IVe Réunion Plénière de l’Association pour l’Étude Taxonomique de la Flore d’Afrique
Tropicale (Lisbonne et Coïmbre, 16–23 septembre, 1960). Junta de Investigações do Ultramar,
Lisbon, pp 111–121
Mills MSL, Olmos F, Melo M et al (2011) Mount Moco: its importance to the conservation of
Swierstra’s Francolin Pternistis swierstrai and the Afromontane avifauna of Angola. Bird
Müller F (2014) About 150 years after Welwitsch – a first more extensive list of new bryophyte
records for Angola. Nova Hedwigia 100:487–505
Müller F, Sollman P, Lautenschläger T (2018) A new synonym of Weissia jamaicensis (Pottiaceae,
Bryophyta) and an extension of the range of the species from the Neotropics to the Palaeotropics.
Plant Fungal Syst 63(1):1–5
Neinhuis C, Lautenschläger T (2014) The potentially natural vegetation in Uíge province and its
current status – arguments for a protected area in the Serra do Pingano and adjacent areas.
Unpublished Report to Ministry of Environment, Luanda, 64 pp
NGOWP (National Geographic Okavango Wilderness Project) (2018) Initial findings from explo-
ration of the upper catchments of the Cuito, Cuanavale, and Cuando Rivers, May 2015 to
December 2016. Report prepared for and submitted to the Ministério do Ambiente of the
Republic of Angola, the Ministry of Environment, Wildlife and Tourism Botswana, and the
Ministry of Environment and Tourism of the Republic of Namibia. Available (as Report 1)
from: http://www.wildbirdtrust.com/owp-publications/
Oldeland J, Erb C, Finckh M, Jürgens N (eds) (2013) Environmental assessments in the Okavango
region. Biodivers Ecol 5:1–418
Rejmánek M, Huntley BJ, le Roux JJ, Richardson DM (2017) A rapid survey of the invasive plant
species in western Angola. Afr J Ecol 55:56–69
Revermann R, Gonçalves FM, Gomes AL et al (2017) Woody species of the miombo woodlands
and geoxylic grasslands of the Cusseque area, south-central Angola. Check List 13:2030
Revermann R, Oldenland J, Gonçalves FM et al (2018) Dry tropical forests and woodlands of the
Cubango basin in southern Africa – First classification and assessment of their woody species
diversity. Phytocoenologia 48:23–50
Smith GF, Figueiredo E (2017) Determining the residence status of widespread plant species: stud-
ies in the flora of Angola. Afr J Ecol 55:710–713
Soares M, Abreu J, Nunes H et al (2009) The Leguminosae of Angola: diversity and endemism.
Syst Geogr Plants 77:141–212
Sosef MSM, Dauby G, Blach-Overgaard A et al (2017) Exploring the floristic diversity of tropical
Africa. BMC Biol 15:15
Teixeira JB (1968a) Angola. In: Hedberg I, Hedberg O (eds.) Conservation of vegetation in
Africa south of the Sahara. Proceedings of a symposium held at the 6th plenary meeting of
the “Association pour l’Etude Taxonomique de la Flore d’Afrique Tropicale” (A.E.T.F.A.T.) in
Uppsala, Sept. 12th–16th, 1966. Acta Phytogeographica Suecica 54:193–197
Teixeira JB (1968b) Parque Nacional do Bicuar. Carta da Vegetação (1ª aproximação) e Memória
Descritiva. Instituto de Investigação Agronómica de Angola, Nova Lisboa
Thiers B (continuously updated). Index Herbariorum: A global directory of public herbaria and
associated staff. New York Botanical Garden’s Virtual Herbarium. http://sweetgum.nybg.org/
science/ih/
Wallenfang J, Finckh M, Oldeland J et al (2015) Impact of shifting cultivation on dense tropical
woodlands in southeast Angola. Trop Conserv Sci 8:863–892
Warburg O (1903) Kunene-Sambesi-Expedition. Kolonial-Wirtschaftliches Komitee, Berlin
White F (1983) The Vegetation of Africa – A Descriptive Memoir to Accompany the UNESCO/
AETFAT/UNSO Vegetation Map of Africa. UNESCO, Paris 356 pp
Chapter 6
Vegetation Survey, Classification
and Mapping in Angola
Rasmus Revermann and Manfred Finckh
Abstract Spatial information about plant species composition and the distribution
of vegetation types is an essential baseline for natural resource management plan-
ning. In Angola, the first countrywide vegetation map was elaborated by Gossweiler
in 1939. Subsequently, Barbosa published a revised map with much higher detail in
1970 and his work has remained the main reference for the vegetation of Angola
until today. However, these early maps were expert drawn and were not based on
systematic surveys. Instead, the delimitation of vegetation units was based on many
years of field observations and also incorporated results of local studies carried out
by other authors. In spite the rich history of the scientific exploration of Angola’s
vegetation in colonial times, quantitative and plot based studies were rare. After the
end of the armed conflict, new vegetation surveys making use of new methodologi-
cal developments in numerical approaches to vegetation classification in combina-
tion with modern remote sensing imagery have provided spatial information of
unprecedented detail. However, vast areas of the country still remain seriously
understudied. At the same time, sustainable land management strategies are urgently
needed due to the increasing pressure on natural resources driven by socio-economic
development and global change, thus calling for a new era of vegetation surveys that
will enable data-based landuse and conservation planning in Angola.
Keywords Conservation · Landuse planning · Natural resources · Plant communi-

ties · Remote sensing
R. Revermann (*) · M. Finckh

Institute for Plant Science and Microbiology, University of Hamburg, Hamburg, Germany
e-mail: rasmus.revermann@gmail.com; manfred.finckh@uni-hamburg.de

https://doi.org/10.1007/978-3-030-03083-4_6
98 R. Revermann and M. Finckh
Introduction
Knowledge on the spatial distribution of vegetation and its species composition is

paramount for any kind of natural resource management and conservation planning.
Vegetation serves as habitat for other organismic groups and is the source of energy
in the ecosystem. As such, vegetation integrates many ecological processes and
reflects patterns of topography, geology, soil, hydrology and climate. Thus, vegeta-
tion classification is ideal to provide an aggregated image of the landscape and its
ecological communities.
Historical Exploration of Vegetation Patterns in Angola
First reports on the vegetation of Angola were directly linked to the floristic explora-
tion of the country, as outlined by Goyder and Gonçalves (2019). Scientific mis-
sions during colonial times in Angola served several purposes: on the one hand they
should chart the potential for economic exploitation and development while on the
other hand they may also have been used to demonstrate the supremacy of the colo-
nial power (Gago et al. 2016). The expedition by the geographer Jessen (1936)
provided a first sketch of the vegetation along the routes of his transects through
western Angola. Jessen’s work remains a classic as he was among the first to docu-
ment the landscape and ecosystem properties of the region. However, it is hardly
read today as it is only available in German.
The systematic descriptions of the vegetation of Angola started with Gossweiler
and Mendonça’s (1939) phytogeographical map of Angola. The often-cited English
summary by Shaw (1947) contributed much to the recognition of Gossweiler’s work
internationally. The map is based on the combined structural and ecological
approach to vegetation classification developed by Brockmann-Jerosch and Rübel
(1912) in Zurich. Thus, in a first level of classification they categorised the vegeta-
tion according to woodiness and persistence into the three categories Lignosa
(woody), Herbosa (herbaceous) and Deserta (land surfaces without permanent veg-
etation cover). The next step of the classification included climatic and edaphic
factors, as well as leaf traits, leading for instance to five sub-categories of woody
vegetation called Pluviilignosa, Laurilignosa, Durilignosa, Ericilignosa,
Aestililignosa and Hiemilignosa. Stand structure was the main criterion for the next
categories, dividing the afore-mentioned categories between tall forests (-silva) and
dense but low forests (-fruticeta) (e.g. Pluviisilva vs. Pluviifruticeta or Durisilva vs.
Durifruticeta). Below this third level we finally find floristically defined vegetation
units, albeit mostly named after one or two dominant species. Similar structural
criteria were used for the sub-classification of the Herbosa and Deserta.
The vegetation map used this rather rigid classification scheme for the 19 main
mapping units. However, the resulting map apparently did not fully satisfy the
authors, who then applied 29 additional symbols to indicate occurrence of small-
6 Vegetation Survey, Classification and Mapping in Angola 99
scale vegetation units, of transition zones and of species that appeared to be of

special interest to the authors – a very nice real world example of dealing with rigid
mapping manuals. However, neglecting these small methodological inconsisten-
cies, the map by Gossweiler & Mendonça presented the first overall picture of the
vegetation of Angola, a first approach towards a systematic compilation of observa-
tions of phytogeographical patterns and a first attempt at ecological interpretation.
While many of the mapped polygons seem outdated in times of modern earth
observation, the number of observed details in remote parts of Angola is still sur-
prising for today’s botanists. The authors were probably the first to report for Angola
on invasive species, on seed dispersal by bats, on the morphological plasticity of the
genus Syzygium and on many other current scientific topics. Also quite astonishing
was the classification of the suffrutex grassland within the woody vegetation types
(Ericifruticeta), more than 30 years before White (1976) published the ground-
breaking paper on the ‘Underground forests of Africa’.
The next important integrating step towards a synthesis on the vegetation units of
Angola and their spatial distribution was Grandvaux Barbosa’s phytogeographical
map of Angola (Barbosa 1970). His work can be seen as a continuation and exten-
sion of the Gossweiler approach. The map clearly benefited from several regional
studies that had been carried out in the meantime (see below) and of course also
from Barbosa’s own knowledge gained during several field missions throughout the
country and his extensive experience of similar vegetation types found in
Mozambique. As ancillary information Barbosa included descriptions of the main
soil types and climatic zones of Angola.
The mapping approach adopted by Barbosa was to some degree harmonised with
the parallel efforts of the Flora Zambesiaca map and the UNESCO initiative map-
ping the vegetation of Africa. The first level of classification differentiates the veg-
etation based on the formation, i.e. deals with the physiognomy of the vegetation
such as closed forests, forest savanna mosaics, woodlands etc. and beyond that
includes azonal edaphic vegetation units such as mangrove stands and coastal dune
vegetation. In the second level of classification vegetation types are distinguished
according to dominant species. In total, the map by Barbosa displays 32 main veg-
etation types and the descriptive text accompanying the map provides details on
over 100 subordinate types (for a brief summary in English see Barbosa 1971).
The result was a good overview of the main vegetation types of Angola, in terms
of spatial patterns much superior to the first attempt by Gossweiler and Mendonça
(1939). Until today the vegetation units of the map by Barbosa (1970) constitute the
foundation for the Angolan part of most continental or global scale vegetation maps
(see below). However, due to Barbosa’s floristic rather than ecological emphasis,
the report on the vegetation units did neither contribute much to a better understand-
ing of the ecology of the main vegetation patterns, nor did it make use of a modern
classification concept based on plant communities.
Shortly after Barbosa’s vegetation map, Diniz (1973) published a monograph on
the physical properties of the agricultural zones of Angola. Included in this mono-
graph, are soil and vegetation maps for 36 agricultural zones, albeit in rather frag-
mented components and without an overall map of the country. The vegetation
classification scheme he used is not clearly defined, somewhere in between those of

Gossweiler and Barbosa, but sometimes with more detail than Barbosa (1970). The
main achievement of Diniz (1973) is that he assembled sound environmental infor-
mation (with a focus on geology and soils) for all delimited agricultural zones.
However, due to the lack of a seamless map and his unclear classification approach
his contribution to the knowledge on the vegetation of Angola did not receive much
attention in the subsequent scientific literature and due to the violent conflicts
following Angola’s independence in 1975, Barbosa’s work has remained the main
reference on the vegetation of Angola.
I ntegration of the Vegetation Map of Angola

Within Continental Scale Maps
The next important step for a better understanding of Angolan vegetation was the
UNESCO/AETFAT/UNSO initiative for a Vegetation Map of Africa (UNESCO/
AETFAT/UNSO 1981), compiled and described by White (1983). For Angola the
continental map is largely based on the units supplied by Barbosa (1970) but they
were subject to further generalisation resulting in only 14 mapping units compared
to Barbosa’s 32 vegetation types. However, the important achievement of White’s
map lies in the fact that it inserted Angolan vegetation in a common conceptual and
methodological framework with the vegetation of neighbouring countries and the
African continent as a whole. As such, the UNESCO (UNESCO/AETFAT/UNSO
1981)/(UNESCO 1981) map and White’s (1983) description established the now
widely used term ‘miombo woodlands’ in our scientific and geographic frame-
works, allowing thus for the comparison of Angola’s ecosystems with similar veg-
etation types throughout Africa. Although Barbosa and White provided seamless
maps covering the entire country, the level of information supporting the mapping
units varies strongly and for some units, especially in the more remote eastern parts
of the country, barely any details are given. All these early maps are based on expert
knowledge and no quantitative data was involved in the process of map making.
The Vegetation Map of Africa then again was the main baseline for the WWF’s
approximation of the world’s terrestrial ecoregions (Olson et al. 2001) in so far as
the African continent was concerned. Although without a presentation of a system-
atic biogeographical database, the map of the terrestrial ecoregions currently consti-
tutes the most used baseline map for strategic conservation planning on a continental
and subcontinental scale (e.g. MacKinnon et al. 2016). The availability of modern
remote sensing techniques has allowed the generation of continental or global land
cover products, i.e. GlobCover, MODIS/Terra Land Cover, GlobLand30 or the map
of African ecosystems by Sayre et al. (2013). However, these maps display struc-
tural vegetation types only and provide no floristic information.
Regional and Local Studies on Vegetation Composition
Early plot based studies were conducted by Ilse von Nolde on the Planalto de Quela
(von Nolde 1938a, b, c). Since the mid-1950s, several local studies based on mis-
sions assessing natural resources were carried out at the regional level in Angola.
Monteiro studied the forest resources in Moxico (Monteiro 1957), in the northern
Maiombe and Dembos forests (Monteiro 1962, 1965a, b, 1967), and in Bié
(Monteiro 1970a) contributing to our knowledge on species composition in the
respective forest types. Monteiro’s (1970a, b) work in Bié needs to be highlighted
as for Angola he implemented new methods in mapping the vegetation. His map of
the woody vegetation of the province of Bié is not drawn based on pure observations
but is based on quantitative vegetation plot data. He collected data on species com-
position in 144 vegetation relevés sized 30 m × 30 m that were subject to a vegeta-
tion classification based on vegetation tables. The mapping process was guided by
aerial photography, quite an advanced approach for its time. Menezes (1965, 1971)
undertook phytosociological studies and produced local vegetation maps in pastoral
ecosystems of the Cunene Province. Teixeira elaborated vegetation maps for two of
the main protected areas of Angola, the Quiçama and Bicuar national parks (Teixeira
et al. 1967; Teixeira 1968). A few years later, Huntley produced a much more
detailed map of the Quiçama National Park in 1972 at a scale of 1:100000 depicting
28 plant communities (Huntley 1972). Aguiar and Diniz (1972) mapped the vegeta-
tion of the western plateau of Cela. Coelho explored the potential of forestry in
Cuando Cubango and elaborated a classification of the lower Cubango Basin into 32
forestry zones (Coelho 1964, 1967). Santos (1982) used a transect approach, so
called ‘itinerários florísticos’, in order to generate an expert-drawn vegetation map
for Cuando Cubango Province (Fig. 6.1).
Modern Approaches to Vegetation Mapping and Classification
This early period of vegetation mapping and classification was followed by the
absence of any such activities for the coming decades due to the long-lasting armed
conflict in the country. During this period significant methodological advances were
made in vegetation ecology and phytosociology as well as in remote sensing tech-
niques. The advent of computers allowed the development of new methodological
tools to semi-automatically classify large amounts of multivariate vegetation plot
data based on objective criteria. As such, vegetation classification moved away from
the subjective assignments of vegetation types to more formalised data analysis.
Similarly, remote sensing imagery became readily available often at no cost and in
unprecedented temporal and spatial resolution. Thus, new numerical methods
together with modern remote sensing products have the potential to provide a much
more detailed and objective picture of vegetation and plant diversity patterns than
expert drawn maps and arbitrarily assigned vegetation types of earlier times
(Fig. 6.2).
15°O 20°O
Cambinda (3)
5°S
Quiçama NP (12) Altiplano da Quela (1)
Quiçama NP (6)
10°S
Planalto occidental da Cela (10)
S76
Bié Province (8)
Moxico (2)
S74 Cusseque (15)
Okavango Basin (16)
Serra da Leba (13) Cubango Basin (18)
15°S
S72 Bicuar NP (7) Western Cuando Cubango (4)
S75
Mucope (9) Cuando Cubango Province (11)
Candelela (14)
S73
Savate (17)
S71 Chitado (5)
1938 - 1945 1960 - 1975 2002 - 2018

km
1945 - 1960 1975 - 2002 0 100 200 300 400 500
Biodiversity Observatories
Fig. 6.1 Location of regional and local studies on vegetation composition, vegetation classifica-
tion or vegetation mapping approaches according to the year the study was published. The country-
wide maps by Gossweiler and Mendonça (1939), Barbosa (1970) and Diniz (1973) are not
depicted. (1) von Nolde 1938a, b, c (2) Monteiro 1957 (3) Monteiro 1962 (4) Coelho 1964 (5)
Menezes 1965 (6) Teixeira et al. 1967, Huntley 1972 unpublished (7) Teixeira 1968 (8) Monteiro
1970a (9) Menezes 1971 (10) Diniz and Aguiar (1968) (11) dos Santos 1982 (12) De Bruyn and
Eberle 2001 (13) Cardoso et al. 2006 (14) Revermann and Finckh 2013a (15) Revermann et al.
2013, Schneibel et al. 2013, Gonçalves et al. 2017 (16) Revermann and Finckh 2013b, Stellmes
et al. 2013 (17) Wallenfang et al. 2015 (18) Revermann 2016, Revermann et al. 2018a (19)
Chisingui et al. 2018. Furthermore, the six biodiversity observatories installed by the SASSCAL
project are shown: Espinheira (S71), Tundavala (S72), Candelela (S73), Cusseque (S74), Bicuar
National Park (S75), Cameia National Park (S76)
Fig. 6.2 Maps for the Okavango Basin located in southeast Angola and extending into Namibia
and northern Botswana. (a) shows the ecoregions as defined by Olson et al (2001) which are
largely based on the vegetation maps of Barbosa (1970) and White (1983), (b) Vegetation map
produced for the same area by The Future Okavango project based on unsupervised classification
of land surface phenology metrics derived from 16–day MODIS EVI time series from the years
2000–2011 (Stellmes et al. 2013) and interpreted using the information of vegetation plots stored
in the vegetation database of the Okavango Basin (Revermann and Finckh 2013b; Revermann et al.
2016a). For an explanation of the vegetation units depicted in the maps please refer to the original
publications
Recent years have seen increasing activity in the investigation of vegetation pat-
terns at the local and regional scale. During the years 1995–2002 some vegetation
surveys were carried out in the Quiçama National Park south of Luanda, for which
the map elaborated by Huntley in the year 1972 served as a baseline. The activities
aimed at gathering data for the re-establishment of the national park and to develop
management strategies (Jeffery et al. 1996). De Bruyn and Eberle (2001) studied a
small fenced of area in the north of the park where they collected 74 relevés and
identified four plant communities including eight subcommunities. Additional
quantitative data was collected to investigate grazing and browsing capacities.
Cardoso et al. (2006) studied the vegetation communities along the steep altitudinal
gradient of the Serra da Leba near Lubango.
Within The Future Okavango (TFO, www.future-okavango.org) project detailed
investigations have been carried out in the Okavango (Cubango) River Basin. The
project team assembled a vegetation database containing vegetation relevé data on
all terrestrial vegetation types within the Okavango Basin (Revermann et al. 2016a).
The plot design followed the standards implemented for woodland vegetation in the
southern neighbour countries, i.e. a nested plot design of one small 10 m × 10 m plot
in a large 20 m × 50 m plot (Strohbach 2001; Jürgens et al. 2012). Based on this
data, classifications for local study sites based on numerical classification approaches
have been published (Revermann and Finckh 2013a; Wallenfang et al. 2015) and a
first classification of the terrestrial vegetation of the entire Cubango Basin was elab-
orated (Revermann et al. 2018a).
The vegetation database of the Okavango Basin was also the foundation to pro-
duce a first vegetation map based on quantitative ground data for the Okavango
Basin (Fig. 6.2b Revermann and Finckh 2013b; Stellmes et al. 2013) and allowed
modelling the α-diversity of vascular plants for the same region (Revermann et al.
2016b).
Based on vegetation relevés various studies have investigated the impact of land
use on vegetation (Revermann et al. 2017) and studied the regeneration of the
vegetation after land use had ceased (Wallenfang et al. 2015; Gonçalves et al. 2017,
2018).
Presently a number of vegetation classification and mapping initiatives are
underway in the framework of the research project SASSCAL (Southern African
Science Service Centre for Climate Change and Adaptive Land Management). For
a compilation of project outcomes see Revermann et al. 2018b, e.g. in the Huíla
Province (Chisingui et al. 2018) and along the coastal plain from the Cunene River
to Benguela including Iona National Park (Jürgens et al. in prep.). The same project
includes six newly implemented biodiversity observatories (http://www.sasscalob-
servationnet.org/), depicted on Fig. 6.1. The standardised monitoring of the 1 km2
sites (Jürgens et al. 2012) will allow the long term monitoring of changes in plant
species composition and plant diversity. Zigelski et al. (2018) present first analyses
of the data gathered on such a biodiversity observatory in the Cameia National Park.
Outlook: A Call for a New Vegetation Survey of Angola
Vegetation and natural resources in general are under strong pressure from the
increasing demands of a growing population and the transition from traditional life-
styles to modern consumerism (cf. Pröpper et al. 2015). The main drivers of defor-
estation and degradation of woodlands and the general loss of pristine vegetation
cover in Angola are the clearing of new fields for shifting cultivation, industrialised
agricultural schemes and the production of charcoal (Cabral et al. 2010; Hansen
et al. 2013; Schneibel et al. 2013, 2016, 2018; Röder et al. 2015; Wallenfang et al.
2015; Mendelsohn 2019). Without adequate knowledge of the spatial distribution
and extent of vegetation types, their species composition and the environmental
drivers of vegetation patterns (climate, geology, soils, landuse) sound landuse man-
agement is not feasible. Thus, a nationwide vegetation survey based on quantitative,
plot level data is urgently needed. Combined with remote sensing data and ecologi-
cal modelling tools an accurate vegetation map can be produced serving the needs
of conservationists, planners, entrepreneurs and scientists alike. A successful vege-
tation survey however relies on good taxonomic knowledge, current flora compen-
dia and plant identification guides. Functioning and strengthened herbaria are also
of great importance building the capacity of the future generation of field ecologists
and environmental scientists.
References
Aguiar FQB, Diniz AC (1972) Carta de Vegetação do Planalto Ocidental da Cela: Estudo
Interpretativo. Série Científica N° 26. Instituto de Investigação Agronómica de Angola, Nova
Lisboa
Barbosa LAG (1971) Phytogeographical map of Angola. Mitteilungen der Botanischen
Staatssammlung München 10:114–115
Brockmann-Jerosch H, Rübel E (1912) Die Einteilung der Pflanzengesellschaften nach okologisch-
physiognomischen Gesichtspunkten. Engelmann, Leipzig, 72 pp
Cabral AIR, Vasconcelos MJ, Oom D et al (2010) Spatial dynamics and quantification of defores-
tation in the central-plateau woodlands of Angola (1990–2009). Appl Geogr 31(3):1185–1193
Cardoso J, Duarte M, Costa E et al (2006) Communidades vegetais da Serra da Leba. In: Moreira
I (ed) Angola: Agricultura, Recursos Naturais e Desenvolvimento. ISA Press, Lisbon,
pp 205–223
Chisingui AV, Gonçalves FMP, Tchamba JJ et al (2018) Vegetation survey of the woodlands of
Huíla province. In: Revermann R, Krewenka KM, Schmiedel U et al (eds) Climate change and
adaptive land management in southern Africa – assessments, changes, challenges, and solu-
tions, Biodivers & Ecol, vol 6. Klaus Hess Publishers, Göttingen & Windhoek, pp 426–437
Coelho H (1964) Contribuição para o Conhecimento da Composição Florística e Possibilidades
de uma Zona Compreendida entre os rios Cubango, Cueio e Quatir. Agronomia Angolana
20:49–82
Coelho H (1967) Zonagem Florestal do Distrito do Cuando-Cubango. Primeiros elementos.
De Bruyn PJN, Eberle D (2001) An ecological study of the plant communities of the fenced sector
of the Quiçama National Park, Angola, with Management Recommendations. B.Sc. (Hons)
Thesis. University of Pretoria, Pretoria
Diniz AC (1973) Características mesológicas de Angola. Missão de Inquéritos Agrícolas de
Angola, Nova Lisboa, 482 pp
Diniz AC, Aguiar FB (1968) Regiões Naturais de Angola. Série Científica N° 2. Instituto de
Investigação Agronómica de Angola, Nova Lisboa, 6 pp + 1 map
Gago MM, Macedo M, Castelo C (2016) Surveying Angola, São Tomé and Timor: experts and
transnational practices. In: Serrão JV, Freire D, Fernández L (eds) Old and new worlds: the
global challenges of rural history, Conference eBook. ISCTE – Instituto Universitário de
Lisboa. Centro de Investigação e Estudos de Sociologia, Lisboa
Gonçalves FMP, Revermann R, Gomes AL, et al. (2017) Tree species diversity and composition of
Miombo woodlands in south-central Angola, a chronosequence of forest recovery after shifting
cultivation. Int J For Res 2017(6202093), 13 pp.
Gonçalves FMP, Revermann R, Cachissapa MJ, et al (2018) Species diversity, population structure
and regeneration of woody species in fallows and mature stands of tropical woodlands of SE
Angola. J For Res. Published online 13 January 2018
Lisbon, 242 pp
Hansen MC, Potapov PV, Moore R et al (2013) High-resolution global maps of 21st-century forest
cover change. Science 342(6160):850–853
Huntley BJ (1972) Parque Nacional da Quiçama. Carta da Vegetação, 1° Aproximação Julho
1972. Ecologist’s report 22. Repartição Técnica da Fauna, Serviços de Veterinária, Luanda,
Mimeograph report
Jeffery RF, van der Waal C, Radloff F (1996) An ecological evaluation with management guide-
lines for the re-establishment of the Quiçama National Park, Angola. B.Sc. (Hons) thesis.
University of Pretoria, Pretoria
Jessen O (1936) Reisen und Forschungen in Angola. Dietrich Reimer Verlag, Berlin, 397 pp
Jürgens N, Schmiedel U, Haarmeyer DH et al (2012) The BIOTA biodiversity observatories in
Africa-a standardized framework for large-scale environmental monitoring. Environ Monit
Assess 184(2):655–678
MacKinnon J, Aveling C, Olivier R et al (2016) Inputs for an EU strategic approach to wildlife
conservation in Africa – regional analysis. European Commission, Directorate-General for
International Cooperation And Development, Brussels, 494 pp
Cham
Menezes JA (1965) Estudo fitosociológico e características das pastagens da região do Chitado.
Boletim do Instituto de Investigação Científica de Angola 2(2):137–181
Menezes JA (1971) Estudo fitoecológico da região de Mucope e carta da vegetação. Boletim
Instituto de Investigação Científica de Angola 8(2):7–54
Monteiro RFR (1957) Aspectos da exploração florestal no distrito do Moxico. Garcia de Orta
5(1):129–146
Monteiro RFR (1962) Le massif forestier du Mayombe angolais. Revue Bois et Forêts des
Tropiques 82:3–17
Monteiro RFR (1965a) A formação florestal dos Dembos. Boletim do Instituto de Investigação
Científica de Angola 2(1):71–82
Monteiro RFR (1965b) Correlação entre as florestas do Maiombe e dos Dembos. Indicação de fac-
tores predominantes. Boletim do Instituto de Investigação Científica de Angola 1(2):257–265
Monteiro RFR (1967) Essências Florestais de Angola. Estudo das Suas Madeiras. Espécies do
Maiombe. Instituto de Investigação Científica de Angola, Luanda
Monteiro RFR (1970a) Estudo da Flora e da Vegetação das Florestas Abertas do Plantalto do Bié.
Instituto de Investigação Científica de Angola, Luanda, 352 pp
Monteiro RFR (1970b) Alguns Elementos de Interese Ecológico da Flora Lenhosa do Planalto do
Bié (Angola). Instituto de Investigação Científica de Angola, Luanda, 166 pp
Olson DM, Dinerstein E, Wikramanayake ED et al (2001) Terrestrial ecoregions of the world: a
new map of life on earth. Bioscience 51(11):933–938
Pröpper M, Gröngröft A, Finckh M et al (2015) The future Okavango – findings, scenarios and rec-
ommendations for action. Research project final synthesis report 2010-2015. Hamburg, 190 pp
Revermann R (2016) Analysis of vegetation and plant diversity patterns in the Okavango basin at
different spatial scales – integration of field based methods, remote sensing information and
ecological modelling. PhD thesis, University of Hamburg Hamburg, 295 pp
Revermann R, Finckh M (2013a) Caiundo – vegetation. Biodivers Ecol 5:91–96
Revermann R, Finckh M (2013b) Okavango basin – vegetation. Biodivers Ecol 5:29–35
Revermann R, Gomes A, Gonçalves FM et al (2013) Cusseque – vegetation. Biodivers Ecol
5:59–63
Revermann R, Gomes AL, Gonçalves FM et al (2016a) Vegetation database of the Okavango basin.
Phytocoenologia 46(1):103–104
Revermann R, Finckh M, Stellmes M et al (2016b) Linking land surface phenology and vegetation-
plot databases to model terrestrial plant alpha diversity of the Okavango Basin. Remote Sens
8:370
Revermann R, Wallenfang J, Oldeland J et al (2017) Species richness and evenness respond to
diverging land-use patterns – a cross-border study of dry tropical woodlands in southern Africa.
Afr J Ecol 55:152–161
Revermann R, Oldeland J, Gonçalvess FM et al (2018a) Dry tropical forests and woodlands of the
Cubango Basin in southern Africa – first classification and assessment of their woody species
diversity. Phytocoenologia 48(1):23–50
Revermann R, Krewenka KM, Schmiedel U et al (eds) (2018b) Climate change and adaptive land
management in southern Africa – assessments, changes, challenges, and solutions. Biodivers
Ecol 6:1–497
Röder A, Pröpper M, Stellmes M et al (2015) Assessing urban growth and rural land use transfor-
mations in a cross-border situation in northern Namibia and southern Angola. Land Use Policy
42:340–354
Santos RM (1982) Itinerários Florísticos e Carta da Vegetacão do Cuando Cubango. Instituto de

Investigação Científica Tropical, Lisbon, 265 pp
Sayre R, Comer P, Hak J et al (2013) A new map of standardized terrestrial ecosystems of Africa.
Association of American Geographers, Washington, DC, 24 pp
Schneibel A, Stellmes M, Revermann R et al (2013) Agricultural expansion during the post-civil
war period in southern Angola based on bi-temporal Landsat data. Biodivers Ecol 5:311–320
Schneibel A, Stellmes M, Röder A et al (2016) Evaluating the trade-off between food and timber
resulting from the conversion of Miombo forests to agricultural land in Angola using multi-
temporal Landsat data. Sci Total Environ 548-549:390–401
Schneibel A, Röder A, Stellmes M et al (2018) Long-term land use change analysis in south-
central Angola. Assessing the trade-off between major ecosystem services with remote sensing
data. Biodivers Ecol 6:360–367
Shaw HKA (1947) The vegetation of Angola. J Ecol 35(1):23–48
Stellmes M, Frantz D, Finckh M et al (2013) Okavango basin – earth observation. Biodivers Ecol
5:23–27
Strohbach BJ (2001) Vegetation survey of Namibia. J Namibia Sci Soc 49:93–124
Teixeira JB (1968) Parque Nacional do Bicuar. Carta da vegetação (1ª aproximação) e Memória
Descritiva. Instituto de Investigação Agronómica de Angola, Nova Lisboa
Teixeira JB, Matos GC, Sousa JNB (1967) Parque Nacional da Quiçama. Carta da Vegetação e
Memória Descritiva. Instituto de Investigação Agronómica de Angola, Nova Lisboa
UNESCO/AETFAT/UNSO (1981) Vegetation map of Africa – scale 1:5 000 000. In: White F (ed)
UNESCO, Paris
von Nolde I (1938a) Probeflächen verschiedener Savannenformationen im Hochland von Quela
in Angola. Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem 14:298–311
von Nolde I (1938b) Probeflächen verschiedener Waldformationen aus dem Hochland von Quela
in Angola. Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem 14:483–486
von Nolde I (1938c) Botanische Studie über das Hochland von Quela in Angola. Feddes
Repertorium Beihefte 101:35
woodlands in southeast Angola. Trop Conserva Sci 8(4):863–892
White F (1976) The underground forests of Africa: a preliminary review. Gard Bull Singapore
11:57–71
White F (1983) The vegetation of Africa – a descriptive memoir to accompany the Unesco/
AETFAT/UNSO vegetation map of Africa. UNESCO, Paris, 356 pp
Zigelski P, Lages F, Finckh M (2018) Seasonal changes of biodiversity patterns and habitat condi-
tions in a flooded savanna – the Cameia national park biodiversity observatory in the upper
Zambezi catchment, Angola. In: Revermann R, Krewenka KM, Schmiedel U et al (eds) Climate
change and adaptive land management in southern Africa – assessments, changes, challenges,
and solutions, Biodivers Ecol, vol 6. Klaus Hess Publishers, Göttingen & Windhoek, pp
438–447
Chapter 7
Suffrutex Dominated Ecosystems
in Angola
Paulina Zigelski, Amândio Gomes, and Manfred Finckh
Abstract A small-scale mosaic of miombo woodlands and open, seasonally inun-

dated grasslands is a typical aspect of the Zambezian phytochorion that extends into
the eastern and central parts of Angola. The grasslands are home to so-called ‘under-
ground trees’ or geoxylic suffrutices, a life form with massive underground wooden
structures. Some (but not all) of the geoxylic suffrutices occur also in open woodland
types. These iconic dwarf shrubs evolved in many plant families under similar envi-
ronmental pressures, converting the Zambezian phytochorion into a unique evolu-
tionary laboratory. In this chapter we assemble the current knowledge on distribution,
diversity, ecology and evolutionary history of geoxylic suffrutices and suffrutex-
grasslands in Angola and highlight their conservation values and challenges.
Keywords Endemism · Geoxyles · Miombo · Phytochorion · Underground forests

· Vegetation
Introduction
Open grassy vegetation is a common aspect of Angolan landscapes and is a charac-

teristic part of the Zambezian phytochorion. Grasses are the most conspicuous ele-
ment of these landscapes towards the end of the rainy season, whereas at the onset
of the rainy season many woody species of so called geoxylic suffrutices or ‘under-
ground trees’ (Davy 1922; White 1976) dominate the aspect of the vegetation. Thus,
in vast areas of central and eastern Angola, the open ‘grasslands’ are de-facto
co-dominated by grasses and geoxylic suffrutices. Closely intertwined with miombo
P. Zigelski (*) · M. Finckh

e-mail: paulina.meller@gmx.de; paulina.meller@studium.uni-hamburg.de;
manfred.finckh@uni-hamburg.de
A. Gomes
Faculty of Sciences, Agostinho Neto University, Luanda, Angola
e-mail: amandiogomes2@hotmail.com

https://doi.org/10.1007/978-3-030-03083-4_7
110 P. Zigelski et al.
woodlands and with wetlands, suffrutex-grasslands constitute one of the main and
most particular ecosystem types of Angola. According to Mayaux et al. (2004), they
cover at least 70,080 km2 or 5.6% of the Angolan territory (not including the small
scale woodland suffrutex-grassland mosaics of the central Angolan plateau).
The geoxylic suffrutex life form is marked by proportionally massive under-
ground woody organs, in literature often termed as lignotuber, xylopodia or woody
rhizomes. Annual shoots sprout readily from the buds on these perennial woody
organs, bearing leaves, inflorescences and fruits before they die back after the end
of the rainy season. Coexistence of grasses and suffrutices is made possible by
occupation of different ecological niches together with phase-delayed activity peri-
ods (i.e. main assimilation/flowering/fruiting time) that reduces competition.
Exploration of Geoxylic Grasslands
The first authors who indicated the distribution and ecological particularity of
suffrutex-grasslands in Angola were Gossweiler and Mendonça (1939), who classi-
fied them as heathland-like woodlands (‘Ericilignosa’). They already noted the
main differentiation between the Cryptosepalum spp. dominated suffrutex commu-
nities (‘Anharas de Ongote’) on ferralitic and psammoferralitic soils and the vegeta-
tion types characterised by Parinari capensis and the Apocynaceae Landolphia
thollonii and L. camptoloba on leached sandy soils (‘Chanas da Borracha’). They
had also already observed the strong thermic oscillations of which at least the
‘Anharas de Ongote’ are subject (see below) and commented on the generative
cycle of Cryptosepalum maraviense from flowering to fruiting in the dry season
(and thus, being inverse to the generative cycle of the C4-grasses).
Using a different mapping and classification approach, typical suffrutex-
grasslands mostly on sandy soils were again mapped and described by Barbosa
(1970) as ‘Chanas da Borracha’ (alluding to the presences of species of the genus
Landolphia), ‘Chanas da Cameia’, and ‘Anharas do Alto’. The Cryptosepalum spp.
dominated ‘Anharas de Ongote’ on ferralitic soils are described (but not depicted on
the map) as being inserted in the main miombo types of the Angolan plateau.
However, he describes the typical spatial pattern, i.e. how they appear close to the
headwaters of the small tributaries and then follow the watercourses in narrow or
broad fringes downstream. Gossweiler and Mendonça (1939) as well as Barbosa
(1970), treated these ecosystems as particular site specific plant communities
closely linked to woodland ecosystems, and not as grass-dominated savannas.
White (1983), however, mapped and described only the sandy ‘Chanas’ as
‘Kalahari and dambo-edge suffrutex grassland’ in the context of the ‘Zambezian
edaphic grassland’, but did not refer to the ‘Anharas de Ongote’ which constitute a
key (but small scale) element of the miombo ecosystems of the Angolan Plateau.
Even in his prominent suffrutex review, White (1976) focuses solely on the ‘Chanas’
in the range of the Zambezi Graben and neither mentions (psammo-) ferralitic
7 Suffrutex Dominated Ecosystems in Angola 111
‘Anharas’, nor lists their dominant key species Cryptosepalum maraviense and C.
exfoliatum ssp. suffruticans in his suffrutex list. He certainly recognises a transition
zone between Zambezian and Guineo-Congolian floras that spans over central and
northern Angola (where the ‘Anharas’ are included) (White 1983). However, he did
not recognise the importance and floristic singularity of the ferralitic suffrutex-
grasslands dominated by Cryptosepalum spp.
Suffrutex Flora and Endemism
The suffrutex life form appears in many different floristic groups and obviously
evolved convergently. A similar center of geoxyle diversity has been reported from
the Brazilian Cerrado. Today, 198 species from 40 families are listed for the western
Zambezian phytochorion (White 1976; Maurin et al. 2014, own data), but an even
higher number is expected as floristic exploration of the region is still poor and new
species might be found (see Goyder and Gonçalves 2019). In some cases suffrutices
are considered a dwarf variety or subspecies of a closely related tree species (e.g.
Gymnosporia senegalensis var. stuhlmanniana, Syzygium guineense ssp. huillense)
and hence classified as such and not as one species, although the genetic relatedness
between tree and dwarf form is rarely investigated. On the other hand, not all dwarf
forms are obligate suffrutices; some can facultatively outgrow the dwarf state if
protected from environmental stressors (White 1976), for instance Oldfieldia dacty-
lophylla or Syzygium guineense ssp. macrocarpum (Zigelski et al. 2018).
Within the suffrutex communities of the Zambezian phytochorion, the Rubiaceae
have the highest number of described taxa (46), followed by Anacardiaceae (22) and
Lamiaceae (14). Table 7.1 lists all families with known geoxylic suffrutex taxa
occurring in Angola and gives examples of common geoxyles for each family.
Furthermore, Fig. 7.1 shows some examples and aspects of suffrutex species given
in Table 7.1. The unique Zambezian geoxylic flora with a high number of endemic
species (Brenan 1978; White 1983; Frost 1996) is a consequence of challenging
environmental conditions, as illustrated further below. According to Figueiredo and
Smith’s catalogue of Angolan plants (2008) and our list of suffrutices (Table 7.1),
121 of the 198 suffrutex species occurring in the Zambezian phytochorion are
known from Angola (61%). Of these 121 species 12 are endemic to Angola (10%).
nvironmental Conditions of Suffrutex-Grasslands

E
Through the Year
The substrate strongly influences the species composition of the suffrutex-

grasslands. In Angola geoxylic suffrutices occur on (a) well-drained arenosols
which are found as seasonally flooded savannas in the Zambezi Graben of the
Table 7.1 List of plant families with geoxylic suffrutices in the Zambezian phytochorion
Plant family N° Species common in Angola Angolan endemics
Rubiaceae 46 Pygmaeothamnus zeyheri (Sond.) Robyns, 2, e.g. Leptactina
Pachystigma pygmaeum (Schltr.) Robyns prostrata
Anacardiaceae 22 Lannea edulis (Sond.) Engl., Rhus arenaria 3, e.g. Lannea
Engl. gossweileri
Lamiaceae 14 Clerodendrum ternatum Schinz, Vitex
madiensis ssp. milanjensis (Britten) F.White
Fabaceae- 13 Erythrina baumii Harms, Abrus 3, e.g. Adenodolichos
Papilionioideae melanospermum ssp. suffruticosus Hassk. mendesii
Proteaceae 11 Protea micans ssp. trichophylla (Engl. & 1, Protea paludosa
Gilg) Chisumpa & Brummitt (Hiern) Engl.
Ochnaceae 9 Ochna arenaria De Wild. & T. Durand,
Ochna manikensis De Wild.
Passifloraceae 7 Paropsia brazzaeana Baill.
Fabaceae- 6 Cryptosepalum maraviense Oliv., C.
Detarioideae exfoliatum ssp. suffruticans (P.A.Duvign.)
Apocynaceae 5 Chamaeclitandra henriquesiana (Hallier f.) 1, Landolphia
Pichon gossweileri
Ebenaceae 5 Diospyros chamaethamnus Mildbr, Euclea
crispa (Thunb.) Gürke
Celastraceae 4 Gymnosporia senegalensis var.
stuhlmanniana Loes.
Dichapetalaceae 4 Dichapetalum cymosum (Hook.) Engl.
Fabaceae- 4 Entada arenaria Schinz
Caesalpinioideae
Myrtaceae 4 Syzygium guineense ssp. huillense, (Hiern)
F. White Eugenia malangensis (O.Hoffm.)
Nied.
Tiliaceae 4 Grewia herbaceae Hiern
Combretaceae 3 Combretum platypetalum Welw. ex M. A. 2, e.g. Combretum
Lawson argyrotrichum
Euphorbiaceae 3 Sclerocroton oblongifolius (Müll.Arg.) Kruijt
& Roebers
Loganiaceae 3 Strychnos gossweileri Exell
Annonaceae 2 Annona stenophylla ssp. nana (Exell)
N. Robson
Apiaceae 2 Steganotaenia hockii (C. Norman)
C. Norman
Chrysobalanaceae 2 Parinari capensis Harv., Magnistipula sapinii
De Wild.
Meliaceae 2 Trichilia quadrivalvis C.DC.
Moraceae 2 Ficus pygmaea Welw. ex Hiern
Myricaceae 2 Morella serrata (Lam.) Killick
Phyllanthaceae 2 Phyllanthus welwitschianus Müll.Arg.
Ranunculaceae 2 Clematis villosa DC.
(continued)
Plant family N° Species common in Angola Angolan endemics
Achariaceae 1 Caloncoba suffruticosa (Milne-Redh.) Exell
& Sleumer
Anisophyllaceae 1 Anisophyllea quangensis Engl. ex Henriq.
Clusiaceae 1 Garcinia buchneri Engl.
Dilleniaceae 1 Tetracera masuiana De Wild. & T. Durand
Fabaceae- 1 Bauhinia mendoncae Torre & Hillc.
Caesalpinioideae
Hypericaceae 1 Psorosperum mechowii Engl.
Ixonanthaceae 1 Phyllocosmus lemaireanus (De Wild. &
T. Durand) T. Durand & H. Durand
Lecythidaceae 1 Napoleonaea gossweileri Baker f.
Linaceae 1 Hugonia gossweileri Baker f. & Exell
Malpighiaceae 1 Sphedamnocarpus angolensis (A. Juss.)
Planch. ex Oliv.
Malvaceae 1 Hibiscus rhodanthus Gürke
Melastomaceae 1 Heterotis canescens (E. Mey. ex Graham)
Jacq.-Fél.
Picrodendraceae 1 Oldfieldia dactylophylla (Welw. ex Oliv.)
J.Léonard
Rhamnaceae 1 Ziziphus zeyheriana Sond.
Urticaceae 1 Pouzolzia parasitica (Forssk.) Schweinf.
N°: overall number of Suffrutex species in the Zambezian phytochorion; examples of species
occurring in Angola are given for each family. Compilation of families and species according to
White (1976), Maurin et al. (2014) and own data
Moxíco province or as sandy alluvial deposits on fossil river terraces along the val-
leys of the southern slopes of the Angolan plateau (Fig. 7.2a); (b) on psammo- fer-
ralitic plinthisols as they frequently occur on the Bíe Plateau in central Angola. The
suffrutex-grasslands on ferralitic soils mostly occur on mid- and foot-slopes and are
embedded within a matrix of miombo woodland (Fig. 7.2b).
Environmental conditions in suffrutex-grasslands change dramatically through-
out the year. The most perceived stresses are man-made fires in the dry season
(May–October) which are mostly deployed to induce resprouting for livestock fod-
der or to facilitate hunting (Hall 1984). Depending on fire intensity, which in turn
depends mostly on fuel load, ambient temperature and wind (Govender et al. 2006),
such fires can completely burn unprotected aboveground biomass.
Another abiotic stress occurring mostly in the early dry season (June–August) is
nocturnal frost, peaking immediately before sunrise. At this time of year masses of
cold dry air from southern latitudes intrude into south-central Africa (Tyson and
Preston-Whyte 2000). As depressions accumulate confluent cold air, the undulating
topography of the Angolan highlands facilitates frequent radiation frost especially
in valleys (Revermann and Finckh 2013; Finckh et al. 2016). Up to 44 frost events
per dry season (with a minimum temperature of −7.5 °C) were recorded by Finckh
et al. (2016), with a temperature span of up to 40 degrees within 12 h. Most woody
Fig. 7.1 Common Angolan suffrutex species. (a) Ochna arenaria (Ochnaceae), fruiting and grow-
ing on sandy sediments of the Bíe Plateau. (b) Syzygium guineense ssp. huillense (Myrtaceae)
flowering in the dry season and growing on sandy soils of the Bíe Plateau. (c) Lannea edulis
(Anacardiaceae), bearing edible fruits, growing on Kalahari sands in southeast Angola. (d)
Hibiscus rodanthus (Malvaceae), growing on Kalahari sands in southeast Angola and flowering in
the rainy season. (e) Landolphia gossweileri (Apocynaceae), typical element of the ‘Chanas da
Borracha’, growing on sandy soils of the Bíe Plateau and bearing edible fruits. (f) Phyllanthus
welwitschianus (Phyllanthaceae), growing on sandy soils of the Bíe Plateau and flowering in the
rainy season. (g) Cryptosepalum exfoliatum ssp. suffruticans (Fabaceae – Detarioideae) with exca-
vated rootstocks, typical element of the ‘Anharas de Ongote’, growing on psammoferralitic soils of
the Bíe Plateau. (h) Parinari capensis (Chrysobalanaceae), typical element of the ‘Chanas da
Borracha’, growing on slightly elevated termite mounds in flooded savannas of the Cameia
National Park, Moxico Province
Fig. 7.2 Typical geoxylic suffrutex grasslands of Angola. (a) ‘Chanas da Cameia’ in the Cameia
National Park, Moxíco Province, during dry season in June. The slightly elevated termite mounds
provide habitat for several geoxyle species that avoid the low-lying areas that are waterlogged from
January to May. (b) ‘Anharas de Ongote’ in the Sovi Valley on the southern slopes of the Bíe
Plateau, in August. The mid- and footslopes are dominated by suffrutex-grassland with the charac-
teristic reddish and green patches of the fresh leaves of Cryptosepalum maraviense, whereas the
wetlands in the drainage lines are covered mostly by Cyperaceae (background, in dark green)
species from tropical background (including geoxylic suffrutices) are sensitive to

frost, their leaves wilt or their shoots die-off entirely.
The geoxylic suffrutex species seem to be triggered by the destruction of their
shoots by frost and/or fire, as they readily resprout after these disturbances and in
most cases already start flowering in the dry season. The suffrutices therefore have
often already finished their generative cycle when the grasses start to cover them.
The suffrutex-grasslands of the sandy plains in eastern Angola are furthermore
subject to seasonal flooding in the late rainy and early dry season (January–May),
leading, for example, in the Cameia National Park to standing water up to 0.5 m
deep. Whereas grass species dominate the sites which are inundated for several
months, suffrutex species seem to avoid fully waterlogged sites and grow patchily
on slightly elevated termite mounds (Fig. 7.2a) or other well drained sites.
The dominant grass species seem to profit from inundation. Their tufts develop
massively in the middle of the rainy season and they flower and bear fruits through-
out the flooding season (own observations).
nowledge Gaps on the Evolution of the Geoxylic Suffrutices

K
and the Formation of Suffrutex-Grasslands
A common observation within suffrutex ecosystems is the resemblance (Meerts

2017) and assumed close relatedness of suffrutex species to tree species that occur
in forests and woodlands. The indigenous people (e.g. the Chokwe in eastern
Angola) in many cases recognise the similarity and relatedness and use similar local
names for such pairs, for instance Muhaua and Mupaua for the tree and suffrutex
forms of Syzygium guineense Willd. DC. The striking fact that the suffrutex life
form was developed by several plant families independently and at roughly the
same time (Maurin et al. 2014) indicates a common driver that triggered its conver-
gent evolution.
Grassy biomes emerged in Africa in the late Miocene approximately 10 mya
(Cerling et al. 1997; Keeley and Rundel 2005; Herbert et al. 2016). This period is
characterised by global climatic fluctuations which led to cooler, drier conditions, to
a drop of atmospheric CO2 concentrations and particularly to pronounced precipita-
tion seasonality (i.e. wet and dry seasons) in southern Africa (Pagani et al. 1999). As
a consequence, humid tropical forests retreated to more favorable sites further north
and were replaced by more open dry and seasonal tropical forest ecosystems like the
miombo (Bonnefille 2011). In parts where miombo landscapes prevail today, cano-
pies were disrupted and allowed the establishment of open ecosystems embedded in
woodland matrices. These open ecosystems were then rapidly occupied by light-
demanding C4-grasses and the evolving geoxylic suffrutices.
It is still an open discussion why open suffrutex-grasslands are able to persist
within the woodlands (or vice versa). It is however likely that rainfall seasonality
and the above described abiotic stresses that characterise the suffrutex-grasslands
play a major role in their establishment and maintenance (Sankaran et al. 2005;
Staver et al. 2011).
Savanna ecologists tend to see fire as the main driver for grassland formation. On
the one hand frequent fires prevent tree establishment if saplings cannot outgrow the
reach of the flames and are destroyed therein. For woodlands in eastern South
Africa, a fire free time period of at least 5 years is necessary for many tree species
to escape the ‘fire trap’ (Sankaran et al. 2004; Gignoux et al. 2009). This time win-
dow, allowing for successful reestablishment of trees, is rarely achieved in Angolan
grasslands, at least nowadays (Schneibel et al. 2013; Stellmes et al. 2013).
C4-savanna grasses, however, respond positively to periodic burning and resprout
within weeks (Bond and Keeley 2005), thus being able to colonise seasonally burnt
sites.
Forest ecologists, on the other hand, attribute the frequent short duration frost
events in the dry season for preventing tree recruitment in the open areas (Finckh
et al. 2016). As the list of suffrutices (Table 7.1) shows, mainly (but not exclusively)
tropical families or genera evolved suffrutex life forms. Frost is deleterious to most
tropical tree taxa, as they have not developed physiological adaptations to this ‘un-
tropical’ stress factor, thus showing little or no frost tolerance (Sakai and Larcher
2012). As the suffrutex-grasslands are typically situated in particularly frost prone
sites (depressions), tree taxa that are not adapted to frost are being filtered out of
such environments.
In any case, a promising strategy to cope with seasonally returning thermic stress
(by frost or fire) is to protect sensitive organs (buds) by hiding them underground.
Tree species relocated their woody biomass and regenerative buds belowground at
the expense of growth height and were thus able to cope with frost and fire prone
sites (White 1976; Maurin et al. 2014; Finckh et al. 2016). Even shallow soil depths
of less than 10 cm are sufficient to alleviate thermic stresses (Revermann and Finckh
2013). The high number of tropical genera and families that contribute to the suf-
frutex flora show how successful this strategy is for frost sensitive and fire suscep-
tible taxa, in order to survive the adverse conditions of the open grasslands.
Concomitantly other evolutionary advantages of the geoxylic life form have been
discussed, for instance poor edaphic conditions, as favoured by White (1976). He
considered the low nutrient status of the leached and locally seasonal waterlogged
soils on Kalahari sands as a likely cause for the lack of regular trees and the suffru-
tication of them as means of compensation. However, trees as well as suffrutices
often grow on the same or similarly poor soils, with comparable physical and chem-
ical properties (Gröngröft et al. 2013); forests and grasslands are not separated by
edaphic boundaries but follow topographic rather than edaphic logics.
The waterlogging argument on the other hand would imply that the woody
underground organs show adaptations to inundation, for instance aerenchymatic tis-
sue or adventitious roots (Parolin 2008). Anatomical analyses of the rootstocks of
four common suffrutex species however did not provide any support for aerenchy-
matic tissue nor other adaptations to inundation (Sanguino 2015). Moreover, in sea-
sonally flooded savannas suffrutices avoid inundated sites. This is even the case for
Syzygium guineense ssp. huillense, a suffrutex closely related to a tree species that
grows along and in rivers and floodplains (Coates Palgrave 2002; Meerts and Hasson
2016).
To summarise, so far the main environmental driver for the astonishing radiation
of geoxylic suffrutices has not been conclusively identified. The emergence of the
suffrutex grassland at the end of the Pliocene and the peak of radiation at the begin-
ning of the Pleistocene is clearly related to climatic seasonality and pronounced dry
seasons. Dry seasons, however, did not only provide the necessary dry fuel for fire
but also provided the atmospheric conditions for nocturnal frost events – the season-
ality argument, thus, does not tip the balance toward fire or frost.
Conservation Value and Conservation Challenges
Various studies recognise the high floristic singularity of the Zambezian phytocho-
rion and suffrutex-grasslands with its unique life forms contribute prominently to its
high number of endemic species (Clayton and Cope 1980; White 1983). The high
degree of suffrutex-grassland endemics within the Zambezian phytochorion as well
as within Angola is a consequence of a unique setting of environmental drivers like
nutrient poor soils, frequent frosts and fires or precipitation seasonality in a small-
scale heterogeneous landscape (Linder 2001). Thus, the Zambezian phytochorion
can be seen as an evolutionary laboratory that promoted the evolution of many spe-
cialised plant species, e.g. suffrutices, orchids and grasses.
Suffrutex-grasslands are sometimes misunderstood as ‘degraded forests’, over-
looking their naturalness. Through this misconception they are listed as sites for
reforestation in order to recover presumably lost forests and to sequestrate atmo-
spheric CO2 (Parr et al. 2014). However, the well-intentioned act of reforestation
would in fact destroy biodiverse natural ecosystems (Bond 2016). A lack of under-
standing, however, frustrates the development of appropriate conservation measures
for the suffrutex grasslands today and in the future. The rebuilding process in Angola
also has risks, happening at a rapid pace and shaping the landscape to human
demands with limited consideration for sustainable management (Pröpper et al.
2015). Flooded savannas in the Moxíco Province for instance are targeted for large-
scale agro-industrial development (ANGOP 2017). Not even National Parks offer
adequate protection to ecosystems in this area, as the first rice schemes emerged
during 2016 within the limits of Cameia National Park (own observation).
Deficiencies in communication and cooperation between different ministries and
governance levels aggravate such problems.
Outlook
Many questions still remain to be answered around the enigmatic life form of the
geoxylic suffrutices. In order to efficiently safeguard suffrutex-grasslands, we need
to understand the evolutionary drivers and evolutionary processes shaping these
ecosystems. For instance, a thorough understanding of the evolutionary drivers and
the response of suffrutices to them would help to assess how current environmental
conditions affect the Zambezian ecosystems and how landscape shaping processes
work. Moreover, investigations about genetic patterns of suffrutices and close tree-
relatives would give insight to speciation processes, means of propagation (clonal or
sexual) and evolutionary history. Also, ecophysiological or morphological measure-
ments would contribute another perspective from which to assess how suffrutices
react to environmental stresses and change processes. All these facets are currently
the subjects of incipient research.
References
ANGOP (2017) Cameia prepara mais de mil hectares para cultivar arroz. Agência Angola Press,
Moxico, 16.07.2017. http://www.angop.ao/angola/pt_pt/noticias/economia/2017/6/28/
Moxico-Cameia-prepara-mais-mil-hectares-para-cultivar-arroz,be825477-10c7-472d-b6e3-
cb1ed5d25974.html. Accessed 22 Aug 2017
Barbosa LAG (1970) Carta fitogeográfica de Angola. Instituto de Investigação Científica de
Angola, Luanda
Bond WJ (2016) Ancient grasslands at risk. Science 351(6269):120–122
Bond W, Keeley J (2005) Fire as a global ‘herbivore’: the ecology and evolution of flammable
ecosystems. Trends Ecol Evol 20(7):387–394
Bonnefille R (2011) Rainforest responses to past climatic changes in tropical Africa. In: Tropical
rainforest responses to climatic change. Springer, Berlin/Heidelberg, pp 125–184
Brenan JP (1978) Some aspects of the phytogeography of tropical Africa. Ann Mo Bot Gard
65(2):437–478
Cerling TE, Harris JM, MacFadden BJ et al (1997) Global vegetation change through the Miocene/
Pliocene boundary. Nature 389(6647):153–158
Clayton WD, Cope TA (1980) The chorology of Old World species of Gramineae. Kew Bull
35(1):135–171
Coates Palgrave K (2002) Trees of southern Africa, 3rd edn. New edition revised and updated by
Meg Coates Palgrave, Struik, Cape Town, pp 833–836
Davy JB (1922) The suffrutescent habit as an adaptation to environment. J Ecol 10(2):211–219
Figueiredo E, Smith G (eds) (2008) Plants of Angola: Plantas de Angola. Strelitzia 22:1–279
Finckh M, Revermann R, Aidar MP (2016) Climate refugees going underground–a response to
Maurin et al. (2014). New Phytol 209(3):904–909
Frost P (1996) The ecology of miombo woodlands. In: Campbell B (ed) The Miombo in transition:
woodlands and welfare in Africa. CIFOR, Jakarta, pp 11–57
Gignoux J, Lahoreau G, Julliard R et al (2009) Establishment and early persistence of tree seed-
lings in an annually burned savanna. J Ecol 97(3):484–495
Gossweiler J, Mendonça FA (1939) Carta fitogeografica de Angola. Ministério das Colónias,
Lisboa, p 242
Govender N, Trollope WS, Van Wilgen BW (2006) The effect of fire season, fire frequency,
rainfall and management on fire intensity in savanna vegetation in South Africa. J Appl Ecol
43(4):748–758
Goyder DJ, Gonçalves FPM (2019) The Flora of Angola: collectors, richness and endemism. In:
tion: a modern synthesis. Springer
Gröngröft A, Luther-Mosebach J, Landschreiber L et al (2013) Cusseque – soils. Biodivers Ecol
5:51–54
Hall M (1984) Man’s historical and traditional use of fire in southern Africa. In: de Booysen PV,
Tainton NM (eds) Ecological effects of fire in South African ecosystems. Springer, Berlin/
Heidelberg, pp 40–52
Herbert TD, Lawrence KT, Tzanova A et al (2016) Late Miocene global cooling and the rise of
modern ecosystems. Nat Geosci 9(11):843–847
Keeley JE, Rundel PW (2005) Fire and the Miocene expansion of C4 grasslands: miocene C4
grassland expansion. Ecol Lett 8(7):683–690
Linder HP (2001) Plant diversity and endemism in sub-Saharan tropical Africa. J Biogeogr
28(2):169–182
Maurin O, Davies TJ, Burrows JE et al (2014) Savanna fire and the origins of the ‘underground
forests’ of Africa. New Phytol 204(1):201–214
Mayaux P, Bartholomé E, Fritz S et al (2004) A new land-cover map of Africa for the year 2000.
J Biogeogr 31(6):861–877
Meerts P (2017) Geoxylic suffrutices of African savannas: short but remarkably similar to trees.
J Trop Ecol 33(4):1–4
Meerts PJ, Hasson M (2016) Arbres et arbustes du Haut-Katanga. National Botanic Garden of
Belgium
Pagani M, Freeman KH, Arthur MA (1999) Late Miocene atmospheric CO2 concentrations and
the expansion of C4 grasses. Science 285(5429):876–879
Parolin P (2008) Submerged in darkness: adaptations to prolonged submergence by woody species
of the Amazonian floodplains. Ann Bot 103(2):359–376
Parr CL, Lehmann CER, Bond WJ et al (2014) Tropical grassy biomes: misunderstood, neglected,
and under threat. Trends Ecol Evol 29(4):205–213
Pröpper M, Gröngröft A, Finckh M et al (2015) The future Okavango: findings, scenarios and
recommendations for action: research project final synthesis report 2010–2015. University of
Hamburg-Biocentre Klein Flottbek, pp 53–129
Revermann R, Finckh M (2013) Cusseque – micro-climatic conditions. Biodivers Ecol 5:47–50
Sakai A, Larcher W (2012) Frost survival of plants: responses and adaptation to freezing stress, vol
62. Springer, New York, pp 138–173
Sanguino G (2015) Wood anatomy and adaptation strategies of suffrutescent shrubs in South-
Central Angola. MSc. thesis, Universität Hamburg, Hamburg, 68 pp
Sankaran M, Ratnam J, Hanan NP (2004) Tree-grass coexistence in savannas revisited – insights
from an examination of assumptions and mechanisms invoked in existing models. Ecol Lett
7(6):480–490
Sankaran M, Hanan NP, Scholes RJ et al (2005) Determinants of woody cover in African savannas.
Nature 438(7069):846–849
Schneibel A, Stellmes M, Frantz D et al (2013) Cusseque – earth observation. Biodivers Ecol
5:55–57
Staver AC, Archibald S, Levin SA (2011) The global extent and determinants of savanna and forest
as alternative biome states. Science 334(6053):230–232
Stellmes M, Frantz D, Finckh M et al (2013) Okavango basin – earth observation. Biodivers Ecol
5:23–27
Tyson PD, Preston-Whyte RA (2000) Weather and climate of Southern Africa, 2nd edn. Oxford
University Press Southern Africa, Cape Town Chapter 10–12
White F (1976) The underground forests of Africa: a preliminary review. Gard Bull Singap
29:57–71
White F (1983) The Zambezian regional centre of endemism. In: White F (ed) The vegetation of
Africa – a descriptive memoir to accompany the UNESCO/AETFAT/UNSO vegetation map of
Africa. UNESCO, Paris, 356 pp
Zigelski P, Lages F, Finckh M (2018) Seasonal changes of biodiversity patterns and habitat condi-
tions in a flooded savanna – the Cameia National Park Biodiversity Observatory in the Upper
Zambezi catchment, Angola. Biodivers Ecol 6:438–447
Chapter 8
Landscape Changes in Angola
John M. Mendelsohn
Abstract Landscape changes in Angola are dominated by woodland and forest

losses due to clearing for crops, bush fires (which convert woodland into shrubland)
and the harvesting of fuel (as wood and charcoal) and timber. Rates of clearing for
small-scale dryland crops are high over much of Angola as a result of poor soil fer-
tility. Erosion is also a severe problem, which has caused widespread losses of top-
soil, soils nutrients and ground water. Rates of erosion are greatest in areas with
steep slopes, sparse plant cover and high numbers of people, as well as around dia-
mond mines in Lunda-Norte. Patterns of river flow and water quality have been
changed, largely as a result of soil erosion and plant cover loss, as well as large
irrigation schemes and dams. High rates of urban growth and the production of
untreated urban waste have led to large concentrations of contamination around
towns. Further research is needed, for example to assess the environmental impacts
of the fishing and petroleum industries offshore, the effects of large volumes of
urban waste being washed into and down major rivers to the sea, and landscape
changes in an around areas of highland forests and grasslands that support popula-
tions of rare and endemic species.
Keywords Bushmeat · Charcoal · Deforestation · Fire · Land transformation ·

Mining impacts · River flows · Shifting cultivation · Soil erosion · Urbanisation
Introduction
Angola is a developing country, its development occurring in multiple ways in dif-

ferent areas of the country and affecting a variety of natural resources. Some changes
and developments are likely to accelerate as the country seeks to diversify its econ-
omy and reduce dependence on revenues from oil and diamonds. It is also likely that
the changes will contribute to such global trends as loss of biodiversity and land
degradation.
J. M. Mendelsohn (*)
RAISON (Research & Information Services of Namibia), Windhoek, Namibia
e-mail: john@raison.com.na

https://doi.org/10.1007/978-3-030-03083-4_8
124 J. M. Mendelsohn
This brief review provides perspectives and information on changes to Angola’s

terrestrial landscapes, particularly in the southern half of the country. There are
three sections to the chapter, the first of which describes the major kinds of land-
scape change. The second is an account of conditions that drive changes, both ulti-
mately and proximately. Finally, areas most affected by major changes are identified
in the third part of the paper.
Major Changes
Woodland and Forest Loss
Losses of woodland are by far the most obvious and conspicuous of changes in
Angola. Much of this has been due to clearing for small-scale crop farming, particu-
larly of dry-land crops, and large-scale commercial agriculture (including relatively
small areas of exotic tree plantations). Other losses have come from the harvesting
of charcoal, wood fuel, timber production (both for commercial and domestic uses),
and runaway bush fires. On a smaller scale, swathes of riverine forest have been
removed to give miners access to alluvial diamonds in rivers in Lunda-Norte.
As a result of all these losses, large areas of forest and savanna are now grass-
lands or shrublands. For example, the greater part of Huambo and Angola’s central
planalto was originally wooded, and 78.4% of the province of Huambo was covered
in miombo woodland in 2002. In 13 years that figure had dropped in 2015 to 48.3%,
amounting to the loss of some 1.265 million ha, 63.2% of which was converted from
forest to crop land (Palacios et al. 2015). Similar losses in western Cuando Cubango,
eastern Huíla and eastern Huambo have been documented by Schneibel et al.
(2013), and elsewhere in Huíla and the Cuvelai drainage in Cunene (Mendelsohn
and Mendelsohn 2018).
A countrywide perspective on the loss of forest or tree canopy cover is presented
in Fig. 8.1. Several relevant features are visible in this image. First is the open,
deforested expanse stretching southwest to northeast across western Huíla, south-
western Huambo and western Bié. Much of this area of highlands was cleared for
crops between the 1950s and 1970s, although grasslands (anharas do alto) probably
always dominated high altitude areas of the central planalto above about 1900 m
above sea level. Substantial areas were cleared at the same time in parts of Cuanza-
Norte, Cuanza-Sul and Malange, but their boundaries are not easily defined.
Second is the clearing of woodlands around urban areas. Many had already been
cleared of tree cover by 2000, after which clearings expanded as trees were removed
progressively further from the town centres, a trend illustrated by Schneibel et al.
(2018). Examples of recent clearings between 2000 and 2015 are conspicuous as
‘red bands’ around Dundo, Menongue, Luena, Malange, Cafunfo, Cubal and
Caimbambo in Fig. 8.1. Much of the deforestation is clear-felling for dryland fields
by residents, while other trees are removed for charcoal production, wood fuel and
timber.
8 Landscape Changes in Angola 125
% forest cover in 2000

0-20
20-40
40-60
60-80
80-100
forest loss 2000−2015
expanse mostly cleared

Dundo in the mid 20th Century
Cafunfo
Malange clearings along

major road
Luau
Luena
Cazombo
Bié
Cubal
Huambo
Caimbambo
clearings
along major
road
Huíla
Bush fires Menongue
Bush fires
Savate
Fig. 8.1 Forest or tree canopy loss between 2000 and 2015 derived from data described by Hansen
et al. (2013), updated and available from http://earthenginepartners.appspot.com/science-
2013-global-forest. Percentage forest cover in the year 2000 is shown in shades of green. Red areas
are those which, by 2015, had lost all the forest or canopy cover that still remained in 2000.
(Source: Hansen/UMD/Google/NASA)
Third, the concentration of clearings along major roads where many rural fami-
lies choose to settle is visible, but requires closer inspection of Fig. 8.1. Most losses
of tree cover here are also due to clear-felling for dryland crops. Local residents also
produce charcoal on a large scale, particularly along roads frequently travelled by
trucks that can transport large volumes of charcoal to urban markets. However, the
effects of charcoal – and timber – harvesting are seldom visible in satellite images
of tree or canopy cover because harvesters typically remove only larger, taller trees,
leaving smaller trees and shrubs which present a seemingly intact canopy of
oodland when viewed from high above. After some years of regrowth, harvesters
w
return to fell those bigger individual trees that produce good charcoal.
Timber has been harvested on a substantial scale for many years. Most of it has
been used for the construction of domestic homes, palisades and fences, or sold as
exported hardwood. The harvesting of selected species and large individual trees
has evidently increased substantially in recent years, and further increases are to be
expected (ANGOP 2017). Conversely, the use of poles for houses, palisades and
fences may be declining, at least in certain areas where people increasingly build
with home-made or bought bricks, and fence with wire (Calunga et al. 2015).
Trees were evidently harvested in large numbers to fuel railway engines running
between Benguela and Huambo, and perhaps elsewhere, in the early twentieth cen-
tury (Silva 2008). There are also reports of Zambezi teak Baikiaea plurijuga and
Marquesia macroura timber being used for sleepers on the Caminho do Ferro
Moçâmedes (CFM) and Caminho de Ferro Benguela (CFB) lines, respectively,
while indigenous woodlands were cleared to make way for the many eucalyptus
plantations established along the CFB line.
Bush fires have major effects on woodlands, particularly in limiting the growth of
trees and shrubs in savannas. Indeed, fires maintain the ‘balance’ between grass and
trees that characterise savannas. However, hot, intense runaway fires set by people
are seemingly more frequent than before. The fiercest of fires kill all plants, old size-
able trees being burnt and scarred year after year until they eventually die. Large
areas have thus been converted from woodland and forest into shrubland, particu-
larly in southern Angola (Fig. 8.2). Much of Cuando Cubango and parts of Moxico
are mosaics of open woodland separated along sharp margins from dense woodland
and forest. As a probable result of fire, the edges of the dense cover are smoothed
and often rounded, in some cases creating circular patches of forest (Fig. 8.3).
Soil Loss (Bulk and Nutrients)
At least three areas appear to have lost large volumes of soil and soil nutrients. The
first is the central planalto and surrounding higher areas of ferralsol soils. In the
catchment of the Cunene River, erosion has been greatest in areas that are densely
populated, extensively cultivated with dryland crops, largely cleared of plant cover
and that have at least moderate slopes (Fig. 8.4). The catchments of other major riv-
ers (Cuando, Queve, Quicombo, Catumbela, Guvrire and Coporolo, for example)
that drain the central catchment are likewise eroded, particularly where slopes are
steep and plant cover is sparse. Similar, more concentrated effects are seen in cities
where inadequate management of storm water has led to the formation of erosion
gullies, many of them damaging urban roads, houses and other infrastructure.
Second is in Lunda-Norte where open-cast mining leads to considerable volumes
of soil (probably also ferralsols) being washed into rivers that flow north into the
Congo Basin (Fig. 8.5; see Ferreira-Baptista et al. 2018).
Fig. 8.2 An example of woodlands converted by repeated hot fires into shrublands in Bicuar
National Park. The fires normally start in the grassy drainage lines (mulolas) from where they
spread into the surrounding woodlands. With the same areas being burnt by fierce fires every few
years, large areas of woodland (dark greenish zones) have progressively been turned into shrub-
lands (pale areas). These satellite images from Google Earth (LandSat/Copernicus) were taken
between 1984 and 2016, and viewed from about 15.3 South, 14.4 East. The red line marks the
western border of Bicuar National Park. (From Mendelsohn and Mendelsohn 2018)
Fig. 8.3 Patches of open woodland (pale grey areas) and dense miombo forest (dark green)
between the Longa and Sovi rivers in Cuando Cubango. The forest margins have probably been
sharpened and smoothed by bush fires. Isolated blocks are so rounded and reminiscent of the
Namib Desert’s fairy circles that they may be called ‘fairy forests’. (The image was taken from
Google Earth (LandSat/Copernicus) as viewed from about 15.4 South, 18.9 East)
Fig. 8.4 The distribution of erosion gullies in relation to vegetation cover (Enhanced Vegetation
Index – EVI) and population density in the catchment of the Cunene River between Huambo in the
north and Xangongo in the south. (Adapted from Mendelsohn and Mendelsohn 2018)
Fig. 8.5 Mining impacts on Angolan rivers. Top left and right: The confluence of the clear Cassai
River and the turbid Lubembe River carrying suspended sediments from open-cast diamond min-
ing in Lunda-Norte. The confluence is in the DRC about 80 km north of Angola’s border. The left
photograph was taken on 30 May 2007, while the right image from Google Earth was taken
10 years later on 21 May 2017, viewed from 6.62 South, 21.07 East. Bottom left: The confluence
of the Calonga and Cunene rivers at Quiteve (16.02 South, 15.20 East), showing the volumes of
eroded sediments from upstream in the Cunene catchment. By contrast, the clear waters of the
Calonga mainly come from areas where arenosols predominate, where few people live and where
large areas of woodland have not been cleared for dryland agriculture. Bottom right: Erosion from
open-cast mining along the Luachimo River 22 km north of Lucapa. (The image from Google
Earth was taken in May 2017 as viewed from 8.23 South, 20.77 East)
There is a likely net loss of certain soil nutrients in the third area, which is where
bush fires are frequent and/or intense, predominantly so in Cuando Cubango,
Moxico and the Lunda provinces (Figs. 8.6 and 8.7). Fires often result in the loss of
nitrogen, phosphorus and organic carbon, although cooler fires also facilitate the
release of nutrients from plant matter into the soil (Jain et al. 2008). A study com-
paring open and dense woodland near Savate (see Fig. 8.1), found much lower nutri-
ent levels in open than dense woodland soils (Wallenfang et al. 2015). This stark
difference was probably a consequence of the open areas being burnt often and
intensely, while the dense woodlands were seldom burnt (Stellmes et al. 2013).
Fig. 8.6 The frequency of fires expressed as the number of years each area of 500 by 500 m burnt
between 2000 and 2010. (From Archibald et al. (2010) and data available at http://wamis.meraka.
org.za/products/firefrequency-map)
Water Flows and Quality
Discharges and the quality of water have changed significantly in certain rivers, and
in a number of ways. The most obvious changes are in the heavy sediment loads
which impair the functioning of aquatic animals and plants that require well-lit
waters, and reduce the capacity of dams. For example, eroded sediments washed
down the Cunene River have evidently accumulated in Gove and Matala dams to
such an extent that their production of hydro-electricity has declined (António 2017).
Fig. 8.7 Left: the seasonality of fires, reflected by the average period of the year when fires were
recorded. Right: the average size of fires. (From Archibald et al. (2010) and data available at http://
wamis.meraka.org.za/products/firefrequency-map)
River flows, soil moisture levels and groundwater recharge have been affected by
losses of plant cover. Sheets of surface flows after heavy rain have increased in bare
areas, causing higher river flows and probabilities of flooding, especially in seasons
with above average rainfall. For example, the clearing of plant cover in the catch-
ment of the Guvrire River around Caimbambo and Cubal (Fig. 8.1) is considered to
have increased the risk and frequency of flooding at the river mouth in the city of
Benguela (Development Workshop 2016).
A different impact of plant cover loss and erosion may affect the Cuvelai. Many
residents there believe that surface flows down the floodplains (chanas) are now
slower and wider than before because eroded sediments deposited in the shallow
channels have further reduced their depths and slopes (Calunga et al. 2015).
Reductions in plant cover result in lower volumes of rain water being trapped or
impeded, thus reducing seepage into the top soil to replenish soil moisture and
recharge local aquifers. With lower soil moisture, seepage to sustain river flows dur-
ing the dry season also declines. This is a likely – and at least partial – explanation for
flows of the Cunene River at Ruacana dropping to less than 10 cubic metres/second
in September 2017. Such low levels were only recorded previously during extreme
drought years in 1993–1994 and 1994–1995 (Mendelsohn and Mendelsohn 2018).
Contamination
Quantitative assessments of the magnitude of environmental contamination from

urban waste are apparently not available for Angola. However, substantial volumes
of waste are generated, particularly in Luanda (now with more than seven million
residents) and other major cities with populations approaching a million or more
people, such as Cabinda, Lubango, Lobito, Huambo and Benguela. Solid waste is
not collected in many middle to low-income bairros, which also lack sewage sys-
tems. The resulting volumes and concentrations of untreated waste from these large
cities have significant impacts on human and environmental health (Development
Workshop 2016).
Drivers of Landscape Change
Population Growth and Natural Resource Exploitation
As elsewhere in the world, but particularly in developing countries, most changes

have been driven by rising demands for natural resources to meet the needs of
Angola’s growing population and increasing consumption per capita. The country’s
population rose from about 6 million people in 1970 to almost 26 million in 2014,
which amounts to an annual growth rate of 3.4%. Over the same period a high pro-
portion of the population shifted from rural to urban areas and their accompanying
economies. Most urban residents live in low income areas where they generally lack
piped water supplies, electricity, secure tenure, sewage systems and solid waste col-
lection services. For example, 85% of Luanda residents live in such areas, with
similar percentages in Lobito (90%), Cabinda (86%) and Benguela (92%)
(Development Workshop 2016). Similar conditions and proportions hold in
Huambo, Malange, Cuito and Ndalatando.
Urban consumption patterns differ from those in rural areas, but one difference
of particular interest concerns the use of fuels for cooking. Rural homes generally
use wood collected from around their homes, whereas the majority of urban people
use purchased charcoal because alternative fuels are more expensive or not available
in towns. Their supplies of charcoal all come from rural areas, in particular from
poor families that harvest and then sell bags of charcoal along roads leading to
major urban areas. The supply of charcoal is therefore an informal one that gener-
ates incomes for many rural homes. For both consumers and suppliers this seems to
be an ideal market, providing affordable fuel for urban consumers and incomes for
rural families, that often being their only monetary income. Similar market arrange-
ments hold for supplies of bush meat from rural suppliers to urban consumers.
Another new, sometimes surprising economic link between urban and rural areas
involves investments in cattle by wealthier town folk. Their cattle (and sometimes
goats and sheep) are placed in rural areas where they are normally tended by
relatives and held as savings or capital, with the best returns coming from large
stock numbers (Gomes 2012). Owners are thus encouraged to have as many animals
as possible, which places added pressures on forage, water and the limited resources
available to poor rural residents (who seldom have other incomes).
Food Production
An abundance of wealth, much of it derived from the boom in oil revenues, has
provided resources to develop large-scale agricultural projects, often with limited or
no environmental impact assessments. For example, several new irrigation schemes
have been developed along the Cunene River. If and when the farms are fully devel-
oped, downstream stretches of the Cunene could be dry for much of the year.
Elsewhere, tens of thousands of hectares of woodland and forest have been cleared
in recent years, one example being the Angola Biocom project which has 70,106 ha
allocated to produce sugar, ethanol fuel and electricity south of Malange (Angola
Biocom 2017).
Clearing for small-scale dryland crop production has caused much of the loss of
woodland and forest in Angola. The rate at which trees are cleared is however driven
by four related, but arguably separate factors. First is the need to feed a growing
number of rural residents. Second is the need for farmers to abandon their fields
after several years of use and to clear new fields (which will produce better yields
than those that have had their supplies of nutrients exhausted). Third is the general
low-input/low-output crop production strategy adopted and adapted for dryland
agriculture, which means that fertilisers are seldom used to replenish soil nutrients.
Fourth is the poor quality of soils available for dryland farming (Ucuassapi and Dias
2006; Asanzi et al. 2006; Wallenfang et al. 2015). Indeed, the relative lack of nutri-
ents and moisture in soils is arguably the most important factor driving the rapid rate
at which Angolan woodlands and forests are cleared, as well as the very slow rate of
recovery.
Grassland and Woodland Fires
Much of Angola’s vegetation has been moulded by frequent fire. This is particularly
true for savanna woodlands, the grasslands of the anharas de ongote in the central
highlands and grassy chanas da borrachas in the Lunda provinces. Most woody
plants in the latter habitats and many in open woodlands are geoxylic suffrutices,
their growth forms adapted to survive frequent hot fires (see Zigelski et al. 2019).
Fire therefore has major impacts on Angola’s vegetation, and any changes in fire
regimes are likely to result in landscape changes. Against that background, and the
widely held assumption that burning has increased in frequency, the following
information is provided on fires in Angola.
Fires are recorded most frequently in grasslands of the Lunda provinces,

Malange, and the Bulozi Floodplains in Moxico, and in open, savanna woodlands in
Cuando Cubango (Fig. 8.6). Additionally, fires are frequent in highland grasslands
(anharas do alto) distributed between Serra da Chela (near Lubango) in the south,
the Benguela, Huambo and Huíla highlands and higher elevations in northern
Cuanza-Sul and southwestern Malange.
Almost all fires are in the dry season between late April and early November.
However, those in northern Angola and the central highlands burn considerably
earlier than in the south (Fig. 8.7). Grass fuel is likely to contain more moisture in
June than later in August, with the likely result that the earlier fires are cooler, less
intense and probably less damaging to vegetation than later, hotter burns in southern
Angola. That seems true for the large fires in the tall grasses that comprise the wide-
spread chanas da borrachas in the Lunda provinces (Huntley 2017). Similar trends
were found by Stellmes et al. (2013) within the river basins of the Cubango and
Cuito Rivers where fires in the northern catchment areas were both earlier and less
intense than those in the south. That trend also roughly corresponded to land cover,
with dense miombo woodland in the northern areas and open savanna woodland,
often called Baikiaea-Burkea woodland in the southern zones.
Areas of Major and Widespread Landscape Change
Figure 8.8 provides perspectives on the distribution of the major landscape changes
described in this chapter. The majority of changes are in and around the central
plateau (planalto) and to the north in parts of Cuanza-Norte, Bengo and Uíge. The
effects of fire are probably most severe in Cuando Cubango, although the large (but
probably cooler) fires that are so frequent in Lunda provinces may too have major
effects on those extensive grasslands.
The landscape changes around towns are limited to the 18 provincial capitals
shown as dark red circles in Fig. 8.8. But landscape changes around many other
large towns need to be recorded.
Future Needs for Research and Documentation
Large volumes of waste are washed into the Atlantic, both close to major coastal
cities – such as Luanda, Benguela and Cabinda – and down large rivers that drain
large areas of the country, such as the Cunene, Cuanza and Queve Rivers. As far as
is known, the volumes, nature and impacts of the waste have not been assessed. The
same is true for impacts on populations of fish and other marine animals which are
harvested from Angolan and foreign vessels that operate offshore, where their activ-
ities and impacts are not monitored.
Fig. 8.8 Areas in Angola where substantial landscape changes have occurred in recent years as a
result of woodland and forest clearing, and of bush fires, and in areas around main roads and towns
where major changes have occurred, or are likely to occur. Woodland and forest clearings are large
contiguous areas shown in Fig. 8.1. Severe fire areas are those where fires burnt in five or more
years between 2000 and 2010, where fires normally burn in August and September when grass is
driest and where fires are normally large (> 20 km2), as derived from data shown in Figs. 8.6 and
8.7. Zones where people settle, farm and harvest wood are usually within 10 km either side of
roads or within 15 km of major towns
The construction of very large dams on the Cuanza River is likely to have affected
the functioning of that river. However, I am not aware of assessments of those
effects, either by individual dams or the cumulative impounding of large volumes of
water.
There are other activities and areas of concern, for example the offshore impacts
of exploration and exploitation by the petroleum industry; large-scale logging in
Cabinda and more recently Moxico and Cuando Cubango; pollution of river water
used for washing and other domestic uses, especially where rivers flow through
large towns; and pesticide contamination from crop farming, particularly from big
commercial farms where large volumes of agricultural poisons are applied.
Finally, the fragmentary patchwork of mini-landscapes that support many spe-
cies and which deserve special conservation measures, requires more study and
documentation. These include the forests of the Escarpment Zone (Faixa subplan-
altica) and Marginal Mountain Chain (Cadeia Marginal de Montanhas).
Considerable numbers of rare and endemic plants and animal species are concen-
trated in these highland forests, many of which are small, covering no more than a
few hundred hectares (Huntley and Matos 1994; Cáceres et al. 2014). The forests
have shrunk, and continue to do so as a result of clearing for crops, harvesting of
timber and charcoal, and grassland fires that kill trees on the forest edges. None of
the forests are legally protected, and all are surrounded by substantial numbers of
rural residents. Some forests are privately owned and their owners should be encour-
aged to manage them for conservation. Likewise, private ownership and manage-
ment could be encouraged for the protection of other forests and areas of special
value.
References
Angola Biocom (2017) http://www.biocom-angola.com/en/company. Accessed 30 Nov 2017

ANGOP (2017) http://www.angop.ao/angola/pt_pt/noticias/economia/2017/5/24/Angola-Mais-
228-mil-madeira-serao-explorados-este-ano,40579b4d-10d3-4bed-b2e5-2751edb213eb.html
António PS (2017) Ponto de Situação Albufeira do Gove 2012–2017. Relatório de PRODEL –
Empresa Pública de Produção de Electricidade, Luanda
Archibald S, Scholes R, Roy D et al (2010) Southern African fire regimes as revealed by remote
sensing. Int J Wildland Fire 19:861–878
Asanzi C, Kiala D, Cesar J et al (2006) Food production in the Planalto of southern Angola. Soil
Sci 171:81–820
Calunga P, Haludilu T, Mendelsohn J et al (2015) Vulnerabilidade na Bacia do Cuvelai/Vulnerability
in the Cuvelai Basin, Angola. Development Workshop, Luanda
Development Workshop (2016) Water resource management under changing climate in Angola’s
coastal settlements. Project Number: 107025–001. Final technical report to the International
Development Research Centre (IDRC), Canada
Ferreira-Baptista L, Manuel J, Aguiar PF et al (2018) Impact of mining on the environment and
water resources in northeastern Angola. Biodivers Ecol 6:155–159
Gomes AF (2012) O Gado na Agricultura Familiar Praticada no Sudoeste de Angola – Meios de
Vida e Vulnerabilidade dos Grupos Domésticos Pastoralistas e Agro-pastoralistas. PhD thesis,
Technical University of Lisbon, Lisbon
cover change. Science 342:850–853
Huntley BJ (2017) Wildlife at war in Angola: the rise and fall of an African Eden. Protea Book
1:53–74
Jain TB, Gould W, Graham RT et al (2008) A soil burn severity index for understanding soil-fire
relations in tropical forests. Ambio 37:563–568
Mendelsohn JM, Mendelsohn S (2018) Sudoeste de Angola: um Retrato da Terra e da Vida. South
West Angola: a Portrait of Land and Life. Raison, Windhoek
Palacios G, Lara-Gomez M, Márquez A et al (2015) Spatial dynamic and quantification of defor-
estation and degradation in Miombo Forest of Huambo Province (Angola) during the period
2002–2015. SASSCAL Proceedings, Huambo, 182 pp
Schneibel A, Stellmes M, Revermann R et al (2013) Agricultural expansion during the post-civil
war period in southern Angola based on bi-temporal Landsat data. Biodivers Ecol 5:311–319
Schneibel A, Röder A, Stellmes M et al (2018) Long-term land use change analysis in south-
central Angola. Assessing the trade-off between major ecosystem services with remote sensing
data. Biodivers Ecol 6:360–367
Silva ERS (2008) Companhia do Caminho de Ferro de Benguela: uma História Sucinta da sua
Formação e Desenvolvimento. Lisbon. https://sites.google.com/site/cfbumahistoriasucinta/
Stellmes M, Frantz D, Finckh M et al (2013) Fire frequency, fire seasonality and fire intensity
within the Okavango region derived from MODIS fire products. Biodivers Ecol 5:351–362
Ucuassapi AP, Dias JCS (2006) Acerca da fertilidade dos solos de Angola. In: Moreira I (ed)
woodlands in southeast Angola. Trop Conserv Sci 8:863–892
Russo V, Lages F, Ferrand N (eds) Biodiversity of Angola. Science & conservation: A modern
synthesis. Springer, Cham
Part III
Invertebrate Diversity: Environmental
Indicators
Chapter 9
The Dragonflies and Damselflies of Angola:
An Updated Synthesis
Jens Kipping, Viola Clausnitzer, Sara R. F. Fernandes Elizalde,

and Klaas-Douwe B. Dijkstra
Abstract Prior to 2012, only 158 species of Odonata were known from Angola.
Surveys in 2012 and 2013 added 76 species and further additions in 2016 brought
the national total to 236 species. This was published earlier in 2017 as the checklist
of the dragonflies and damselflies (Odonata) of Angola by the same authors (Kipping
et al. Afr Invertebr 58 (I):65–91. https://africaninvertebrates.pensoft.net/arti-
cle/11382/, 2017) on which this chapter is based. Records obtained in 2017 and
2018 and a survey by two of the authors in December 2017 led to the discovery of
25 additional species, of which several are undescribed. We provide a revised check-
list here comprising 260 species and discuss the history of research, the biogeogra-
phy of the fauna with endemism and the potential for further discoveries. The
national total is likely to be above 300 species. This would make Angola one of the
richest countries for Odonata in Africa.
Keywords Africa · Biogeography · Checklist · Conservation · Endemism
J. Kipping (*)
BioCart Ökologische Gutachten, Taucha/Leipzig, Germany
e-mail: biocartkipping@email.de
V. Clausnitzer
Senckenberg Museum for Natural History, Görlitz, Görlitz, Germany
e-mail: Viola.Clausnitzer@senckenberg.de
S. R. F. Fernandes Elizalde
SASSCAL – BID GBIF, Instituto de Investigação Agronómica, Huambo, Angola
e-mail: kikas.sara@gmail.com
K.-D. B. Dijkstra
Naturalis Biodiversity Center, Leiden, The Netherlands
e-mail: kd.dijkstra@naturalis.nl

https://doi.org/10.1007/978-3-030-03083-4_9
142 J. Kipping et al.
Introduction
Given the country’s size, diverse landscapes, climatic regimes and habitats Angola
is likely to be one of the richest in Odonata species in Africa. However, Angola’s
biodiversity is very poorly known, with comparatively limited research before inde-
pendence in 1975 halting altogether in the three decades of civil war and unrest that
followed. Research coverage is also limited for Odonata, with much of the north and
east never surveyed at all (Clausnitzer et al. 2012). The potentially very species-rich
highland catchments of the Congo, Cuanza, Cubango (Okavango) and Zambezi riv-
ers are almost unknown and may hold many undescribed species. The whole
Angolan part of the extensively marshy Cuando River and almost the entire Cuito
River system are also largely unsurveyed.
History of Odonata Research in Angola
Research on Odonata began in July 1928, when the Swiss zoologist Albert Monard
embarked on the first of his two expeditions to Angola, which lasted until February
1929. Monard was a curator at the Natural Museum of La-Chaux-de-Fonds in
Switzerland with a broad interest in nature who mainly collected vertebrates and
plants. Ris (1931) identified 27 and described four species from Monard’s first
expedition.
With the death of Ris, Monard submitted the Odonata from his second expedi-
tion (April 1932 to October 1933) to Cynthia Longfield at the British Museum (now
the Natural History Museum) in London, who had published several records
obtained by Karl Jordan from Mount Moco in 1934 (Longfield 1936). Longfield
(1947) identified 77 species from Monard’s new material and described 13 new spe-
cies and two new genera. She also dealt with the Odonata held at the Dundo Museum
in northern Angola, first revising the genus Orthetrum based on the long series
available (Longfield 1955) and later listing 61 species from the collection, including
three new species (Longfield 1959).
Elliott CG Pinhey (1961a, b) described five new species of Gomphidae from
northern Angola received from António de Barros Machado of the Dundo Museum.
While Longfield (1959) stated that the Dundo collection “shows the usual scarcity
of the genera Gomphidae”, Pinhey (1961a) noted it “was particularly notable for the
number of Gomphids.” Possibly Machado split the material between the two
authors. It is uncertain whether the material was collected in Dundo or only held
there, as most records lack details on collector, date and precise locality. However,
Pinhey (1961b) did detail collecting in localities around Dundo, suggesting that all
material came from this part of Lunda-Norte Province. The collector was probably
Machado himself. No-one has worked on this collection since and its state is thus
unknown.
9 The Dragonflies and Damselflies of Angola: An Updated Synthesis 143
Elliot Pinhey was curator at the National Museum of Zimbabwe from 1955 until
1975 and while he collected intensively in adjacent countries, he only visited Angola
twice (Vick et al. 2001). In April and May 1963 Pinhey participated in an expedition
to northwestern Zambia, also visiting an area east of Caianda and the Lutchigena
River in Angola directly adjacent to the Ikelenge Pedicle of Zambia, where he
recorded 26 species (Pinhey 1964, 1974, 1984). His second excursion into Angola
went to an area between Luanda and the Duque de Bragança Falls on the Lucala
River (now known as Calandula Falls) in October 1964 with records of 32 species
(Pinhey 1965).
Pinhey further treated the material of three collectors, describing a species in
honour of each of them. Edward S Ross of the California Academy of Sciences col-
lected between Cuchi and Dondo in 1957 and 1958 (Pinhey 1966), the American
expert of mammal behaviour Richard Estes in central Angola in 1970 (Pinhey
1971a), and Ivan Bampton around the Serra da Chela and Tundavala in 1973 (Pinhey
1975). In the 1975 paper he also repeated records from Pinhey (1964, 1965) and
Longfield (1947), and provided a gazetteer, causing confusion about the precise
locality of some sites. The correct historic collecting sites could be verified with the
gazetteer of Mendes et al. (2013).
Various collectors gathered about 1000 specimens in the collection of the
Instituto de Investigação Agronómica in Huambo between 1950 and 1974. These
records were never published but this will be done shortly by Sara F Elizalde and
David Elizalde as a GBIF dataset.
After Angola’s independence in 1975 there was a long break in field research,
with only a few records by various collectors. Namely in the two decades between
1980 and 2000 not a single record of Odonata is available. Some years after the end
of the civil war a renaissance of research began, resulting in a growing number of
records (Fig. 9.1). All localities with available Odonata records distinguished in the
three periods (a) pre-independence 1928–1974, (b) after independence 1975–2001
and (c) after the end of civil war 2002-today are shown in Fig. 9.2.
Fig. 9.1 Number of Odonata records from Angola over past decades
Fig. 9.2 Records of Odonata from Angola before 1975, before 2002 and up to 2018
Origin of Recent Data
In January 2009, an expedition led by Brian Huntley visited the Serra da Chela in
southwestern Angola and the Namib Desert to the south. During that survey Warwick
Tarboton collected and photographed Odonata around Humpata (7 field days).
Jens Kipping surveyed the upper catchment of the Okavango (Cubango) River
on the SAREP (Southern African Regional Environmental Program) Expedition
from 5 to 22 May 2012 (18 field days). A second SAREP survey visited southeast-
ern Angola with the Cubango and Cuando River floodplains in April 2013.
Viola Clausnitzer and K-D B Dijkstra in collaboration with the Universidade

Kimpa Vita (Uíge) and the Technical University of Dresden (Germany) surveyed
around Uíge, Negage and Ndalatando in northern Angola in the wet season from 13
November to 1 December 2012 (19 days). Dijkstra revisited this area in the dry
season, from 26 September to 5 October 2013 (10 days).
From 27 November to 10 December 2016 (14 field days), Manfred Haacks and
colleagues of SASSCAL (Southern African Science Service Centre for Climate
Change and Adaptive Land Management) visited Bicuar NP and a few other places
in southern Angola.
Sara F Elizalde and David Elizalde, Chris Hines, André Günther, Raik Moritz
and Jens Kipping surveyed the Serra da Chela around Lubango and the mountain
range stretching from Huambo northwards to Gabela from 30 November to 19
December 2017 (20 field days).
Sara F Elizalde, Chris Hines, Rogério Ferreira and other experts provided many
photographic records from 2016 to 2018.
The National Geographic Okavango Wilderness Project (NGOWP 2018) gath-
ered scattered data on Odonata which has not yet been fully considered, except for
some field photographs and exceptional records provided by John Mendelsohn.
Apart from the field surveys the authors also examined the Angolan collections
and type material in the Natural History Museum in London, the National History
Museum of Zimbabwe in Bulawayo (Dijkstra 2007a, b), the Royal Museum for
Central Africa in Tervuren, Belgium and the Instituto de Investigação Agronómica
in Huambo, Angola. All records are kept in the Odonata Database of Africa – ODA
(Kipping et al. 2009) and mapped per species on African Dragonflies and Damselflies
Online – ADDO (visit http://addo.adu.org.za/ also for further information about all
mentioned species).
Odonata Species Recorded in Angola
From all the historic sources mentioned above, 152 species of Odonata were known
to occur in Angola until 2009. Some of the formerly published species had to be
deleted from the country list in the light of new taxonomic knowledge and after
careful validation of all records (see Kipping et al. 2017).
In 2009 Warwick Tarboton recorded 47 species of Odonata at the Serra da Chela
of which five were recorded in Angola for the first time and one was new to science
(Tarboton 2009, Dijkstra et al. 2015). The first SAREP Expedition in 2012 yielded
87 species, 17 of them new to the country list and two new to science (Kipping
2012, Dijkstra et al. 2015). One additional species new for the country came from a
second SAREP Expedition in April 2013 of which all collected specimens were
examined. The first expedition to Uíge, Negage and Ndalatando resulted in 138 spe-
cies, of which 43 were recorded for the first time in Angola and five were new to
science. The second visit produced 86 species, adding another 15 to the national list.
With the surveys from 2009 to 2013 and a careful review of the historic data, the
known odonate fauna of Angola had increased from 152 species in the year 2009 to
234 species in 2013: an increase of about one-third with only 54 days in the field.
Two species were added in 2016 by photographs made by Chris Hines and speci-
mens from the collection of the Instituto de Investigação Agronómica in Huambo
provided by Sara F Elizalde. The state of knowledge at the end of 2016 was pub-
lished as the checklist of the dragonflies and damselflies of Angola by Kipping et al.
in early 2017 (free download: https://africaninvertebrates.pensoft.net/
article/11382/).
The SASSCAL expedition in November–December 2016 recorded 44 species,
amongst them one new species for Angola. The latest survey in December 2017
yielded 88 species of which 10 were new for the country list, amongst them proba-
bly three species new to science. A further 14 species new for Angola were recorded
only in 2017 and early 2018 by Chris Hines and colleagues mostly in northern
Angola.
The updated checklist of the Odonata of Angola, now of 260 species, is provided
in Appendix 1. The ODA database now holds about 4900 Angolan records from
more than 400 localities. All species of Appendix 1 are reliably recorded from
Angolan territory. Footnotes will give further information about the 25 additional
species to the updated country list and one species that was deleted from it.
There are 15 more species listed in Appendix 2 that are known to occur at rivers
bordering the country with Namibia and Zambia. The Namibian bank of the
Okavango River is very well surveyed (Suhling and Martens 2007, 2014) and most
of the mentioned species derive from this river. These species were technically not
found on the Angolan riverbank and therefore not included in the checklist. But
naturally they belong to the country’s fauna.
Composition
Angola’s rich dragonfly fauna expresses its geographic position, size and diversity.
Its territory, especially in the north, falls within a region with an estimated highly
diverse fauna (Fig. 9.3). Dijkstra et al. (2011) observed that roughly half of tropical
African species occur predominantly within the extensive lowland forests of west-
ern and central Africa, a quarter is associated with the eastern and southern part
dominated by highlands, while the remaining quarter occurs in open habitats
throughout much of the Afrotropics. Indeed, about half of Angola’s species are
widespread across the continent and its exceptional diversity can be attributed to
two major sources. Almost 30% are confined to forest habitats in the north, mostly
below 1000 m altitude. Nine species confined to the Lower Guinea, the forest area
that stretches between the Congo Basin and Atlantic Ocean from Cameroon to
Gabon and western Congo, reaching their southern limit in northwestern Angola.
Nearly 20% favour the swamps, grasslands, miombo woodlands and gallery forests
that stretch eastwards, mostly above 1000 m asl. This fauna is concentrated in
Katanga and northern Zambia but has now been proven to extend across to the
Fig. 9.3 Spatial estimation of Odonata diversity in continental Africa, based on the summation of
the inferred ranges of all 770 species known; mapped as the number of species per Hydro1K basin.
(Adapted from Clausnitzer et al. 2012). Angola is outlined in white
Angolan upland. This was confirmed by the discovery of Orthetrum kafwi at two
localities in Cuanza-Sul in December 2017. The species was until then only known
from Upemba NP in southern DRC. A number of palustrine species, e.g. Anax ban-
gweuluensis, Pinheyagrion angolicum, Pseudagrion deningi and P. rufostigma, pre-
fer larger marsh areas and swamps as in the Okavango Delta of Botswana (see
Kipping 2010) and spreading north into Bié highlands with the headwaters of that
river system. The discovery of Trithemis integra near Uíge is also of special interest,
as it had seemed to be endemic to the Albertine Rift, being known previously only
from western Tanzania and Uganda and eastern DRC.
Endemism
Seventeen valid species and several recently discovered undescribed species have so
far only been found in Angola (Figs. 10.4, 10. 5 and 10.6 for examples). With the
exception of two known only from their alleged type localities in far northeastern
Angola, all are limited to the central plateau: the type locality for Platycypha rubri-
ventris is questionable as it may be that of Pseudagrion dundoense, which could
also be a river species from the very poorly sampled southern Congo Basin. No
endemics have been found below 1200 m asl in the east, although some drop down
to about 500 m west of the escarpment. While the proportion of endemics (7%) is
lower than for Ethiopia (12 endemics; 11%) and South Africa (30 endemics; 18%),
countries that also enclose distinct highland areas, this still ranks Angola as one of
Africa’s greatest centres of endemism for Odonata, rivalling the highlands of
Cameroon (13 endemics) and the Albertine Rift, Eastern Arc and Katanga. Moreover,
the number is expected to increase, as almost two-fifths were described since explo-
ration was reinitiated and undescribed species of Platycypha, Paragomphus and
Tetrathemis are already known to us.
Only Platycypha presents an endemic radiation. While Chlorocypha (the fami-
ly’s other large Afrotropical genus) has diversified with almost 30 species largely
in the forested lowlands of west and central Africa, Platycypha is ecologically
more diverse, with species adapted to open, submontane and lake habitats as well.
The Angolan endemics are found mainly between 1300 and 1800 m altitude in
open habitats. The widespread P. angolensis replaces the common P. caligata,
which extends from South Africa to Ethiopia but only peripherally into Angola.
Platycypha bamptoni is probably confined to Serra da Chela; a similar undescribed
species appears more widespread. Platycypha crocea is typical of very small
streams in the Bié highlands and escarpment mountains whereas the other two
inhabit larger streams and rivers. A local radiation of a group that has otherwise
diversified in the highlands to the east, and forests to the north, fits the overall
affinities of Angola’s endemic Odonata both geographically and ecologically
(Kipping et al. 2017) (Fig. 9.4).
The four endemic Pseudagrion species have separate origins but similar links:
the nearest relatives of P. angolense and P. estesi appear to be the rainforest species
P. grilloti Legrand, 1987 and P. kibalense respectively. The former is limited to
Congo and Gabon, but the latter extends to Cameroon and Uganda. P. sarepi is
closely related to P. fisheri and P. greeni, both of which extend from Angola into
Zambia. While these species belong to the genus’s A-group, the B-group species P.
dundoense is known only from Dundo and may not be endemic at all (see above).
Notogomphus kimpavita is the sister-species of N. praetorius found in highlands
across southern Africa (including Angola), while Eleuthemis eogaster is nearest to
an unnamed species from Gabon (Dijkstra et al. 2015).
Molecular data for Umma femina and Onychogomphus rossii are not available
yet. By its unusual habitus and colouration Umma femina (see Fig. 9.5) is a very
distinct member of the genus. It is definitely the odonate flagship species of the
Fig. 9.4 Photographs of some of Angola’s (near) endemic dragonflies and damselflies. (a) Sarep
Sprite (Pseudagrion sarepi), (b) Blue Wisp (Agriocnemis angolensis) that just extends into
Namibia and Zambia, (c) Angola Claspertail (Onychogomphus rossii), (d) Angola Longleg
(Notogomphus kimpavita), (e) Sunrise Firebelly (Eleuthemis eogaster), (f) Angola Micmac
(Micromacromia flava). (All males, photographs a–d by J Kipping, e–f by K-DB Dijkstra)
Angolan highlands and probably also most threatened. The morphology of O. rossii
is close to other pale Onychogomphus species from the open plateaus stretching
from Angola to Zambia and Katanga.
Thus, like the majority of Angola’s Odonata, most endemics probably originated
quite recently and proximally from the forests to the north and open habitats to the
east. However, some affinities are unresolved and potentially more distant:
Agriocnemis toto and especially A. canuango have no obvious close relatives
(Dijkstra et al. 2015), while the near-endemic A. angolensis and A. bumhilli are
probably related to each other but even more distinct overall (Kipping et al. 2017).
Fig. 9.5 Photographs of some of Angola’s endemic damselflies. (a) Angola Blue Jewel (Platycypha
crocea), (b) Highland Blue Jewel (Platycypha bamptoni), (c) undescribed Blue Jewel (Platycypha
sp. nov.), (d) Angola Dancing Jewel (Platycypha angolensis), (e) Angola Sparklewing (Umma
femina), (f) Stout Threadtail (Elattoneura tarbotonorum), (g) Angola Sprite (Pseudagrion ango-
lense), (h) Estes’s Sprite (Pseudagrion estesi). (All males, photographs by J Kipping)
These data suggest that Angola may be the centre of diversification of this genus,
which includes Africa’s smallest damselflies. The morphologically very distinct
Aciagrion rarum is only known from very few specimens from Lunda Sul Province
in the northeast and molecular data is not available yet.
Micromacromia flava is morphologically nearest M. miraculosa (Förster 1906),
known only from the East Usambara Mountains of north-eastern Tanzania and the
only one of four Micromacromia species adapted to non-forest habitats, being
strongly pruinose with maturity. Elattoneura tarbotonorum may be closest to
E. frenulata of southwestern South Africa (Dijkstra et al. 2015): after its discovery
at the Serra da Chela in 2009 it was found more widespread in December 2017
along the mountain range stretching north into Cuanza-Sul.
Potential for Discovery
If we compare the tallies for the well-studied neighbouring countries of Zambia and
Namibia, the total number of species in Angola should lie somewhat above 300,
meaning that less than 80% of the fauna is currently known. All Odonata expedi-
tions in modern times surveyed areas that are easily accessible.
Additions can be expected throughout the country, but especially in the remote
regions on the eastern and particularly northern border, as species diversity is
expected to be extraordinarily high in the transition to the Congolian rainforest
(Fig. 9.3). The province of Lunda-Norte with only 92 recorded species and 162
records should be the richest area for discovery, around Dundo where exploration
began in the 1950s. Generally, all the northern and eastern provinces are largely
unsurveyed and the discrepancy between the amount of available data, the number
of known species and the expected diversity is extremely high. This applies also to
the provinces of Lunda-Sul (10 species, 11 records), Zaire (17 species, 21 records),
Malanje (35 species, 152 records) and Moxico (46 species, 51 records). An excep-
tion is Uíge where recent surveys increased the number of known species to 145
from 820 records gathered.
The central highlands can also yield more surprises, like the discovery of addi-
tional endemic species, with three areas being especially notable. Firstly, despite
having most records, the north-south directed mountain range that lies entirely
above 1600 m asl and includes the Serra do Chilengue, Serra da Chela and Angola’s
highest peak at Mount Moco (2620 m asl) is poorly sampled as the large gaps in
Fig. 9.2 illustrates.
Secondly, except for its extreme northern and southern ends, the western escarp-
ment has only been surveyed recently, which already led to the discovery of an
undescribed Paragomphus species from Cumbira Forest. Even more easily acces-
sible provinces such as Bengo and Cuanza-Sul will prove to be much richer in spe-
cies than currently known. The potential of these mountains is illustrated by the
discovery of a spectacular and unique but unknown species by Chris Hines and
Rogério Ferreira in May 2018 (Fig. 9.6). Two males at a stream that flows off the
Fig. 9.6 An undescribed species that probably belongs to Trithemis, although the extensive mark-
ings and dense veins in the wings are unusual even for that highly diverse genus. Two males were
observed at a stream running off Namba Mountains in Cuanza-Sul. (Photograph by R Ferreira)
Namba Mountains in Cuanza-Sul were photographed but not collected. These

mountains reach over 2000 m in altitude and harbour larger pockets of Afromontane
forest than Mt. Moco (Mills et al. 2013). They are known for their plant endemism
and fieldwork there will definitely lead to the discovery of more endemic Odonata.
Thirdly, an extensive plateau at 1200–1600 m altitude stretches east from the Bié
Highlands. Except for its southern edge, this area shared between Bié and Moxico
Provinces, which is almost as large as Uganda (or the United Kingdom), has almost
no records. A few collections from the NGS Okavango Wilderness Project suggest
that more new species for the country and for science can be expected here. These
deep Kalahari sands are the ‘watertower’ of Angola and its neighbours, incorporat-
ing the headwaters of the Cuito, Cuando, Chicapa, Cuango, Cuanza and large tribu-
taries of the Congo and Zambezi such as the Cassai and Lungué-Bungo. The sources
of the vast catchments of the Congo, Cuanza, Okavango and Zambezi meet in a
small area between Munhango and Cangonga. Watersheds are prone to endemism
(Dijkstra et al. 2011) and this region is the top priority for further research.
Studying insect collections of Angola’s museums will be also a valuable source
of more records and possibly even to get insight into past conditions in the light of
the recent landscape change. Of special interest is the Dundo Museum that holds
many interesting specimens of Odonata, some of which have been published. This
remarkable collection has not been studied since independence but survived the
civil war. There is also material dispersed over several museums in Europe and
probably also in private collections, mainly in Portugal.
New species are most likely to be found among genera prone to narrow (high-
land) ranges, i.e. with known Angolan endemics like Platycypha and Pseudagrion,
but also Agriocnemis, Elattoneura, Notogomphus and Paragomphus. Also possible
is the discovery of endemics in genera that are well represented across the country
and continent, and that have highland endemics elsewhere but not in Angola, such
as Africallagma, Neodythemis and Orthetrum. However, given the biogeographic
diversity of Angola’s fauna and endemics, we could expect greater surprises. Among
forest genera with no known Angolan endemic, Allocnemis seems most likely to
reveal one, e.g. on the escarpment. The presence (or local endemism) of distinctly
Lower Guinean genera like Neurolestes, Africocypha, Pentaphlebia and Stenocnemis
seems less likely, but the Lower Guinean Stenocypha gracilis (Karsch 1899) has
four endemic relatives in the Albertine Rift and the sister-taxon of the Upper and
Lower Guinean Tragogomphus is Nepogomphoides stuhlmanni (Karsch 1899) in
the Eastern Arc, suggesting an Angolan taxon is possible.
Some typical African highland genera are notably absent from Angola.
Atoconeura is most likely to be present above 1400 m asl, being found in Zambia,
Katanga, the Lower Guinea and Albertine Rift. However, its absence also from
South Africa suggests historical factors may have been limiting, e.g. that the high-
lands were too harsh in cooler periods and too isolated when habitats were suitable
(Dijkstra 2006).
This might not apply to Proischnura, present in South Africa as well as Cameroon
and the Albertine Rift. However, that genus is absent from Katanga and northern
Zambia, which lies lower and thus possibly provided no stepping-stone to the
mountains of Angola. Kipping et al. (2017) also noted the absence of Zosteraeschna
and Pinheyschna, which have a similar range (although the latter does occur in
Katanga and northern Zambia), but isolated populations of Z. minuscula (McLachlan
1895) and P. subpupillata (McLachlan 1896) were discovered in the Serra da Chela
in southern Angola in December 2017.
Conservation
Our findings show that Angola’s wealth of aquatic habitats harbours a rich freshwa-
ter fauna. Although large areas are relatively untouched, Angola’s rapid economic
and population growth will have a tremendous impact on the environment and thus
human well-being in the future. In the light of this, Angola’s development should
consider (1) the establishment of sewage works in cities and larger villages; (2) a
stop to deforestation, especially along stream courses; (3) restoration of deforested
water catchments; (4) village-level awareness campaigns for sustainable use of
freshwater sources, e.g. no detergents and waste dumping in rivers; (5) biodiversity
surveys and monitoring to feed into a national conservation plan.
With the exception of four species, all endemics are currently considered Data
Deficient for the IUCN Red List of Threatened Species. Platycypha angolensis,
Pseudagrion angolense and Micromacromia flava are Near Threatened because,
while they seem fairly widespread, their dependence on relatively natural habitats
may put them at risk as human development progresses. Only Umma femina is now
listed as threatened. It is currently known from only a few sites in the fairly densely
populated highlands around Lubango and seems to inhabit exclusively the smaller
and cooler highland streams. There is much development in this densely settled
region and increasing pressure on those habitats by grazing, deforestation and
urbanisation. As it seems to prefer cool mountain streams we can assume additional
risks from climate change and it is therefore thought to be Vulnerable to extinction.
More research on all endemic species’ statuses and ecology is urgently required.
Angola has an exceptional fauna of dragonflies and damselflies, as well as many
valuable rivers and wetlands. Odonata are excellent indicators of the health and
biodiversity of both the freshwater and terrestrial realm. As the biological survey of
Angola advances, they should be a priority taxon.
Acknowledgements We are grateful to Her Excellency Madame Minister of Environment Dr.

Paula C Francisco Coelho (MINAMB) for making the SAREP survey in southern Angola possible,
to Dr. Chris Brooks of SAREP for the preparation and organization of the 2012 survey, to Marta
Alexandre Zumbo (MINAMB), Maria Helena Loa (MINAMB), Julius Bravo (MINAMB),
Francisco de Almeida (INIP), Manuel Domingos (INIP) and Gabriel Cabinda (Agriculture and
Rural Development and Fisheries) for their help in organization and management on the 2012 tour,
and to Vince Shacks and Werner Conradie for collecting specimens on the second SAREP survey
in 2013. We thank Alvaro Bruno Toto Nienguesso, the driving force behind biodiversity research
in Uíge Province, Angola, Prof Dr. Neinhuis and Dr. Thea Lautenschlaeger from TU Dresden for
inviting us to the field survey in Uíge province. Part of the fieldwork in Angola was supported by
a travel fund from the German Academic Exchange Service (DAAD). We thank Dr. Aristófanes
Pontes, director of the Instituto Nacional da Biodiversidade e Áreas de Conservação (INBAC) for
the support during the December 2017 expedition, by providing the necessary permits. Chris Hines
provided many valuable photographic records. Further records were provided by Warwick
Tarboton, Dr. Manfred Haacks (SASSCAL), John Mendelsohn (RAISON) and Rogério Ferreira.
The latter also gave permission to use his wonderful photograph.
Appendices
Appendix 1
Checklist of Odonata recorded from Angola.

#
– see Taxonomic comments in Kipping et al. (2017); ## – see footnotes follow-
ing this table;
(V) Validation of species: “1!” new national record made by the authors; “1!!”
new national record made by the authors and addendum to Kipping et al. (2017);
“1” records obtained by authors and confirming existing records; “2” specimens
kept in collections (identification confirmed or primary types); “3” literature records,
regarded as reliable because specimens were described well or location agrees with
known biogeographic pattern; “4!!” new national record made by other persons and
addendum to Kipping et al. (2017); ** – range restricted to Angola; * – range
restricted to Angola with very few exceptions (see Endemism in the discussion).
(RL) Global conservation status according to the IUCN Red List of Threatened
Species (2016): CR (Critically Endangered), DD (Data-Deficient), EN (Endangered),
NT (Near-Threatened), VU (Vulnerable), LC (Least Concern), NE (Not Evaluated)
Scientific name English name V RL

Lestidae
Lestes amicus (Martin, 1910) Yellow-winged Spreadwing 1 LC
Lestes dissimulans (Fraser, 1955) Cryptic Spreadwing 1 LC
Lestes pallidus (Rambur, 1842) Pallid Spreadwing 1 LC
Lestes pinheyi (Fraser, 1955) Pinhey’s Spreadwing 1 LC
Lestes plagiatus (Burmeister, 1839) Highland Spreadwing 1 LC
Lestes tridens (McLachlan, 1895) Spotted Spreadwing 1 LC
Lestes virgatus (Burmeister, 1839) Smoky Spreadwing 3 LC
Calopterygidae
Phaon camerunensis (Sjöstedt, 1900) Emerald Demoiselle 1! LC
Phaon iridipennis (Burmeister, 1839) Glistening Demoiselle 1 LC
Sapho orichalcea (McLachlan, 1869)# Mountain Bluewing 1! LC
Umma electa (Longfield, 1933) Metallic Sparklewing 1 LC
Umma femina (Longfield, 1947) Angola Sparklewing 1** VU
Umma longistigma (Selys, 1869) Bare-bellied Sparklewing 1 LC
Umma mesostigma (Selys, 1879) Hairy-bellied Sparklewing 1! LC
Chlorocyphidae
Chlorocypha aphrodite (Le Roi, 1915)## Blue Jewel 4!! LC
Chlorocypha cancellata (Selys, 1879) Exquisite Jewel 1! LC
Chlorocypha curta (Hagen in Selys, 1853) Blue-tipped Jewel 1! LC
Chlorocypha cyanifrons (Selys, 1873) Blue-fronted Jewel 1! LC
Chlorocypha fabamacula (Pinhey, 1961) Spotted Jewel 1 LC
Chlorocypha victoriae (Förster, 1914) Victoria’s Jewel 1 LC
Platycypha angolensis (Longfield, 1959) Angola Dancing Jewel 1** NT
Platycypha bamptoni (Pinhey, 1975)# Highland Blue Jewel 1** NE
Platycypha cf. bamptoni (Pinhey, 1975)# (near Highland Blue Jewel) 1!** NE
Platycypha caligata (Selys, 1853)# Common Dancing Jewel 2 LC
Platycypha crocea (Longfield, 1947)# Angola Blue Jewel 1** LC
Platycypha rubriventris (Pinhey, 1975)# Red-bellied Blue Jewel 2** DD
Platycypha rufitibia (Pinhey, 1961) Beautiful Jewel 1 LC
Platycnemididae
Allocnemis nigripes (Selys, 1886) Rainbow Yellowwing 1 LC
Allocnemis pauli (Longfield, 1936) Orange-tipped Yellowwing 1! LC
Copera congolensis (Martin, 1908) Congo Featherleg 1! LC
Elattoneura acuta (Kimmins, 1938) Red Threadtail 1! LC
Elattoneura cellularis (Grünberg, 1902)# Zambezi Threadtail 3 LC
Elattoneura cf. glauca (Selys, 1860)# (near Common Threadtail) 1 LC
Elattoneura lliba (Legrand, 1985) Eastern Stream Threadtail 1! LC
Elattoneura tarbotonorum (Dijkstra, 2015)# Stout Threadtail 1** DD
Mesocnemis singularis (Karsch, 1891)## Common Riverjack 1!! LC
Mesocnemis cf. singularis (Karsch, 1891)# (near Common Riverjack) 1! NE
Coenagrionidae
Aciagrion africanum (Martin, 1908) Blue Slim 1 LC
Aciagrion macrootithenae (Pinhey, 1972) Awl-tipped Slim 3 DD
(continued)

Aciagrion nodosum (Pinhey, 1964) Cryptic Slim 1! LC
Aciagrion rarum (Longfield, 1947) Tiny Slim 2** DD
Aciagrion steeleae (Kimmins, 1955) Swamp Slim 3 LC
Aciagrion zambiense (Pinhey, 1972) Zambia Slim 3 DD
Africallagma fractum (Ris, 1921) Slender Bluet 1 LC
Africallagma glaucum (Burmeister, 1839) Swamp Bluet 1 LC
Africallagma sinuatum (Ris, 1921)## Peak Bluet 4!! LC
Africallagma subtile (Ris, 1921)## Fragile Bluet 1!! LC
Africallagma vaginale (Sjöstedt, 1917) Forest Bluet 1! LC
Agriocnemis angolensis (Longfield, 1947) Blue Wisp 1* LC
Agriocnemis bumhilli (Kipping, Suhling & Martens, Bumhill Wisp 1!* LC
2012)
Agriocnemis canuango (Dijkstra, 2015) Bog Wisp 1!** DD
Agriocnemis exilis (Selys, 1872) Little Wisp 1 LC
Agriocnemis forcipata (Le Roi, 1915) Greater Pincer-tailed Wisp 1 LC
Agriocnemis gratiosa (Gerstäcker, 1891)## Gracious Wisp 4!! LC
Agriocnemis cf. maclachlani (Selys, 1877)# (near Forest Wisp) 1! LC
Agriocnemis pinheyi (Balinsky, 1963)## Pinhey’s Wisp 1!! LC
Agriocnemis ruberrima (Balinsky, 1961) Orange Wisp 1! LC
Agriocnemis toto (Dijkstra, 2015) Bruno’s Wisp 1!** DD
Agriocnemis victoria (Fraser, 1928) Lesser Pincer-tailed Wisp 1 LC
Azuragrion nigridorsum (Selys, 1876) Sailing Bluet 1 LC
Ceriagrion annulatum (Fraser, 1955) Green-eyed Citril 1! LC
Ceriagrion bakeri (Fraser, 1941) Blue-fronted Citril 3 LC
Ceriagrion corallinum (Campion, 1914) Green-fronted Citril 1 LC
Ceriagrion glabrum (Burmeister, 1839) Common Citril 1 LC
Ceriagrion junceum (Dijkstra & Kipping, 2015) Spikerush Citril 1! LC
Ceriagrion platystigma (Fraser, 1941) Variable Citril 1 LC
Ceriagrion sakejii (Pinhey, 1963) Cream-sided Citril 1! LC
Ceriagrion suave (Ris, 1921) Plain Citril 1 LC
Ceriagrion whellani (Longfield, 1952) Yellow-faced Citril 1! LC
Ischnura senegalensis (Rambur, 1842) Tropical Bluetail 1 LC
Pinheyagrion angolicum (Pinhey, 1966) Pinhey’s Bluet 1 LC
Pseudagrion (A) angolense (Selys, 1876) Angola Sprite 1** NT
Pseudagrion (A) coeruleipunctum (Pinhey, 1964) Pretty Sprite 3 LC
Pseudagrion (A) estesi (Pinhey, 1971) Estes’s Sprite 1** LC
Pseudagrion (A) fisheri (Pinhey, 1961) Dark-tailed Sprite 3 LC
Pseudagrion (A) greeni (Pinhey, 1961) Clasper-tailed Sprite 1 LC
Pseudagrion (A) inconspicuum (Ris, 1931) Little Sprite 1 LC
Pseudagrion (A) kersteni (Gerstäcker, 1869) Powder-faced Sprite 1 LC
Pseudagrion (A) kibalense (Longfield, 1959) Forest Sprite 1 LC
Pseudagrion (A) makabusiense (Pinhey, 1950) Green-striped Sprite 3 LC
Pseudagrion (A) melanicterum (Selys, 1876) Farmbush Sprite 1 LC
Pseudagrion (A) salisburyense (Ris, 1921) Slate Sprite 1 LC
(continued)

Pseudagrion (A) sarepi (Kipping & Dijkstra, 2015) Sarep Sprite 1!** DD
Pseudagrion (A) serrulatum (Karsch, 1894) Superb Sprite 1! LC
Pseudagrion (A) simonae (Legrand, 1987) Wide-striped Sprite 1! LC
Pseudagrion (A) simplicilaminatum (Carletti & Blue Slim Sprite 4!! LC
Terzani, 1997)##
Pseudagrion (B) acaciae (Förster, 1906) Acacia Sprite 1 LC
Pseudagrion (B) camerunense (Karsch, 1899)## Yellow-fronted Sprite 4!! LC
Pseudagrion (B) coeleste (Longfield, 1947) Catshead Sprite 1 LC
Pseudagrion (B) deningi (Pinhey, 1961) Dark Sprite 1! LC
Pseudagrion (B) dundoense (Longfield, 1959) Dundo Sprite 2** DD
Pseudagrion (B) glaucescens (Selys, 1876) Blue-green Sprite 1 LC
Pseudagrion (B) hamoni (Fraser, 1955) Swarthy Sprite 1! LC
Pseudagrion (B) helenae (Balinsky, 1964) Little Blue Sprite 1! LC
Pseudagrion (B) isidromorai (Compte Sart, 1967) Large Blue Sprite 1! LC
Pseudagrion (B) massaicum (Sjöstedt, 1909) Masai Sprite 1 LC
Pseudagrion (B) rufostigma (Longfield, 1947) Ruby Sprite 1 LC
Pseudagrion (B) sjoestedti (Förster, 1906) Variable Sprite 1 LC
Pseudagrion (B) sublacteum (Karsch, 1893) Cherry-eye Sprite 1 LC
Aeshnidae
Afroaeschna scotias (Pinhey, 1952) Shadow Hawker 1! LC
Anaciaeschna triangulifera (McLachlan, 1896)## Evening Hawker 4!! LC
Anax bangweuluensis (Kimmins, 1955)## Swamp Emperor 4!! NT
Anax congoliath (Fraser, 1953) Dark Emperor 1! LC
Anax ephippiger (Burmeister, 1839) Vagrant Emperor 1 LC
Anax imperator (Leach, 1815) Blue Emperor 1 LC
Anax speratus (Hagen, 1867) Eastern Orange Emperor 1 LC
Anax tristis (Hagen, 1867) Black Emperor 1 LC
Gynacantha (A) sextans (McLachlan, 1896) Dark-rayed Duskhawker 3 LC
Gynacantha (A) vesiculata (Karsch, 1891) Lesser Girdled Duskhawker 3 LC
Gynacantha (B) bullata (Karsch, 1891) Black-kneed Duskhawker 1 LC
Gynacantha (B) manderica (Grünberg, 1902) Little Duskhawker 3 LC
Heliaeschna cynthiae (Fraser, 1939)## Blade-tipped Duskhawker 4!! LC
Heliaeschna fuliginosa (Karsch, 1893) Black-banded Duskhawker 1 LC
Heliaeschna ugandica (McLachlan, 1896) Uganda Duskhawker 3 LC
Pinheyschna subpupillata (McLachlan, 1896)## Stream Hawker 1!! LC
Zosteraeschna minuscula (McLachlan, 1895)## Friendly Hawker 1!! LC
Gomphidae
Crenigomphus cf. cornutus (Pinhey, 1956)# (near Horned Talontail) 1! LC
Diastatomma selysi (Schouteden, 1934) Common Hoetail 3 LC
Diastatomma soror (Schouteden, 1934) Painted Hoetail 3 LC
Gomphidia quarrei (Schouteden, 1934) Southern Fingertail 3 LC
Ictinogomphus dundoensis (Pinhey, 1961) Swamp Tigertail 1 LC
Ictinogomphus ferox (Rambur, 1842) Common Tigertail 1 LC
Ictinogomphus regisalberti (Schouteden, 1934) Congo Tigertail 3 LC
(continued)

Lestinogomphus calcaratus (Dijkstra, 2015) Spurred Fairytail 1! LC
Libyogomphus tenaculatus (Fraser, 1926) Large Horntail 1! LC
Mastigogomphus chapini (Klots, 1944)# Western Snorkeltail 2 LC
Mastigogomphus dissimilis (Cammaerts, 2004)## Southern Snorkeltail 2 LC
Microgomphus cf. nyassicus (Grünberg, 1902)# (near Eastern Scissortail) 1! LC
Neurogomphus alius (Cammaerts, 2004) Large Siphontail 1! LC
Notogomphus kimpavita (Dijkstra & Clausnitzer, Angola Longleg 1!** DD
2015)
Notogomphus praetorius (Selys, 1878) Yellowjack Longleg 2 LC
Notogomphus spinosus (Karsch, 1890) Jungle Longleg 1! LC
Onychogomphus rossii (Pinhey, 1966) Angola Claspertail 1** DD
Onychogomphus cf. styx (Pinhey, 1961)# (near Northern Dark 1! LC
Claspertail)
Paragomphus abnormis (Karsch, 1890) Humdrum Hooktail 1! LC
Paragomphus cognatus (Rambur, 1842) Rock Hooktail 1!! LC
Paragomphus cf. darwalli (Dijkstra, Mézière & (near Darwall’s Hooktail) 1! DD
Papazian, 2015)#
Paragomphus genei (Selys, 1841) Common Hooktail 1 LC
Paragomphus machadoi (Pinhey, 1961) Forest Hooktail 2 LC
Paragomphus cf. nigroviridis (Cammaerts, 1969)# (near Black-and-green 1! LC
Hooktail)
Paragomphus sabicus (Pinhey, 1950)## Flapper Hooktail 1!! LC
Paragomphus sp. nov. ## (Hooktail, undescribed 1!!** NE
species)
Phyllogomphus annulus (Klots, 1944) Crested Leaftail 1 LC
Phyllogomphus selysi (Schouteden, 1933) Bold Leaftail 3 LC
Macromiidae
Phyllomacromia aureozona (Pinhey, 1966) Golden-banded Cruiser 1! LC
Phyllomacromia contumax (Selys, 1879) Two-banded Cruiser 1! LC
Phyllomacromia hervei (Legrand, 1980) River Cruiser 1! LC
Phyllomacromia melania (Selys, 1871) Sombre Cruiser 1 LC
Phyllomacromia overlaeti (Schouteden, 1934) Clubbed Cruiser 3 LC
Phyllomacromia paula (Karsch, 1892) Greater Double-spined 3 LC
Cruiser
Phyllomacromia picta (Hagen in Selys, 1871) Darting Cruiser 3 LC
Phyllomacromia unifasciata (Fraser, 1954) Golden-eyed Cruiser 3 LC
Libellulidae
Acisoma inflatum (Selys, 1882) Stout Pintail 1 LC
Acisoma trifidum (Kirby, 1889) Pied Pintail 1 LC
Aethiothemis bequaerti (Ris, 1919) Skimmer-like Flasher 1 LC
Aethiothemis ellioti (Lieftinck, 1969) Plump Flasher 1! LC
Aethiothemis mediofasciata (Ris, 1931)# Orange Flasher 2 LC
Aethiothemis solitaria (Martin, 1908) Pearly Flasher 1 LC
Aethriamanta rezia (Kirby, 1889) Pygmy Basker 1 LC
Brachythemis lacustris (Kirby, 1889) Red Groundling 1 LC
(continued)

Brachythemis leucosticta (Burmeister, 1839) Southern Banded 1 LC
Groundling
Bradinopyga strachani (Kirby, 1900)## Red Rockdweller 1!! LC
Chalcostephia flavifrons (Kirby, 1889) Inspector 1! LC
Crocothemis brevistigma (Pinhey, 1961) Spotted Scarlet 1! LC
Crocothemis divisa (Baumann, 1898) Rock Scarlet 1 LC
Crocothemis erythraea (Brullé, 1832) Broad Scarlet 1 LC
Crocothemis sanguinolenta (Burmeister, 1839) Little Scarlet 1 LC
Cyanothemis simpsoni (Ris, 1915) Bluebolt 1! LC
Diplacodes deminuta (Lieftinck, 1969) Little Percher 1 LC
Diplacodes lefebvrii (Rambur, 1842) Black Percher 1 LC
Diplacodes luminans (Karsch, 1893) Barbet Percher 1 LC
Diplacodes pumila (Dijkstra, 2006) Dwarf Percher 1! LC
Eleuthemis eogaster (Dijkstra, 2015) Sunrise Firebelly 1!** DD
Eleuthemis libera (Dijkstra & Kipping, 2015) Free Firebelly 1! DD
Hadrothemis camarensis (Kirby, 1889) Saddled Jungleskimmer 3 LC
Hadrothemis coacta (Karsch, 1891) Robust Jungleskimmer 1! LC
Hadrothemis defecta (Karsch, 1891) Scarlet Jungleskimmer 3 LC
Hemistigma albipunctum (Rambur, 1842) African Piedspot 1 LC
Malgassophlebia bispina (Fraser, 1958) Ringed Leaftipper 1! LC
Micromacromia camerunica (Karsch, 1890) Stream Micmac 1! LC
Micromacromia flava (Longfield, 1947) Angola Micmac 1** NT
Neodythemis afra (Ris, 1909) Seepage Junglewatcher 1! LC
Neodythemis klingi (Karsch, 1890) Stream Junglewatcher 1! LC
Nesciothemis cf. farinosa (Förster, 1898)# (near Eastern Blacktail) 1 LC
Nesciothemis fitzgeraldi (Longfield, 1955) Lesser Peppertail 1! LC
Notiothemis jonesi (Ris, 1919)## Eastern Forestwatcher 1!! LC
Notiothemis robertsi (Fraser, 1944) Western Forestwatcher 1! LC
Olpogastra lugubris (Karsch, 1895) Bottletail 1 LC
Orthetrum abbotti (Calvert, 1892) Little Skimmer 1 LC
Orthetrum austeni (Kirby, 1900) Giant Skimmer 1 LC
Orthetrum brachiale (Palisot de Beauvois, 1817) Banded Skimmer 1 LC
Orthetrum caffrum (Burmeister, 1839) Two-striped Skimmer 1 LC
Orthetrum chrysostigma (Burmeister, 1839) Epaulet Skimmer 1 LC
Orthetrum guineense (Ris, 1910) Guinea Skimmer 1 LC
Orthetrum hintzi (Schmidt, 1951) Dark-shouldered Skimmer 1 LC
Orthetrum icteromelas (Ris, 1910) Spectacled Skimmer 1 LC
Orthetrum julia (Kirby, 1900) Julia Skimmer 1 LC
Orthetrum kafwi (Dijkstra, 2015)## Bog Skimmer 1!! DD
Orthetrum machadoi (Longfield, 1955) Highland Skimmer 1 LC
Orthetrum macrostigma (Longfield, 1947) Sharkfin Skimmer 1 LC
Orthetrum microstigma (Ris, 1911) Farmbush Skimmer 1 LC
Orthetrum monardi (Schmidt, 1951) Woodland Skimmer 1 LC
Orthetrum robustum (Balinsky, 1965) Robust Skimmer 1! LC
(continued)

Orthetrum saegeri (Pinhey, 1966) Eastern Mushroom 1! LC
Skimmer
Orthetrum stemmale (Burmeister, 1839) Bold Skimmer 1 LC
Orthetrum trinacria (Selys, 1841) Long Skimmer 1 LC
Oxythemis phoenicosceles (Ris, 1910) Pepperpants 1! LC
Palpopleura albifrons (Legrand, 1979) Pale-faced Widow 1! LC
Palpopleura deceptor (Calvert, 1899) Deceptive Widow 3 LC
Palpopleura jucunda (Rambur, 1842) Yellow-veined Widow 1 LC
Palpopleura lucia (Drury, 1773) Lucia Widow 1 LC
Palpopleura portia (Drury, 1773) Portia Widow 1 LC
Pantala flavescens (Fabricius, 1798) Wandering Glider 1 LC
Porpax asperipes (Karsch, 1896) Powdered Pricklyleg 1 LC
Porpax risi (Pinhey, 1958) Highland Pricklyleg 1 LC
Rhyothemis fenestrina (Rambur, 1842) Skylight Flutterer 1 LC
Rhyothemis mariposa (Ris, 1913) Butterfly Flutterer 2 LC
Rhyothemis cf. notata (Fabricius, 1781)## (near Veiled Flutterer) 4!! LC
Rhyothemis semihyalina (Desjardins, 1832) Phantom Flutterer 1! LC
Sympetrum fonscolombii (Selys, 1840) Nomad 1 LC
Tetrathemis camerunensis (Sjöstedt, 1900) Forest Elf 2 LC
Tetrathemis fraseri (Legrand, 1977) Treefall Elf 1! LC
Tetrathemis polleni (Selys, 1869) Black-splashed Elf 2 LC
Tetrathemis sp. nov. ## (Elf, undescribed species) 4!!** NE
Thermochoria equivocata (Kirby, 1889) Dash-winged Piedface 1! LC
Tholymis tillarga (Fabricius, 1798) Twister 1 LC
Tramea basilaris (Palisot de Beauvois, 1817) Keyhole Glider 1 LC
Trithemis aconita (Lieftinck, 1969) Halfshade Dropwing 1! LC
Trithemis aenea (Pinhey, 1961)## Bronze Dropwing 4!! LC
Trithemis annulata (Palisot de Beauvois, 1807) Violet Dropwing 1 LC
Trithemis anomala (Pinhey, 1956) Striped Dropwing 1! LC
Trithemis apicalis (Fraser, 1954) Furtive Dropwing 1! LC
Trithemis arteriosa (Burmeister, 1839) Red-veined Dropwing 1 LC
Trithemis basitincta (Ris, 1912) Jungle Dropwing 1! LC
Trithemis dichroa (Karsch, 1893) Black Dropwing 1 LC
Trithemis dorsalis (Rambur, 1842) Highland Dropwing 1 LC
Trithemis cf. dubia (Fraser, 1954)# (near Sleek Dropwing) 1! DD
Trithemis furva (Karsch, 1899) Navy Dropwing 1 LC
Trithemis imitata (Pinhey, 1961)# Northern Fluttering 1! LC
Dropwing
Trithemis integra (Dijkstra, 2007) Albertine Dropwing 1! LC
Trithemis kirbyi (Selys, 1891) Orange-winged Dropwing 1 LC
Trithemis leakeyi (Pinhey, 1956) Mealy Dropwing 1! LC
Trithemis monardi (Ris, 1931)# Southern Fluttering 1 LC
Dropwing
Trithemis nuptialis (Karsch, 1894) Hairy-legged Dropwing 1 LC
Trithemis palustris (Damm & Hadrys, 2009)# Marsh Dropwing 1! LC
(continued)

Trithemis pluvialis (Förster, 1906) Russet Dropwing 1 LC
Trithemis pruinata (Karsch, 1899) Cobalt Dropwing 1! LC
Trithemis stictica (Burmeister, 1839) Jaunty Dropwing 1 LC
Trithemis werneri (Ris, 1912) Elegant Dropwing 3 LC
Trithemis sp. nov. (Fig 9.6)## (Dropwing, undescribed 4!!** NE
species)
Urothemis assignata (Selys, 1872) Red Basker 1 LC
Urothemis edwardsii (Selys, 1849) Blue Basker 1 LC
Urothemis venata (Dijkstra & Mézière, 2015)## Red-veined Basker 4 !! LC
Zygonoides fuelleborni (Grünberg, 1902) Southern Riverking 3 LC
Zygonyx denticulatus (Dijkstra & Kipping, 2015) Pale Cascader 1! LC
Zygonyx eusebia (Ris, 1912) Imperial Cascader 3 LC
Zygonyx flavicosta (Sjöstedt, 1900)## Ensign Cascader 1 LC
Zygonyx natalensis (Martin, 1900) Blue Cascader 1 LC
Zygonyx regisalberti (Schouteden, 1934) Regal Cascader 1 LC
Zygonyx torridus (Kirby, 1889) Ringed Cascader 1 LC
Notes on new country records (by J Kipping and S F Elizalde unless stated
otherwise)
Chlorocypha aphrodite – male photographed by C Hines near Lucala north of
Uíge in June 2017.
Mesocnemis singularis – first record of true M. singularis (see Kipping et al.
2017) from the Angolan bank of the Cunene River in December 2017.
Africallagma sinuatum – single male photographed by C Hines near Cambondo,
Cuanza-Norte Province in February 2017.
Africallagma subtile – several collected at marshy floodplains of the Yevedula
River, 20 km northwest of Caconda, Benguela Province in December 2017.
Agriocnemis gratiosa – several collected by M Haacks from Bicuar NP, Huíla
Province in December 2016.
Agriocnemis pinheyi – several collected at a marsh northwest of Caconda,
Benguela Province in December 2017.
Pseudagrion (A) simplicilaminatum – male photographed by C Hines near
Lucala north of Uíge in June 2017.
Pseudagrion (B) camerunense – male photographed by C Hines in Cuanza River
floodplains south of Luanda in January 2018.
Anaciaeschna triangulifera – female photographed by C Hines in Cuanza River
floodplains south of Luanda in June 2017. Westernmost record; nearest locality is
Ikelenge in northwestern Zambia, about 1200 km to the east.
Anax bangweuluensis – teneral male photographed by J Mendelsohn at Lake
Saliakembo, Moxico Province in October 2017. The Cuito River links to the nearest
known population in the Okavango Delta, Botswana, about 750 km away.
Heliaeschna cynthiae – female and two males recorded by C Hines at the Rio
Nzadi and near Quicunga in Uíge Province in June 2017.
Pinheyschna subpupillata – many observed and collected at Tchiamena River
near Lubango and Neve River near Humpata on Serra da Chela, Huíla Province in
December 2017. Presumably isolated population; widespread in South Africa, with

another isolated population on border of Mozambique and Zimbabwe. With the new
finding of this species a former record of a female P. rileyi (Calvert, 1892) from
Tundavala (Pinhey 1975) became more doubtful and the species is therefore deleted
from the country list.
Zosteraeschna minuscula – male collected at the Tchiamena River near Lubango
on Serra da Chela, Huíla Province in December 2017. Northernmost record; wide-
spread in South Africa but with scattered records in Namibia and eastern Botswana.
Mastigogomphus dissimilis – the Instituto de Investigação Agronómica in
Huambo has one male from Nova Sintra (Catabola), Bié Province from October
1973, coll. L Amorim.
Paragomphus cognatus – presence in Angola was uncertain due to lack of reli-
able material (Kipping et al. 2017), but several males collected at Tchiamena, Leba
and Neve Rivers in the Serra da Chela in December 2017.
Paragomphus sabicus – common at the Rio Coporolo, north of Chongoroi,
Benguela Province in December 2017.
Paragomphus sp. nov. – two males collected at the Uiri River near Conda,
Cuanza-Sul Province in December 2017 belong to an undescribed species similar to
P. cognatus but darker and with stouter paraprocts and more curved cerci.
Bradinopyga strachani – the Instituto de Investigação Agronómica in Huambo
has three males from Ndalatando, Cuanza-Norte Province from March 1973, coll. U
Passos. Several also collected at Rio Mussenju, south of Quilengues, Benguela
Province in December 2017 and photographed by R Ferreira at Calandula Falls,
Lunda-Norte Province in June 2018.
Notiothemis jonesi – male was collected in Lubango, Huíla Province in December
2017.
Orthetrum kafwi – several males and females collected at boggy streams and
bogs in the highlands around Cassongue, Cuanza-Sul Province in December 2017.
Previously know only from the type locality in the Upemba National Park in
Katanga, which lies 1400 km to the east.
Rhyothemis cf. notata – male photographed by J Mendelsohn at Sacangombe
near the Cuito River source in Moxico Province in November 2011. The black
markings in the forewings reach only to the nodus and in the hindwings halfway the
nodus and pterostigma, which is much less than even the palest variation of R.
notata illustrated by Dijkstra & Clausnitzer (2014). The habitat is open, while true
R. notata favour rainforest conditions. This species therefore needs to be verified
with specimens.
Tetrathemis sp. nov. – several males photographed by C Hines in dry forest near
Cambondo, Cuanza-Norte Province in March 2017. Differs from T. fraseri by the
smoky wings and shape of the very hairy cerci.
Trithemis aenea – photographed by C Hines near Lucala north of Uíge in June
2017.
Trithemis sp. nov. – see Fig. 9.6 and main text.
Urothemis venata – photographed by Carel van der Merwe in the Cuango area,
Cuanza-Norte Province in May 2017.
Appendix 2
Odonata recorded from rivers bordering Angola that most likely also occur in
Angola.
Scientific name English name Nearest occurrence

Coenagrionidae
Pseudagrion (A) spernatum Upland Sprite At Jimbe and other rivers in Ikelenge Pedicle
(Selys, 1881) of north-western Zambia.
Pseudagrion (B) assegaii Assegai Sprite Cuando River in Namibian Caprivi Strip.
(Pinhey, 1950)
Pseudagrion (B) sudanicum (Le Blue-sided Okavango and Cuando Rivers in Namibian
Roi, 1915) Sprite Caprivi Strip.
Gomphidae
Crenigomphus kavangoensis Kavango Okavango River in Namibia.
(Suhling & Marais, 2010) Talontail
Lestinogomphus angustus Common Cunene, Okavango and Cuando Rivers in
(Martin, 1911) Fairytail northern Namibia.
Lestinogomphus silkeae Silke’s One locality on the southern bank of the
(Kipping, 2010) Fairytail Okavango River near Rundu, Namibia.
Paragomphus cataractae Cataract Waterfalls and rapids of the Cunene and
(Pinhey, 1963) Hooktail Okavango Rivers in northern Namibia.
Paragomphus elpidius (Ris, Corkscrew Cunene, Okavango and Cuando River in
1921) Hooktail northern Namibia and the Ikelenge Pedicle of
Zambia.
Neurogomphus cocytius Kokytos Okavango River in northern Namibia.
Cammaerts, 2004 Siphontail
Libellulidae
Parazyxomma flavicans Banded Okavango and Cuando Rivers in northern
(Martin, 1908) Duskdarter Namibia.
Trithemis aequalis (Lieftinck, Swamp Okavango and Cuando Rivers in the
1969) Dropwing Namibian Caprivi.
Trithemis donaldsoni (Calvert, Denim Okavango and Cunene Rivers in northern
1899) Dropwing Namibia.
Trithemis hecate (Ris, 1912) Silhouette Common along the Cunene, Okavango and
Dropwing Cuando Rivers in northern Namibia.
Trithemis morrisoni (Damm & Rapids Okavango and Cuando Rivers in the
Hadrys, 2009) Dropwing Namibian Caprivi.
Trithetrum navasi (Lacroix, Fiery Darter Cunene, Okavango and Cuando Rivers in
1921) northern Namibia.
References
Clausnitzer V, Koch R, Dijkstra K-DB et al (2012) Focus on African freshwaters: hotspots of drag-
onfly diversity and conservation concern. Front Ecol Environ 10:129–134
Damm S, Hadrys H (2009) Trithemis morrisoni sp. nov. and Trithemis palustris sp. nov. from
the Okavango and Upper Zambezi Floodplains previously hidden under T. stictica (Odonata:
Libellulidae). Int J Odonatol 12(1):131–145
Dijkstra K-DB (2006) The Atoconeura problem revisited: taxonomy, phylogeny and biogeogra-
phy of a dragonfly genus in the highlands of Africa (Odonata, Libellulidae). Tijdschrift voor
Entomologie 149:121–144
Dijkstra K-DB (2007a) The name-bearing types of Odonata held in the Natural History Museum
of Zimbabwe, with systematic notes on Afrotropical taxa. Part 1: introduction and Anisoptera.
Int J Odonatol 10(1):1–29
Dijkstra K-DB (2007b) The name-bearing types of Odonata held in the Natural History Museum
of Zimbabwe, with systematic notes on Afrotropical taxa. Part 2: Zygoptera and description of
new species. Int J Odonatol 10(2):137–170
Dijkstra K-DB, Clausnitzer V (2014) The dragonflies and damselflies of Eastern Africa: hand-
book for all Odonata from Sudan to Zimbabwe. . Studies in afrotropical zoology 298. Royal
Museums for Central Africa, Tervuren, 263 pp
Dijkstra K-DB, Kipping J, Mézière N (2015) Sixty new dragonfly and damselfly species from
Africa (Odonata). Odonatologica 44(4):447–678
Dijkstra K-DB, Boudot J-P, Clausnitzer V et al (2011) Chapter 5. Dragonflies and damselflies of
Africa (Odonata): history, diversity, distribution, and conservation. In: Darwall WRT, Smith
KG, Allen DJ et al (eds) The diversity of life in African freshwaters: under water, under threat.
An analysis of the status and distribution of freshwater species throughout mainland Africa.
IUCN, Cambridge and Gland, 347 pp
Kipping J (2010) The dragonflies and damselflies of Botswana – an annotated checklist with notes
on distribution, phenology, habitats and Red List status of the species (Insecta: Odonata).
Mauritiana (Altenburg) 21:126–204
Kipping J (2012) Southern African Regional Environmental Program (SAREP) – first biodiver-
sity field survey upper Cubango (Okavango) catchment, Angola, May 2012 – Dragonflies &
Damselflies (Insecta: Odonata). Expert Report:1–108
Kipping J, Clausnitzer V, Fernandes Elizalde SRF et al (2017) The dragonflies and damselflies
(Odonata) of Angola. Afr Invertebr 58(I):65–91 https://africaninvertebrates.pensoft.net/
article/11382/
Kipping J, Dijkstra K-DB, Clausnitzer V et al (2009) Odonata Database of Africa (ODA). Agrion
13:20–23
Longfield C (1936) Studies on African Odonata, with synonymy and descriptions of new species
and subspecies. Trans R Entomol Soc Lond 85:467–499
Longfield C (1947) The Odonata of South Angola: results of the Mission Scientifiques Suisses
1928–29, 1932–33. Arquivos do Museu Bocage 16:1–31
Longfield C (1955) The Odonata of North Angola, part 1. Publicações Culturais, Companhia de
Diamantes de Angola 27:11–64
Longfield C (1959) The Odonata of North Angola, part 2. Publicações Culturais, Companhia de
Mendes LF, Bivar-de-Sousa A, Figueira R et al (2013) Gazetteer of the Angolan localities known
for beetles (Coleoptera) and butterflies (Lepidoptera: Papilionoidea). Boletim da Sociedade
Portuguesa de Entomologia 228(VIII–14):257–292
Mills MSL, Melo M, Vaz A (2013) The Namba mountains: new hope for Afromontane forest birds
NGOWP – National Geographic Okavango Wilderness Project (2018) Initial Findings from
Exploration of the Upper Catchments of the Cuito, Cuanavale and Cuando Rivers in Central
Pinhey ECG (1961a) A collection of Odonata from Dundo, Angola with the descriptions of two new
species of Gomphids. Publicações Culturais, Companhia de Diamantes de Angola 56:71–78
Pinhey ECG (1961b) Some dragonflies (Odonata) from Angola and descriptions of three new
species of the family Gomphidae. Publicações Culturais, Companhia de Diamantes de Angola
56:79–86
Pinhey ECG (1964) Dragonflies (Odonata) of the Angola-Congo borders of Rhodesia. Publicações
Culturais, Companhia de Diamantes de Angola 63:97–130
Pinhey ECG (1965) Odonata from Luanda and the Lucala River, Angola. Revista de Biologia
5:159–164
Pinhey ECG (1966) New distributional records for African Odonata and notes on a few larvae.
Arnoldia Rhodesia 2(26):1–5
Pinhey ECG (1971a) Notes on the genus Pseudagrion Selys (Odonata: Coenagrionidae). Arnoldia
Rhodesia 5(6):1–4
Pinhey ECG (1971b) Odonata collected in Republique Centre-Africaine by R. Pujol. Arnoldia
Rhodesia 5(18):1–16
Pinhey ECG (1974) A revision of the African Agriocnemis Selys and Mortonagrion Fraser
(Odonata: Coenagrionidae). Occasional Papers of the National Monuments of Rhodesia B
5/4:171–278
Pinhey ECG (1975) A collection of Odonata from Angola. Arnoldia Rhodesia 7(23):1–16
Pinhey ECG (1984) A check-list of the Odonata of Zimbabwe and Zambia. Smithersia 3:1–64
Ris F (1931) Odonata aus Süd-Angola. Revue Suisse Zoologie 38(7):97–112
Suhling F, Martens A (2007) Dragonflies and damselflies of Namibia. Gamsberg Macmillan,
Windhoek, 280 pp
Suhling F, Martens A (2014) Distribution maps and checklist of Namibian Odonata. Libellula
Suppl 13:107–175
Tarboton W (2009) A dragonfly survey of the Humpata District. In: Huntley BJ (ed) Projecto de
estudo da biodiversidade de Angola. (Biodiversity Rapid Assessment – Huíla /Namibe) Report
on Pilot project. SANBI, Cape Town, 3 pp
Vick GS, Chelmick DG, Martens A (2001) In memory of Elliot Charles Gordon Pinhey (10 July
1910 – 7 May 1999). Odonatologica 30:1–11
Chapter 10
The Butterflies and Skippers (Lepidoptera:
Papilionoidea) of Angola: An Updated
Checklist
Luís F. Mendes, A. Bivar-de-Sousa, and Mark C. Williams
Abstract Presently, 792 species/subspecies of butterflies and skippers (Lepidoptera:

Papilionoidea) are known from Angola, a country with a rich diversity of habitats,
but where extensive areas remain unsurveyed and where systematic collecting pro-
grammes have not been undertaken. Only three species were known from Angola in
1820. From the beginning of the twenty-first century, many new species have been
described and more than 220 faunistic novelties have been assigned. As a whole, of
the 792 taxa now listed for Angola, 57 species/subspecies are endemic and almost
the same number are known to be near-endemics, shared by Angola and by one or
another neighbouring country. The Nymphalidae are the most diverse family. The
Lycaenidae and Papilionidae have the highest levels of endemism. A revised check-
list with taxonomic and ecological notes is presented and the development of knowl-
edge of the superfamily over time in Angola is analysed.
Keywords Africa · Conservation · Ecology · Endemism · Taxonomy
L. F. Mendes (*)
Museu Nacional de História Natural e da Ciência, Universidade de Lisboa, Lisboa, Portugal
CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Vairão, Portugal
e-mail: luisfmendes22@gmail.com
A. Bivar-de-Sousa
Museu Nacional de História Natural e da Ciência, Universidade de Lisboa, Lisboa, Portugal
Sociedade Portuguesa de Entomologia, Lisboa, Portugal
e-mail: abivarsousa@gmail.com
M. C. Williams
Pretoria University, Pretoria, South Africa
e-mail: lepidochrysops@gmail.com

https://doi.org/10.1007/978-3-030-03083-4_10
168 L. F. Mendes et al.
Introduction
Angola is a large country of 1,246,700 km2, notable for its great diversity of physi-
ography, climates, habitats and resultant biodiversity). The country includes seven
biomes and 15 ecoregions, ranging from equatorial rainforests of the northwest
(Cabinda) and along the northern border with the Democratic Republic of Congo,
through the moist miombo woodlands and savannas of the central plateaus, to the
dry forests and woodlands of the southeast, and to the arid shrublands and Namib
Desert of the southwest. Isolated forests with Congolian affinities are found along
the Angolan Escarpment, and similar remnant patches of Afromontane forests are
found on some of the highest mountains such as Mount Moco and Mount Namba.
Despite the fact that at the beginning of the nineteenth century only a few species
of butterflies and skippers (Insecta: Lepidoptera: Papilionoidea) were recorded from
Angola, today a large number of taxa (at least 792 species and subspecies: Fig. 10.1,
Table 10.1 and Appendix) are known to occur in the country. However, extensive
areas are still poorly surveyed for butterflies, or have not been surveyed at all
(Fig. 10.2). This applies in particular to the southern provinces of Namibe, Cunene
and Cuando Cubango and the northwestern province of Zaire as well as most of
southern Moxico. Furthermore, the Baixa de Cassanje (Malanje), separated from
surrounding areas by steep escarpments, appears to have distinctive vegetation and
may produce some interesting butterflies. Although most of the localities where but-
Fig. 10.1 Cumulative number of species/subspecies of Papilionoidea reported from Angola from
1801–1819 (first records) to the recent decade – 2011–2017 – according to Appendix. For practical
reasons, species which first reference to the country was untraceable (marked in Appendix with a
▲) were included in the decade 2001–2010; species that are now assigned as faunistic novelties
to Angola (marked in Appendix with a ◘) are included in the last decade (2011–2017)
10 The Butterflies and Skippers (Lepidoptera: Papilionoidea) of Angola: An Updated… 169
Table 10.1 Number of species of Papilionoidea families and subfamilies known to occur in the
Afrotropical Region and Angola (with % of Afrotropical species present in the country), and
number of species endemic to Angola (with % of endemism shown)
Family Afrotropical Angola Endemism
Subfamily N° N° | % N|%
HESPERIIDAE 618 134 | 22 5 | 3.7
Coeliadinae 21 7 | 33
Pyrginae 216 48 | 22
Heteropterinae 27 5 | 19
Hesperiinae 334 74 | 22
PAPILIONIDAE 101 33 | 33 3 | 9.1
PIERIDAE 200 67 | 34 5 | 7.5
Pseudopontiinae 5 2 | 40
Coliadinae 14 8 | 57
Pierinae 181 57 | 32
LYCAENIDAE 1837 210 | 11 18 | 8.6
Miletinae 119 11 | 9
Poritiinae 658 53 | 8
Theclinae 301 42 | 14
Aphnaeinae 260 21 | 8
Polyommatinae 496 83 | 17
RIODINIDAE 15 4 | 27 0|0
NYMPHALIDAE 1634 344 | 21 26 | 7.6
Libytheinae 5 2 | 40
Danainae 26 9 | 35
Satyrinae 347 50 | 14
Charaxinae 190 56 | 30
Apaturinae 3 1 | 33
Nymphalinae 73 35 | 48
Cyrestinae 1 1 | 100
Biblidinae 31 16 | 52
Limenitidinae 702 97 | 14
Heliconiinae 256 77 | 30
TOTALS 4405 792 | 18
terflies and skippers have been collected in Angola have been determined (Mendes
et al. 2013b), some localities previously reported for a few species remain untraced
despite searches by us, using the detailed maps of the Junta de Investigações do
Ultramar (JIU 1948–1963).
The accumulation of knowledge in regard to Angolan butterflies has been con-
strained by several factors. The two largest Angolan entomological collections,
deposited in the Museu do Dundo (Lunda-Norte) and in the Instituto de Investigação
Agronómica (Huambo), have never been studied in detail. In addition little field-
work was carried out in Angola during the post-independence period because of the
protracted civil war. Finally, the vastness of the country and the difficulty in access-
ing many remote regions has impeded progress.
Fig. 10.2 Map of Angola showing, marked in orange, the known areas surveyed for the
Papilionoidea from the beginning of their study, in the nineteenth century to the present day – each
square ca. 33 × 33 km. The collecting pressure varies across the country, from ‘squares’ where
samples were obtained only once, in passing, to others where the collectors were based for months
Until recently the Hesperiidae (skippers) were placed in the superfamily

Hesperioidea, separate from the rest of the butterflies, which were placed in the
superfamily Papilionoidea. However, today the skippers and butterflies are all
placed in the Papilionoidea (e.g. Heikkilä et al. 2012). The classification used for
butterflies in this chapter is based on Williams (2018), Espeland et al. (2018) and
Dhungel and Wahlberg (2018). Six families of butterflies are represented in Angola,
namely Papilionidae, Hesperiidae, Pieridae, Riodinidae, Lycaenidae and
Nymphalidae.
History of Research on the Papilionoidea of Angola
The first known reference to butterflies obtained in Angola is by Latreille and Godart
(1819), who reported the presence of Colotis euippe (Linnaeus, 1758) and described
Acraea parrhasia servona. In the decade between 1871 and 1880, Druce (1875)
reported about 90 species from Angola for the first time, a number of these being
descriptions of species new to science. By the end of the nineteenth century a total
of 214 butterfly taxa were known from Angola.
The first contributions to our knowledge of Angolan butterflies by Portuguese

researchers were only made in the middle nineteen hundreds. These were the result
of the activities in Angola of the Centro de Zoologia (CZ) of the Junta de
Investigações do Ultramar, coordinated by its first director Fernando Frade. In 2014
this research institution was renamed the Instituto de Investigação Científica
Tropical (IICT). Working from these large zoological collections, as well as from
further specimens obtained in Angola by Amélia Bacelar (1948, 1956, 1958a, b,
1961) and Miguel Ladeiro (1956), considerably expanded the list of Angolan but-
terflies. Most of this material, obtained during colonial times, was stored in Lisbon,
with corrections to the published identifications only being made recently. All of
this material has now been integrated into the collections of the Museu Nacional de
História Natural e da Ciência (MUHNAC). Albert Monard (1956) of the La Chaux-
de-Fonds Swiss Museum also studied other material obtained by the CZ missions.
Significant contributions in the twentieth century were also made by Weymer (1901)
on the southern Angolan species, and by Evans (1937) on the Hesperiidae. All of the
then known Angolan butterflies were listed by Aurivillius (in Seitz) in 1928. All of
the Angolan Charaxinae were dealt with by Henning in his 1988 book on the African
taxa of this family. The 339 taxa added to the faunal list during the twentieth century
brought the total to 553 known butterfly taxa for Angola.
During the first 18 years of the twenty-first century, 239 further taxa were added
to the total. In the first decade of the present century, most of the new information
was due to several contributions by Libert (1999, 2000, 2004) on the Lycaenidae,
and by Gardiner (2004). The latter author listed taxa from the southeastern Cuando
Cubango province, which borders the Caprivi Strip of Namibia. Cuando Cubango
and the easternmost province of Moxico are the only provinces in Angola with
Zambezian fauna. To these taxa we add our own contributions (Bivar-de-Sousa and
Mendes 2006, 2007, 2009a, b; Mendes and Bivar-de-Sousa 2006a, b, 2007a, b,
2009a, b, c, d). Over the last 8 years 33 species were described as new or recorded
for the first time from Angola by Mendes and Bivar-de-Sousa (2012, 2017) Mendes
et al. (2013a, 2017, 2018), Bivar-de-Sousa and Mendes (2014) and Bivar-de-Sousa
et al. (2017), Turlin and Vingerhoedt (2013) and Pierre and Bernaud (2013). Finally,
66 further taxa are now recorded as faunistic novelties for the country (Appendix).
The current total number of butterfly taxa for Angola now stands at 792.
Sources Consulted for the Checklist
In preparing this revised checklist of the Papilionoidea, the following collections of

Angolan butterflies held by institutions in Portugal were examined: Museu Nacional
de História Natural e da Ciência (MUHNAC) in Lisbon, Museu de História Natural
da Universidade do Porto (MHNC-UP), Liceu Nun’Álvares in the Caldas da Saúde
and the Singeverga Order of St Benedict Abbey in Areias. Major contributions to
these collections were made by A Bivar-de-Sousa (Luanda district and Cuanza-
Norte, Cuanza-Sul and Moxico Provinces), António Figueira, (northwestern
Angola), Mário Macedo (northern Angola), Passos de Carvalho (Huambo and
Cuanza-Norte Provinces), Carneiro Mendes and Pessoa Guerreiro. The Angolan

insect collection of Nozolino de Azevedo (mainly Huambo Province), maintained
and made available by his widow, was also studied.
The collections in the MUHNAC in Lisbon were destroyed by a fire in March
1978. However, prior to the fire BS had studied some of the material and published
his findings. In 1995 LM studied the collections, mainly of Barros Machado and
Luna de Carvalho, in the Dundo Museum in Angola but there was insufficient time
to do a detailed analysis. We did not inspect the entomological collections in the
former Instituto de Investigação Agronómica de Angola, collected mainly by Passos
de Carvalho, but they are apparently in good condition. No entomological collec-
tions were found by LM, in 1995 and 2013, at the Museu de História Natural de
Luanda. Material collected from 2010 to 2014 by Ruben Capela and Carmen Van-
Dúnen Santos of Agostinho Neto University, Luanda and Artur Serrano of the
Faculty of Sciences, Lisbon University, was examined by us.
In addition, images of live specimens published by Lautenschläger and Neinhhuis
(2014) were examined, as were several images presented by Jorge Palmeirim of
Lisbon University and Pedro Vaz Pinto of the Kissama Foundation.
Taxa Excluded from the Checklist
A number of taxa have erroneously been reported to occur in Angola. This was due
mainly to probable misidentifications or mislabelled specimens. Some older records
are omitted because the known range of the taxon is unlikely to include Angola. A
list of the omitted taxa is given below.
• Hesperiidae: Eretis djaelaelae (Wallengren, 1857), Metisella metis (Linnaeus,
1764), Kedestes chaca (Trimen, 1873), Platylesches chamaeleon (Mabille,
1891).
• Papilionidae: Papilio menestheus Drury, 1773, Graphium taboranus (Oberthür,
1886), Graphium (Arisbe) junodi (Trimen, 1893).
• Pieridae: Eurema brigitta (Stoll, 1780), Colotis chrysonome (Klug, 1829),
Colotis ephyia (Klug, 1829), Belenois theora (Doubleday, 1846), Mylothris
rubricosta (Mabille, 1890), Mylothris similis Lathy, 1906.
• Lycaenidae: Telipna acraea (Westwood, 1851), Cooksonia abri Collins &
Larsen, 2008, Mimacraea darwinia Butler, 1872, Liptena bassae Bethune-Baker,
1926, Aethiopana honorius honorius (Fabricius, 1793), Stempfferia uniformis
(Kirby, 1887), Stempfferia dorothea (Bethune-Baker, 1904), Oxylides faunus
(Drury, 1773), Dapidodigma hymen (Fabricius, 1775), Aloeides molomo (Trimen,
1870), Leptomyrina lara (Linnaeus, 1764), Deudorix livia (Klug, 1834),
Neurellipes onias (Hulstaert, 1924), Zintha hintza (Trimen, 1864).
• Riodinidae: Afriodinia caeca semicaeca (Riley, 1932), Afriodinia gerontes
(Fabricius, 1781).
• Nymphalidae: Bicyclus milyas (Hewitson, 1864), Ypthima congoana Overlaet,
1955, Charaxes jahlusa argynnides Westwood, 1864, Junonia touhilimasa
Vuillot, 1892, Neptis continuata Holland, 1892, Neptis strigata Aurivillius,

1894, Evena oberthueri (Karsch, 1894), Euriphene atrovirens (Mabille, 1878),
Bebearia mardania (Fabricius, 1793), Euphaedra morini Hecq, 1983, Euphaedra
xypete (Hewitson, 1865), Euphaedra campaspe (Felder & Felder, 1867),
Euphaedra inanum (Butler, 1873), Euphaedra eupalus (Fabricius, 1781).
A Revised Checklist of the Papilionoidea of Angola
A revised and annotated checklist of the Papilionoidea of Angola (Appendix) con-

firms the presence of at least 792 taxa in the country. Their presence is based mainly
on verification by the authors of this chapter. Some taxa, recorded by other authors,
are accepted because they were, with rare exceptions, reported by more than one
author, are based on reliable literature records, or because Angola falls within their
putative geographical range. In the Checklist, the first reference to their occurrence
in Angola is given, followed by the sources of validation of the record and their
preferred habitat(s). Occasionally more than one subspecies of a particular species
occurs in the country. This is due to both the size and ecological diversity of Angola.
A number of forests, especially gallery-forests, are independent of each other as are
the fragmented forests of the Angolan Escarpment. In addition, the southeastern
parts of Moxico and the Cuando Cubango provinces are part of the Zambesi Basin;
consequently their fauna has affinities with that of eastern Africa.
As far as habitats are concerned the great majority of the Angolan Papilionoidea,
as might be expected, occur in forest, both wet and dry (Appendix). However, the
Hesperiidae and Pieridae appear to be almost as diverse in moist woodland (miombo)
and dry woodland as they are in forest. The number of Pieridae in dry woodland and
miombo is similar, while the number of species in dry woodland, arid shrubland and
grassland surpasses that of wet forest. The subfamily Nymphalinae is more diverse
in miombo than forest and equally diverse in savanna. The Heliconiinae
(Nymphalidae) in savanna are almost as diverse as they are in wet forest.
Composition, Diversity and Endemism
All six families and all of the subfamilies (except the Lycaeninae, Leach, 1815) of
Afrotropical butterflies are represented in Angola (Table 10.1).
One genus and 56 species/subspecies of Papilionoidea are endemic to Angola,
many of which were described over the last few decades. The endemic genus
Mashunoides, Mendes and Bivar-de-Sousa 2009a, b, c, d (Nymphalidae:
Satyrinae) is confined to Cuando Cubango Province, in the ecotone between
miombo and savanna/dry woodland mosaic. Endemism rates for Angolan butter-
fly families are highest for the Papilionidae and Lycaenidae and lowest for the
Hesperiidae and Riodinidae (Tables 10.1 and 10.2). Examples of endemic species
are illustrated in Fig. 10.3.
Table 10.2 Endemic butterfly species and subspecies in Angola

Family Endemic species Endemic subspecies
Hesperiidae Eagris multiplagata Calleagris jamesoni ansorgei
Abantis bergeri Eretis herewardi rotundimacula
Spialia colotes colotes
Papilionidae Papilio bacelarae Papilio macinnoni benguellae
Papilio chitondensis
Pieridae Mylothris carvalhoi Appias epaphia angolensis
Appias phaola uigensis
Appias sylvia ribeiroi
Mylothris spica gabela
Lycaenidae Alaena rosei Liptena homeyeri straminea
Cooksonia nozolinoi Falcuna libyssa angolensis
Falcuna lacteata Cigarits modestus modestus
Deloneura barca Leptomyrina henningi
Aloeides angolensis angolensis
Zeritis krystyna
Cupidesthes vidua
Uranothauma nozolinoi Lepidochrysops
ansorgei
Lepidochrysops flavisquamosa
Lepidochrysops fulvescens
Lepidochrysops hawker
Lepidochrysops nacrescens
Lepidochrysops reichenowi
Nymphalidae Brakefieldia angolensis Amauris crawshayi angola,
Brakefieldia ochracea Amauris dannfelti dannfelti
Neita bikuarica Charaxes fulvescens
Mashunoides carneiromendesi rubenarturi,
Charaxes figuerai Charaxes macclouni carvalhoi,
Charaxes ehmckei Charaxes lucretius saldanhai
Precis larseni Charaxes jahlusa angolensis
Bebearia hassoni Charaxes minor karinae
Euphaedra divoides Charaxes trajanus bambi
Euphaedra uigensis Palla ussheri hassoni
Acraea bellona Sevenia occidentalium penricei
Acraea lapidorum, Euphaedra harpalyce
Acraea onerata commineura
Acraea violarum anchietai
Conservation
Because butterflies are sensitive to changing environmental conditions and are taxo-
nomically well known, they are valuable as indicators of ecological dynamics. They
are also key drivers of ecological processes. In particular, adult butterflies are active
pollinators of many plants and the imagos and larvae are an important source of
nutrition for a diverse range of vertebrate and invertebrate predators and insect para-
sitoids. Their conservation importance is also due to their positive and occasionally
negative economic impacts. Although humans utilise mainly moth caterpillars as a
Fig. 10.3 The holotype specimens of endemic Angolan papilionoidea: Left to right, top to bottom
(V Ventral, D Dorsal): 1. Abantis bergeri male D (Mendes and Bivar-de-Sousa 2009a, b, c, d), 2.
Eagris multiplagata male V (Bivar-de-Sousa and Mendes 2007), 3. Cooksonia nozolinoi female D
(Mendes and Bivar-de-Sousa 2007), 4. Papilio bacelarae male D (Bivar-de-Sousa and Mendes
2009a, b), 5. Mashunoides carneiromendesi male V (Mendes and Bivar-de-Sousa 2009a, b, c, d),
6. Charaxes jahlusa angolensis male D (Mendes et al. 2017), 7. Euxanthe trajanus bambi male D
(Bivar-de-Sousa and Mendes 2006), 8. Euphaedra (Euphaedrana) divoides male V (Bivar- de-
Sousa and Mendes 2018)
food source, the larvae of the skipper Coeliades libeon is much appreciated. A lim-
ited number of butterfly species are agricultural pests, including Papilio demodocus
(young citrus orchards), Lampides boeticus (cultivated Leguminosae) and Acraea
acerata (sweet-potatoes). A few species, such as Pyrrhochalcia iphis and Zophopetes
dysmephila, may cause damage in coconut and oil-palm plantations.
In terms of species of conservation concern, information on the status of Angolan
butterflies is very limited. Many species of Angolan butterflies are obviously abun-
dant and widespread, both within and outside the country. Those taxa that appear to
be rare and/or more localised may be genuinely rare or local but this may simply
reflect a paucity of information. This makes it difficult or impossible to propose
rational conservation measures at present. The urgent need for more fieldwork, par-
ticularly in regard to the endemic taxa, is thus highlighted. In the meantime habitat
conservation, especially with respect to isolated forest patches, can be considered as
part of a wider effort to conserve both the fauna and flora of the country.
Potential Future Discoveries and Research
Considering the number of taxa new to science described in the last few decades
there are almost certainly further undiscovered butterfly taxa in Angola. Vast areas
of the country remain unexplored, mainly because of inaccessibility and post-
independence political instability. Not only will new taxa be found but also known
taxa from bordering countries will be added to the list of Angolan butterflies during
future fieldwork. This work will also improve our knowledge in regard to the distri-
bution of the taxa in the country. Finally, almost nothing is known about the habi-
tats, behaviours, early stages and larval host plants of Angolan butterflies, making
these fertile areas for future research on the fauna. More information concerning all
the endemic taxa is urgently needed in order to determine conservation priorities.
Appendix
Checklist of Papilionoidea recorded from Angola (by Family | Sub-family).

For the species’ authors and references, Bivar-de-Sousa is abbreviated as BS and
Mendes as M; (*) = Taxa with Angola as type locality; ◘ = Taxa now reported as
new to Angola; ▲ = Previous references existing but not traced (several species
when reported from Angola were assigned under names today considered syn-
onyms, others at species level – the corresponding Angolan endemics were
described later).
V = Validation of the taxon – 1: Endemic (restricted to Angola); 2: Restricted to
Angola and to the neighbouring northern countries – Gabon, Congo and/or DRC; 3:
Restricted to Angola and to the neighbouring eastern countries – southern DRC
(former Shaba) and/or Zambia; 4: Restricted to Angola and to the southern and
south-eastern neighbouring countries – Namibia and/or Botswana; 5: Species/sub-

species with material studied by the authors; 6: Taxa exclusively known in the coun-
try from previously collected unstudied material – known from bibliographic
references only. H – Preferred habitat: A: Humid forest – primary and secondary
wet forest, gallery and riverine forest, forest edge; B: Dry forest, including dry for-
est and savanna mosaics; C: Brachystegia woodland (miombo) and other woodland;
D: Mixed savanna with or without trees; E: Swampy areas, including the northeast-
ern moist thicket and savanna mosaic; F: Arid shrubland and grassland; G: Rocky
hillsides; H: Ubiquitous or almost ubiquitous; X: Caterpillars (almost) monopha-
gous, the imagos range dependent on the presence of host-plants
Taxon First reference for Angola V H

Hesperiidae | Coeliadinae
Coeliades bixana (Evans, 1940) Evans, 1937, as C. bixae 6 A
Coeliades c. chalybe (Westwood, 1852) Evans, 1937 5 A
Coeliades libeon (Druce, 1875) Druce, 1875 (*) 5 B,C
Coeliades f. forestan (Stoll, 1782) Ladeiro, 1959 5 H
Coeliades hanno (Plötz, 1879) Evans, 1937 6 A
Coeliades pisistratus (Fabricius, 1793) Bacelar, 1948 6 D
Pyrrhochalcia iphis dejongi (Collins & Bacelar, 1956, as P. iphis 2,5 A
Larsen, 2008)
Hesperiidae | Pyrginae
Apallaga rutilans (Mabille, 1877) M et al., 2013a 5 A
Apallaga h. homeyeri (Plötz, 1880) Plötz, 1880 (*) 6 A
Celaenorrhinus p. proxima (Mabille, M & BS, 2009a, b, c, d 5 A
1877)
Tagiades flesus (Fabricius, 1781) Evans, 1937 5 A,B
Eagris lucetia (Hewitson, 1875) Aurivillius, 1928 5 A
Eagris decastigma fuscosa (Holland, M et al., 2013a 5 A
1893
Eagris tigris liberti (Collins & Larsen, Evans, 1937, as E. tigris 6 A
2005)
Eagris h. hereus (Druce, 1875) Druce, 1875 (*) 6 A
Eagris t. tetrastigma (Mabille, 1891) M et al., 2013a 5 A
Eagris multiplagata (BS & M, 2007) BS & M, 2007 (*) 1,5 A
Ortholexis hollandi (Druce, 1909) f. Evans, 1937 6 A
karschi (Evans, 1937)
Calleagris hollandi (Butler, 1897) Evans, 1937 5 C
Calleagris jamesoni ansorgei (Evans, Weymer, 1901 (*), as C. jamesoni 1,5 C
1951)
Calleagris l. lacteus (Mabille, 1877) Bacelar, 1961 6 A
Eretis lugens (Rogenhofer, 1891) Larsen, 2005 5 D
Eretis herewardi rotundimacula (Evans, Evans, 1937 (*) 1,5 C
1937)
Eretis melania (Mabille, 1891) Evans, 1937 5 C,D
(continued)

Sarangesa loelius (Mabille, 1877) Bacelar, 1948 5 C
Sarangesa l. lucidella (Mabille, 1891) M et al., 2013a 5 D
Sarangesa motozi (Wallengren, 1857) Aurivillius, 1928 5 A,C
Sarangesa phidyle (Walker, 1870) Evans, 1937 5 B,D
Sarangesa s. seineri (Strand, 1909) Evans, 1937 5 D
Sarangesa p. pandaensis (Joicey & Evans, 1937 3,5 C
Talbot, 1921)
Sarangesa bouvieri (Mabille, 1877) Evans, 1937 6 B
Sarangesa brigida sanaga (Miller, 1964) M et al., 2013a 5 A
Sarangesa maculata (Mabille, 1891) M et al., 2013a 5 A,B,C
Triskelionia tricerata (Mabille, 1891) M et al., 2013a 6 A
Caprona cassualala (Bethune-Baker, Bethune-Baker, 1911 (*) 5 D,F
1911)
Caprona pillaana (Wallengren, 1857) M & BS, 2013a 5 D,F
Netrobalane canopus (Trimen, 1864) M & BS, 2013a 5 D,A
Leucochitonea levubu (Wallengren, 1857) Weymer, 1901 5 C,D
Abantis tettensis (Hopffer, 1855) Aurivillius, 1928 5 C,D
Abantis bergeri (M & BS, 2009a, b, c, d) M & BS, 2009a, b, c, d (*) 1,5 C
Abantis paradisea (Butler, 1870) Weymer, 1901 6 C
Abantis zambesiaca (Westwood, 1874) Weymer, 1901 5 D,C
Abantis contigua (Evans, 1937) Evans, 1937 5 C
Abantis venosa (Trimen, 1889) M et al., 2013a 5 C
Abantis vidua (Weymer, 1901) Weymer, 1901 3 C
Spialia m. mafa (Trimen, 1870) Weymer, 1901 6 D
Spialia spio (Linnaeus, 1764) Weymer, 1901 5 D
Spialia delagoae (Trimen, 1898) Larsen, 1996 5 D
Spialia c. colotes (Druce, 1875) Druce, 1875 (*) 1,5 D,B
Spialia colotes transvaaliae (Trimen, ▲ 5 D
1889)
Spialia ferax (Wallengren, 1863) M et al., 2013a 5 C,D
Spialia dromus (Plötz, 1884) Weymer, 1901 5 C,D
Spialia p. ploetzi (Aurivillius, 1891) Evans, 1937, as S. rebeli 6 A
Spialia secessus (Trimen, 1891) Trimen, 1891 (*) 5 F
Gomalia e. elma (Trimen, 1862) Aurivilius, 1928 5 D
Hesperiidae | Heteropterinae
Metisella m. midas (Butler, 1894) Monard, 1956 5 E
Metisella a. angolana (Karsch, 1896) Karsch, 1896 (*) 5 B
Metisella willemi (Wallengren, 1857) M et al., 2013a, b 5 C
Metisella meninx (Trimen, 1873) Evans, 1937 6 E
Lepella lepeletier (Latreille, 1824) Druce, 1875 5 F
Hesperiidae | Hesperiinae
Astictopterus abjecta (Snellen, 1872) Snellen, 1872 (*) 6 A
Astictopterus punctulata (Butler, 1895) M et al., 2013a 5 C
Kedestes mohozutza (Wallengren, 1857) Monard, 1956 5 D
(continued)

Kedestes nerva paola (Plötz, 1884) Plötz, 1884 (*) 5 A
Kedestes brunneostriga (Plötz, 1884) Plötz, 1884 (*) 5 C
Kedestes straeleni (Evans, 1956) M et al., 2013a 5 C
Kedestes l. lema (Neave, 1910) Evans, 1937 3 C
Kedestes callicles (Hewitson, 1868) Aurivillius, 1928 5 C
Gorgyra mocquerysii (Holland, 1896) Evans, 1937 5 A
Gorgyra diversata (Evans, 1937) Evans, 1937 6 A
Ceratrichia nothus makomensis (Strand, M et al., 2013a 5 A
1913)
Ceratrichia punctata (Holland, 1896) Evans, 1937 6 A
Teniorhinus harona (Westwood, 1881) Weymer, 1901, as Oxypalpus ruso 5 C
Teniorhinus ignita (Mabille, 1877) Monard, 1956 5 B
Pardaleodes edipus (Stoll, 1781) Bacelar, 1948 6 A
Pardaleodes i. incerta (Snellen, 1872) Evans, 1937 5 A,D
Pardaleodes sator pusiella (Mabille, Mabille, 1877 (*) 5 A
1877)
Pardaleodes t. tibullus (Fabricius, 1793) M & BS, 2009 5 A
Acada biseriata (Mabille, 1893) Evans, 1937 5 C
Parosmodes lentiginosa (Holland, 1896) Evans, 1937 5 A
Parosmodes m. morantii (Trimen, 1873) Weymer, 1901 5 C/D
Osmodes laronia (Hewitson, 1868) Druce, 1875 6 A
Osmodes thora (Plötz, 1884) Evans,1937 6 A
Acleros mackenii olaus (Plötz, 1884) Druce, 1875, the species 5 A
Acleros nigrapex (Strand, 1913) M et al., 2013a 5 A
Acleros ploetzi (Mabille, 1889) M & BS, 2009 5 A
Semalea arela (Mabille, 1891) M et al., 2013a 5 A
Semalea pulvina (Plötz, 1879) M & BS, 2009 5 A
Semalea sextilis (Plötz, 1886) M et al., 2013a 5 A
Hypoleucis o. ophiusa (Hewitson, 1866) M & BS, 2009 5 A
Meza indusiata (Mabille, 1891) Larsen, 2005 5 A,B
Meza meza (Hewitson, 1877) Hewitson, 1877 (*) 5 A
Meza c. cybeutes (Holland, 1894) Evans, 1937 6 A
Meza mabillei (Holland, 1893) M et al., 2013a 5 A
Paronymus ligora (Hewitson, 1876) Hewitson, 1876 (*) 6 A
Andronymus n. neander (Plötz, 1884) Evans, 1937 5 A
Andronymus c. caesar (Fabricius, 1793) Aurivillius, 1928 as A. caesar 5 A
Andronymus caesar philander (Hopffer, Aurivillius, 1928 as A. caesar 6 A
1855)
Andronymus hero (Evans, 1937) Evans, 1937 5 A
Andronymus helles (Evans, 1937) Evans, 1937 5 A
Chondrolepis niveicornis (Plötz, 1882) Plötz, 1882 (*) 5 E
Zophopetes dysmephila (Trimen, 1868) Aurivillius, 1928, as Z. schultzi 6 A,D
Zophopetes cerymica (Hewitson, 1867) M & BS, 2009 5 X
Gamia shelleyi (Sharpe, 1890) M et al., 2013a 5 A
(continued)

Gretna cylinda (Hewitson, 1876) Aurivillius, 1928 (*) 5 A,C
Gretna waga (Plötz, 1886) M et al., 2013a 5 A,C
Pteroteinon laufella (Hewitson, 1868) Druce, 1875 5 B,C
Pteroteinon caenira (Hewitson, 1867) M & BS, 2009 5 B
Pteroteinon concaenira (Belcastro & M et al., 2013a 6 B
Larsen, 1996)
Leona maracanda (Hewitson, 1876) Hewitson, 1876 (*) 5 A
Caenides dacela (Hewitson, 1876) Williams, 2007 5 A
Monza cretacea (Snellen, 1872) Evans, 1937 5 B
Fresna nyassae (Hewitson, 1878) Aurivillius, 1928 5 C,D
Platylesches langa (Evans, 1937) M et al., 2013a 6 C
Platylesches moritili (Wallengren, 1857) Trimen, 1891 5 C,D
Platylesches robustus (Neave, 1910) M et al., 2013a 6 C
Platylesches cf. batangae (Holland, 1894) ▲ 5 B
Brusa allardi (Berger, 1967) M et al., 2013a 6 C
Zenonia zeno (Trimen, 1864) Plötz, 1883, as Hesperia coanza 5 A,C
Pelopidas m. mathias (Fabricius, 1798) Evans, 1937 5 C
Pelopidas thrax (Hübner, 1821) Gardiner, 2004 5 D
Borbo fallax (Gaede, 1916) M & BS, 2009a, b, c, d 5 D
Borbo fanta (Evans, 1937) Evans, 1937 5 D
Borbo sirena (Evans, 1937) M et al., 2013a 5 A,B,C
Borbo b. borbonica (Boisduval, 1833) ▲ 5 D
Borbo detecta (Trimen, 1893) Weymer, 1901 5 B
Larsenia gemella (Mabille, 1884) Evans, 1937 5 D
Borbo micans (Holland, 1896) M & BS, 2009 5 E
Larsenia perobscura (Druce, 1812) M et al., 2013a 5 D
Borbo f. fatuellus (Hopffer, 1855) M & BS, 2009 5 B,C
Larsenia holtzi (Plötz, 1883) Plötz, 1883 (*) 5 D
Parnara monasi (Trimen, 1889) Evans, 1937 5 E
Afrogegenes hottentota (Latreille, 1824) Weymer, 1901 5 C,D
Afrogegenes letterstedti (Wallengren, Evans, 1937 5 D
1857)
Gegenes pumilio gambica (Mabille, 1878) M & BS, 2009 5 D
Papilionidae
Papilio a. antimachus (Drury, 1782) Carvalho, 1962 5 A
Papilio zalmoxis (Hewitson, 1864) BS, 1983 5 A
Papilio bacelarae (BS & M., 2009) BS & M, 2009 (*) 1,5 A
Papilio f. filaprae (Suffert, 1904) Druce, 1875, as P. cypraeophila 5 A
Papilio m. mechowi (Dewitz, 1881) Dewitz, 1881 (*) 5 A
Papilio mechowianus (Dewitz, 1885) Aurivillius, 1928 6 A
Papilio zenobia (Fabricius, 1775) BS & Fernandes, 1966 5 A
Papilio cynorta (Fabricius, 1793) Druce, 1875 5 A
Papilio echerioides homeyeri (Plötz, Plötz, 1880 (*) 5 A,C
1880)
(continued)

Papilio chitondensis (BS & Fernandes, BS & Fernandes, 1966 (*) 1,5 B,C
1966)
Papilio chrapkowskoides nurettini Bacelar, 1956, as P. bromius 5 A
(Koçak, 1983)
Papilio n. nireus (Linnaeus, 1758) Druce, 1875 5 H
Papilio nireus lyaeus (Doubleday, 1845) Ladeiro, 1956 5 A,D
Papilio sosia pulchra (Berger, 1950) BS & Fernandes, 1966 5 A
Papilio mackinnoni benguellae (Jordan, Jordan, 1908 (*) 1,5 A
1908)
Papilio d. dardanus (Brown, 1776) Druce, 1875 5 C
Papilio phorcas congoanus (Rothschild, BS & Fernandes, 1964 5 A
1896)
Papilio h. hesperus (Westwood, 1843) Aurivillius, 1928 5 A
Papilio l. lormieri (Distant, 1874) Bacelar, 1956 5 A
Papilio d. demodocus (Esper, 1798) Weymer, 1901 5 H
Graphium a. angolanus (Goeze, 1779) Goeze, 1779 (*) 5 H
Graphium schaffgotschi (Niepelt, 1927) Villiers, 1979 4,5 D
Graphium ridleyanus (White, 1843) Druce, 1875 5 A
Graphium latreillianus theorini Aurivillius, 1928 5 A
(Aurivillius,1881)
Graphium tynderaeus (Fabricius, 1793) BS & Fernandes, 1964 5 A
Graphium a. almansor (Honrath, 1884) Aurivillius, 1928 5 A,C
Graphium ucalegonides (Staudinger, Smith & Vane-Wright, 2001 2 A
1884)
Graphium h. hachei (Dewitz, 1881) Dewitz, 1881 (*) 5 A
Graphium poggianus (Honrath, 1884) Aurivillius, 1928 3 A
Graphium u. ucalegon (Hewitson, 1865) Bacelar, 1956 5 A
Graphium l. leonidas (Fabricius, 1793) Druce, 1875 5 H
Graphium antheus (Cramer, 1779) Druce, 1875 5 A,D
Graphium p. policenes (Cramer, 1775) Druce, 1875 5 A,C
Graphium p. porthaon (Hewitson, 1865) Gardiner, 2004 6 B,D
Pieridae | Pseudopontinae
Pseudopontia paradoxa (Felder & Felder, ◘ 5 A
1869)
Pseudopontia australis (Dixey, 1923) Snellen, 1882, as P. paradoxa 3,5 A
Pieridae | Coliadinae
Catopsilia florella (Fabricius, 1775) Weymer, 1901 5 H
Colias e. electo (Linnaeus, 1763) Gardiner, 2004 6 D
Colias electo hecate (Strecker, 1905) Bacelar, 1948 5 F
Eurema b. brigitta (Stoll, 1780) Butler, 1871 5 D,H
Eurema desjardinsi regularis (Butler, Mabille, 1877 5 D,C
1876)
Eurema floricola leonis (Butler, 1886) Trimen, 1891, as E. floricola 6 B
Eurema hapale (Mabille, 1882) Ladeiro, 1956 5 A,B
Eurema hecabe solifera (Butler, 1875) Butler, 1875 (*) 5 D
(continued)

Eurema senegalensis (Boisduval, 1836) Butler, 1871 5 A
Pieridae | Pierinae
Pinacopteryx e. eriphia (Godart, 1819) Butler, 1871 5 B,D
Nepheronia a. argia (Fabricius, 1775) Druce, 1875 5 A
Nepheronia b. buquetii (Boisduval, 1836) Druce, 1875 5 B,D
Nepheronia p. pharis (Boisduval, 1836) Aurivillius, 1928 5 B
Nepheronia thalassina verulanus (Ward, Bacelar, 1958a, b 5 B
1871)
Eronia cleodora Hübner, 1823 Aurivillius, 1928 6 D
Afrodryas leda (Boisduval, 1847) Bacelar, 1961 5 B
Teracolus a. agoye (Wallengren, 1857) Weymer, 1901 5 D
Colotis calais williami (Henning & Willis, 2009 6 D
Henning, 1994)
Colotis antevippe gavisa (Wallengren, Trimen, 1891 5 D
1857)
Colotis celimene pholoe (Wallengren, Talbot, 1939 4,5 F
1860)
Colotis annae walkeri (Butler, 1884) Butler, 1884 (*) 4,5 F
Colotis doubledayi (Hopffer, 1862) Hopffer, 1862 (*) 5 D
Colotis e. euippe (Linnaeus, 1758) Latreille & Godart, 1819 5 B
Colotis euippe mediata (Talbot, 1939) Talbot, 1939 5 C,D
Colotis evagore antigone (Boisduval, Druce, 1875 5 B
1836)
Colotis e. evenina (Wallengren, 1857) Trimen, 1891 5 F
Colotis ione (Godart, 1819) Bacelar, 1961 6 D
Colotis regina (Trimen, 1863) Trimen, 1891 5 D
Colotis vesta rhodesinus (Butler, 1894) Bacelar, 1958a, b 5 D
Teracolus e. eris (Klug, 1829) Druce, 1875 5 D
Teracolus subfasciatus (Swainson, 1833) Aurivillius, 1928 5 C
Belenois aurota (Fabricius, 1793) Trimen, 1891 5 H
Belenois calypso dentigera (Butler, 1888) Druce, 1875, as B. calypso 5 A
Belenois welwitschii welwitschii Rogenhofer, 1890 (*) 5 C
(Rogenhofer, 1890)
Belenois crawshayi (Butler, 1894) Aurivillius, 1928 5 C,D
Belenois creona severina (Stoll, 1781) Butler, 1871 5 H
Belenois g. gidica (Godart, 1819) ▲ 5 D
Belenois r. rubrosignata (Weymer, 1901) Weymer, 1901 (*) 3,5 C
Belenois s. solilucis (Butler, 1874) Butler, 1874 (*) 5 A
Belenois sudanensis mayumbana (Berger, ◘ 2,5 A
1981)
Belenois sudanensis pseudodentigera ◘ 5 C
(Berger, 1981)
Belenois theuszi (Dewitz, 1889) Dewitz, 1889 (*) 5 A
Pieridae | Pierinae
Belenois t. thysa (Hopffer, 1855) ◘ 5 C
(continued)

Belenois thysa meldolae (Butler, 1872) Butler, 1872 (*) 5 A,B
Dixeia capricornus falkensteinii (Dewitz, Dewitz, 1879 (*) 2,5 B,C
1879)
Dixeia sp. ◘ 5 A
Dixeia pigea (Boisduval, 1836) Aurivillius, 1928 5 C
Pontia h. helice (Linnaeus, 1764) Willis, 2009 5 D
Appias epaphia angolensis (M & BS, M & BS, 2006 (*) 1,5 A,B
2006)
Appias perlucens (Butler, 1898) Butler, 1898 (*) 5 A
Appias phaola uigensis (M & BS, 2006) M & BS, 2006 (*) 1,5 A
Appias s. sabina (Felder & Felder, 1865) Druce, 1875 5 A
Appias sylvia nyassana (Butler, 1897) Druce, 1875, as Belenois 6 A?
Appias sylvia ribeiroi (M & BS, 2006) M & BS, 2006 (*) 1,5 A
Leptosia a. alcesta (Stoll, 1782) Druce, 1875 5 A
Leptosia h. hybrida (Bernardi, 1952) ◘ 5 A,C
Leptosia n. nupta (Butler, 1873) Butler, 1873 (*) 5 A
Leptosia wigginsi pseudalcesta (Bernardi, ◘ 5 A
1965)
Mylothris carvalhoi (M & BS, 2009) M & BS, 2009 (*) 1,5 A
Mylothris mavunda (Hancock & Heath, Koçak & Kemal, 2009 3 A?
1985)
Mylothris a. agathina (Cramer, 1779) Trimen, 1891 5 C
Mylothris asphodelus (Butler, 1888) Aurivillius, 1928 5 A
Mylothris elodina diva (Berger, 1954) Berger, 1981 2,5 C
Mylothris poppea (Cramer, 1777) Druce, 1875 5 A
Mylothris rembina (Plötz, 1880) Talbot, 1944 6 A
Mylothris rhodope (Fabricius, 1775) Talbot, 1944 5 D
Mylothris rueppellii rhodesiana (Riley, Talbot, 1944 5 C
1921)
Mylothris spica gabela (Berger, 1979) Berger, 1979 (*) 1,6 A
Mylothris sulphurea (Aurivillius, 1895) ◘ 5 A
Mylothris y. cf. yulei (Butler, 1897) ◘ 5 A
Lycaenidae | Miletinae
Euliphyra mirifica (Holland, 1890) Larsen, 2005 6 A
Aslauga m. marshalli (Butler, 1899) Larsen, 2005 6 A
Megalopaplpus zymna (Westwood, 1851) Ackery et al., 1995 5 A
Spalgis l. lemolea (Druce, 1890) Ladeiro, 1956 5 A,C
Lachnocnema angolanus (Libert, 1996) Libert, 1996 b (*) 5 A,D
Lachnocnema bamptoni (Libert, 1996) Libert, 1996 b (*) 6 C
Lachnocnema bibulus (Fabricius, 1793) Libert, 1996 b 5 A,C,D
Lachnocnema emperamus (Snellen, 1872) Snellen, 1872 5 A
Lachnocnema intermedia (Libert, 1996) Ladeiro, 1956, as L. durbani (*) 5 C
Lachnocnema laches (Fabricius, 1793) Libert, 1996 a 5 A,C
Lachnocnema r. regularis (Libert, 1996) Libert, 1996 c 6 C?
(continued)

Lycaenidae | Poritiinae
Alaena amazoula congoana (Aurivillius, Aurivillius, 1914 (*) 6 G
1914)
Alaena rosei (Vane-Wright, 1980) Vane-Wright, 1980 (*) 1,5 G
Pentila maculata pardalena (Druce, 1910) Stempffer & Bennett, 1961 6 A
Pentila amenaida (Hewitson, 1873) Hewitson, 1873 (*) 5 A
Pentila pauli benguellana (Stempffer & Stempffer & Bennett, 1961 (*) 5 A,B
Bennett, 1961)
Pentila t. tachyroides (Dewitz, 1879) Dewitz, 1879 (*) 5 A
Telipna acraeoides (Grose-Smith & Grose-Smith & Kirby, 1890 (*) 5 A
Kirby, 1890)
Telipna a. albofasciata (Aurivillius, 1910) Libert, 2005 6 A
Telipna atrinervis (Hulstaert, 1924) ◘ 5 A
Telipna cuypersi (Libert, 2005) Libert, 2005 6 A
Telipna nyanza katangae (Stempffer, Libert, 2005 6 A
1961)
Telipna s. sanguinea (Plötz, 1880) Aurivillius, 1928 5 A
Ornipholidotus gabonensis (Stempffer, ▲ 6 A
1947)
Ornipholidotus perfragilis (Holland, Libert, 2005 6 A
1890)
Ornipholidotus ugandae goodi (Libert, Libert, 2005 6 A
2000)
Cooksonia nozolinoi (M & BS, 2007) M & BS, 2007 (*) 1,5 C
Mimacraea charmian (Grose-Smith & Grose-Smith & Kirby, 1890 (*) 6 A
Kirby, 1890)
Mimacraea landbecki (Druce, 1910) Libert, 2000 b 5 A
Mimacraea marshalli (Trimen, 1898) Libert, 2000 b 5 A
Mimeresia debora deborula (Aurivillius, ◘ 5 A
1899)
Eresiomera osheba (Holland, 1890) ▲ 5 A
Citrinophila e. erastus (Hewitson, 1866) Aurivillius, 1928 6 A
Cnodontes vansomereni (Stempffer & ▲ 5 D
Bennett, 1953)
Liptena evanescens (Kirby, 1887) ◘ 5 A
Liptena fatima (Kirby, 1890) ◘ 5 A
Liptena h. homeyeri (Dewitz, 1884) ▲ 6 A
Liptena homeyeri straminea (Stempffer, Stempffer, Bennett & May, 1974 (*) 1,6 A
Bennett & May, 1974)
Liptena parva (Kirby, 1887) ◘ 5 A
Liptena undularis (Hewitson, 1866) Druce, 1875 5 A
Liptena xanthostola xantha (Grose-Smith, Larsen, 2005 6 A
1901)
Falcuna h. hollandii (Aurivillius, 1895) Ackery et al., 1995 6 A
Falcuna lacteata (Stempffer & Bennett, Stempffer & Bennett, 1963 (*) 1,6 A
1963)
(continued)

Falcuna libyssa angolensis (Stempffer & Stempffer & Bennett, 1963 (*) 1,5 A
Bennett, 1963)
Falcuna s. synesia (Hulstaert, 1924) Stempffer & Bennett, 1963 (*) 2 A
Tetrarhanis ilala etoumbi (Stempffer, ◘ 5 A
1964)
Tetrarhanis i. ilma (Hewitson, 1873) Hewitson, 1873 (*) 6 A
Larinopoda lircaea (Hewitson, 1866) Stempffer, 1957 6 A
Larinopoda tera (Hewitson, 1873) Aurivillius, 1928 5 A
Hewitsonia bitjeana (Bethune-Baker, ◘ 5 A
1915)
Hewitsonia k. kirbyi (Dewitz, 1879) Dewitz, 1879 (*) 6 A
Cerautola ceraunia (Hewitson, 1873) Larsen, 2005 6 A
Cerautola crowleyi leucographa (Libert, Libert, 1999 6 A
1999)
Hewitola hewitsonii (Mabille, 1877) Mabille, 1877 (*) 6 A
Cerautola miranda vidua (Talbot, 1935) Bacelar, 1958a, b 5 A
Epitola posthumus (Fabricius, 1793) Bacelar, 1956 5 A
Epitola urania (Kirby, 1887) Libert, 1999 6 A
Hypophytala h. hyetta (Hewitson, 1873) Hewitson, 1873 (*) 2 A
Stempfferia cercene (Hewitson, 1873) Hewitson, 1873 (*) 6 A
Stempfferia cinerea (Berger, 1981) Libert, 1999 2 A
Stempfferia michelae centralis (Libert, Libert, 1999 5 A
1999)
Deloneura barca (Grose-Smith, 1901) Grose-Smith, 1901 (*) 1,6 C?,D?
Deloneura cf. subfusca (Hawker-Smith, ◘ 5 F
1933)
Epitolina dispar (Kirby, 1887) Larsen, 2005 6 A
Epitolina melissa (Druce, 1888) Larsen, 2005 6 A
Lycaenidae | Theclinae
Myrina s. silenus (Fabricius, 1775) Druce, 1875 5 B,D
Myrina silenus ficedula (Trimen, 1879) Gardiner, 2004 6 D
Oxylides binza (Berger, 1981) Druce, 1875 as O. faunus 2,5 A
Lycaenidae | Theclinae
Oxylides feminina stempfferi (Berger, Libert, 2004 6 A
1981)
Syrmoptera amasa (Hewitson, 1869) Libert, 2004 6 A
Syrmoptera homeyerii (Dewitz, 1879) Dewitz, 1879 (*) 6 A
Dapidodigma demeter nuptus (Clench, Larsen, 2005 3,5 A
1961)
Lycaenidae | Aphnaeinae
Lipaphneus a. cf. aderna (Plötz, 1880) ◘ 5 A
Crudaria leroma (Wallengren, 1857) Gardiner, 2004 6 D
Aloeides angolensis (Tite & Dickson, Tite & Dickson, 1973 (*) 1,6 F
1973)
Aphnaeus erikssoni (Trimen, 1891) Trimen, 1891 (*) 5 C
(continued)

Aphnaeus orcas (Drury, 1782) Larsen, 2005 6 A
Aphnaeus affinis (Riley, 1921) Libert, 2013
Erikssonia acraeina (Trimen, 1891) Trimen, 1891 (*) 6 D
Pseudaletis a. agrippina (Druce, 1888) ▲ 5 A
Cigaritis ella (Hewitson, 1865) Gardiner, 2004 6 D
Cigaritis phanes (Trimen, 1873) Weymer, 1901 6 F
Cigaritis homeyeri (Dewitz, 1887) Aurivillius, 1928 5 C
Cigaritis m. modestus (Trimen, 1891) Trimen, 1891 (*) 1,4,5 A,C
Cigaritis mozambica (Bertoloni, 1850) ▲ 6 D
Cigaritis natalensis (Westwood, 1851) Ladeiro, 1956 6 C,D
Cigaritis trimeni congolanus (Dufrane, ▲ 2,5 A
1954)
Zeritis fontainei (Stempffer, 1956) Willis, 2009 6 C,G?
Zeritis krystyna (D’Abrera, 1980) D’Abrera, 1980 (*) 1,6 C?
Zeritis sorhagenii (Dewitz, 1879) Dewitz, 1879 (*) 6 C?
Axiocerces a. amanga (Westwood, 1881) Trimen, 1891 5 C
Axiocerces bambana orichalcea (Henning ◘ 5 B
& Henning, 1996)
Axiocerces amanga baumi (Weymer, Weymer, 1901 (*) ,5 C
1901)
Axiocerces t. tjoanae (Wallengren, 1857) Henning & Henning, 1996 6 B
Iolaus hemicyanus barnsi (Joicey & ◘ 2,5 A
Talbot, 1921)
Iolaus i. iasis (Hewitson, 1865) Larsen, 2005 6 A
Iolaus mimosae rhodosense (Stempffer & Gardiner, 2004 6 D
Bennett, 1959)
Iolaus obscura (Aurivillius, 1923) ◘ 4,5 D
Iolaus violacea (Riley, 1928) Riley, 1928 (*) 5 C
Iolaus pallene (Wallengren, 1857) Gardiner, 2004 6 D
Iolaus trimeni (Wallengren, 1875) Ackery et al., 1995 5 C,D
Iolaus iturensis (Joicey & Talbot, 1921) ▲ 6 C
Iolaus parasilanus mabillei (Riley, 1928) Riley, 1928 (*) 2 A
Iolaus s. silarus (Druce, 1885) Gardiner, 2004 5 C,D
Iolaus t. timon (Fabricius, 1787) Ackery et al., 1995 6 A
Hemiolaus vividus (Pinhey, 1962) Aurivillius, 1928, as caeculus 5 C,D
Stugeta bowkeri maria (Suffert, 1904) Druce, 1875, as bowkeri 3,5 B
Stugeta bowkeri tearei (Dikson, 1980) Gardiner, 2004 6 A,C,E
Hypolycaena a. antifaunus (Westwood, Druce, 1875 5 A
1851)
Hypolycaena h. hatita (Hewitson, 1865) Druce, 1875 5 A
Hypolycaena l. lebona (Hewitson, 1865) Druce, 1875 6 A
Hypolycaena naara (Hewitson, 1873) Hewitson, 1873 (*) 6 A
Hypolycaena nigra (Bethune-Baker, M & BS, 2012 5 A
1914)
(continued)

Hypolycaena p. philippus (Fabricius, Druce, 1875 5 D
1793)
Hypolycaena buxtoni spurcus (Talbot, M & BS, 2012 5 F?
1929 M & BS., 2012)
Pilodeudorix badhami (Carcasson, 1961) Libert, 2004 6 ?
Pilodeudorix caerulea (Druce, 1890) Libert, 2004 5 C,D
Pilodeudorix pseudoderitas (Stempffer, Larsen, 2005 6 A
1964)
Pilodeudorix zeloides (Butler, 1901) Libert, 2004 6 C
Paradeudorix cobaltina (Stempffer, 1964) Larsen, 2005 6 A
Leptomyrina henningi angolensis (M & M & BS, 2009 (*) 1,5 B
BS, 2009)
Pilodeudorix deritas (Hewitson, 1874) Hewitson, 1874 (*) 5 A
Pilodeudorix m. mera (Hewitson, 1873) Hewitson, 1873 (*) 5 A
Pilodeudorix otraeda genuba (Hewitson, ▲ 6 A
1875)
Hypomyrina nomenia (Hewitson, 1874) Larsen, 2005 5 A
Deudorix antalus (Hopffer, 1855) Bacelar, 1948 5 D,G
Deudorix caliginosa (Lathy, 1903) ▲ 6 C
Deudorix dinochares (Grose-Smith, 1887) Gardiner, 2004 5 C,D
Deudorix cf. diocles (Hewitson, 1869) Libert, 2004 5 C
Deudorix lorisona coffea (Jackson, 1966) Libert, 2004 5 A,C
Capys c. connexiva (Butler, 1896) Henning & Henning, 1988 6 X
Lycaenidae | Polyommatine
Anthene akoae (Libert, 2010) Libert. 2010 6 ?
Anthene alberta (Bethune-Baker, 1910) Aurivillius, 1928 5 A,C
Anthene a. amarah (Guérin-Méneville, Stempffer, 1957 5 D
1849)
Anthene lvida livida (Trimen, 1881) Gardiner, 2004 6 D
Anthene c. crawshayi (Butler, 1899) ◘ 5 D
Anthene d. definita (Butler, 1899) Gardiner, 2004 5 A
Anthene larydas (Cramer, 1780) Weymer, 1901 5 A
Anthene l. ligures (Hewitson, 1874) Hewitson, 1874 (*) 6 A
Anthene liodes (Hewitson, 1874) Aurivillius, 1909 6 A,D
Anthene l. lunulata (Trimen, 1894) Trimen, 1894 (*) 5 D
Anthene nigropunctata (Bethune-Baker, Gardiner, 2004 6 ?
1910)
Anthene princeps (Butler, 1876) Gardiner, 2004 5 A,D
Anthene r. rubricinctus (Holland, 1891) Aurivillius, 1928 6 A
Anthene sylvanus (Drury, 1773) Aurivillius, 1928 6 A
Anthene talboti (Stempffer, 1936) Libert, 2010 6 D
Neurellipes flavomaculatus (Grose-Smith Aurivillius, 1928 6 A
& Kirby, 1893)
Neurellipes lachares (Hewitson, 1878) Larsen, 2005 6 A
Neurellipes onias (Hulstaert, 1924) Willis, 2009 6 ?
(continued)

Neurellipes pyroptera (Aurivillius, 1895) Libert, 2010 6 A
Neurypexina lyzanius (Hewitson, 1874) Druce, 1875 6 A
Triclema lacides (Hewitson, 1874) Hewitson, 1874 (*) 6 A
Triclema lucretilis (Hewitson, 1874) Stempffer, 1957 6 A
Triclema cf. nigeriae (Aurivillius, 1905) Libert, 2010 5 D
Monile g. gemmifera (Neave, 1910) Libert, 2010 6 A
Cupidesthes vidua (Talbot, 1929) Talbot, 1929 (*) 1,6 C?
Pseudonacaduba aethiops (Mabille, Mabille, 1877 (*) 5 A
1877)
Pseudonacaduba s. sichela (Wallengren, Weymer, 1901 5 D
1857)
Lampides boeticus (Linnaeus, 1767) Butler, 1871 5 H
Uranothauma falkensteinii (Dewitz, Dewitz, 1879 (*) 5 A?
1879)
Uranothauma antinorii cf. felthami BS & M, 2007 5 A
(Stevenson, 1934)
Uranothauma h. heritsia (Hewitson, Stempffer, 1957 5 A,D
1876)
Uranothauma nozolinoi (BS & M., 2007) BS & M, 2007 (*) 1,5 C
Uranothauma poggei (Dewitz, 1879) Dewitz, 1879 (*) 5 A,C,D
Uranothauma c. cyara (Hewitson, 1876) Hewitson, 1876 (*) 5 A?
Cacyreus lingeus (Stoll, 1782) Gardiner, 2004 5 B,D
Cacyreus marshalli (Butler, 1898) Gardiner, 2004 6 D
Cacyreus virilis (Aurivillius, 1924) Aurivillius, 1924 (*) 6 D
Leptotes babaulti (Stempffer, 1935) Stempffer, 1957 5 A,D
Leptotes brevidentatus (Tite, 1958) Tite, 1958 (* – in part) 5 A,C,D
Leptotes jeanneli (Stempffer, 1935) Stempffer, 1957 5 A,D
Leptotes p. pirithous (Linnaeus, 1767) Snellen, 1882 5 D
Leptotes p. pulchra (Murray, 1874) Gardiner, 2004 5 E
Tuxentius calice (Hopffer, 1855) Gardiner, 2004 5 E
Tuxentius c. carana (Hewitson, 1876) Hewitson, 1876 (*) 5 A
Tuxentius margaritaceus (Sharpe, 1892) Larsen, 2005 6 A
Tuxentius m. melaena (Trimen, 1887) Aurivillius, 1928 6 C,D
Tarucus sybaris linearis (Aurivillius, Aurivillius, 1928 6 D
1924)
Actizera lucida (Trimen, 1883) Stempffer, 1957 5 F
Eicochrysops eicotrochilus (Bethune- ▲ 6 C
Baker, 1924)
Eicochrysops hippocrates (Fabricius, Gardiner, 2004 5 A,B,D
1793)
Eicochrysops messapus mahallakoaena Weymer, 1901, as messapus 6 C
(Wallengren, 1857)
Cupidopsis c. cissus (Godart, 1824) Ladeiro, 1956 5 E
Cupidopsis j. jobates (Hopffer, 1855) Stempffer, 1957 5 D
Euchrysops barkeri (Trimen, 1893) Aurivillius, 1928 6 C
(continued)

Euchrysops malathana (Boisduval, 1833) Stempffer, 1957 5 C,D
Euchrysops osiris (Hopffer, 1855) Druce, 1875 5 C,D
Euchrysops subpallida (Bethune-Baker, Gardiner, 2004 6 F,G
1923)
Lepidochrysops abyssiniensis loveni
(Aurivillius, 1921)
Lepidochrysops ansorgei (Tite, 1959) Tite, 1959 (*) 1,5 C,D
Lepidochrysops chlouages (Bethune- ▲ 5 A,D
Baker, 1923)
Lepidochrysops flavisquamosa (Tite, Tite, 1959 (*) 1,6 C
1959)
Lepidochrysops fulvescens (Tite, 1961) Tite, 1961 (*) 1,6 C
Lepidochrysops g. glauca (Trimen, 1887) Bacelar, 1948 5 D
Lepidochrysops hawkeri (Talbot, 1929) Talbot, 1929 (*) 1,5 C?
Lepidochrysops nacrescens (Tite, 1961) Tite, 1961 (*) 1,6 C
Lepidochrysops reichenowi (Dewitz, Dewitz, 1879 (*) 1,6 ?
1879)
Thermoniphas distincta (Talbot, 1935) ◘ 5 C
Thermoniphas p. plurilimbata (Karsch, ◘ 5 A
1895)
Thermoniphas t. togara (Plötz, 1880) ▲ 6 A
Oboronia guessfeldtii (Dewitz, 1879) Larsen, 1991 5 B
Oboronia pseudopunctatus (Strand, 1912) ▲ 6 A
Oboronia punctatus (Dewitz, 1879) 5 A
Actizera lucida (Trimen, 1883) Willis, 2009 6 C
Brephidium metophis (Wallengren, 1860) Willis, 2009 6 ?
Azanus isis (Drury, 1773) Bacelar, 1948 5 A
Azanus jesous (Guérin-Méneville, 1849) Trimen, 1891 6 D
Azanus mirza (Plötz, 1880) Stempffer, 1957 5 D,A
Azanus moriqua (Wallengren, 1857) Weymer, 1901 5 D
Azanus natalensis (Trimen, 1887) Bacelar, 1948 5 D
Azanus ubaldus (Stoll, 1782) Bacelar, 1948 5 F
Chilades trochylus (Freyer, 1844) Ladeiro, 1956 5 D
Zizeeria k. knysna (Trimen, 1862) Ladeiro, 1956 5 D
Zizina otis antanossa (Mabille, 1877) ▲ 5 D
Zizula hylax (Fabricius, 1775) Gardiner, 2004 5 H
Riodinidae
Afriodinia dewitzi (Aurivillius, 1899) ▲ 6 A
Afriodinia intermedia (Aurivillius, 1895) Larsen, 2005 6 A
Afriodinia r. rogersi (Druce, 1878) Druce, 1878 (*) 5 A
Afriodinia tantalus caerulea (Riley, 1932) Druce, 1875 as tantalus 6 A
Nymphalidae | Libytheinae
Libythea labdaca (Westwood, 1851) Snellen, 1882 5 A
Libythea laius (Trimen, 1879) Gardiner, 2004 5 A
(continued)

Nymphalidae | Danainae
Danaus c. orientis (Aurivillius, 1909) Butler, 1871 5 H
Tirumala petiverana (Doubleday, 1847) Butler, 1866, as Danais leonora 5 D
Amauris n. niavius (Linnaeus, 1758) Aurivillius, 1928 5 A,B,D
Amauris t. tartarea (Mabille, 1876) Mabille, 1876 (*) 5 A
Amauris crawshayi angola (Bethune- Bethune-Baker, 1914 (*) 1,6 A
Baker, 1914)
Amauris h. hecate (Butler, 1866) ◘ 5 A
Amauris d. dannfelti (Aurivillius, 1891) Aurivillius, 1891 (*) 1,5 A,C
Amauris h. hyalites (Butler, 1874) Butler, 1874 (*) 5 A
Amauris vashti (Butler, 1869) ◘ 5 A
Nymphalidae | Satyrinae
Gnophodes betsimena parmeno Aurivillius, 1928 5 A
(Doubleday, 1849)
Gnophodes chelys (Fabicius, 1793) Aurivillius, 1928 5 A
Melanitis leda (Linnaeus, 1758) Bacelar, 1948 5 B,H
Elymnias b. bammakoo (Westwood, 1851) Druce, 1875 5 A
Bicyclus iccius (Hewitson, 1865) Larsen, 2005 6 A
Bicyclus sebetus (Hewitson, 1877) Aurivillius, 1928 5 A
Bicyclus s. saussurei (Dewitz, 1879) Dewitz, 1879 (*) 3,5 A
Bicyclus s. suffusa (Riley, 1921) ▲ 3,5 C
Bicyclus taenias (Hewitson, 1877) ◘ 5 A,B
Bicyclus nachtetis (Condamin, 1965) ▲ 3 A
Bicyclus technatis (Hewitson, 1877) Larsen, 2005 6 A
Bicyclus vulgaris (Butler, 1868) Druce, 1875 5 A,D
Bicyclus moyses (Condamin & Fox, 1964) Condamin & Fox, 1964 5 A
Bicyclus sandace (Hewitson, 1877) Bacelar, 1958a, b 5 A,D
Bicyclus auricruda fulgida (Fox, 1963) Aurivillius, 1928, as auricruda 5 A
Bicyclus collinsi (Aduse-Poku et al., Hewitson, 1873 5 B,D
2009)
Bicyclus angulosa selousi (Trimen, 1895) Condamin, 1963 5 C,D
Bicyclus campus (Karsch, 1893) ▲ 6 A
Bicyclus a. anynana (Butler, 1879) Gardiner, 2004 6 D
Bicyclus anynana centralis (Condamin, Condamin, 1968 5 A
1968)
Bicyclus cottrelli (van Son, 1952) ▲ 5 C
Bicyclus s. safitza (Westwood, 1850) Butler, 1871, as Mycalesis caffra 5 D
Bicyclus funebris (Guérin-Méneville, Ladeiro, 1956 5 A,B,D
1844)
Bicyclus istaris (Plötz, 1880) ▲ 5 A
Bicyclus lamani (Aurivillius, 1900) Bacelar, 1958a, b 5 A
Bicyclus golo (Aurivillius, 1893) Monard, 1956 5 A
Bicyclus s. smithi (Aurivillius, 1899) ▲ 5 A
Bicyclus vansoni (Condamin, 1965) Condamin, 1965 5 C
(continued)

Bicyclus buea (Strand, 1912) Larsen, 2005 5 A
Bicyclus sanaos (Hewitson, 1866) Druce, 1875 5 A
Hallelesis asochis congoensis (Joicey & Druce, 1875, as asochis 6 E
Talbot, 1921)
Brakefieldia angolensis (Kielland, 1994) Kielland, 1994 (*) 1,6 C
Brakefieldia p. phaea (Karsch, 1894) Kielland, 1994 5 C
Brakefieldia simonsii (Butler, 1877) Gardiner, 2004 5 D,F
Brakefieldia centralis (Aurivillius, 1903) Ackery et al., 1995 3 C
Brakefieldia ochracea (Lathy, 1906) Lathy, 1906 (*) 1,5 C
Brakefieldia eliasis (Hewitson, 1866) Druce, 1875 3,5 A,B
Mashuna upemba (Overlaet, 1955) ◘ 5 E
Ypthima a. asterope (Klug, 1832) Druce, 1875 5 D
Ypthima asterope hereroica (van Son, Gardiner, 2004 6 D
1955)
Ypthima c. condamini (Kielland, 1982) Larsen, 2005 6 C,D
Ypthima granulosa (Butler, 1883) Ladeiro, 1956 6 C?,D
Ypthima recta (Overlaet, 1955) Kielland, 1982 6 A
Ypthima doleta (Kyrby, 1880) Aurivillius, 1928 5 A,B
Ypthima i. impura (Elwes & Edwards, Aurivillius, 1928 5 A,D
1893)
Ypthima impura paupera (Ungemach, Gardiner, 2004 6 A,D
1932)
Ypthima praestans (Overlaet, 1954) ▲ 5 A
Ypthima pulchra (Overlaet, 1954) ▲ 5 A
Ypthima diplommata (Overlaet, 1954) Kielland, 1982 3,6 New
data
Ypthimomorpha itonia (Hewitson, 1865) Aurivillius, 1928 5 D,E
Neita bikuarica (M & BS, 2006) M & BS, 2006 (*) 1,5 C
Neocoenyra cooksoni (Druce, 1907) ▲ 6 C
Mashunoides carneiromendesi (M & BS, M & BS, 2009 (*) 1,5 D
2009)
Nymphalidae | Charaxinae
Charaxes fulvescens rubenarturi (BS & BS et al., 2017 (*) 1,5 A
M, 2017)
Charaxes varanes vologeses (Mabille, Mabille, 1876, sub Palla (*) 5 B,D
1876)
Charaxes candiope (Godart, 1824) Druce, 1875 5 B,D
Charaxes cynthia kinduana (Le Cerf, Aurivillius, 1928 5 A,D
1923)
Charaxes macclounii (Butler, 1895) van Someren, 1970 6 A
Charaxes macclouni carvalhoi (BS, 1983) BS, 1983 (*) 1,5 A
Charaxes protoclea protonothodes (van Aurivillius, 1928, as protoclea 5 A,B
Someren, 1971)
Charaxes lucretius saldanhai (BS, 1983) BS, 1983 (*) 1,5 A
(continued)

Charaxes brutus angustus (Rothschild, Druce, 1875, as brutus 5 A,B,D
1900)
Charaxes brutus natalensis (Staudinger, van Someren, 1970 6 A,B,D
1885)
Charaxes c. castor (Cramer, 1775) Druce, 1875 5 A,B
Charaxes druceanus proximans (Joicey & Aurivillius, 1928 5 A,D
Talbot, 1922)
Charaxes eudoxus mechowi (Rothschild, Rothschild, 1900 (*) 5 A,D
1900)
Charaxes eudoxus mitchelli (Plantrou & ◘ 5 A,D
Howarth, 1977)
Charaxes s. saturnus (Butler, 1866) Druce, 1875, as C. pelias 5 A,D
brunnescens
Charaxes p. pollux (Cramer, 1775) Druce, 1875 5 A,C
Charaxes numenes aequatorialis (van Aurivillius, 1928, as numenes 5 A
Someren, 1972)
Charaxes tiridates tiridatinus (Röber, Druce, 1875, the species 5 A
1936)
Charaxes ameliae amelina (Joicey & ◘ 5 C
Talbot, 1925)
Charaxes b. bohemani (Felder & Felder, Druce, 1875 5 D
1859)
Charaxes p. pythodoris (Hewitson, 1873) Hewitson, 1873 (*) 5 B
Charaxes smaragdalis leopoldi van Someren, 1964 2 A
(Ghesquiére, 1933)
Charaxes zingha (Stoll, 1780) Bacelar, 1958a, b 5 A
Charaxes a. achaemenes (Felder & van Someren, 1970 5 C,D
Felder, 1867)
Charaxes e. etesipe (Godart, 1824) 5 A,E
Charaxes p. penricei (Rothschild, 1900) Rothschild, 1900 (*) 5 C,D
Charaxes penricei dealbata (van van Someren, 1966 (*) 2,5 A
Someren, 1966)
Charaxes jahlusa angolensis (M & BS, BS et al., 2017 (*) 1,5 A
2017)
Charaxes eupale latimargo (Joicey & Druce, 1875 as eupale 5 A
Talbot, 1921)
Charaxes minor karinae (Bouyer, 1999) Bouyer, 1999 (*) 1,5 A
Charaxes anticlea proadusta (van Aurivillius, 1928, as anticlea 5 A
Someren, 1971)
Charaxes h. hildebrandti (Dewitz, 1879) Dewitz, 1879 (*) 5 A
Charaxes hildebrandti katangensis BS & M, 2014 5 A
(Talbot, 1928)
Charaxes g. guderiana (Dewitz, 1879) Dewitz, 1879 (*) 5 A,D
Charaxes brainei (van Son, 1966) Henning, 1988 4 D
Charaxes catachrous (van Someren & ◘ 5 A
Jackson, 1952)
(continued)

Charaxes cedreatis (Hewitson, 1874) Aurivillius, 1928 5 A,C
Charaxes diversiforma (van Someren & van Someren, 1969 5 A
Jackson, 1957)
Charaxes etheocles silvestris (Turlin, Druce, 1875, as C. ephyra 5 A,B
2011)
Charaxes figueirai (BS & M, 2014) BS & M, 2014 (*) 1,5 C,D
Charaxes fulgurata (Aurivillius, 1899) Aurivillius, 1899 (*) 5 C
Charaxes howarthi (Minig, 1976) Henning, 1988 5 C
Charaxes phaeus (Hewitson, 1877) Gardiner, 2004 6 C,D
Charaxes variata (van Someren, 1969) ◘ 3,5 C
Charaxes p. paphianus (Ward, 1871) Aurivillius, 1928 5 A
Charaxes pleione congoensis (Plantrou, van Someren 1974, as pleione 5 A
1989)
Charaxes ehmckei (Homeyer & Dewitz, Homeyer & Dewitz, 1882 (*) 1,5 A,B
1882)
Charaxes zoolina (Westwood, 1850) Gardiner, 2004 6 A
Charaxes kahldeni (Homeyer & Dewitz, Homeyer & Dewitz, 1882 (*) 5 A
1882)
Charaxes n. nichetes (Grose-Smith, 1883) Aurivillius, 1928 5 A
Charaxes nichetes pantherinus Rousseau- ◘ 5 A
(Decelle, 1934)
Charaxes lycurgus (Fabricius, 1793) Plantrou, 1978 5 A
Charaxes zelica rougeoti (Plantrou, 1978) Plantrou, 1978 5 A
Charaxes doubledayi (Aurivillius, 1899) Bacelar, 1956 5 A
Palla decius (Cramer, 1777) Druce, 1875 5 A
Palla publius centralis (van Someren, ◘ 5 A
1975)
Palla ussheri hassoni (Turlin & Turlin & Vingerhoedt, 2013 (*) 1,5 A
Vingerhoedt, 2013)
Palla violinitens coniger (Butler, 1896) Aurivillius, 1928 5 A
Charaxes c. crossleyi (Ward, 1871) Aurivillius, 1928 6 A
Charaxes eurinome ansellica (Butler, Butler, 1870 (*) 5 B
1870)
Charaxes trajanus bambi BS & M, 2007 BS & M, 2007 (*) 1,5 A
Nymphalidae | Apaturinae
Apaturopsis c. cleocharis (Hewitson, Hewitson, 1873 (*) 6 A
1873)
Nymphalidae | Nymphalinae
Kallimoides rumia jadyae (Fox, 1968) Druce, 1875, as rumia 5 A
Vanessula milca buechneri (Dewitz, 1887) ▲ 6 A
Antanartia d. delius (Drury, 1782) ▲ 5 A
Vanessa cardui (Linnaeus, 1758) Snellen, 1882 5 C,D
Precis antilope (Feisthamel, 1850) Monard, 1956 5 C,D
Precis a. archesia (Cramer, 1779) Aurivillius, 1928 5 C,D
Precis c. ceryne (Boisduval, 1847) Druce, 1875 5 A,B,E
(continued)

Precis coelestina (Dewitz, 1879) Dewitz, 1879 (*) 5 A
Precis octavia sesamus (Trimen, 1883) Druce, 1875, as 5 C
Precis pelarga (Fabricius, 1775) Ladeiro, 1956 5 A,C
Precis actia (Distant, 1880) Aurivillius, 1928 5 A,C
Precis s. sinuata (Plötz, 1880) Bacelar, 1956 5 A,C
Precis rauana silvicola (Schultze, 1916) ◘ 5 A
Precis larseni manuscript name – (M M et al., 2018 (*) 1,5 A,C
et al., 2018)
Hypolimnas a. anthedon (Doubleday, Druce, 1875, sub Diadema 5 A,C
1845)
Hypolimnas misippus (Linnaeus, 1764) Druce, 1875 5 H
Hypolimnas d. dinarcha (Hewitson, 1865) Bacelar, 1958a, b 5 A
Hypolimnas m. monteironis (Druce, 1874) Druce, 1874 (*) 5 A
Hypolimnas s. salmacis (Drury, 1773) Druce, 1875 5 A
Salamis c. cacta (Fabricius, 1793) Ackery et al., 1995 5 A
Protogoniomorpha anacardii ansorgei Rothschild, 1904 (*) 5 A,C
(Rothschild, 1904)
Protogoniomorpha parhassus (Drury, Druce, 1875, as Diadema salamis 5 A,C
1782)
Protogoniomorpha t. temora (Felder & Aurivillius, 1928 6 A
Felder, 1867)
Junonia artaxia (Hewitson, 1864) Aurivillius, 1928 5 C,D
Junonia hierta crebrene (Trimen, 1870) Butler, 1871 5 A,C,D
Junonia n. natalica (Felder & Felder, Gardiner, 2004 6 D
1860)
Junonia natalica angolensis (Rothschild, Rothschild, 1918 (*) 3,5 A,C
1918)
Junonia o. oenone (Linnaeus, 1758) Butler, 1871 5 H
Junonia orythia madagascariensis Bacelar, 1948 5 A,C,D
(Guenée, 1865)
Junonia sophia infracta (Butler, 1888) Bacelar, 1948 5 A
Junonia stygia (Aurivillius, 1894) Aurivillius, 1928 5 A
Junonia w. westermanni (Westwood, Hewitson, 1873 5 A
1870)
Junonia terea elgiva (Hewitson, 1864) Aurivillius, 1928 5 A
Junonia ansorgei (Rothschild, 1899) ◘ 5 A
Junonia cymodoce lugens (Schultze, Aurivillius, 1909, as cymodoce 5 A
1912)
Catacroptera c. cloanthe (Stoll, 1781) Butler, 1871 5 A,C,D
Nymphalidae | Cyrestinae
Cyrestis c. camillus (Fabricius, 1781) Ladeiro, 1956 5 A
Nymphalidae | Biblidinae
Biblia anvatara crameri (Aurivillius, Bacelar, 1948 5 C,D
1894)
Byblia ilithyia (Drury, 1773) Snellen, 1882 5 A,D
(continued)

Mesoxantha ethosea ethoseoides (Rebel, Aurivillius, 1928 5 A
1914)
Ariadne albifascia (Joicey & Talbot, ▲ 5 A
1921)
Ariadne enotrea archeri (Carcasson, Carcasson, 1958 (*) 5 A
1958)
Neptidiopsis ophione nucleata (Grünberg, Aurivillius, 1928 5 A
1911)
Eurytela dryope angulata (Aurivillius, Butler, 1871, as dryope 5 A,C,D
1899)
Eurytela h. hiarbas (Drury, 1782) Druce, 1875 5 A
Sevenia boisduvali omissa (Rothschild, ▲ 5 A
1918)
Sevenia occidentalium penricei Rothschild & Jordan, 1903 (*) 1,5 A
(Rothschild & Jordan, 1903)
Sevenia consors (Rothschild & Jordan, Rothschild & Jordan, 1903 (*) 5 C
1903)
Sevenia t. trimeni (Aurivillius, 1899) Aurivillius, 1899, as Crenis 5 A,C
natalensis var. trimeni (*)
Sevenia umbrina (Karsch, 1892) ▲ 5
Sevenia amulia intermedia (Carcasson, Aurivillius, 1928, as amulia 5 A,C,E
1961)
Sevenia benguelae (Chapman, 1872) Aurivillius, 1928 5 A,C,D
Sevenia p. pechueli (Dewitz, 1879) Dewitz, 1879 (*) 5 C
Nymphalidae | Limenitidinae
Harma theobene superna (Fox, 1968) Fox, 1968 5 A
Cymothoe o. oemilius (Doumet, 1859) Bacelar, 1958a, b 5 A
Cymothoe b. beckeri (Herrich-Schaeffer, Druce, 1875 5 A
1858)
Cymothoe haynae fumosa (Staudinger, Bacelar, 1956 2,5 A
1896)
Cymothoe confusa (Aurivillius, 1887) Bacelar, 1956 5 A
Cymothoe lucasii cloetensi (Seeldrayers, ◘ 5 A
1896)
Cymothoe h. harmilla (Hewitson, 1874) Larsen, 2005 6 A
Cymothoe h. hesiodotus (Hewitson, 1869) Aurivillius, 1928 6 A
Cymothoe h. hypatha (Hewitson, 1866) Druce, 1875 5 A
Cymothoe lurida hesione (Weymer, 1907) Druce, 1875, as lurida 5 A
Cymothoe altisidora (Hewitson, 1869) Aurivillius, 1898 5 A
Cymothoe capella (Ward, 1871) Bacelar, 1956 5 A
Cymothoe caenis (Drury, 1773) Druce, 1874 5 A
Cymothoe jodutta ciceronis (Ward, 1871) Bacelar, 1956 5 A
Cymothoe jodutta ehmckei (Dewitz, 1887) ◘ 5 A
Cymothoe cf. c. coccinata (Hewitson, ◘ 5 A
1874)
(continued)

Cymothoe excelsa deltoides (Overlaet, D’Abrera, 1980 3,5 A
1944)
Cymothoe s. sangaris (Godart, 1824) Druce, 1875 5 A
Pseudoneptis bugandensis ianthe Snellen, 1882, as bugandensis 5 A
(Hemming, 1964)
Pseudacraea eurytus eurytus (Linnaeus, Druce, 1875 5 A
1758)
Pseudacraea d. dolomena (Hewitson, Aurivillius,1928 6 A
1865)
Pseudacraea b. boisduvalii (Doubleday, Druce, 1875 5 A
1845)
Pseudacraea kuenowii gottbergi (Dewitz, Williams, 2007, as kuenowii 6 A
1884)
Pseudacraea lucretia protracta (Butler, Butler, 1874 (*) 5 A
1874)
Pseudacraea poggei (Dewitz, 1879) Gardiner, 2004 5 A,D
Pseudacraea semire (Cramer, 1779) Druce, 1875 5 A
Neptis saclava marpessa (Hopffer, 1855) Druce, 1875 5 A,B,C
Neptis nemetes margueriteae (Fox, 1968) Butler, 1871, the species 6 C,D
Neptis gratiosa (Overlaet, 1955) ▲ 5 C
Neptis jordani (Neave, 1910) Gardiner, 2004 6 D
Neptis kiriakoffi (Overlaet, 1955) ◘ 5 A,C
Neptis laeta (Overlaet, 1955) Gardiner, 2004 5 C
Neptis morosa (Overlaet, 1955) Larsen, 2005 5 A,C
Neptis s. serena (Overlaet, 1955) Gardiner, 2004 5 A,C,E
Neptis alta (Overlaet, 1955) Gardiner, 2004 5 C
Neptis constantiae kaumba (Condamin, ◘ 5 A
1966)
Neptis nysiades (Hewitson, 1868) Aurivillius, 1928 5 A
Neptis nicomedes (Hewitson, 1874) Hewitson, 1874 (*) 5 A
Neptis quintilla (Mabille, 1890) Larsen, 2005 5 A
Neptis a. agouale (Pierre-Baltus, 1978) ◘ 5 A
Neptis melicerta (Drury, 1773) Aurivillius, 1928 5 A,D
Neptis nebrodes (Hewitson, 1874) Hewitson, 1874 (*) 6 A
Neptis nicoteles (Hewitson, 1874) Hewitson, 1874 (*) 6 A
Neptis e. exaleuca (Karsch, 1894) ◘ 5 A
Evena crithea (Drury, 1773) Fox, 1968 5 A
Evena angustatum (Felder & Felder, ◘ 5 A
1867)
Euryphura c. chalcis (Felder & Felder, Bacelar, 1956, as E. fulminea 5 A
1860)
Euryphura plautilla Hewitson, 1865 Druce, 1875 5 A
Euryphura concordia (Hopffer, 1855) Aurivillius, 1928 5 C,E
Hamanumida daedalus (Fabricius, 1775) Druce, 1875, as Aterica meleagris 5 C,D,F
Pseudargynnis hegemone (Godart, 1819) Druce, 1874, as Aterica clorana 5 E
(continued)

Aterica g. extensa (Heron, 1909) Druce, 1875, as Aterica cupavia 5 A,B,C
Cynandra opis bernardii (Lagnel, 1967) Druce, 1875, as Aterica afer 5 A
Euriphene barombina (Aurivillius, 1894) Larsen, 2005 5 A
Euriphene iris (Aurivillius, 1903) ▲ 5 C
Euriphene plagiata (Aurivillius, 1897) Larsen, 2005 6 A
Euriphene saphirina trioculata (Talbot, ▲ 6 A
1927)
Euriphene t. tadema (Hewitson, 1866) ◘ 5 A
Euriphene gambiae gabonica (Bernardi, Bacelar, 1958a, b 6 A
1966)
Bebearia phantasia concolor (Hecq, Druce, 1875 6 A
1988)
Bebearia languida (Schultze, 1920) ▲ 5 A
Bebearia a. absolon (Fabricius, 1793) Bacelar, 1958a, b 5 A
Bebearia micans (Aurivillius, 1899) ◘ 5 A
Bebearia zonara (Butler, 1871) ◘ 5 A
Bebearia oxione squalida (Talbot, 1928) Aurivillius, 1909, as oxione A
Bebearia cocalia katera (van Someren, ◘ 5 A
1939)
Bebearia guineensis (Felder & Felder, Holmes, 2001 6 A
1867)
Bebearia sophus aruunda (Overlaet, Druce, 1875, as sophus 5 A
1955)
Bebearia plistonax (Hewitson, 1874) Hewitson, 1874 (*) 5 F
Bebearia hassoni (Hecq, 1998) Hecq, 1998 (*) 1 A
Euphaedra medon celestis (Hecq, 1986) Butler, 1871, as medon 5 A
Euphaedra z. zaddachii (Dewitz, 1879) Dewitz, 1879 (*) 5 A
Euphaedra cf. sinuosa (Hecq, 1974) ◘ 6 A
Euphaedra diffusa diffusa (Gaede, 1916) ◘ 6 A
Euphaedra ansorgei (Rothschild, 1918) ◘ 5 A
Euphaedra p. permixtum (Butler, 1873) Bacelar, 1956 5 A
Euphaedra divoides (BS & M, 2018) Staudinger, 1886, as E. themis var. 1,5 A
(Manuscript name) innocentia (?)
Euphaedra adonina spectacularis (Hecq, Bacelar, 1956 5 A
1997)
Euphaedra ceres electra (Hecq, 1983) Butler, 1871, as ceres 5 A
Euphaedra fontainei (Hecq, 1977) ▲ 5 A
Euphaedra v. viridicaerulea (Bartel, 1905) ◘ 5 A
Euphaedra preussiana robusta (Hecq, ◘ 5 A
1983)
Euphaedra rezia (Hewitson, 1866) Bacelar, 1956 5 A
Euphaedra albofasciata (Berger, 1981) ◘ 5 A
Euphaedra disjuncta virens (Hecq, 1984) ◘ 5 A
Euphaedra mayumbensis (Hecq, 1984) ◘ 5 A
(continued)

Euphaedra p. preussi (Staudinger, 1891) Aurivillius, 1928 6 A
Euphaedra uigensis (BS & M, 2017) BS & M, 2017 (*) 1,5 A
Euphaedra c. castanoides (Hecq, 1985) ◘ 5 A,F
Euphaedra coprates (Druce, 1875) Druce, 1875 (*) 5 A
Euphaedra e. eleus (Drury, 1782) Aurivillius, 1928 5 A
Euphaedra simplex (Hecq, 1978) ◘ 5 A
Euphaedra ruspina (Hewitson, 1865) Druce, 1875 5 A,C
Euphaedra harpalyce comminuera (Hecq, Hecq, 1999 (*) 1.5 A
1999)
Euphaedra harpalyce spatiosa (Mabille, Mabille, 1876 (*) 6 A
1876)
Euphaedra losinga wardi (Druce, 1874) Druce, 1874 5 A
Euphaedra losinga limita (Hecq, 1978) ◘ 3,5 A
Euptera mocquerysi (Staudinger, 1893) Bacelar, 1956, as E. falsathyma 5 A
Nymphalidae | Heliconiinae
Acraea r. rogersi (Hewitson, 1873) Hewitson, 1873 (*) 5 B,D
Acraea e. egina (Cramer, 1775) Druce, 1875 5 A,C
Acraea acrita ambigua (Trimen, 1891) Weymer, 1901, as A. bella 6 C,D
Acraea bellona (Weymer, 1908) Weymer, 1908 (*) 1,5 C
Acraea periphanes (Oberthür, 1893) Le Doux, 1923 5 C,E
Acraea asema (Hewitson, 1877) Trimen, 1891 6 C
Acraea o. omrora (Trimen, 1894) Trimen, 1894 (*) 1,6 C
Acraea violarum anchietai (M & BS, M & BS, 2017 (*) 1,5 C,D
2017)
Acraea buettneri (Rogenhofer, 1890) Trimen, 1891, as A. felina 5 A,C
Acraea cepheus (Linnaeus, 1758) Eltringham, 1912 5 A,C
Acraea atolmis (Westwood, 1881) Aurivillius, 1928, morph acontias 5 C
Acraea bailundensis (Wichgraf, 1918) Wichgraf, 1918 (*) 3,5 C
Acraea diogenes (Suffert, 1904) ▲ 3,5 D
Acraea guillemei (Oberthür, 1893) Aurivillius, 1928 6 C
Acraea lapidorum (Pierre, 1988) Pierre, 1988 (*) 1,5 C
Acraea onerata (Trimen, 1891) Trimen, 1891 (*) 1,4 C,D
Acraea aglaonice (Westwood, 1881) Gardiner, 2004 6 D
Acraea atergatis (Westwood, 1881) Aurivillius, 1928 5 D
Acraea axina (Westwood, 1881) Aurivillius, 1928 5 C
Acraea c. caldarena (Hewitson, 1877) Gardiner, 2004 5 C
Acraea ella (Eltringham, 1911) Eltringham, 1911 (*) 4,5 C
Acraea lygus (Druce, 1875) Druce, 1875 (*) 5 C, D
Acraea natalica (Boisduval, 1847) Aurivillius, 1928 5 A,C
Acraea oncaea (Hopffer, 1855) ◘ 6 C
Acraea pseudegina (Westwood, 1852) Druce, 1875 5 A
Acraea pudorella (Aurivillius, 1899) ▲ 5 C
Acraea stenobea (Wallengren, 1860) Trimen, 1891 6 C,D
Acraea anemosa (Hewitson, 1865) Aurivillius, 1928 5 C,D
(continued)

Acraea p. pseudolycia (Butler, 1874) Butler, 1874 (*) 3,5 A,C
Acraea acara melanophanes (Le Cerf, Pierre & Bernaud, 2013 5 D
1927)
Acraea z. zetes (Linnaeus, 1758) Druce, 1875 5 C,D
Acraea admatha (Hewitson, 1865) Pierre, 1979 6 A
Acraea endoscota (Le Doux, 1928) Larsen, 2005 6 A
Acrae l. leucographa (Ribbe, 1889) Larsen, 2005 6 A
Acraea q. quirina (Fabricius, 1781) ▲ 5 A
Acraea camaena (Drury, 1773) Larsen, 2005 6 B
Acraea n. neobule (Doubleday, 1843) Druce, 1875 5 A,C,D
Acraea eugenia ochreata (Grünberg, Larsen, 2005 5 A
1910)
Acraea brainei (Henning, 1986) Ackery et al., 1995 6 ?
Acraea e. epaea (Cramer, 1779) Aurivillius, 1928 5 A
Acraea formosa (Butler, 1874) Butler, 1874 (*) 5 A,B
Acraea l. leopoldina (Aurivillius, 1895) ▲ 3 A
Acraea p. poggei (Dewitz, 1879) Dewitz, 1879 (*) 6 C
Acraea alcinoe camerunica (Aurivillius, ▲ 6 A
1893)
Acraea umbra macarioides (Aurivillius, Aurivillius, 1928 5 A,B
1893)
Acraea consanguinea intermedia Le Doux, 1937 2,5 A
(Aurivillius, 1899)
Acraea excisa (Butler, 1874) Bacelar, 1948 5 A
Acraea pseuderyta (Godman & Salvin, Aurivillius, 1928 5 A
1890)
Acraea vestalis congoensis (Le Doux, ◘ 5 A
1937)
Telchinia p. perenna (Doubleday, 1847) Aurivillius, 1928 5 A
Telchinia p. penelope (Staudinger, 1896) ◘ 5 A
Telchinia o. oreas (Sharpe, 1891) Lathy, 1906 6 C
Telchinia circeis (Drury, 1782) Larsen, 2005 6 A
Telchinia parrhasia servona (Godart, Godart, 1819 (*) 6 A
1819)
Telchinia peneleos pelasgia (Grose- Larsen, 2005 5 A,C
Smith, 1900)
Telchinia p. pharsalus (Ward, 1871) Aurivillius, 1928 5 A,B
Telchinia encedana (Pierre, 1976) Pierre, 1976 6 E
Telchinia e. encedon (Linnaeus, 1758) Druce, 1875 5 A,C,D
Telchinia alciope (Hewitson, 1852) Bacelar, 1956 5 A
Telchinia a. aurivillii (Staudinger, 1896) ▲ 6 A
Telchinia esebria (Hewitson, 1861) Butler, 1874 5 B,C
Telchinia j. jodutta (Fabricius, 1793) Druce, 1875 5 A
Telchinia lycoa (Godart, 1819) Druce, 1875 5 A,C
Telchinia serena (Fabricius, 1775) Butler, 1871 5 A,C,D
(continued)

Telchinia v. ventura (Hewitson, 1877) Monard, 1956 6 E
Telchinia acerata (Hewitson, 1874) Snellen, 1882 5 A,C
Telchinia oberthueri (Butler, 1895) Bacelar, 1948 6 A
Telchinia sotikensis karschi (Aurivillius, Aurivillius, 1928 5 A
1899)
Telchinia b. bonasia (Fabricius, 1775) Druce, 1875 5 A,C,D
Telchinia uvui balina (Karsch, 1892) Larsen, 2005 6 A
Telchinia o. orestia (Hewitson, 1874) Snellen, 1882 5 A
Telchinia p. pentapolis (Ward, 1871) ▲ 6 A
Telchinia induna imduna (Trimen, 1895) 5 C
Telchinia r. rahira (Boisduval, 1833) Druce, 1875 5 C,D,E
Telchnia mirifica (Lathy, 1906) Lathy, 1906 (*) 3 C
Lachnoptera anticlia (Hübner, 1819) Bacelar, 1958a, b, as L. iole 6 A
Phalanta e. eurytis (Doubleday, 1847) Bacelar, 1956 5 A
Phalanta phalantha aethiopica Druce, 1875 5 C,D
(Rothschild & Jordan, 1903)
References
Ackery PR, Smith CR, Vane Wright RI (1995) Carcasson’s African Butterflies. An annotated cata-
logue of the Papilionoidea and Hesperioidea of the Afrotropical Region. CSIRO, Victoria, i–xii
+ 803 pp
Aduse-Poku K, Vingerhoedt E, Wahlberg N (2009) Out-of-Africa again: A phylogenetic hypoth-
esis of the genus Charaxes (Lepidoptera: Nymphalidae) based on five genes region. Mol
Phylogenet Evol 53:463–478
Aurivillius C In Seitz A (1928) Les Macrolepidopteres du Globe. Les Macrolepidoptères de la
Faune Ethiopienne 13(4):615 pp + 80 pl. E. Le Moult, Paris
Bacelar A (1948) Lepidópteros de África principalmente das colónias portuguesas (colecção do
Museu Bocage). Arquivos do Museu Bocage 19:165–207
Bacelar A (1956) Lepidópteros (Rhopalocera) de Buco Zau, enclave de Cabinda, Angola. Anais da
Junta de Investigações do Ultramar 11(3):175–197
Bacelar A (1958a) Alguns Lepidópteros (Rhopalocera) do enclave de Cabinda. Revista portuguesa
de Zoologia e Biologia Geral 1(2/3):197–217
Bacelar A (1958b) Alguns Lepidópteros (Rhopalocera) da África Ocidental portuguesa. Revista
portuguesa de Zoologia e Biologia Geral 1(4):311–330
Bacelar A (1961) Lepidópteros do Bié (Rhopalocera) da colecção do Colégio de São Bento, em
Luso (Angola). Memórias da Junta de Investigações do Ultramar (2)23:61–81
Berger LA (1979) Espèces peu connues et descriptions de nouvelles sous-espèces de Mylothris
(Lepidoptera Pieridae). Revue de Zoologie Africaine 93(1):1–9
Bethune-Baker GT (1914) Notes on the taxonomic value of genital armature in Lepidoptera. Trans
Entomol Soc London 1914:314–337
Bivar-de-Sousa A (1983) Contribuição para o conhecimento dos lepidópteros de Angola (3ª nota).
Dados sobre a ocorrência do género Charaxes (Lep. Nymphalidae) em Angola (1ª parte). Actas
do I Congresso Ibérico de Entomologia 1:107–119
Bivar-de-Sousa A, Fernandes JA (1964) Contribuição para o conhecimento dos Lepidópteros de
Angola. Boletim da Sociedade Portuguesa de Ciências Naturais (2) 10(25):104–115
Bivar-de-Sousa A, Mendes LF (2006) On the genus Euxanthe Hübner, 1819 in Angola, with
description of a new subspecies (Lepidoptera, Nymphalidae, Charaxinae). Nouvelle Revue
d’Entomologie 23(4):369–376
Bivar-de-Sousa A, Mendes LF (2007) New data on the Uranothauma from Angola, with description
of a new species (Lepidoptera: Lycaenidae: Polyommatinae). Boletin Sociedad Entomológica
Aragonesa 41:73–76
Bivar-de-Sousa A, Mendes LF (2009a) On a new species of the genus Princeps Hübner, (1807)
from Cabinda (Angola) (Lepidoptera: Papilionidae). SHILAP, Revista de Lepidopterologia
37(147):313–318
Bivar-de-Sousa A, Mendes LF (2009b) New data on the Angolan Charaxes of the “etheocles
group” with description of a new species (Lepidoptera, Nymphalidae, Charaxinae). Boletim da
Sociedade portuguesa de Entomologia 8(15)(229):293–309
Bivar-de-Sousa A, Mendes LF (2014) New data on the Angolan Charaxesof the “etheocles group”
with description of a new species (Lepidoptera, Nymphalidae, Charaxinae). Boletim da
Sociedade portuguesa de Entomologia 229(8–15):293–309
Bivar-de-Sousa A, Mendes LF, Vasconcelos S (2017) Description of one new species and one new
subspecies of Nymphalidae from Angola (Lepidoptera: Papilionoidea). SHILAP, Revista de
Lepidopterologia 45(178):227–236
Bivar-de-Sousa A, Mendes LF (2018, in press) The “themis group” of Euphaedra (Euphaedrana)
in Angola. Revision and description of one new species (Lepidoptera: Nymphalidae:
Limenetidinae). Boletim da Sociedade portuguesa de Entomologia
Bouyer T (1999) Note sur les Charaxes du «groupe eupale» avec description d’une nouvelle sous-
espèce (Lepidoptera Nymphalidae). Entomologia Africana 4(1):37–40
Butler AG (1872) On a small collection of butterflies from Angola. Proc Zool Soc London
1871:721–725
Carvalho EL (1962) Alguns Papilionídeos da Lunda (Lepidoptera). Publicações Culturais da
Companhia dos Diamantes de Angola 60:163–170
Condamin M (1966) Mise au point sur les Neptis au facies d’ «14avann» (Lepidoptera:
Nymphalidae). Bulletin de l’Institut Fondamental de l’Afrique Noire 28(A)(3):1008–10029
D’Abrera B (1980) Butterflies of the afrotropical region. Lansdowe Ed., Melbourne, pp i–xx +
1–593
Dhungel B, Wahlberg N (2018) Molecular systematics of the subfamily Limenitidinae (Lepidoptera:
Nymphalidae). PeerJ 6:e4311
Druce H (1875) A list of the collections of diurnal Lepidoptera made by J. J. Monteiro in Angola
with description of some new species. Proc Zool Soc London 27:406–417
Eltringham H (1912) A monograph of the African species of the genus Acraea Fab., with a supple-
ment of those of the Oriental Region. Trans Entomol Soc London 1912(1):1–374
Espeland M, Breinholt J, Willmott KR et al (2018) A comprehensive and dated phylogenomic
analysis of butterflies. Curr Biol 28:1–9
Evans WH (1937) Catalog of African Hesperiidae (Indicating the classification and nomenclature
adopted in the British Museum). British Museum, London, 212pp + 30 pl
Gardiner A (2004) Chapter 10. Butterflies of the four corners area. In: Timberlake JR, Childers SL
(eds) Biodiversity of the Four Corners Area: Technical Review, vol 15. Occasional Publications
on Biodiversity, Bulawayo, pp 381–397
Hecq J (1997) Euphaedra. Lambillionea & Hecq, Tervuren & Mont-Sur-Marchienne, 121 pp +
48 pl
Heikkilä M, Kaila L, Mutanen M et al (2012) Cretaceous origin and repeated Tertiary diversifica-
tion of the redefined butterflies. Proc R Soc Lond B 279:1093–1099
Henning SF (1988) The Charaxes Butterflies of Africa. Aloe Books, Johannesburg, 457 pp
Holmes CWN (2001) A reaprisal of the Bebearia mardania complex (Lepidoptera, Nymphalidae).
Trop Zool 14:31–62
Homeyer A, Dewitz H (1882) Drei neue westafrikanische Charaxes. Berliner Entomologisches
Zeitschrift 26(II):381–383
JIU (1948–1963) Cartas do levantamento aerofotogramétrico de Angola (escala 1: 250,000). Junta
de Investigações do Ultramar: folhas:8–471
Kielland J (1982) Revision of the genus Ypthima in the Ethiopian Region excluding Madagascar
(Lepidoptera, Satyridae). Tijdschrift voor Entomologie 125(5):99–154
Kielland J (1994) A revision of the genus Henotesia (excluding Madagascar and other Indian
Ocean islands) (Lepidoptera Satyridae). Lambillionea 94(2):235–273
Ladeiro JM (1956) Lepidópteros de Angola (estudo de uma colecção oferecida ao Museu Zoológico
de Coimbra). Anais da Junta de Investigações do Ultramar 11(3):151–172
Larsen TB (2005) Butterflies of West Africa. Text volume: 1–595. Apollo Books, Stenstrup
Latreille PA, Godart JB (1819) Encyclopédie Méthodologique. Histoire Naturelle (Zoologie), 9,
Entomologie, Paris, i–iv + 1–328
Lautenschläger T, Neinhuis C (eds) (2014) Riquezas Naturais do Uíge – Uma breve Introdução
sobre o estado atual. A utilização, a Ameaça e a Preservação da Biodiversidade. Technische
Universität Dresden, Dresden, 125 pp
Libert M (1999) Revision des Epitola (ls). Révision des genres Epitola Westwood, Hypophytala
Clench et Stempfferia Jackson et description de trois nouveaux genres (Lepidoptera
Lycaenidae). ABRI & Lambillionea, Nairobi/Tervuren, pp 1–219, pl. I–XVI
Libert M (2000) Révision du genre Mimacraea Butler, avec description de quatre nouvelles espèces
et deux nouvelles sous-espèces (Lepidoptera, Lycaenidae). Lambillionea & ABRI, Tervuren/
Nairobi, pp 1–73
Libert M (2004) Revision du genre Oxylides Hübner (Lepidoptera, Lycaenidae). Lambillionea
104(2):143–158
Mendes LF, Bivar-de-Sousa A (2006a) A new species of Neita van Son (Nymphalidae, Satyrinae)
from southern Angola. Boletin Sociedad Entomológica Aragonesa 39:95–96
Mendes LF, Bivar-de-Sousa A (2006b) Notes and descriptions of Afrotropical Appias butterflies
(Lepidoptera: Pieridae). Boletin Sociedad Entomológica Aragonesa 39:151–160
Mendes LF, Bivar-de-Sousa A (2007a) New species of Cooksonia Druce, 1905 from Angola
(Lepidoptera: Lycaenidae, Lipteninae). SHILAP, Revista de Lepidopterologia 35(138):265–268
Mendes LF, Bivar-de-Sousa A (2007b) On the genus Eagris Guenée, 1863 in Angola (Lepidoptera:
Hesperiidae). SHILAP, Revista de Lepidopterologia 35(139):311–316
Mendes LF, Bivar-de-Sousa A (2009a) Description of a new species of Mylothris from northern
Angola (Lepidoptera Pieridae). Bolletino della Societá Entomológica Italiana 141(1):55–58
Mendes LF, Bivar-de-Sousa A (2009b) New account on the butterflies of Angola. The genus
Leptomyrina (Lepidoptera Lycaenidae). Bolletino della Societá Entomológica Italiana
141(2):109–112
Mendes LF, Bivar-de-Sousa A (2009c) On a new south-eastern Angolan Satyrine butterfly belong-
ing to a new genus (Lepidoptera, Nymphalidae). Entomologia Africana 14(2):5–8
Mendes LF, Bivar-de-Sousa A (2009d) The genus Abantis Hopffer, 1855 in Angola and description
of a new species (Lepidoptera: Hesperiidae, Pyrginae). SHILAP, Revista de Lepidopterologia
37(147):313–318
Mendes LF, Bivar-de-Sousa A (2012) Notes on the species of Hypolycaena (Lepidoptera,
Lycaenidae, Theclinae) known to occur in Angola. Boletin Sociedad Entomológica Aragonesa
50:193–197
Mendes LF, Bivar-de-Sousa A, Figueira R (2013a) Butterflies of Angola / Borboletas diurnas de
Angola. Lepidoptera. Papilionoidea, I. Hesperiidae, Papilionidae. IICT and CIBIO, Lisboa and
Porto, 288 pp
Mendes LF, Bivar-de-Sousa A, Figueira R et al (2013b) Gazetteer of the Angolan localities known
for beetles (Coleoptera) and butterflies (Lepidoptera: Papilionoidea). Boletim da Sociedade
Portuguesa de Entomologia 8(14/228):257–290
Mendes LF, Bivar-de-Sousa A, Vasconcelos S et al (2017) Description of two new subspecies
and notes on Charaxes Ochenheimer, 1816 of Angola (Lepidoptera: Nymphalidae). SHILAP,
Revista de Lepidopterologia 45(178):299–315
Mendes LF, Bivar-de-Sousa A, Vasconcelos S (2018, in press) On the butterflies of genus Precis
Hübner, 1819 from Angola and description of a new species (Lepidoptera: Nymphalidae:
Nymphalinae). SHILAP Revista de Lepidopterologia
Monard A (1956) Compendium Entomologicum Angolae – 1. Insecta. VI – Ord. Lepidoptera.
Anais da Junta de Investigações do Ultramar 11(3):119–128
NHM – Natural History Museum (2004). Wallowtails. Available at: http://internet.nhm.ac.uk/cgi-

bin/perth/wallowtails/list.dsml
Pierre J (1988) Les Acraea du super-group «egina»: Révision et phylogénie (Lepidoptera :
Nymphalidae). Annales de la Société Entomologique de France 24(3):263–287
Pierre J, Bernaud D (2013) Butterflies of the world. NymphalidaeXXIII. Acraea subgenus Acraea.
Goecke & Evers, Keltern, 39:1–8, pl. 1–28
Rothschild W, Jordan K (1903) Lepidoptera collected by Oscar Neumann in north-east Africa.
Novitates Zoologicae 10(3):491–542
Smith CR, Vane-Wright RI (2001) A review of the Afrotropical species of the genus Graphium
(Lepidoptera: Rhopalocera: Papilionidae). Bull Nat His Mus London (Entomol) 70(2):503–719
Stempffer H (1957) Les Lépidoptères de l’Afrique noire française. Lycaenidés. Initiations
Africaines 14(3):1–228
Stempffer H, Bennett NH (1963) A new genus of Lipteninae (Lepidoptera: Lycaenidae). Bull Brit
Mus (Nat His) (Entomol) 13:171–194
Talbot G (1944) A preliminary revision of the genus Mylothris Hübn. (1819) (Lep. Rhop. Pieridae).
T Roy Ent Soc London 94(2):155–185
Tite GE (1959) New species and notes on the genus Lepidochrysops (Lepidoptera, Lycaenidae).
The Entomologist 92:158–163
Tite GE (1961) New species of the genus Lepidochrysops (Lepidoptera, Lycaenidae). The
Entomologist 94:21–25
Tite GE, Dickson CGC (1973) The genus Aloeides and allied genera. Bull Brit Mus (Nat His)
(Entomol) 29:227–280
Trimen R (1891) On butterflies collected in tropical south-western Africa by Mr. A. W. Eriksson.
Proc Zool Soc 1891:59–107
Turlin B, Vingerhoedt E (2013) Butterflies of the World, supl. 23. Les Charaxinae de la faune
Afrotropicale. Les genres Palla et Euxanthe. Nymphalidae: Charaxinae: Pallini et Euxanthini.
Goecke & Evers, Keltern
Weymer G (1901) Beitrag zur Lepidopterofauna von Angola. Entomologischen Zeitschriften,
Stuttgart 15(17):61–64, 65–67, 69–70
Weymer G (1908) Einige neuer Lepidopteren des Deutschen Entom. National-Museums, gesam-
melt von Dr. F. Cr. Wellman in Benguella. Deutsche Entomologische Zeitschrift 1908:507–413
Willis CK (2009) Amist the butterflies of southwestern Angola. Metamorphosis 20(3):74–88
Williams MC (2018) Afrotropical butterflies. Available at: http://www.lepsocafrica.org/?p=publi
cations&s=atb
Part IV
Vertebrates: Distribution and Diversity
Chapter 11
The Freshwater Fishes of Angola
Paul H. Skelton
Abstract The discovery and exploration of Angolan freshwater fishes was largely
effected by foreign scientists on expeditions organised by European and North
American parties. Current knowledge of Angolan freshwater fishes is briefly
described according to the main drainage systems that include Cabinda, Lower
Congo, Angolan Coastal region including the Cuanza, the southern Congo tributar-
ies, the Zambezi, Okavango, Cunene and Cuvelai drainages. A biogeographic model
to explain the freshwater fish fauna of Angola is presented. The need for the conser-
vation of Angolan freshwater fishes will rise with rapidly increasing pressures on
aquatic ecosystems from urbanisation, dams for power, agriculture and human
needs, habitat destruction from mining and deforestation, pollution, the introduction
of alien species and overfishing.
Keywords Africa · Cuanza · Cunene · Cuvelai · Okavango · Southern Congo ·

Zambezi
Historical Review
Despite Poll’s work (1967) over a very limited area, Angola remains a poorly known region
in which there remains much to be discovered (Lévêque and Paugy 2017a: 93)
The quotation above sums up the current state of knowledge for the freshwater
fishes of Angola. Poll (1967) is a landmark publication that reviews the historical
literature and records the known species and their distribution within the major river
basins of the country at that time. No other account of Angolan fishes as a whole has
been published. The current situation of a poorly known region is due to a number
of factors including the historical neglect of scientific exploration by the colonial
P. H. Skelton (*)
South African Institute for Aquatic Biodiversity (SAIAB), Grahamstown, South Africa
Wild Bird Trust, National Geographic Okavango Wilderness Project, Hogsback, South Africa
e-mail: P.skelton@saiab.ac.za

https://doi.org/10.1007/978-3-030-03083-4_11
208 P. H. Skelton
authorities, widely dispersed collections in international institutions from various

expeditions, the relative inaccessibility to scientists and collectors of the inland riv-
ers and biologically rich areas, and the difficulties of aquatic exploration relative to
terrestrial fauna. The fact that there is no national Angolan depository for wet col-
lections such as fishes fostered by local expertise is a further hindrance to discovery.
This aspect is fundamental to effective and sustained scientific productivity in any
endeavor such as ichthyology (Skelton and Swartz 2011). This accentuates the situ-
ation for Angolan freshwater fishes when it is recognised that Poll’s (1967) account
rested largely on the collection in the Diamond Company of Angola (DIAMANG)
museum in Dundo, which to a large extent is a product of the industrial diamond
mining activity in the mainly local drainages.
There are four distinct phases of scientific discovery of Angolan freshwater
fishes, Phase 1 – early explorations in the second half of the nineteenth century;
Phase 2 – scientific expeditions in the twentieth century until World War II; Phase
3 – post WWII to Angolan independence in 1975; and, Phase 4 – post independence
investigations.
Although several of Castelnau’s (1861) Lake Ngami fishes occur in the Angolan
reaches of the Okavango River system, the discovery and scientific description of
Angolan freshwater fishes was initiated by Steindachner (1866) describing a collec-
tion derived from the Atlantic coastal rivers. Steindachner’s species include some
iconic species such as his Kneria angolensis, Clarias angolensis and Enteromius
kessleri that help define the Angolan Atlantic coastal fauna. Guimarães (1884)
working with specimens in the Lisbon Museum (subsequently lost in the fire of
1978) (Saldanha 1978) submitted by the Portuguese explorer José Alberto de
Oliveira Anchieta provided detailed descriptions and illustrations of three species
taken from the Cunene and the Curoca Rivers from 1873–1884, viz. Schilbe stein-
dachneri, Mormyrus anchietae and Enteromius mattozi.
The second phase of discovery (early twentieth century) is marked by a series of
expedition reports that include freshwater fishes. Boulenger’s (1909–1916) cata-
logue of fishes in the British Museum (Natural History) provided the basis of
Angolan freshwater fish fauna. Again the fauna included collections such as
Woosnam’s Okavango collection described by Boulenger (1911) that includes spe-
cies which occur in Angolan reaches. Boulenger (1910) described a collection by
Ansorge from the Cuanza and Bengo Rivers that set the scene for considering the
uniqueness of the fauna of these rivers of the Atlantic coast. Other notable expedi-
tions that included descriptions of freshwater fishes are the Vernay Angola
Expedition of 1925 (Nichols and Boulton 1927), the Gray African Expedition of
1929 (Fowler 1930), the Vernay-Lang Kalahari Expedition of 1930 (Fowler 1935),
the Swiss Scientific Mission to Angola 1928–1929 and 1932–1933 (Pellegrin 1936),
and Karl Jordan’s Expedition to South West Africa and Angola of 1933–1934
(Trewavas 1936). All these expeditions realised new species but were somewhat
limited in geographical scope to the southern Atlantic coastal rivers and to the
upland western reaches of the Cubango (Okavango) tributaries, the Oshana-Etosha
system and the plateau reaches of the Cuanza. This restriction emanates from the
11 The Freshwater Fishes of Angola 209
access realised by the Benguella Railway constructed from 1903 to 1928 from
Lobito port to Huambo and beyond (Ball 2015).
The third phase of scientific exploration of the freshwater fishes of Angola after
the WWII up until independence in 1975 is significant in that studies into ecological
aspects as well as the beginnings of a synthesis of the Angolan fauna occurred.
Ladiges and Voelker (1961) studied the fish fauna of the Longa River in the Angolan
watershed highlands. In addition to providing an ecological description and zona-
tion of the river they described a few new species – Kneria maydelli from the
Cunene, Enteromius (as Barbus) roussellei and Chiloglanis sardinhai. Ladiges
(1964) followed up this article with an account of the zoogeography and ecology of
Angolan freshwater fishes based on a present/absent list of fishes in the Angolan
Coastal region, the Cunene, the Okavango Basin, and the Zambezi. Trewavas (1973)
recorded the cichlids of the Cuanza and Bengo Rivers that exposed the independent
derivation of the cichlid fauna of the Cuanza River in terms of the inland and coastal
reaches. An unpublished collection by Graham Bell-Cross from the Okavango and
the Cunene basins was deposited in the NHM in 1965, and this together with collec-
tions made by Mike Penrith from the State Museum in Windhoek, Namibia, pro-
vided specimens essential for Greenwood’s (1984) revision of serranochromine
species. Mike Penrith’s collections from the Cunene and Okavango in the early
1970s led to a few descriptions of new species by Penrith (1970) and Penrith (1973).
A major milestone account of Angolan freshwater fishes was Max Poll’s (1967)
Contribution à la Faune Ichthyologique de l’Angola – based largely on the exten-
sive collections made by Barros Machado and others and lavishly illustrated with
excellent fish drawings as well as a gallery of photographs drawn from the Dundo
Museum in Lunda-Norte. Poll (1967) summarised the history of freshwater ichthy-
ology and provided a full checklist of 264 species in 18 families and 54 genera as
then recorded from the inland waters of the country (excluding the Cabinda enclave).
A faunistic and zoogeographical account considered five ichthyological regions
(see below). Acknowledging a clearly incomplete inventory Poll listed the diversity
of his regions as follows: The Congo tributaries with 121 species are richest and
most diverse with characteristic families and genera known from the Congo Basin.
Next in diversity was the western Atlantic coastal region with 109 species, followed
by the Zambezi (62 species) but Poll pointed out that Bell-Cross had then recently
recorded 77 species from the upper Zambezi, also of tropical diversity but distinct
in character from the Congolean rivers. The Okavango-Cubango (57 species)
reflected its close connections to the Zambezi as well as to the Cunene (55 species)
in the west. The Cunene presented a mixed fauna of both the Zambezian elements
as well as Atlantic coastal nature.
Poll (1967) pointed out and summarised a few notable ichthyological character-
istics of the Angolan fauna – there was no pronounced endemic character to the
fauna as a whole. The occurrence of lungfish (Protopterus) in Angola is known
only from records in Congo tributaries and from Cabinda, but Poll mentions that he
was shown a photograph by Ladiges of Protopterus annectens brieni from the
Cubango region (see this recorded in Ladiges 1964: 265). Such occurrence of lung-
fish in the Cubango or Okavango system has not yet been confirmed in spite of
210 P. H. Skelton
extensive collecting in that drainage. Polypterids are restricted to Congo tributaries

as are freshwater clupeids (however marine or estuarine species also occur in
coastal Atlantic rivers). The presence of kneriids is a distinct feature of the fauna
especially of the escarpment reaches of rivers of the coastal region. Mormyrid
diversity (36 species) is relatively high, most especially in the southern Congo riv-
ers. Characins (17 species) are less diverse but there is an equivalent representation
of Citharinids (16 species). The largest family represented in the country is the
Cyprinidae (79 species) and this is especially notable for the Atlantic coastal drain-
ages (43 species) that is even richer than the Congo tributaries (27 species).
However the chedrins (Raiamas and Opsaridium and Engraulicypris) are poorly
represented – two species in Congo tributaries, one in the Zambezian region and
one in the Atlantic coastal drainages. Of the catfishes, the claroteids (10 species)
are a presence as are the clariids (17 species) of which the majority (11 species) are
represented across different provinces. Other catfish families present include schil-
beids (eight species) mochokids (15 species), amphiliids (six species) and one
malapterurid. Cyprinodonts are relatively few (eight species) but show a particular
relationship across the Cassai-Zambezi watershed. Cichlids (31 species) are well
represented but not as well as the cyprinids. They are however more endemic in
nature, in particular the Atlantic coastal fauna (19 species with eight endemic). The
anabantids (three species) and mastacembelids (three species) are poorly
represented.
The last phase of ichthyological exploration informing on the fishes of Angola,
since independence in 1975, includes several taxonomic or systematic articles (e.g.
Greenwood 1984, Musilová et al. 2013); published river faunal accounts (Skelton
et al. 1985; Hay et al. 1997) and several unrestricted informal fish survey reports
emanating from specific projects (Bills et al. 2012, 2013; Skelton et al. 2016). These
surveys have exposed several new species to the fauna and together with phyloge-
netic studies on a wide range of lineages that include Angolan representatives, have
led to a vastly improved understanding of the distributional nuances that give expla-
nation to improved biogeographical insights.
Freshwater Drainages and Ecoregions of Angola
The drainages of Angola include southern source tributaries of the Congo, western
source tributaries of the Zambezi, coastal rivers to the Atlantic from the Chiloango
in Cabinda to the Cunene in the south, and the endorheic Etosha and Okavango
Basin drainages in the south (Fig. 11.1). The watershed between the Congo system
and the coastal Atlantic and Zambezian rivers is a major ichthyofaunal divide of
considerable biogeographic significance (Poll 1967, Jubb 1967, Skelton 1994,
Snoeks et al. 2011, Paugy et al. 2017).
The freshwater fishes of Angola fall within four major African ichthyological
provinces (Fig. 11.1) – Lower Guinea, Congolese, Angolan (coastal) or Cuanza and
Zambezian (Roberts 1975, Snoeks et al. 2011, Lévêque and Paugy 2017b).
Fig. 11.1 Left: Main drainage basins of Angola. A: Lower Congo, B: Cuanzan or Atlantic
Coastal, C: Cunene, D: Southwest Congo E: (west) Upper Zambezi, F: Okavango, G: Cuvelai.
Chiloango in Cabinda not illustrated. Center: Ichthyological provinces in Angola, modified from
Lévêque and Paugy (2017b) to include upper Cuanza and upper Cunene in the Zambezian Province.
A: Congolian, B: Angolan or Cuanzan, C: Zambezian. Cabinda and the Chiloango River in the
Lower Guinea Province not illustrated. Right: Aquatic ecoregions in Angola, modified, after
Thieme et al. (2005). A – Lower Congo, B – Cassai, C – Cuanza, D – Zambezian headwaters, E –
Okavango and Upper Zambezian floodplains, F – Namib coastal, G – Etosha. Southern West
Coastal Equatorial (Cabinda) not illustrated
Previously Poll (1967) considered the freshwater fishes of Angola to be from five
ichthyological regions drawn along watershed lines – Congo tributaries, Zambezi
region, Angolan (western) coastal drainages excluding the Cunene, the
Cubango-Okavango, and the Cunene. Thieme et al. (2005) defined ecoregions as “a
large area containing a distinct assemblage of natural communities and species,
whose boundaries approximate the original extent of natural communities before
major land use change”, and often reach across watershed lines. The Thieme et al.
(2005) aquatic ecoregion map for Africa included Angolan inland waters within
eight aquatic ecoregions as follows (Fig. 11.1): Floodplains, Swamps and Lakes –
Region 12 Okavango Floodplains, Region 16 Upper Zambezi Floodplains; Moist
Forest Rivers – Region 22 Lower Congo, Region 29 Southern West Coastal
Equatorial; Savannah Dry Forest – Region 63 Cuanza, Region 76 Upper Zambezian
headwaters; Xeric Systems – Region 82 Etosha, Region 88 Namib Coastal. Both the
Ichthyological Provinces and the Ecoregions are convenient categories to consider
the freshwater fishes of Angola.
Cabinda
Pellegrin (1928) recorded 28 species from the Chiloango River from Republic of
Congo (formerly ‘French’ Congo). The freshwater fishes of Lower Guinea, Central
West Africa that includes Cabinda were considered in detail through the two
212 P. H. Skelton
volumes edited by Stiassny et al. (2007). This rich Central West African fauna
includes 555 species in 147 genera and 38 families of which 78 species, 52 genera
and 25 families have been recorded from the Chiloango (also Shiloango) River in
Cabinda (Appendix 1). This Atlantic coastal river is clearly boosted by the large
number of marine and estuarine species that enter freshwaters sporadically or regu-
larly (Whitfield 2007). A number of species from here such as Enteromius holotae-
nia, Enteromius musumbi, Aplocheilichthys spilauchen and Oreochromis angolensis,
and estuarine species of marine origin also occur in the lower reaches of Angolan
Atlantic coastal rivers, some to as far south as the Cunene River (Penrith 1982, Hay
et al. 1997). Fowler (1930) recorded a number of species in the Academy of Natural
Sciences of Philadelphia collection taken from the Chiloango region as it was
known at the time. The fauna from the Chiloango in Angolan territory is likely
underrepresented in most groups due to lack of sampling.
Lower Congo
There are no records available of fishes collected in Angolan waters of the Lower
Congo mainstream or of the southern bank tributaries. The largest of these tributar-
ies is the Inkisi River of which the fish fauna is known from the studies of Wamuini
Lunkayilakio et al. (2010) supplemented by the description of new species described
in association with that work (Wamuini Lunkayilakio and Vreven 2008, 2010).
Based on these studies it is likely that most of the species in the DRC from the
reaches above the Sanga Falls at least are likely to occur in Angola as well. The
nature of the likely fauna of this neglected area of Angola as far as fish exploration
is concerned (Appendix 2) indicates that the species are essentially of Congolian or
Lower Guinean affinity with a few endemic species indicative of the isolation of
fauna in the river reaches above the Sanga Falls. The widespread presence of
Oreochromis niloticus is attributed to introduction for aquaculture (Wamuini
Lunkayilakio et al. 2010).
Cuanza and Atlantic Coastal Rivers
Poll (1967) listed 110 species in 32 genera and 15 families from the Atlantic coastal
region that includes the Cuanza River, which is revised (Appendix 3) to 105 species
in 45 genera and 17 families in the light of more recent surveys in the Cuanza. There
are very few species recorded from the Angolan Coastal rivers other than the
Cuanza, and in areas north of the Bengo to the mouth of the Congo records from
Angola are practically non-existent. Devaere et al. (2007) record Channallabes
apus as being described from this region. Fowler (1930) noted species of the Cuanza
and the Bengo rivers received from the British Museum on exchange, in many
instances as described by Boulenger (1910) or as recorded in Boulenger
(1909–1916). Trewavas (1936) recorded and described seven species from a head-
water stream of the Cuvo River arising on Mount Moco including the only Amphilius
species (Amphilius lentiginosus) described from the region. A second undescribed
Amphilius species has been recorded in the Cuanza (South African Institute for
Aquatic Biodiversity – SAIAB – collection). Both these species differ in key mor-
phological characteristics from the Amphilius of the Zambezian region that indicate
their faunal connections lie primarily with the Lower Guinean or Congolean regions.
Trewavas (1936) also described species from the Longa (Enteromius breviceps), the
Catumbela (Enteromius dorsolineatus, E evansi), and the Balombo (Enteromius
dorsolineatus). Pellegrin (1936) described the fishes collected by two Swiss expedi-
tions (1928–1929 and 1932) made under the direction of Monard from the Musée
d’Histoire Naturelle de la Chaux-de-Fonds included two species, Enteromius kes-
sleri, Clarias dumerilii, that were drawn from the Cueve, with the bulk of the col-
lections coming from the Cunene, the Cuvelai and the Cubango. Ladiges and
Voelker (1961) described Kneria maydelli from the Cunene, and Enteromius rou-
sellei and Chiloglanis sardinhai from the Longa. Poll (1967) described Kneria sjo-
landersi and Chiloglanis angolensis from the Bero, to the north of the Cunene.
Trewavas (1973) recorded Oreochromis angolensis and Tilapia cabrae from the
Bengo. Bills et al. (2012) made a small collection from the upper reaches of the
Cueve River that included species of the following genera – Petrocephalus,
Enteromius, Labeobarbus, Micralestes, Amphilius, Chiloglanis, Clarias,
Pharyngochromis, Thoracochromis, Tilapia, Coptodon, and Mastacembelus. The
list is typically ‘Zambezian’ and the species positively identified are closely linked
to the upper Cuanzan and Cubango fauna. The indication from these references is
therefore that the Angolan Coastal fauna is a mix of Lower Guinean (along the
coastal plain) and Zambezian (above the escarpment) with some Congolean ele-
ments in the upper Cuanza/Lucala (see below).
The ‘Cuanzan or Angolan Coastal ’ ichthyofaunal region is drawn primarily on
what is known of the fishes of the Cuanza River as described by Boulenger (1910)
and in Boulenger’s (1909–1916) catalogue of fishes in the British Museum (Natural
History) now the Natural History Museum (NHM). Fowler’s (1930) account of the
fishes of the Gray African Expedition in 1929 included records from the Bengo and
the lower Cuanza, but also a collection of species from Chouzo on the upper reaches
of the Cutato-Cuanza tributary, that provided a first strong indication that the fauna
of these reaches is ‘Zambezian’ in character and different to those from the coastal
reaches as reported by Boulenger (1910) and others. This association was later reit-
erated by Trewavas (1973) when considering the cichlid species of the Cuanza and
Bengo rivers and has been firmly supported by the extensive surveys conducted by
SAIAB and INIP (Instituto Nacional de Investigação Pesqueira) between 2005 and
2010. The current assessment records at least 102 species, some of which are unde-
scribed (Appendix 3). The collections indicate that the river basin is even more
heterogenous in fish faunal characteristics than simply ‘lower’ and ‘upper’ and the
different zones distinguishable include (1) the lower reaches from the escarpment
base to the sea, (2) the escarpment reaches, (3) the upper Cuanza and (4) the Lucala
214 P. H. Skelton
tributary, itself probably sub-zoned into the middle and upper reaches separated by
the Calandula Falls (formerly ‘Duque de Bragança’ Falls).
Two ecophysiological components derive the fishes of the lower Cuanza: a
diverse Tropical West African or Lower Guinean brackish water or marine
component, and secondly the primary and secondary freshwater fishes. The known
Tropical West African brackish water fishes from the system are generally wide-
spread species and do not include endemics. Some species such as the Bull Shark
(Carcharhinus leucas) and the Atlantic Tarpon (Megalops atlanticus) are well
known as gamefish from this river. Two clupeid species include the recognised
freshwater species (Pellonula vorax and Odaxothrissa ansorgii) and probably other
brackish water forms. One haemulid (Pomadasys sp.) and one polynemid threadfin,
possibly Polydactylus quadrifilis as known from Central West Africa (Snoeks and
Vreven 2007), have been recorded (SAIAB records). Mullets (Mugilidae), as yet
unidentified at species level, are present as are the sleepers (Eleotridae) and gobies
(Awaous and Periopthalmus). Two pipefish have been positively identified:
Enneacampus ansorgii and Microphis brachyurus aculeatus. The tonguefish
Cynoglossus senegalensis was collected in the downstream reaches.
The freshwater species of this lower zone are mostly widespread species that
also occur in coastal reaches of rivers to the north, well into the adjacent Lower
Guinean Province and beyond, and many probably also to the south. An example of
this is Parailia occidentalis that has a range through to the Senegal River in West
Africa (de Vos 1995). The species that occur are found generally throughout the
region to the escarpment, with a few ascending into middle Cuanza sections. Other
characteristic species in this zone include mormyrids of the genera Hippopotamyrus,
Marcusenius and Petrocephalus, the alestid Alestes ansorgii, cyprinids of the genus
Labeo, two Enteromius species (E. holotaenia and E. musumbi), and several distinc-
tive claroteid catfishes (two Chrysichthys species C. acutirostris and C. ansorgii), as
well as Schilbe bocagii, and the widespread Clarias gariepinus. The Chrysichthys
species confirm the West Africa coastal affinities of the assemblage as the genus is
not known from the upper reaches nor from the upper Zambezian floodplain fauna.
The cichlid fauna, as detailed systematically by Trewavas (1973) is in part also
restricted to the zone – Oreochomis angolensis, Hemichromis angolensis and
Tilapia cf. cabrae. The range of the procatopodid lampeye Aplocheilichthys
spilauchen previously known from the Senegal River to the Bengo River has been
extended to the Cuanza. The absence of the anabantid genus Ctenopoma from this
zone is remarkable.
The Escarpment Zone of the Cuanza is characterised by a stepwise series of
rapids, cascades and falls interspersed by rocky pools and runs. The fish fauna of
this important zone for hydropower development is rich but relatively poorly known
or described. The SAIAB-INIP collections are extensive and indicate that few spe-
cies from the coastal zone penetrate high up into the zone. This is most probably
partly an artefact of the Cambambe Dam near the base, in existence for several
decades, which has likely affected the natural penetration of many species. The
major freshwater families are represented; the smaller cyprinids, various catfish
families, and cichlids are particularly well represented. The generic composition
includes: Hippopotamyrus, Petrocephalus, Marcusenius, Parakneria, Enteromius,

Labeobarbus, Labeo, Raiamas, Brycinus, Rhabdalestes, Hepsetus, Schilbe,
Chrysichthys, Clarias, Clariallabes, Parauchenoglanis, Chiloglanis, Synodontis,
Micropanchax, Hemichromis, Pharyngochromis, Pseudocrenilabrus,
Serranochromis, Tilapia, Oreochromis, and Mastacembelus. Only a single
Labeobarbus species was recorded in this zone during the survey and it also occurs
in the Lucala tributary. Boulenger (1910) recorded two Labeobarbus species from
the Cuanza at Dondo – L. rocadasi and L gulielmi. A unique morphotype of
Labeobarbus with an extremely pointed tiny mouth, collected during the Capanda
pre-impoundment surveys is present in the Luanda Museum (pers. obs., Fig. 11.2)
and is likely to be an undescribed species.
The Upper Cuanza extends from a waterfall on the mainstream above the
Capanda dam to the watershed and consists largely of relatively low-gradient flood-
plain rivers on Kalahari sand formations, similar to that of the upper reaches of the
Zambezi and Okavango systems in Angola. Characteristic genera from this zone
are: Hippopotamyrus, Petrocephalus, Marcusenius, Parakneria, Enteromius,
Labeobarbus, Labeo, Brycinus, Rhabdalestes, Hepsetus, Schilbe, Chrysichthys,
Doumea, Clarias, Clariallabes, Parauchenoglanis, Chiloglanis, Synodontis,
Micropanchax, Hemichromis, Pharyngochromis, Pseudocrenilabrus,
Serranochromis, Tilapia, Oreochromis, and Mastacembelus. Fishes from Chouzo in
the upper Cuanza described by Fowler (1930), include species such as Marcusenius
angolensis, Hepsetus cuvieri, Labeo rocadasi, Enteromius evansi (type locality),
Enteromius lujae (identity of this species is still debated but the same species occurs
in the Okavango headwaters), Clarias gariepinus, Clarias theodorae (as C. fou-
loni), Clarias ngamensis (as Dinotopterus prentissgrayi), Ctenopoma machadoi
(type locality), Serranochromis macrocephalus (as Tilapia acuticeps, see Trewavas
1973). Norman (1923) described Synodontis laessoei, synonymised with Synodontis
nigromaculatus by Poll (1971), as the only species of this genus in the Cuanza, a
contrast to the specious lineage in the Okavango-Zambezi region (Day et al. 2009,
Pinton et al. 2013). Few species characteristic of the upper Cuanza are found beyond
the zone within the basin. This agrees with the notion that the fauna in this zone is
historically and biogeographically an integral part of the ‘Zambezian’ fauna
Fig. 11.2 An extraordinary undescribed Labeobarbus species from the Cuanza River in the
Luanda Museum, 2005. (Photo PH Skelton)
216 P. H. Skelton
(Trewavas 1973). Ladiges (1964) and Poll (1967) showed this to be general for the
fauna as a whole, and specific studies on species like Hepsetus cuvieri (Zengeya
et al. 2011) and cichlids like Serranochromis and Tilapia sparrmanii (Musilová
et al. 2013) confirm this relationship. Recent surveys across the watershed between
the Cuanza and the Okavango indicate that a number of other species like Parakneria
fortuita, and several Enteromius species like E mocoensis, E evansi, E breviceps, E
brevidorsalis occur in streams on either side and have helped to define the Upper
Zambezi headwaters ecoregion that embraces this trans-system conformance.
An early indication that the Lucala River, a major tributary that joins the system
in the lower reaches, is exceptional for its fishes was the fauna collected by Ansorge
using a wide range of methods including explosives (Boulenger 1910). It is however
only in the escarpment and upper reaches that such exception occurs. An assem-
blage of large fishes of the genus Labeobarbus in particular is outstanding, and
Boulenger (1910) described 12 species now in Labeobarbus (Vreven et al. 2016),
all of which remain valid at this time. In addition to these species, unpublished bar-
code studies conducted by SAIAB on the fauna indicates that several lineages in the
system are restricted to the Lucala, including an Alestes, Pharyngochromis,
Serranochromis, Tilapia, two Enteromius species, a Parakneria, Hippopotamyrus,
and a undescribed Congoglanis.
The significance of the use of explosives in assembling Ansorge’s collection
described by Boulenger (1910) is that it included a number of large mainstream spe-
cies that otherwise are extremely difficult to collect. The assemblage of large
Labeobarbus described in the paper has defined the Cuanza Basin since that time.
The overall faunal characteristics of the Lucala include species of the following
genera: Hippopotamyrus, Petrocephalus, Kneria, Alestes, Enteromius, Labeobarbus,
Labeo, Raiamas, Amphilius, Congoglanis, Schilbe, Clarias, Chiloglanis, Synodontis,
Micropanchax, Pharyngochromis, Serranochromis, Tilapia, and Mastacembelus.
The fauna of the upper reaches is poorly known. Only a single collection made by
SAIAB was drawn from the Lucala above the Calandula Falls. This limited sample
is not sufficient to gauge the full character of the zone, but does indicate a degree of
continuity with the Middle Lucala zone, and differing through the absence of major
elements like the Labeobarbus species so characteristic of the latter. The physical
character of the upper reaches suggests there is a zonal distinction in the ecological
character and thus the faunal elements. The known fauna includes species of the
following genera: Hippopotamyrus, Petrocephalus, Parakneria, ‘Barbus’,
Enteromius, Amphilius, Congoglanis, Clarias, Micropanchax, Pharyngochromis,
Serranochromis. Little else can be stated at this point except that an investigation
into the fauna is highly desirable given the unique nature of the Middle Lucala.
The Lucala catchment shares its watershed with tributaries of the Congo-Cuango
River and is likely one of the underlying reasons for its unique character. A high
degree of endemicity to this catchment is therefore evident and with further taxo-
nomic investigation likely to be upheld and enhanced.
Cunene
Poll (1967), from the ichthyological perspective, treated the Cunene River system
as a separate entity to the Atlantic coastal region, whereas it has been regarded as
part of the Zambezian Province by Roberts (1975), part of the ‘Angola’ ichthyofau-
nal province by Lévêque and Paugy (2017a, b), and divided as part of the Namib
aquatic ecoregion and part upper Zambezian headwaters ecoregions by Thieme
et al. (2005). The reason for these varied treatments is that the river system is geo-
eco-historically complex. Thus it has a dual geomorphological origin (the upper
reaches being a natural part of the Kalahari Basin that has been captured by an
Atlantic coastal river) and environmentally the lower reaches sit within the ‘xeric’
Namib region and the inland upper reaches within the savanna dry forest environs.
The fishes of the Cunene River are relatively well documented, starting with
Schilbe steindachneri (a synonym of S. intermedius) and Mormyrus anchietae (a
synonym of M. lacerda) described by Guimarães (1884), and summarised in the
most recent checklist by Hay et al. (1997). Excluding the more strictly marine fami-
lies there are 82 species recorded from the Cunene (Appendix 3). Hay et al. (1997)
also record the broad distribution of species within the system according to three
sections, the upper reaches down to Ruacana Falls, a middle section down to Epupa
Falls and the lower river from below Epupa Falls to the mouth. Of 65 species
recorded above Ruacana Falls 13 are restricted to that section. At least one species,
Marcusenius deserti, is restricted to the lower reaches close to the coast (Kramer
et al. 2016). Apart from the several marine species recorded in the extreme lower
reaches by Penrith (1970) and Hay et al. (1997) that reflect a southernmost exten-
sion of the tropical (Lower Guinean) fauna, the general composition is clearly
Zambezian in character. There are few representatives indicative of the Angolan
(Cuanzan) Province, e.g. Enteromius mattozi (described by Guimarães (1884) from
the Curoca River to the north of the Cunene). Pellegrin’s (1921) Enteromius (for-
merly Barbus) rohani, probably a synonym of E. mattozi, was likely taken from the
Caculovar River, a tributary of the Cunene, and not from the Lomba (neither the
Zambezi as Pellegrin claimed, nor the Longa coastal Atlantic as suggested by Poll
1967). Enteromius argenteus is another minnow that has been reported from the
Cunene but whose identity is unconfirmed – and is likely to be juveniles of E. mat-
tozi (Skelton Unpublished Data).
There are also several isolated endemics from the system such as Marcusenius
deserti, Marcusenius magnoculis, Marcusenius multisquamatus, Hippopotamyrus
longilateralis, Engraulicypris howesi, Zaireichthys cuneneensis, Orthochromis
machadoi, Thoracochromis albolabrus, Thoracochromis buysi, that suggests a
degree of isolation probably reflecting older biogeographic connections. The
absence of certain conspicuous families or genera such as Parakneria, Labeobarbus,
Opsaridium, Hydrocynus, Parauchenoglanis, Amphilius, Hemichromis, and
Mastacembelus is also noteworthy and perhaps indicative of a lack of more recent
connections with the Zambezian and Cuanza systems.
218 P. H. Skelton
Cassai and Southern Congo Rivers
Collections from the Lulua River, a tributary of the Cassai in Congo by Fowler
(1930) whilst not strictly in Angola, probably pertain to Angola as well. Thus
although not the only source, Poll (1967) is the current practical published source
for the fishes of the southern Congo river tributaries in Angola. There are three main
tributaries draining the region, from the east the Cassai including the Luangwe, the
Cuilu (or Kwilu) and the Cuango. Poll (1967: 18–23) plotted the records of the
fishes of each of these in his distribution table, recording 108, 28 and 37 species
respectively and in the addendum supplemented the Cassai with three species and
the Cuango with 24 species. The figure for the southern catchments of the Congo in
Angola is now estimated at around 162 species (Appendix 3). The Cuilo and Cuango
faunas are most evidently far from well explored. The Cassai fish fauna is better
represented but still poorly explored, and includes species both typical of the Congo
(e.g. Polypterus ornatipinnis, Channallabes apus, several mormyrid species,
Bryconaethiops microstoma, Alestes grandisquamis, Distichodus fasciolatus,
Distichodus lusosso, Mastacembelus congicus), and many species found also in the
Upper Zambezi or the Okavango (e.g. Hydrocynus vittatus, Hepsetus cuvieri,
Pollimyrus castelnaui, Enteromius brevidorsalis, Parauchenoglanis ngamensis,
Clarias stappersii, Clarias theodorae, Schilbe yangambianus, Micropanchax katan-
gae, Oreochromis andersonii, Coptodon rendalli, Tilapia sparrmanii, Tilapia
ruweti, Hemichromis elongatus, Serranochromis microcephalus, Serranochromis
robustus jallae, Pseudocrenilabrus philander, Ctenopoma multispine,
Microctenopoma intermedium). The presence of Dundocharax bidentatus in the
Cassai and the rare Zambezian endemic not yet found in Angola, Neolebias lozii are
further good indicators of geographical connection. The strong Cassai-Zambezian
faunal association is attributed to the clear evidence of hydrological pattern that the
upper Cassai was formerly part of the Upper Zambezi system (Bell-Cross 1965).
Zambezian-Cuando-Cubango Headwaters and Floodplains
There is sufficient direct connection between the Zambezi, the Cuando and the
Okavango river basins and similarity of the fish faunas in each to consider these
under a single heading.
The Zambezi headwaters in Angola drain the Kalahari sand formation over an
extensive divide with the Cassai to form major floodplains known as the Bulozi
Floodplains. There are a number of lakes associated with the drainage including the
largest freshwater lake in Angola, Lake Dilolo. The Okavango drainage is divided
into two branches, the Cuito-Cuanavale in the east and the Cubango in the west. The
Cuito-Cuanavale drains Kalahari sand formations giving rise to extensive low-
gradient seepage bog and floodplain rivers in slump valleys extending into miombo
savanna woodlands in the upper reaches. There are several lakes in these headwaters.
The Cubango branch arises as several relatively steep gradient rocky rivers in the
Angolan highlands on the Bié plateaux before descending to the low-gradient
reaches along the Namibian border to join with the Cuito before crossing to
Botswana and forming the mostly endorheic Okavango Delta. The watershed of the
system is shared with the Cuando, the Zambezi (mainly the Lungwe-Bungo), the
Cueve-Cuanza, and the Cuanza as well as the Cunene and Cuvelai oshanas in the
west.
The fishes of the Upper Zambezi are well studied and documented (e.g. Jackson
1961, Jubb 1961, 1967, Balon 1974, Bell-Cross and Minshull 1988, Tweddle 2010)
with numbers now estimated at around 100–120 species (Appendix 3; Tweddle
et al. 2004), possibly with as many as 20–25 undescribed. However published
records from the Angolan territory are sparse, and limited in the published literature
to Poll’s (1967) 41 species (against his checklist of 62) taken mostly from two
localities close to the watershed (Lagoa Calundo and the Longa-Luena tributary).
Recent collections from the source reaches of Zambezian tributaries in Angola
made by the National Geographic Okavango Wilderness Project (NGOWP 2018)
are still being assessed but include 39 species from 12 families that have been
included in the checklist of fishes from this region (Appendix 3). One notable new
record is Enteromius chiumbeensis described by Poll (1967) from the Chiumbe
River a tributary of the Cassai, reinforcing the close connections between these
adjacent trans-watershed systems.
The upper Zambezian fish fauna is distinctive in several respects, most notably
for the relatively speciose endemic Synodontis catfishes and the serranochromine
cichlids (Trewavas 1964, Bell-Cross 1975, Greenwood 1993, Day et al. 2009;
Pinton et al. 2013). To a large extent, in Angola, the fauna is ecologically tuned to
the extensive seepage and floodplain drainages within a band of miombo savanna
woodland on Kalahari sand deposits. Overall the known Angolan Upper Zambezi
fish fauna is similar to that of the better-studied (in Angola) Okavango Basin fishes
(often with the same or closely related species e.g., mormyrids of the genera
Hippopotamyrus, Marcusenius, Petrocephalus, Pollimyrus – Kramer et al. 2003,
2004, 2012, 2014, and Zaireichthys species – Eccles et al. 2011). Whilst there are a
few endemics, only one, Paramormyrops jacksoni Poll 1967 is restricted to Angola.
The isolated Neolebias lozii is known only from the Barotse floodplains in Zambia.
Fishes of the Cuando-Linyanti-Chobe system have not been reported on from the
Angolan section of that Zambezi tributary but van der Waal and Skelton (1984)
provided a checklist of fishes in the Cuando River in Namibian waters. The 56 spe-
cies recorded were all also found in the Zambezi system in Namibia. The Pallid
Sand Catlet, Zaireichthys pallidus Eccles et al. (2011) is described from the Cuando
but is not restricted to that system. Kramer et al. (2014) described a new species of
Pollimyrus from the Cuando, a species possibly endemic to that tributary. Recent
collections by the National Geographic Okavango Wilderness Project (NGOWP/
SAIAB) from the upper reaches of the Cuando in Angola further inform the list of
species (Appendix 3).
The fishes of the Okavango Basin have been studied and reported on in the litera-
ture for over 150 years since Castelnau (1861) described 14 species from Lake
220 P. H. Skelton
Ngami, including the iconic Tigerfish (Hydrocynus vittatus) the Southern African
Pike (Hepsetus cuvieri), the large Blunttooth Catfish (Clarius ngamensis) and the
Three Spot Bream (Oreochromis andersonii). Fifty years later Boulenger (1911)
reported on a collection from the Okavango-Lake Ngami made by RB Woosnam
and described six new species including one named for Castelnau – Pollimyrus
castelnaui. These fishes were all included in Gilchrist and Thompson (1913, 1917)
and Boulenger (1909–1916). Fowler (1935) described a collection made from the
Delta by the Vernay-Lang Expedition of 1930. Pellegrin (1936) described fishes
collected by two Swiss expeditions of 1929 and 1933 from the Cunene, the Cuvelai
and the Cubango. Barnard (1948) described in detail a collection from Rundu,
Namibia. The results of all these efforts were summarised in checklists published by
Poll (1967), Jubb (1967), Jubb and Gaigher (1971) and Skelton et al. (1985). More
recently surveys of Angolan Okavango Basin rivers have been made (Bills et al.
2012, 2013, Skelton et al. 2016) that have reached little-explored areas, encountered
additional species and provide for a more complete assessment of the fishes and
their intra-basin distributions.
The additional species recently discovered include new species of Clariallabes,
several serranochromine cichlids, and a dwarf climbing perch (Microctenopoma
sp). Recent distribution records extend the range of several species from the Congo
tributaries or in the case of Clypeobarbus bellcrossi from Zambezi headwaters in
Zambia to the Okavango. Congolean species such as Marcusenius moorii (Günther)
and Enteromius chicapaensis (Poll), and Nannocharax lineostriatus (Poll), and sev-
eral Micropanchax as M. luluae, M. nigrolateralis, M. lineolateralis. The known
range of a number of species from the Atlantic Coastal and Cuanza systems has
been extended to the Okavango, e.g. Enteromius breviceps, E. brevidorsalis, E.
evansi, E. mocoensis, E. greenwoodi. A new understanding of the complex distribu-
tion of the twin species Enteromius trimaculatus and E. poechii has also been
reached – the former being found in the Cunene and the extreme upper reaches of
the Cubango in place of the latter which is widespread in the downstream floodplain
reaches of the Okavango and Upper Zambezi system.
Cuvelai
The Cuvelai drainage lies in a triangle between the Okavango in the east and the
Cunene in the west and the streams known as ‘iishanas’ are intermittent, only flow-
ing during periods of sustained rainfall into the endorheic Etosha Pan in Namibia
(van der Waal 1991, Hipondoka et al. 2018). The 1929 and 1932–1933 Swiss expe-
ditions to Angola collected the following species from Mupa (Pellegrin 1936):
Marcusenius altidorsalis (?), Mormyrus lacerda, Enteromius paludinosus, Tilapia
sparrmanii, and Pseudocrenilabrus philander. Seventeen species, all conforming to
Cunene fauna, have been confirmed from the western iishanas of the system by
Hipondoka et al. (2018), and connections with the Cunene substantiated through
remote sensing techniques. Four widespread pioneering species are consistently
present in collections, viz., Clarias gariepinus, Enteromius paludinosus,

Oreochromis andersonii and Pseudocrenilabrus philander and several others are
common – Clarias ngamensis, Schilbe intermedius and Enteromius trimaculatus.
Biogeography
The biogeography of Angolan freshwater fishes is closely tied to the geomorphol-

ogy and the geomorphological history of the territory. In brief, Angola consists of a
narrow coastal plain, a distinct escarpment and an interior plateau that is being
eroded most rapidly from the Congo Basin. The coastal plain consists of a series of
rivers flowing from the escarpment or – in the case of the Congo in the north, the
Cuanza in the middle and the Cunene in the south – where the escarpment has been
penetrated, from the interior plateau or the Congo Basin. The fish fauna of the
coastal plain is primarily a southern extension of the tropical coastal fauna of West
Africa and Central West Africa. River connections along this narrow strip are either
via sea-level fluctuations or via river captures between watersheds, either as adja-
cent systems or via extended reaches through captured inland drainages that are not
determined by the coastal gradients and processes. According to Lévêque and Paugy
(2017a,b) the primary direction of dispersal of the coastal west African fauna was
northwards from the Congo. Present day ocean currents off Angola are counter
clockwise (http://oceancurrents.rsmas.miami.edu) and it is possibly only inshore
counter currents that might have facilitated faunal dispersal southwards from the
Congo, especially after the capture and penetration of the Congo Basin by the Lower
Congo in the late Cretaceous (Flügel et al. 2015). Such would certainly explain
much of the marine derived elements of the region. Given favourable currents it is
likely also that the considerable volumes of freshwater entering the sea from the
Congo at various times would facilitate even freshwater fishes down the coast and
might explain the presence of such species as Enteromius musumbi, Physailia occi-
dentalis, Chysichthys spp, Oreochromis angolensis and Aplocheilichthys spilauchen
in the Angolan region. An alternative and complementary explanation for some
freshwater faunal elements such as Marcusenius deserti and Raiamas ansorgii of
the Angolan Coastal reaches is that it is primarily derived via the Cuanzan and
Cunene gateways through capture of portions of the Kalahari Basin drainage. It is
not only the Cuanza and the Cunene that have breached the escarpment but also the
Cuvo and the Longa and possibly others, as is evident in the list of freshwater fishes
reported from these lesser rivers (see above).
The evolution of the extensive Kalahari Basin is certainly key to understanding
the majority of the freshwater fish fauna of Angola. Haddon and McCarthy (2005),
Key et al. (2015), Moore and Larkin (2001), and Moore et al. (2012) sketch the
evolution of the Kalahari Basin and its drainage since the breakup of Gondwanaland
and the isolation of Africa in the late Cretaceous. Following rifting, the continental
margins were probably elevated and this formed an escarpment that separated the
narrow coastal plain from the elevated Kalahari sedimentary basin that was drained
222 P. H. Skelton
primarily by the palaeo-upper Zambezi, the predominant system in the Angolan

region (Fig. 11.3). The western portion of the system flowed from the escarpment
highlands of the extreme northwest of the basin, now part of the Cuanza, generally
southeast through the Limpopo valley to the Indian Ocean. The eastern parts of the
upper Zambezi reached northeastwards to as far as pre-rift East African plateaus and
Fig. 11.3 A diagrammatic model for the post-Cretaceous biogeography of Angolan freshwater
fishes. Angolan border – fine dotted line; present day drainage – thin lines; present day inter basin
watersheds – open dotted lines; paleo drainage lines – thick extended arrows; paleo and present
escarpment retreat – dashed arrows; paleo and present south and southwestern Congo Basin water-
shed – thick dashed lines; Angolan escarpment – right slanted hash; gateway drainage captures –
large open bi-directional arrows. Coastal dispersal of fishes – large left-slanted bi-directions arrow.
The model is based on geomorphological interpretations by Flügel et al. (2015), Haddon and
McCarthy (2005), Moore and Larkin (2001), Moore et al. (2012), and others
included the proto-Luangwa and the proto-Chambeshi-Kafue-upper Zambezi as

well as the Okavango. These drained into an interior basin to form, at times, a mega
palaeo lake – Palaeolake Magadigadi (Burroughs et al. 2009, Moore and Larkin
2001, Moore et al. 2012, Podgorski et al. 2013). The proto-Cunene consisted of an
upper portion draining endorheically to the Etosha basin. The most significant
events in the history of the Kalahari Basin were firstly the downwarping and back-
tilting of drainage coupled with upwarping along the southern margins that severed
the initial Indian Ocean outlet via the Limpopo; the tapping of the Congo Basin by
the lower Congo River that advanced the erosion and southern retreat of the north-
ern watershed of the basin, especially in the northeast (Luapula-Chambeshi) and, in
the Angola area, the Cassai-Zambezi. The dismemberment and tapping of drainage
portions from the Kalahari Basin to coastal outlets including the Cuanza, the
Cunene, and the Zambezi also affects the biogeographical history significantly
(Moore and Larkin 2001, Moore et al. 2012, Key et al. 2015).
The most profound biogeographic significance to emerge from this geomorpho-
logical narrative is that the Kalahari Basin has been an evolutionary basin for fishes
over a long period of time. The evidence is exemplified in the serranochromine
cichlid radiation and the clade of Synodontis catfish and the radiation of several
mormyrid genera that characterise the Zambezian fauna (Bell-Cross 1975,
Greenwood 1984, Kramer et al. 2003, 2004, Day et al. 2009, Kramer and Swartz
2010, Kramer et al. 2012, Schwarzer et al. 2012, Pinton et al. 2013, Kramer and
Wink 2013). The strong identity of the upper Zambezian fauna further exemplifies
this notion. That the fauna has been supplemented with species from neighbouring
ichthyological provinces, especially the Congo, is also evident in species or genera
with internally restricted distributions such as Hepsetus cuvieri, Hydrocynus
vittatus, Parauchenoglanis ngamensis, Mastacembelus, Hemichromis elongatus,
Amphilius and others. The broader distributions of some species into basins like
the east coast rivers (e.g. Enteromius bifrenatus, Microctenopoma intermedium,
Clarias theodorae, Brycinus lateralis) gives biological credence to the former east
coast linkage and subsequent drainage dismemberment on the proto-upper Zambezi
(Skelton 1994, 2001).
There are other emerging details of biogeographical interest to Angola that will
in time lead to a detailed accounting of the origins and development of the freshwa-
ter fishes. Thus the presence of doumeine catfishes in the Cuanza, southwest of the
Congo, indicates clearly insemination from the Congo. The flock of Labeobarbus
species in the Lucala-Cuanza probably also indicates a Congolian insemination.
However the assumption that all traffic was from the Congo is not necessarily cor-
rect and Neolebias bidentatus in the Cassai, for example, as with other ‘Zambezian’
elements in that system, more likely reflects a Zambezian (i.e. Kalahari) insemina-
tion to the Congo. This, in essence, is the basis of the ‘Upper Zambezi headwater’
freshwater ecoregion (Fig. 11.1: basin C).
224 P. H. Skelton
Conservation
Angola is an emerging African economy with a rapidly growing human population

and increasing demand on freshwater resources. The rapid population growth and
expansion of urban areas in places such as Luanda but also in the more rural districts
(Mendelsohn and Weber 2015) is placing an ever increasing stress on the environ-
ment, especially that of the rivers for which such urban growth centres are depen-
dent on for water and power. Although many Angolan rivers are relatively
unregulated there are dams on several systems such as the three major hydroelectric
dams on the Cuanza. A further four hydroelectric dams are planned for the escarp-
ment section of this system alone. In the case of certain transboundary rivers like the
Okavango, the threat of increased river regulation is of serious concern to the integ-
rity of the Okavango Delta in Botswana, a World Heritage and Ramsar site (King
and Chonguic 2016).
Diamond mining activities along the southern Congo tributaries have had envi-
ronmental impacts of unknown severity as practically no public investigations or
information is available.
With human populations, urbanisation and development comes pollution and
other direct threats to aquatic life such as fishing and the introduction of invasive
alien species. Few alien fishes have been recorded from Angola, but two species that
have been introduced are Oreochromis mossambicus (SAIAB, in the Cuanza) and
Oreochromis niloticus in Cabinda and, as recently confirmed, in the upper Cubango.
The threats these particular species pose as aliens is well documented (e.g. Wise
et al. 2007, Zengeya et al. 2013, Bbole et al. 2014). This is the first record of an alien
species with high impact potential in the Okavango system and the threat posed is
transboundary in nature. Potential transboundary threats from outside Angola
include that of alien crayfish from the Zambezi (Nunes et al. 2016).
Indigenous fishery practices in Angola include a range of gear ranging from sim-
ple traps to elaborate fishing fences and walls (Poll 1967, Mendelsohn and Weber
2015). In places such traditional practices are still in evidence (Fig. 11.4 top), but
elsewhere traditional practices are being replaced by modern gear such as monofila-
ment gillnets and mosquito-net seines (Fig. 11.4 bottom) that are excessively
destructive and unsustainable (Tweddle et al. 2015).
The current IUCN redlist assessments for Angolan freshwater fishes (Appendix
3) reflects the relatively weak knowledge of the species – a third of the known spe-
cies are either not assessed or are Data Deficient (DD). One species (Oreochromis
lepidurus) is listed as Endangered (see Moelants 2010), three are Vulnerable (1%)
and 185 (65%) are Least Concern. The endangered species is a Lower Congo
endemic found mainly in the DRC and is primarily threatened by oil pollution
derived from boats. The Vulnerable species are also cichlids of the genus
Oreochromis – O. andersonii (see Marshall and Tweddle 2007) and O. macrochir
(see Marshall and Tweddle 2007), both are threatened through hybridisation from
the alien invasive species Oreochromis niloticus. The latter species has recently
been confirmed as present in Angola, within the Okavango catchment and its impact
Fig. 11.4 Top – traditional fishing fence on the Cacuchi River, 2012 (Photo PH Skelton). Bottom –
Drying fish caught with monofilament gillnets on the Cuito River, 2015. (Photo G Neef)
226 P. H. Skelton
on the native Oreochromis is now an imminent threat. Given the situation of rapidly
escalating changes to the natural aquatic environment in Angola it is likely that the
IUCN redlist score for the country will rise rapidly.
Acknowledgements I am supported in my research by the Director and staff of SAIAB, in par-

ticular Roger Bills and members of the collections division, administration staff, and by Maditaba
Meltaf in the library for the provision of literature. Steve Boyes and John Hilton of the Wild Bird
Trust have provided me with excellent opportunity to study fishes in Angola since 2015. I have
been supported in the field and laboratory by Adjani Costa, Roger Bills, Ben van der Waal, Götz
Neef and others of the National Geographic Okavango Wilderness Project. SAIAB engagement
with Angolan fishes was initiated in 2005 in partnership with INIP (Instituto Nacional de
Investigação Pesqueira). Ernst R Swartz (SAIAB) and D Neto (INIP) were instrumental in opening
the channels of new knowledge on Angolan freshwater fishes.
Appendices
Appendix 1
Freshwater and brackish water fishes of Shiloango River, Cabinda, as recorded by

Stiassny et al. (2007)
Species Author & Date

Clupeidae
Pellonula vorax Günther, 1868
Mormyridae
Isichthys henryi Gill, 1863
Marcusenius moorii Günther, 1863
Paramormyrops kingsleyae (Günther, 1863)
Brienomyrus brachyistius (Gill, 1862)
Hepsetidae
Hepsetus lineatus (Pellegrin, 1926)
Alestidae
Brycinus longipinnis (Günther, 1864)
Brycinus macrolepidotus Valenciennes, 1850
Brycinus kingsleyae (Günther, 1896)
Nannopetersius ansorgii (Boulenger, 1910)
Distichodontidae
Distichodus notospilus Günther, 1867
Eugnathichthys macroterolepis Boulenger, 1899
Nannaethiops unitaeniatus Günther, 1872
Nannocharax parvus Pellegrin, 1906
Neolebias ansorgii Boulenger, 1912
Neolebias spilotaenia Boulenger, 1912
(continued)

Cyprinidae
Enteromius carens (Boulenger, 1912)
Enteromius jae (Boulenger, 1903)
Enteromius guirali (Thominot, 1886)
Enteromius callipterus (Boulenger, 1907)
Enteromius camptacanthus (Bleeker, 1863)
Enteromius rubrostigma (Poll & Lambert, 1964)
Enteromius holotaenia (Boulenger, 1904)
Labeobarbus aspius (Boulenger, 1912)
Labeobarbus cardozoi (Boulenger, 1912)
Labeobarbus roylii (Boulenger, 1912)
Labeobarbus batesii (Boulenger, 1903)
Labeobarbus sandersi (Boulenger, 1912)
Labeo batesii Boulenger, 1911
Labeo lukulae Boulenger, 1902
Opsaridium ubangiense (Pellegrin, 1901)
Ariidae
Arius latiscutatus Günther, 1864
Claroteidae
Anaspidoglanis macrostoma (Pellegrin, 1909)
Parauchenoglanis altipinnis (Boulenger, 1911)
Chrysichthys auratus (Geoffroy Saint-Hilaire, 1809)
Chrysichthys nigrodigittatus (Lacépède, 1803)
Schilbeidae
Parailia occidentalis (Pellegrin, 1901)
Pareutropius debauwi (Boulenger, 1900)
Clariidae
Clarias angolensis Steindachner, 1866
Clarias gabonensis Günther, 1867
Malapteruridae
Malapterurus beninensis Murray, 1855
Procatopodidae
Aplocheilichthys spilauchen (Duméril, 1861)
Plataplochilus loemensis (Pellegrin, 1924)
Nothobranchiidae
Epiplatys singa (Boulenger, 1899)
Aphyosemion escherischi (Ahl, 1924)
Anabantidae
Ctenopoma nigropannosum Reichenow, 1875
Microctenopoma ansorgii (Boulenger, 1912)
Microctenopoma nanum (Günther, 1896)
Microctenopoma congicum (Boulenger, 1887)
(continued)
228 P. H. Skelton

Cichlidae
Pelvicachromis subocellatus (Günther, 1872)
Chilochromis duponti Boulenger, 1902
Coptodon tholloni (Sauvage, 1884)
Pelmatolapia cabrae (Boulenger, 1899)
Coptodon guineensis (Günther, 1862)
Oreochromis schwebischi (Sauvage, 1884)
Sarotherodon nigripinnis (Guichenot, 1861)
Lutjanidae
Lutjanus dentatus (Duméril, 1861)
Monodactylidae
Monodactylus sebae (Cuvier, 1829)
Polynemidae
Polydactylus quadrifilis (Cuvier, 1829)
Mugilidae
Mugil bananensis (Pellegrin, 1927)
Neochelon falcipinnis (Valenciennes, 1836)
Chelon dumerili (Steindachner, 1870)
Eleotridae
Eleotris daganensis Steindachner, 1870
Eleotris senegalensis Steindachner, 1870
Eleotris vittata Duméril, 1861
Bostrychus africanus (Steindachner, 1879)
Dormitator lebretonis (Steindachner, 1870)
Gobiidae
Periopthalmus barbarus (Linnaeus, 1766)
Gobionellus occidentalis (Boulenger, 1909)
Bathygobius soporator (Valenciennes, 1837)
Bathygobius casamancus (Rochebrune, 1880)
Nematogobius maindroni (Sauvage, 1880)
Microdesmidae
Microdesmus aethiopicus (Chabanaud, 1927)
Mastacembelidae
Mastacembelus shiloangoensis (Vreven, 2004)
Mastacembelus niger Sauvage, 1879
Syngnathidae
Enneacampus ansorgii (Boulenger, 1910)
Microphis aculeatus (Kaup, 1856)
Cynoglossidae
Cynoglossus senegalensis (Kaup, 1858)
Citharichthys stampflii (Steindachner, 1894)
Appendix 2
Freshwater fishes of the Inkisi River DRC, from above the Sangha waterfalls, after
Wamuini Lunkayilakio et al. (2010)

Mormyridae
Hippopotamyrus cf. ansorgii (Boulenger, 1905)
Paramormyrops cf. kingsleyae (Günther, 1896)
Paramormyrops cf. sphekodes (Sauvage, 1879)
Cyprinidae
Enteromius miolepis (Boulenger, 1902)
Enteromius unitaeniatus (Günther, 1867)
Enteromius vandersti (Poll, 1945)
Garra congoensis Poll, 1959
Labeo macrostomus Boulenger, 1898
Labeobarbus sp. nov.
Labeobarbus boulengeri Vreven, Musschoot, Snoeks & Schliewen, 2016
Labeobarbus robertsi (Banister, 1984)
Raiamas kheeli Stiassny, Schelly & Schliewen, 2006
Alestidae
Nannopetersius mutambuei Wamuini Lunkayilakio & Vreven, 2008
Claroteidae
Parauchenoglanis balayi (Sauvage, 1879)
Clariidae
Clarias angolensis Steindachner, 1866
Clarias buthupogon Sauvage, 1879
Clarias camerunensis Lönnberg, 1895
Clarias gariepinus (Burchell, 1822)
Clarias gabonensis Günther, 1867
Schilbeidae
Schilbe zairensis de Vos, 1995
Cichlidae
Haplochromis snoeksi Wamuini Lunkayilakio & Vreven, 2010
Hemichromis elongatus (Guichenot, 1861)
Oreochromis niloticus (Linnaeus, 1758)
Sarotherodon galilaeus (Linnaeus, 1758)
Coptodon tholloni (Sauvage, 1884)
Anabantidae
Ctenopoma nigropannosum Reichenow, 1875
Chanidae
Parachanna obscura (Günther, 1861)
230 P. H. Skelton
Appendix 3
Freshwater fishes of the (A) Cuanza (Atlantic coastal), (C) southern Congo, (Z)
Upper Zambezian, (O) Okavango, and (K) Cunene basins in Angola, after Poll
(1967) with updated adjustments for taxonomy and known records by the author
Species Author & Date A C Z O K Ia

Protopteridae
Protopterus aethiopicus Heckel, 1851 x DD
Protopterus dolloi Boulenger, 1900 x LC
Polypteridae
Polypterus ornatipinnis Boulenger, 1902 x LC
Clupeidae
Pellonula vorax Günther, 1868 x x LC
Odaxothrissa ansorgii Boulenger, 1910 x x LC
Kneriidae
Kneria angolensis Steindachner, 1866 x x ? LC
Kneria maydelli Ladiges & Voelker, 1961 x LC
Kneria polli Trewavas, 1936 x x LC
Kneria sjolandersi Poll, 1967 x DD
Kneria ansorgii (Boulenger, 1910) x x DD
Parakneria marmorata (Norman, 1923) x DD
Parakneria vilhenae Poll, 1965 x DD
Parakneria fortuita Penrith, 1973 x x x DD
Mormyridae
Mormyrops attenuatus Boulenger, 1898 x LC
Mormyrops anguilloides (Linnaeus, 1758) x LC
Petrocephalus okavagoensis Kramer et al., 2012 x x NE
Petrocephalus magnitrunci Kramer et al., 2012 x NE
Petrocephalus magnoculis Kramer et al., 2012 x NE
Petrocephalus longicapitis Kramer et al., 2012 x x NE
Petrocephalus christyi Boulenger, 1920 x NE
Petrocephalus cunganus Boulenger, 1910 x DD
Petrocephalus micropthalmus Pellegrin, 1909 x LC
Petrocephalus simus Sauvage, 1879 x x ? LC
Hippopotamyrus ansorgii (Boulenger, 1905) x x x LC
Hippopotamyrus Kramer & Swartz, 2010 x NE
longilateralis
Pollimyrus brevis (Boulenger, 1913) x LC
Pollimyrus castelnaui (Boulenger, 1911) x x x LC
Pollimyrus cuandoensis Kramer, van der Bank & x NE
Wink, 2013
Pollimyrus marianne Kramer et al., 2003 x NE
Cyphomyrus cubangoensis (Pellegrin, 1936) x x NE
(continued)

Cyphomyrus psittacus (Boulenger, 1897) x LC
Paramormyrops jacksoni (Poll, 1967) x DD
Marcusenius altisambesi Kramer et al., 2007 x x x LC
Hippopotamyrus pappenheimi (Boulenger, 1910) x LC
Heteromormyrus (Steindachner, 1866) x DD
pauciradiatus
Pollimyrus tumifrons (Boulenger, 1902) x NE
Marcusenius desertus Kramer, vanderBank & x NE
Wink, 2016
Marcusenius multisquamatus Kramer & Wink, 2013 x NE
Marcusenius angolensis (Boulenger, 1905) x x x x x LC
Marcusenius cuangoanus (Poll, 1967) x VU
Marcusenius dundoensis (Poll, 1967) x DD
Marcusenius moorii (Günther, 1867) x LC
Marcusenius stanleyanus (Boulenger, 1897) x LC
Campylomormyrus alces (Boulenger, 1920) x LC
Campylomormyrus cassaicus (Poll, 1967) x DD
Campylomormyrus elephas (Boulenger, 1898) x LC
Campylomormyrus numenius (Boulenger, 1898) x LC
Campylomormyrus (David & Poll, 1937) x DD
luapulaensis
Campylomormyrus (Boulenger, 1898) x LC
rhynchophorus
Campylomormyrus tshokwe (Poll, 1967) x LC
Gnathonemus barbatus Poll, 1967 x DD
Gnathonemus petersii (Günther, 1862) x LC
Mormyrus caballus Boulenger, 1898 x NE
Mormyrus lacerda Castelnau, 1861 x x x x LC
Mormyrus rume Valenciennes, 1847 x NE
Cyprinidae
Garra dembeensis (Rüppell, 1835) x LC
Clypeobarbus bellcrossi (Jubb, 1965) x x DD
Coptostomabarbus wittei David & Poll, 1937 x x LC
Enteromius afrovernayi (Nichols & Boulton, 1927) x x x x LC
Enteromius amphigramma (Boulenger, 1903) x
Enteromius ansorgii (Boulenger, 1904) x LC
Enteromius argenteus (Günther, 1868) x x LC
Enteromius barotseensis (Pellegrin, 1920) x x x LC
Enteromius barnardi (Jubb, 1965) x x x LC
Enteromius bifrenatus (Fowler, 1935) x x x LC
Enteromius breviceps (Trewavas, 1936) x x x LC
Enteromius brevidorsalis (Boulenger, 1915) x x x x x LC
Enteromius brevilateralis (Poll, 1967) x x DD
Enteromius caudosignatus (Poll, 1967) x DD
(continued)
232 P. H. Skelton

Enteromius chicapaensis (Poll, 1967) x x LC
Enteromius chiumbeensis (Pellegrin, 1936) x x LC
Enteromius dorsolineatus (Trewavas, 1936) x x LC
Enteromius eutaenia (Boulenger, 1904) x x x x x DD
Enteromius evansi (Fowler, 1930) x x LC
Enteromius fasciolatus (Günther, 1868) x x x x x LC
Enteromius greenwoodi (Poll, 1967) x x DD
Enteromius haasianus (David, 1936) x x x x LC
Enteromius holotaenia (Boulenger, 1904) x x LC
Enteromius kerstenii (Peters, 1868) x x x LC
Enteromius kessleri (Steindachneri, 1866) x x x x LC
Enteromius lineomaculatus (Boulenger, 1903) x x x x LC
Enteromius lujae (Boulenger, 1913) x x x x x DD
Enteromius machadoi (Poll, 1967) x DD
Enteromius mattozi (Guimarães, 1884) x x x LC
Enteromius mediosquamatus (Poll, 1967) x DD
Enteromius miolepis (Boulenger, 1902) x x x LC
Enteromius mocoensis (Trewavas, 1936) x x DD
Enteromius multilineatus (Worthington, 1933) x x x x LC
Enteromius musumbi (Boulenger, 1910) x LC
Enteromius paludinosus (Peters, 1852) x x x x x LC
Enteromius petchkovski (Poll, 1967) x DD
Enteromius poechii (Steindachneri, 1911) ? x x x LC
Enteromius radiatus (Peters, 1853) x x x x x LC
Enteromius rousellei (Ladiges & Voelker, 1961) x DD
Enteromius thamalakanensis (Fowler, 1935) x x x LC
Enteromius trimaculatus (Peters, 1852) x x x x LC
Enteromius unitaeniatus (Günther, 1867) x x x x x LC
Enteromius cf viviparus (Weber, 1897) x x x NE
Enteromius wellmani (Boulenger, 1911) x DD
Labeobarbus caudovittatus (Boulenger, 1902) x LC
Labeobarbus codringtonii (Boulenger, 1908) x x x LC
Labeobarbus ensis (Boulenger, 1910) x LC
Labeobarbus gulielmi (Boulenger, 1910) x DD
Labeobarbus girardi (Boulenger, 1910) x DD
Labeobarbus jubbi (Poll, 1967) x DD
Labeobarbus lucius (Boulenger, 1910) x DD
Labeobarbus marequensis (Smith, 1841) x LC
(Cassai)
Labeobarbus nanningsi de Beaufort, 1933 x x DD
Labeobarbus rhinophorus (Boulenger, 1910) x DD
Labeobarbus rocadasi (Boulenger, 1910) x DD
Labeobarbus rosae (Boulenger, 1910) x DD
(continued)

Labeobarbus ansorgii (Boulenger, 1906) x LC
Labeobarbus ensifer (Boulenger, 1910) x LC
Labeobarbus boulengeri Vreven et al., 2016 x NE
Labeobarbus macrolepidotus (Pellegrin, 1928) x LC
Labeobarbus steindachneri (Boulenger, 1910) x LC
Labeobarbus stenostomata (Boulenger, 1910) x DD
Labeobarbus varicostoma (Boulenger, 1910) x DD
Labeo annectens Boulenger, 1903 x x LC
Labeo ansorgii Boulenger, 1907 x x x LC
Labeo chariensis Pellegrin, 1904 x LC
Labeo cylindricus Peters, 1852 x x x LC
Labeo greeni Boulenger, 1902 x ? LC
Labeo lineatus Boulenger, 1898 x LC
Labeo longipinnis Boulenger, 1898 x LC
Labeo macrostoma Boulenger, 1898 x LC
Labeo parvus Boulenger, 1902 x x LC
Labeo rocadasi Boulenger, 1907 x LC
Labeo ruddi Boulenger, 1907 x LC
Labeo velifer Boulenger, 1898 x NE
Labeo weeksii Boulenger, 1909 x LC
Engraulicypris howesi Ridden, Bills & Villet, 2016 x NE
Opsaridium zambezense (Peters, 1852) x x x LC
Raiamas ansorgii (Boulenger, 1910) x DD
Raiamas christyi (Boulenger, 1920) x LC
Hepsetidae
Hepsetus cuvieri (Castelnau, 1861) x x x x x NE
Alestidae
Bryconaethiops microstoma Günther, 1873 x LC
Alestes macropthalmus Günther, 1867 x LC
Brycinus kingsleyae (Günther, 1896) x LC
Brycinus grandisquamis (Boulenger, 1899) x LC
Brycinus humilis (Boulenger, 1905) x x DD
Brycinus imberi (Peters, 1852) ? x LC
Brycinus lateralis (Boulenger, 1900) x x x x LC
Micralestes acutidens (Peters, 1852) x x x LC
Micralestes argyrotaenia Trewavas, 1936 x LC
Micralestes humilis Boulenger, 1899 x LC
Nannopetersius ansorgii (Boulenger, 1910) x LC
Rhabdalestes maunensis (Fowler, 1935) x x x LC
Hydrocynus vittatus Castelnau, 1861 x x x LC
Distichodontidae
Distichodus fasciolatus Boulenger, 1898 x LC
Distichodus lusosso Schilthuis, 1891 x LC
Distichodus maculatus Boulenger, 1898 x LC
(continued)
234 P. H. Skelton

Distichodus notospilus Günther, 1867 x LC
Distichodus sexfasciatus Boulenger, 1897 x LC
Nannocharax macropterus Pellegrin, 1926 x x x LC
Nannocharax procatopus Boulenger, 1920 x LC
Nannocharax angolensis (Poll, 1967) x LC
Nannocharax lineostriatus (Poll, 1967) x x x DD
Nannocharax machadoi (Poll, 1967) x x x LC
Nannocharax multifasciatus Boulenger, 1923 x x x DD
Dundocharax bidentatus Poll, 1967 x DD
Claroteidae
Chrysichthys ansorgii Boulenger, 1910 x LC
Chrysichthys bocagii Boulenger, 1910 x LC
Chrysichthys cranchii (Leach, 1818) x LC
Chrysichthys delhezi Boulenger, 1899 x LC
Chrysichthys macropterus Boulenger, 1920 x DD
Chrysichthys nigrodigitatus (Lacepède, 1803) x LC
Parauchenoglanis ngamensis (Boulenger, 1911) x x x LC
Amphiliidae
Zaireichthys dorae (Poll, 1967) x DD
Zaireichthys flavomaculatus (Pellegrin, 1926) x DD
Zaireichthys pallidus Eccles, Tweddle & Skelton, x x NE
2011
Zaireichthys conspicuus Eccles, Tweddle & Skelton, x x NE
2011
Zaireichthys kavangoensis Eccles, Tweddle & Skelton, x NE
2011
Zaireichthys kunenensis Eccles, Tweddle & Skelton, x NE
2011
Congoglanis alula (Nichols & Griscom, 1917) x LC
Doumea angolensis Boulenger, 1906 x LC
Congoglanis howesi Vari, Ferraris & Skelton, x NE
2012
Congoglanis sp. x NE
Amphilius lentiginosus Trewavas, 1936 x ? DD
Amphilius cubangoensis Pellegrin, 1936 x x NE
Phractura macrura Poll, 1967 x DD
Phractura scaphyrhynchura (Vaillant, 1886) x LC
Malapteruridae
Malapterurus monsembeensis Roberts, 2000 x LC
Clariidae
Heterobranchus longifilis Valenciennes, 1840 x LC
Channallabes apus (Günther, 1873) x x LC
Clarias angolensis Steindachner, 1866 x x LC
Clarias buthupogon Sauvage, 1879 x LC
(continued)

Clarias dumerilii Steindachner, 1866 x x x LC
Clarias platycephalus Boulenger, 1902 x NE
Clarias gariepinus (Burchell, 1822) x x x x x LC
Clarias ngamensis Castelnau, 1861 x x x x x LC
Clarias nigromarmoratus Poll, 1967 x LC
Clarias stappersii Boulenger, 1915 x x x x x LC
Clarias liocephalus Boulenger, 1898 x x x x LC
Clarias theodorae Weber, 1897 x x x x LC
Clariallabes heterocephalus Poll, 1967 x LC
Clariallabes variabilis Pellegrin, 1926 x LC
Clariallabes platyprosopos Jubb, 1965 x x LC
Clariallabes sp x NE
Platyclarias machadoi Poll, 1977 x DD
Schilbeidae
Parailia occidentalis (Pellegrin, 1901) x LC
Schilbe intermedium Rüppell, 1832 x x x x LC
Schilbe angolensis (De Vos, 1984) x DD
Schilbe ansorgii (Boulenger, 1910) x LC
Schilbe bocagii (Guimarães, 1884) x LC
Schilbe grenfelli (Boulenger, 1900) x LC
Schilbe yangambianus (Poll, 1954) x x LC
Mochokidae
Synodontis laessoei Norman, 1923 x DD
Synodontis leopardinus Pellegrin, 1914 x x x LC
Synodontis longirostris Boulenger, 1902 x LC
Synodontis macrostigma Boulenger, 1911 x x x LC
Synodontis macrostoma Skelton & White, 1990 x x x LC
Synodontis nigromaculatus Boulenger, 1905 x x x x LC
Synodontis ornatipinnis Boulenger, 1899 x x LC
Synodontis thamalakanensis Fowler, 1935 x x x LC
Synodontis woosnami Boulenger, 1911 x x x LC
Synodontis vanderwaali Skelton & White, 1990 x x x LC
Chiloglanis angolensis Poll, 1967 x x DD
Chiloglanis fasciatus Pellegrin, 1936 x x LC
Chiloglanis lukugae Poll, 1944 x LC
Chiloglanis micropogon Poll, 1952 x NE
Chiloglanis sardinhai Ladiges & Voelker, 1961 x LC
Euchilichthys astatodon (Pellegrin, 1928) x LC
Euchilichthys royauxi Boulenger, 1902 x LC
Atopochilus macrocephalus Boulenger, 1906 x DD
Chiloglanis sp. (dark) x x NE
Chiloglanis sp. (gold) x x NE
(continued)
236 P. H. Skelton

Procatopodidae
Aplocheilichthys spilauchen (Duméril, 1861) x LC
Micropanchax hutereaui (Boulenger, 1913) x x x LC
Micropanchax johnstonii (Günther, 1894) x x x x LC
Micropanchax katangae (Boulenger, 1912) x x x x LC
Micropanchax luluae (Fowler, 1930) x x NE
Micropanchax macrurus (Boulenger, 1904) x x x LC
Micropanchax mediolateralis (Poll, 1967) x x LC
Micropanchax myaposae (Boulenger, 1908) x LC
Micropanchax nigrolateralis (Poll, 1967) x x DD
Micropanchax 'pigmy' x x NE
Cichlidae
Hemichromis elongatus (Guichenot, 1861) x x x x LC
Hemichromis angolensis Steindachner, 1865 x NE
Pharyngochromis acuticeps (Steindachner, 1866) x x x x LC
Pseudocrenilabrus philander (Weber, 1897) x x x x x LC
Oreochromis andersonii (Castelnau, 1861) x x x VU
Oreochromis macrochir (Boulenger, 1912) x x x x VU
Oreochromis angolensis (Trewavas, 1973) x LC
Coptodon rendalli (Boulenger, 1897) x x x x x LC
Pelmatolapia cabrae (Boulenger, 1899) x x LC
Oreochromis lepidurus (Boulenger, 1899) x x EN
Oreochromis schwebischi (Sauvage, 1884) x x LC
Tilapia sparrmanii Smith, 1840 x x x x x LC
Tilapia ruweti (Poll & Thys van den x x x LC
Audenaerde, 1965)
Serranochromis altus Winemiller & Kelso- x x LC
Winemiller, 1991
Serranochromis angusticeps (Boulenger, 1907) x x x x LC
Serranochromis longimanus (Boulenger, 1911) x x LC
Serranochromis (Boulenger, 1899) x x x x x LC
macrocephalus
Serranochromis robustus (Boulenger, 1864) x x x x LC
jallae
Serranochromis thumbergi (Castelnau, 1861) ? x x x LC
Sargochromis greenwoodi (Bell-Cross, 1975) x x LC
Sargochromis carlottae (Boulenger, 1905) x x LC
Sargochromis giardi (Pellegrin, 1903) x x x LC
Sargochromis coulteri (Bell-Cross, 1975) x LC
Sargochromis codringtonii (Boulenger, 1908) x x x LC
Thoracochromis lucullae (Boulenger, 1913) x LC
Orthochromis machadoi (Poll, 1967) x LC
Sargochromis thysi (Poll, 1967) x DD
Chetia welwitschi (Boulenger, 1898) x x DD
(continued)

Chetia gracilis (Greenwood, 1984) x LC
Thoracochromis albolabrus (Trewavas & Thys vd x LC
Audenaerde, 1969)
Thoracochromis buysi (Penrith, 1970) x LC
Anabantidae
Ctenopoma machadoi (Fowler, 1930) x LC
Ctenopoma multispine Peters, 1844 x x x x LC
Microctenopoma intermedium (Pellegrin, 1920) x x x LC
Microctenopoma sp. x x NE
Mastacembelidae
Mastacembelus ansorgii Boulenger, 1905 x DD
Mastacembelus niger Sauvage, 1879 x LC
Mastacembelus congicus Boulenger, 1896 x LC
Mastacembelus frenatus Boulenger, 1901 x x LC
Mastacembelus sp. x NE
Eleotridae
Eleotris vittata Duméril, 1861 x LC
Dormitator lebretonis (Steindachner, 1870) x NE
Gobiidae
Awaous lateristriga (Duméril, 1861) x NE
Nematogobius maindroni (Sauvage, 1880) x NE
Ctenogobius lepturus (Pfaff, 1933) x NE
Periophthalmus barbarus (Linnaeus, 1766) x LC
Syngnathidae
Enneacampus ansorgii (Boulenger, 1910) x LC
Enneacampus kaupi (Bleeker, 1863) x LC
TOTALS 104 161 93 103 82
IUCN status (I) as recorded by Darwell et al. (2011) and IUCN (2018). The table is for tentative
indications of distribution and IUCN status
DD data deficient, EN endangered, LC least concern, NE not evaluated, VU vulnerable
a
IUCN Red List Categories Codes
References
Ball P (2015) Benguela – more than just a current. The Heritage Portal, p 13. http://www.the-
heritageportal.co.za/article/Benguela-more-just-current
Balon EK (1974) Fishes from the edge of Victoria Falls, Africa: demise of a physical barrier for
downstream invasions. Copeia 1974(3):643–660
Barnard KH (1948) Report on a collection of fishes from the Okavango River, with notes on
Zambesi fishes. Ann S Afr Mus 36:407–458
Bbole I, Katongo C, Deines AM et al (2014) Hybridization between non-indigenous Oreochromis
niloticus and native Oreochromis species in the lower Kafue River and its potential impacts on
fishery. J Ecol Nat Environ 6(6):215–225
238 P. H. Skelton
Bell-Cross G (1965) Movement of fish across the Congo-Zambezi watershed in the Mwinilunga
district of Northern Rhodesia. Proceedings of the Central African Scientific and Medical
Congress, Lusaka, 1963, pp 415–424
Bell-Cross G (1975) A revision of certain Haplochromis species (Pisces: Cichlidae) of Central
Africa. Occas Pap Natl Mus Monuments Rhod Ser B 5(7):405–464
Bell-Cross G, Minshull JL (1988) The fishes of Zimbabwe. National Museums and Monuments
of Zimbabwe, Harare
Bills IR, Skelton PH, Almeida F (2012) A survey of the fishes of the upper Okavango system in
Angola. SAIAB Investigational Report 73, 61 pp
Bills IR, Mazungula N, Almeida F (2013) A survey of the fishes of upper Okavango River system
in Angola. SAIAB Investigational Report 74, 21 pp
Boulenger GA (1909–1916) Catalogue of the fresh-water of Africa in the British Museum (Natural
History), Vol 1 (1909) Vol 2 (1910), Vol 3 (1915), Vol 4 (1916). Trustees of the British Museum,
London
Boulenger GA (1910) LXI.–on a large collection of fishes made by Dr. W. J. Ansorge in the Quanza
and Bengo Rivers, Angola. Ann Mag Nat Hist 6(36):537–561
Boulenger GA (1911) V. on a collection of fishes from the Lake Ngami Basin, Bechuanaland.
Trans Zool Soc London 18(5):399–418, pls XXXVIII-XLIII
Burrough SL, Thomas DSG, Bailey RM (2009) Mega-lake in the Kalahari: a late Pleistocene
record of the Palaeolake Magadigadi system. Quat Sci Rev 28:1392–1411
Castelnau F (1861) Mémoire sur les Poissons de l’Afrique Australe. J-B Baillière et Fils, Paris, p
78
Day JJ, Bills R, Friel JP (2009) Lacustrine radiation in African Synodontis catfish. J Evol Biol
22:805–817
De Vos LDG (1995) A systematic revision of the African Schilbeidae (Teleostei, Siluriformes).
With an annotated bibliography. Annalen Zoologische Wetenschappen 271:1–450
Devaere S, Adriaens D, Verraes W (2007) Channallabes sanghaensis sp.n. a new anguilliform cat-
fish from the Congo River basin, with some comments on other anguilliform clariids (Teleostei,
Siluriformes). Belg J Zool 137:17–26
Eccles DH, Tweddle D, Skelton PH (2011) Eight new species in the dwarf catfish genus Zaireichthys
(Siluriformes: Amphiliidae). Smithiana Bull 13:3–28
Flügel TJ, Eckardt FD, Cotterill FPD (2015) Chapter 15: the present day drainage patterns of
the Congo river system and their Neogene evolution. In: de Wit MJ et al (eds) Geology and
resource potential of the Congo basin, Regional geology reviews. Springer, Berlin/Heidelberg,
pp 315–337
Fowler HW (1930) The fresh-water fishes obtained by the gray African expedition – 1929. With
notes on other species in the academy collection. Proc Acad Natl Sci Phila 82:27–83
Fowler HW (1935) Scientific results of the Vernay-Lang Kalahari Expedition, March to September,
1930. The freshwater fishes. Ann Transv Mus 16(2):251–293
Gilchrist JDF, Thompson WW (1913) The freshwater fishes of South Africa. Ann S Afr Mus
11(5):321–463
Gilchrist JDF, Thompson WW (1917) The freshwater fishes of South Africa (continued). Ann S
Afr Mus 11(6):465–575
Greenwood PH (1984) The haplochromine species (Teleostei, Cichlidae) of the Cunene and cer-
tain other Angolan rivers. Bull Brit Mus (Nat Hist) 47(4):187–239
Greenwood PH (1993) A review of the serranochromine cichlid fish genera Pharyngochromis,
Sargochromis, Serranochromis and Chetia (Teleostei, Labroidei). Bull Brit Mus (Nat Hist)
59:33–44
Guimarāes ARP (1884) 1. Diagnoses de trois nouveaux poisons d’Angola. J Sci Math Phys Lisboa
37:1–10
Haddon IG, McCarthy TS (2005) The Mesozoic–Cenozoic interior sag basins of Central Africa:
the late-cretaceous–Cenozoic Kalahari and Okavango basins. J Afr Earth Sci 43:316–333
Hay CJ, van Zyl BJ, van der Bank FH et al (1997) A survey of the fishes of the Kunene River,
Namibia. Modoqua 19:129–141
Hipondoka MHT, van der Waal BCW, Ndeutapo MH, Hango L (2018) Sources of fish in the
ephemeral western iishana region of the Cuvelai–Etosha Basin in Angola and Namibia. Afr J
Aquat Sci 43(3):199–214.https://doi.org/10.2989/16085914.2018.1506310
Jackson PBN (1961) The fishes of northern Rhodesia: a checklist of indigenous species. Department
of Game and Fisheries, Lusaka
Jubb RA (1961) An illustrated guide to the freshwater fishes of the Zambezi River, Lake Kariba,
Pungwe, Sabi, Lundi and Limpopo Rivers. Stuart Manning, Bulawayo
Jubb RA (1967) The freshwater fishes of southern Africa. AA Balkema, Cape Town
Jubb RA, Gaigher IG (1971) Checklist of the fishes of Botswana. Arnoldia, Rhodesia 5(97):1–22
Key RM, Cotterill FPD, Moore AE (2015) The Zambezi river: an archive of tectonic events linked
to the amalgamation and disruption of Gondwana and subsequent evolution of the African
plate. S Afr J Geol 118:425–438
King J, Chonguic E (2016) Integrated management of the Cubango-Okavango River basin.
Ecohydrol Hydrobiol 16:263–271
Kramer B, Swartz ER (2010) A new species of slender Stonebasher within the Hippopotamyrus
ansorgii complex from the Cunene River in southern Africa (Teleostei: Mormyriformes). J Nat
Hist 44(35–36):2213–2242
Kramer B, Wink M (2013) East–west differentiation in the Marcusenius macrolepidotus spe-
cies complex in southern Africa: the description of a new species for the lower Cunene River,
Namibia (Teleostei: Mormyridae). J Nat Hist 47(35–36):2327–2362
Kramer B, van der Bank FH, Flint N et al (2003) Evidence for parapatric speciation in the
Mormyrid fish, Pollimyrus castelnaui (Boulenger, 1911), from the Okavango–upper Zambezi
River systems: P. marianne sp. nov., defined by electric organ discharges, morphology and
genetics. Environ Biol Fish 77:47–70
Kramer B, van der Bank FH, Wink M (2004) The Hippopotamyrus ansorgii species complex in
the upper Zambezi River system with a description of a new species, H. szaboi (Mormyridae).
Zool Scr 33:1–18
Kramer B, Bills IR, Skelton PH et al (2012) A critical revision of the churchill snoutfish, genus
Petrocephalus Marcusen, 1854 (Actinopterygii: Teleostei: Mormyridae), from southern and
eastern Africa, with the recognition of Petrocephalus tanensis, and the description of five new
species. J Nat Hist 46:2179–2258
Kramer B, van der Bank H, Wink M (2014) Marked differentiation in a new species of dwarf stone-
basher, Pollimyrus cuandoensis sp. nov. (Mormyridae: Teleostei), from a contact zone with two
sibling species of the Okavango and Zambezi rivers. J Nat Hist 48(7–8):429–463
Kramer B, van der Bank FH, Wink M (2016) Marcusenius desertus sp. nov. (Teleostei:
Mormyridae), a mormyrid fish from the Namib desert. Afr J Aquat Sci 41(1):1–18
Ladiges W (1964) Beiträge zur zoogeographie und Oekologie der süßwasserfische Angolas. Die
Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 61:221–272
Ladiges W, Voelker J (1961) Untersuchungen über die Fishfauna in Gebirgsgewässern des
Wasserscheidenhochlands in Angola. Die Mitteilungen aus dem Hamburgischen Zoologischen
Museum und Institut 59:117–140
Lévêque C, Paugy D (2017a) General characteristics of ichthyological fauna. In: Paugy D, Lévêque
C, Otero O (eds.) The inland water fishes of Africa, diversity, ecology and human use. IRD
Éditions, Paris, & Royal Museum for Central Africa, Tervuren, pp 83–96
Lévêque C, Paugy D (2017b) Geographical distribution and Affinities of African freshwater fishes.
In: Paugy D, Lévêque C, Otero O (eds) The inland water fishes of Africa, diversity, ecology and
human use. IRD Éditions. France, & Royal Museum for Central Africa, Belgium, pp 97–114
Marshall BE, Tweddle D (2007) Oreochromis macrochir. The IUCN Red List of Threatened
Species 2007: e.T63336A12659168
Mendelsohn J, Weber B (2015) An atlas and profile of Moxico, Angola. RAISON, Windhoek
240 P. H. Skelton
Moelants T 2010. Oreochromis lepidurus. The IUCN Red List of Threatened Species2010:
e.T182875A7991695
Moore AE, Larkin PA (2001) Drainage evolution in south-Central Africa since the break-up of
Gondwana. S Afr J Geol 104:47–68
Moore AE, Cotterill FPD, Eckardt FD (2012) The evolution and ages of Makgadikgadi palaeo-
lakes: Consilient evidence from Kalahari drainage evolution. S Afr J Geol 115:385–413
Musilová Z, Kalous L, Petrtýl M et al (2013) Cichlid fishes in the Angolan headwaters region:
molecular evidence of the ichthyofaunal contact between the Cuanza and Okavango-Zambezi
systems. PLoS One 8(5):e65047
NGOWP – National Geographic Okavango Wilderness Project (2018) Initial findings from
exploration of the upper catchments of the Cuito, Cuanavale and Cuando Rivers in Central
Nichols JT, Boulton R (1927) Three new minnows of the genus Barbus, and a new characin from
the Vernay Angola expedition. Am Mus Novit 264:1–8
Norman JR (1923) A new cyprinoid fish from Tanganyika territory, and two new fishes from
Angola. Ann Mag Nat Hist 12(72):694–696
Nunes AL, Douthwaite RJ, Tyser B et al (2016) Invasive crayfish threaten Okavango Delta. Front
Ecol Environ 14(5):237–238
Paugy D, Lévèque C, Otero O (eds) (2017) The inland water fishes of Africa, IRD Éditions.
Institut de Recherche pour de Developpement/RMCA Royal Museum for Central Africa, Paris/
Tervuren
Pellegrin J (1921) Description d'un Barbeau nouveau de l'Angola. Bull Soc Zool Fr 46:118–120
Pellegrin J (1928) Poissons du Chiloango et du Congo receuillis par l’expédition du Dr Schouteden
(1920–1922). Annales du Musée Royal du Congo Belge, Zoologie Série 1 3(1):1–50
Pellegrin J (1936) Contribution à l’ichthyologie de l’Angola. Arquivos do Museu Bocage 7:45–62
Penrith M-L (1970) Report on a small collection of fishes from the Kunene River mouth.
Cimbebasia Series A 1:165–176
Penrith MJ (1973) A new species of Parakneria from Angola (Pisces: Kneriidae). Cimbebasia
Series A 11:131–135
Penrith MJ (1982) Additions to the checklist of southern African freshwater fishes and a gazetteer
of south-western Angolan collecting localities. J Limnol Soc South Afr 8(2):71–75
Pinton A, Agnèse J-F, Paugy D, Otero O (2013) A large-scale phylogeny of Synodontis
(Mochokidae, Siluriformes) reveals the influence of geological events on continental diversity
during the Cenozoic. Mol Phylogenet Evol 66:1027–1040
Podgorski JE, Green AG, Kgotlhang L et al (2013) Paleo-megalake and paleo-megafan in southern
Africa. Geology 11:1155–1158
Poll M (1967) Contribution à la Faune Ichthyologique de l’Angola. Publicaçōes Culturais 75 75.
Companhia dos Diamentes de Angola (DIAMANG), Lisbon, 381 pp
Poll M (1971) Révision des Synodontis Africains (Famille Mochocidae). Annales Musée Royal
de l’Afrique Centrale Serie IN-8 Sciences Zoologiques No. 191. Musée Royal de l’Afrique
Centrale, Tervuren, 497 pp
Roberts TC (1975) Geographical distribution of African freshwater fishes. Zool J Linnean Soc
57(4):249–319
Saldanha L (1978) Museu Bocage. Copeia 1978(4):739–740
Schwarzer J, Swartz ER, Vreven E et al (2012) Repeated trans-watershed hybridization among
haplochromine cichlids (Cichlidae) was triggered by Neogene landscape evolution. Proc R Soc
London, Ser B 279:4389–4398
Skelton PH (1994) Diversity and distribution of freshwater fishes in East and Southern Africa.
Annales Musée Royal de l’Afrique Centrale, Sciences Zoologiques 275:95–131
Skelton PH (2001) A complete guide to the freshwater fishes of Southern Africa. Struik, Cape
Town
Skelton PH, Swartz ER (2011) Walking the tightrope: trends in African freshwater systematic
ichthyology. J Fish Biol 79:1413–1435
Skelton PH, Bruton MN, Merron GS et al (1985) The fishes of the Okavango drainage system
in Angola, South West Africa and Botswana: taxonomy and distribution. Ichthyol. Bull. JLB
Smith Inst Ichthyol 50:1–21
Skelton PH, Neef G, Costa A (2016) Into the wilderness expedition 2015: the fishes. SAIAB
Investigational Report No 75, 49 pp
Snoeks J, Vreven EJ (2007) Chapter 38: Polynemidae, 445-449 in: Stiassny, MLJ, Teugels GG,
Hopkins CD (eds) The fresh and brackish water fishes of lower Guinea, west-Central Africa.
Collection Faune et Flore tropicales 42, vol 2. Institut de recherché pour le développement,
Paris, France/Muséum national d’histoire naturelle, Paris, France/Musée royal de l’Afrique
Centrale, Tervuren
Snoeks J, Harrison IJ, Stiassny MLJ (2011) Chapter 3: The status and distribution of freshwater
fishes. In: Darwall WRT, Smith KG, Allen DJ, Holland RA, Harrison IJ, Brooks EGE (eds)
The diversity of life in African freshwaters: under water, under threat. An analysis of the status
and distribution of freshwater species throughout mainland Africa. IUCN, Cambridge/Gland,
pp 42–73
Steindachner F (1866) Ichthyologische Mittheilungen. (IX.) [With subtitles I-VI.]. Verh Zool Bot
Ges Wien 16:761–796
Stiassny MLJ, Teugels GG, Hopkins CD (eds) (2007) The fresh and brackish water fishes of Lower
Guinea, West-Central Africa. Collection Faune et Flore Tropicales 42, Volume 1 and 2. IRD &
Muséum National d’Histoire Naturelle, Paris & Musée Royal de l’Afrique Centrale, Tervuren
Thieme ML, Abell R, Stiassny ML et al (eds) (2005) Freshwater ecoregions of Africa and
Madagascar, a conservation assessment. Island Press, Washington
Trewavas E (1936) Dr. Karl Jordan’s expedition to south-West Africa and Angola: the fresh-water
fishes. Novitates Zoologicae 40:63–74
Trewavas E (1964) A revision of the genus Serranochromis Regan (Pisces, Cichlidae). Annales
Musée Royal de l’Afrique Centrale Serie IN-8 Sciences Zoologiques No. 125, Musée Royal de
l’Afrique Centrale, Tervuren, 58 pp
Trewavas E (1973) A new species of cichlid fishes of rivers Quanza and Bengo, Angola, with a list
of the known Cichlidae of these rivers and a note on Pseudocrenilabrus natalensis fowler. Bull
Brit Mus (Nat Hist) 25(1):28–37
Tweddle D (2010) Overview of the Zambezi River system: its history, fish fauna, fisheries, and
conservation. Aquat Ecosyst Health Manage 13(3):224–240
Tweddle D, Skelton, PH, van der Waal et al (2004) Aquatic biodiversity survey “four corners”
transboundary natural resources management area. SAIAB Investigational Report No 71
202 pp
Tweddle D, Cowx IG, Peel RA et al (2015) Challenges in fisheries management in the Zambezi,
one of the great rivers of Africa. Fish Manag Ecol 22:99–111
Van der Waal BCW (1991) A survey of the fisheries in Kavango, Namibia. Modoqua 17(2):113–122
Van der Waal BCW, Skelton PH (1984) Checklist of fishes of Caprivi. Modoqua 13(4):303–321
Vreven EJ, Musschoot T, Snoeks J et al (2016) The African hexaploid Torini (Cypriniformes:
Cyprinidae): review of a tumultuous history. Zool J Linnean Soc 177(2):231–305
Wamuini Lunkayilakio S, Vreven E (2010) ‘Haplochromis’ snoeksi, a new species from the Inkisi
River basin, lower Congo (Perciformes: Cichlidae). Ichthyol Explor Freshwaters 21(3):279–287
Wamuini Lunkayilakio SW, Vreven E (2008) Nannopetersius mutambuei (Characiformes:
Alestidae), a new species from the Inkisi River basin, Democratic Republic of Congo. Ichthyol
Explor Freshwaters 19:367–376
Wamuini Lunkayilakio S, Vreven E, Vandewalle P et al (2010) Contribution à la connaissance de
l’ichtyofaune de l’Inkisi au Bas-Congo (RD du Congo). Cybium 34(1):83–91
Whitfield AK (2007) Estuary associated fish species. In: Stiassny MLJ, Teugels GG, Hopkins CD
(eds) The fresh and brackish water fishes of Lower Guinea, West-Central Africa. Collection
242 P. H. Skelton
Faune et Flore Tropicales 42, vol 1. IRD & Muséum National d’Histoire Naturelle, Paris &
Musée Royal de l’Afrique Centrale, Tervuren, pp 46-56
Wise RM, van Wilgen BW, Hill MP et al (2007) The economic impact and appropriate manage-
ment of selected invasive alien species on the African continent. Final report for GISP. CSIR
report number CSIR/RBSD/ER/2007/0044/C
Zengeya TA, Decru E, Vreven EJ (2011) Revalidation of Hepsetus cuvieri (Castelnau, 1861)
(Characiformes: Hepsetidae) from the Quanza, Zambezi and southern part of the Congo ich-
thyofaunal provinces. J Nat Hist 45:1723–1744
Zengeya TA, Robertson MP, Booth AJ et al (2013) Qualitative ecological risk assessment of the
invasive Nile tilapia, Oreochromis niloticus in a sub-tropical African river system (Limpopo
river, South Africa). Aquat Conserv Mar Freshwat Ecosyst 23:51–64
Chapter 12
The Amphibians of Angola: Early Studies
and the Current State of Knowledge
Ninda Baptista, Werner Conradie, Pedro Vaz Pinto, and William R. Branch
Abstract Angolan amphibians have been studied since the mid-nineteenth century
by explorers and scientists from all over the western world, and collections have been
deposited in around 20 museums and institutions in Europe, Northern America, and
Africa. A significant interruption of this study occurred during Angola’s liberation
struggle and civil war for nearly four decades and, as a consequence, knowledge
about the country’s biodiversity became outdated with critical gaps. Since 2009, a
new era in Angolan biodiversity studies started as expeditions scattered in southwest-
N. Baptista (*)
Instituto Superior de Ciências da Educaҫão da Huíla, Rua Sarmento Rodrigues,
Lubango, Angola
e-mail: nindabaptista@gmail.com
W. Conradie
National Geographic Okavango Wilderness Project, Wild Bird Trust, Hogsback, South Africa
School of Natural Resource Management, Nelson Mandela University, George, South Africa
e-mail: werner@bayworld.co.za
P. Vaz Pinto
Fundação Kissama, Luanda, Angola
e-mail: pedrovazpinto@gmail.com
W. R. Branch (deceased)

https://doi.org/10.1007/978-3-030-03083-4_12
244 N. Baptista et al.
ern, northeastern, southeastern, and northwestern Angola lead to exciting discoveries,

including new records for the country, descriptions of new species, range extensions
and taxonomical updates. Currently 111 amphibian species are listed for the country
(of which 21 are endemic), but this number is an underestimate and the various unre-
solved taxonomical issues challenge the study of every other aspect of this group. The
Angolan amphibian fauna remains one of the most poorly known in Africa and much
still has to be done in order to understand its diversity, evolution and conservation
needs. An overview of existing knowledge of Angolan amphibians is presented,
including an updated checklist for the country, comments on problematic groups,
endemic species, biogeography, recent findings, and priority research topics.
Keywords Angolan escarpment · Checklist · Endemism · Herpetology · Research

priorities · Taxonomy
Introduction
Amphibians are a fascinatingly diverse group that plays crucial ecological roles
(Beard et al. 2002; Davic and Welsh 2004; Regester et al. 2006) and are useful as
indicators of ecosystem health (Waddle 2006), thus the relevance of their study
surpasses herpetological curiosity. Despite the fact that the rate of description of
amphibian species in the world is continuously increasing, current taxonomic
research is still insufficient to properly inform conservation planning (Köhler et al.
2005; Brito 2010).
Like other groups presented in this book, Angolan amphibians are among the most
poorly known in Africa (Conradie et al. 2016). To study this group it is necessary to
deal with historical as well as scientific issues including: many species are known from
holotypes collected more than a century ago and which may have been subsequently
lost; collection localities had old colonial names, some no longer used and others con-
fused with homonyms; a considerable amount of early literature is written in diverse
languages (Portuguese, French, German, English and even Latin) and is not easily
accessible; and many names used for Angolan taxa have been lost in synonymies and
their current status remains problematic. Overviews of the history and evolution of the
southern African amphibian taxonomy exist, mentioning Angolan taxa briefly
(Poynton 1964; Channing 1999; Du Preez and Carruthers 2009, 2017). This chapter
focuses on Angola, and the compiled information is intended to serve as a baseline that
facilitates the study of this group. It consists of an essentially chronological summary
of the studies of Angolan amphibians since the very first to the most recent findings,
presents a checklist of species, and identifies some of the most evident challenges and
exciting research priorities. Given the complicated status of many names available for
Angolan taxa, species considered as valid in this review follow Frost (2018). An Atlas
of historical and bibliographic records of Angolan herpetofauna has been released
subsequent to the compilation of information for this chapter (Marques et al. 2018).
12 The Amphibians of Angola: Early Studies and the Current State of Knowledge 245
Early Beginnings
The European exploration and settlement in Africa resulted in the discovery of

strange and wondrous animals. As these were sent in increasing numbers to
European centres of learning and study, they stimulated the departure of expeditions
to explore the Angolan flora and fauna by Portugal and by other nations. The exotic
collections obtained by these explorers were then shipped to their home countries,
and so, in the nineteenth century, the study of amphibians from Angola started in
Europe. This was the case for the rest of southern Africa, the only exception being
South Africa, which in the early 1800s already had Andrew Smith, a British explorer
and researcher, based in the Cape (Channing 1999; Branch and Bauer 2005).
In 1866, José Vicente Barbosa du Bocage made the first list of amphibians and
reptiles from Angola based on assorted specimens deposited in the Natural History
Museum of Lisbon (Bocage 1866a, b). It documented only 19 amphibian species,
eight of which were new to science and which Bocage (1866b) described (Hyperolius
cinnamomeoventris, H. tristis, H. fuscigula, H. quinquevittatus, H. steindachneri,
Rana (=Ptychadena) subpunctata, Rana (=Amietia) angolensis, Bufo funereus
(=Sclerophrys funerea). The material came from two expeditions, one by José de
Anchieta in 1864 to Cabinda, and the other from Duque de Bragança (now
Calandula) by Pinheiro Bayão.
During this period, Europeans were exploring Angola, either on their own initia-
tive, or on behalf of various institutions that promoted scientific expeditions to
Angola. Publications from this era consist essentially of descriptions of new species
and new distribution records for known species. The renowned Austrian explorer
and botanist Friedrich Martin Josef Welwitsch (1806–1872) explored Angola for the
Portuguese government, arriving in 1853 and undertaking almost a decade of stren-
uous exploring and collecting. After his return to Europe his collections were
donated to the British Museum, later shared with Portugal, and the Angolan amphib-
ians were reported on by Günther (1865), who described new species of reed frog
(Hyperolius nasutus, H. parallelus).
Collections from the Austrian frigate Novara were deposited in the Natural
History Museum of Vienna and studied by Steindachner (1867), who described
Ptychadena porosissima and Hyperolius bocagei from no precise locality. Anchieta
persisted in his extensive exploration of Angola, and Bocage (1867, 1873, 1879a, b,
1882, 1893, 1897b) examined his specimens, as well as the herpetological collec-
tions of Capello & Ivens (Bocage 1879a, b), describing Hylambates (=Leptopelis)
anchietae, Hylambates (=Leptopelis) cynnamomeus, and Rappia (=Hyperolius)
benguellensis among other species currently not valid. The German explorers von
Homeyer, who collected in Pungo-Andongo, and von Mechow, who collected in
Malanje and Cuango, had their specimens deposited in the Zoological Museum of
Berlin and studied by Peters (1877, 1882), who described Bufo buchneri from
Cabinda. Boulenger (1882) studied the material from the British Museum and
described Tomopterna tuberculosa, and Rochebrune (1885) described four new
Hyperolius species from Cabinda (H. lucani, H. maestus, H. protchei, H.
rhizophilus).
Bocage (1895a) compiled the extant information about the herpetology of Angola
and Congo, using all the above-mentioned references, except for Rochebrune’s
(1885). A total of 40 species of amphibians were listed for Angola. Even today,
more than a century after its release, this work is still a valuable reference on
Angolan herpetology. After this, Bocage published several other findings (Bocage
1895b, 1896a, b, 1897a, b), mostly from Anchieta’s new collections, with new
locality records for many frogs, and the description of a new pygmy toad, Bufo
(=Poyntonophrynus) dombensis.
From 1898 to 1906, José Júlio Bethencourt Ferreira studied Angolan material
collected by Anchieta, Francisco Newton and Pereira do Nascimento (Ferreira
1897, 1900, 1904, 1906), and described new species (Rappia (=Afrixalus) osorioi,
Arthroleptis carquejai, Rappia (=Hyperolius) nobrei) and some species and variet-
ies that have been subsequently synonymised.
From 1903 to 1905, William John Ansorge collected considerable material in
northern, central and southwestern Angola. The collected amphibians are deposited
in the British Museum, and were studied by Boulenger (1905, 1907a, b). Arthroleptis
(=Phrynobatrachus) parvulus, Arthroleptis xenochirus, Rana (=Ptychadena) ans-
orgii, Rana (=Tomopterna) cryptotis, and Rana (=Ptychadena) bunoderma were all
described from this material.
A number of expeditions in Angola included herpetological surveys, and had
their reptiles studied, but the amphibians were not reported. Examples of this are the
Rohan-Chabot Mission (1912–1914), which explored the south of Angola and had
its specimens deposited in the Paris Natural History Museum, and the Vernay
Angola Expedition (1925), from which the large collection is housed in the American
Museum of Natural History.
Analysing material from the Berlin Zoological Museum, Ahl (1925) described
Hylarthroleptis (=Phrynobatrachus) brevipalmatus from Angola, and several spe-
cies of reed frogs, two of which are endemic to Angola (Hyperolius bicolor,
Hyperolius gularis) and others which have later been synonymised into larger spe-
cies complexes such as Hyperolius parallelus complex (H. angolensis, H. huillen-
sis, H. microstictus), Hyperolius marmoratus complex (H. decoratus, H.
marungaensis), and Hyperolius platyceps complex (H. angolanus).
In 1930–1931, the Pulitzer-Angola Expedition surveyed southwestern and cen-
tral areas of the country. Over 400 specimens of amphibians were collected and
deposited in the Carnegie Museum, in the United States of America. These were
studied by Karl Patterson Schmidt (1936), who reported on 17 species. Although no
new species were described, some were synonymised and others revived from syn-
onymy leading the author to highlight the importance of understanding the Angolan
fauna for clarifying African amphibian taxonomy.
During two trips to central and southern Angola (1928–1929 and 1932–1933)
Albert Monard made important collections of amphibians and reptiles, as well as
other groups. The herpetological material was deposited in the La Chaux-de-Fonds
Museum, Switzerland. Monard (1937) provided an updated compilation of Angolan
amphibians with a revision of the existing literature (including Ahl, Bocage,
Boulenger and Schmidt’s publications), as well as his own findings. Five new spe-
cies of frog were described: Hyperolius cinereus, Cassiniopsis (=Kassina) kuvan-
gensis, Rana (=Ptychadena) keilingi, Hyperolius erythromelanus, Rana

(=Ptychadena) buneli, the last two now considered synonyms of H. paralleus and
Ptychadena bunoderma, respectively. In total, 80 species of amphibians were men-
tioned, meaning that in the four decades since Bocage’s (1895a) first synthesis the
known frog species for Angola had doubled.
In 1933–1934, Karl Jordan’s expedition to South West Africa (now Namibia) and
Angola surveyed localities on the Angolan escarpment (Congulo and Quirimbo)
and afromontane forest (Mount Moco) (Jordan 1936). This material is deposited in
the British Museum and the herpetofauna studied by Parker (1936). One new spe-
cies of treefrog (Leptopelis jordani) and a new subspecies of white-lipped frog
(Rana (=Amnirana) albolabris acustirostris) were described from this expedition.
As the name acustirostris was preoccupied, Mertens (1938b) proposed the replace-
ment name Rana (=Amnirana) albolabris parkeriana, which was later elevated to a
full species by Perret (1977). Both these species remain known only from their type
localities, and are escarpment-endemics.
In the 1930s W Schack visited Angola and made a collection of amphibians
which were deposited in the Natur-Museum Senckenberg, Frankfurt, and studied by
Mertens (1938a), who recorded only eight species, none of which was new.
In 1952–1954, within the scope of the Hamburg Museum expeditions, GA von
Maydell made significant herpetological collections from north to south of Angola.
The reptiles were studied by Walter Hellmich (1957a), but the amphibians have
never been studied until recently (Ceríaco et al. 2014b). Hellmich made a trip to the
Angolan region of Entre-Rios, and reported on new localities for frog species
(Hellmich 1957b), also commenting on the Angolan biogeography.
From 1957 to 1959, the Portuguese Mission of Apiarian Studies of the Overseas
collected amphibians especially in central and eastern Angola (Luando and Cameia),
which were deposited in the Zoology Center of the Institute of Tropical Scientific
Research, in Lisbon. These were studied only decades later, by Clara Ruas (1996,
2002).
Raymond F Laurent worked extensively on the herpetofauna of the Congo Basin.
He studied material from Museu do Dundo, Lunda-Norte, including the extensive
collection made in southwestern Angola by the Museum Director, António Barros
Machado. During this period, he recorded several new frogs for Angola (Laurent
1950, 1954, 1964), and described four new species (Ptychadena grandisonae, P.
guibei, P. perplicata and Hyperolius vilhenai).
In 1971 and 1974, Wulf Haacke, from the then Transvaal Museum, South Africa,
made two trips to Angola to search mainly for geckos, but incidentally collecting
amphibians that were later studied by John Poynton (Poynton and Haacke 1993).
Until the 1970s, zoological expeditions surveyed mostly southwestern and cen-
tral parts of the country, which were more easily accessible than the inland plateau
and the moist forests of the north. Herpetological knowledge about the northeastern
region was greatly improved by Laurent’s studies. The most poorly studied areas of
Angola remained the northwest (the region of Zaire and Uíge provinces, and north-
ern Malange, Bengo, and Cuanza-Norte provinces), followed by the southeastern
‘lands at the end of the world’, a commonly used expression that refers to the very
remote and extensive regions of Moxico and Cuando Cubango provinces.
Recent History and Increase of Information
For almost three decades, in the period between Angola’s independence and the end of
the civil war (1975–2002), the country’s instability precluded virtually all field sur-
veys. Every amphibian publication dating from this period involved taxonomic revi-
sions based on existing literature and museum collections, e.g. Perret’s (1976) revision
of the amphibians, particularly types, deposited in the Lisbon Museum of Natural
History. This has become an extremely valuable work given the subsequent loss of
these important specimens following the fire that destroyed the museum in 1978.
A key for the identification of Angolan amphibians mainly based on literature
revision, including all the species listed for Angola at the time, was published (Cei
1977). With dichotomous keys, drawings, and insights on the Angolan amphibian
biogeography, it was intended to make Angolan amphibian identification more
accessible to the general public and particularly to students. Poynton (1964) pub-
lished a faunal study of the southern African amphibians, which referred to Angolan
material. This was later updated from 1985 to 1991, when Poynton & Broadley
published Amphibia Zambesiaca, a series of papers (Poynton and Broadley 1985a,
b, 1987, 1988, 1991) that addressed in detail all the amphibian families occurring in
the Zambezi drainage region, including many that extend into Angola. The publica-
tion of a toponymic index of the zoological collections made in Angola (Crawford-
Cabral and Mesquitela 1989) was a valuable contribution to the study of vertebrates
of the country. It provided an overview of the zoological collections performed in
Angola and studies related to these expeditions, including a section of type locali-
ties and the list of described vertebrates per locality, which lists amphibian species,
subspecies and varieties.
In 1993, Poynton & Haacke described the first new Angolan amphibian species
in decades: Bufo (= Pontynophrynus) grandisonae, based on Haacke’s expeditions
of the 1970s. In 1996, the re-examination of Monard’s collection of amphibians
from 1928, revealed an ‘enigmatic’ ranid originally identified as Aubria subsigillata
that could not be assigned to any known genus (Perret 1996), but which was later
assigned to Aubria masako (Channing 2001) following features described by Ohler
(1996). A comprehensive revision of the Angolan amphibians and mapping of each
species’ distribution based on museum and literature records was made by Ruas
(1996), providing taxonomic comments on some species, but not addressing the
Hyperoliidae family (then including the current Leptopelinae subfamily). Ruas
(2002) described in detail the contents of the amphibian collection deposited in the
Zoology Center of the Institute of Tropical Scientific Research in Lisbon, again
excluding the Hyperoliidae and Leptopelinae, which are still to be examined.
Channing (1999) discussed aspects of Angolan amphibian taxonomy within a
southern African historical perspective. Blanc and Frétey (2000) analysed the bio-
geography, species richness and endemism of the central African and Angolan
amphibians, based on the number of species per country. They highlighted the dis-
crepancy in species richness among genera in Angola, with Bufo (currently
Mertensophryne, Sclerophrys and Poyntonophrynus), Hyperolius and Ptychadena
being the most specious genera, which totalled 42 species, almost half of the species
known for the country at the time (86).
Only in 2009 did Angolan-international collaboration lead to a new era of field

surveys, initiated with an expedition to Huíla and Namibe provinces in southwest-
ern Angola. This trip, organised by Brian Huntley, can be considered as a historical
landmark for research on Angolan biodiversity. Numerous groups were surveyed
(plants, invertebrates, mammals, birds, reptiles and amphibians). A new escarpment-
endemic reed frog, Hyperolius chelaensis, was described from Serra da Chela
(Conradie et al. 2012), and the colourful ashy reed frog, Hyperolius cinereus Monard
1937 was rediscovered (Conradie et al. 2013). Later in the same year, Alan Channing
and Pedro Vaz Pinto surveyed Cangandala National Park and made a trip to
Calandula, revisiting this important type locality of several amphibian species, and
rediscovered Hyperolius steindachneri Bocage, 1866 in Angola (Channing and Vaz
Pinto Unpublished Data). The material obtained from these trips was important for
a number of taxonomic revisions. The Angolan river frog Amietia angolensis, previ-
ously thought to be widespread in Africa, was found to occur only in Angola
(Channing and Baptista 2013; Channing et al. 2016), reed frogs of the Hyperolius
nasutus complex (Channing et al. 2013) were shown to include numerous cryptic
species, with possibly four occurring in Angola, and the Hyperolius cinnamo-
meoventris complex was split into different sister clades (Schick et al. 2010).
Another Angolan international expedition, again organised by Brian Huntley in
2011, visited the unexplored Lagoa Carumbo, Angola’s second largest freshwater
lake, in Lunda Norte province. Preliminary findings revealed a complex herpeto-
fauna (Branch and Conradie 2015), with the description of the new Hyperolius ray-
mondi (Conradie et al. 2013), and the addition of two new country records: Amnirana
cf. lepus and Hyperolius pardalis.
Two books, Treefroogs of Africa (Schiøtz 1999) and Amphibians of Central and
Southern Africa (Channing 2001) address the Angolan territory, providing species
identification keys, colour photographs, and distribution maps. In 2011, a book on the
central African and Angolan amphibians was released (Frétey et al. 2011). It addressed
Angolan fauna only briefly, providing a species list (without discussion), and synthe-
sis of species and habitat/biogeographical associations. In Tadpoles of Africa
(Channing et al. 2012), the larvae of several species occurring in the country are
described, and the description of Leptopelis anchietae and Ptychadena porosissima
tadpoles are based on Angolan specimens. The popular book Frogs of Southern
Africa – A Complete Guide (Du Preez and Carruthers 2009, 2017) provides descrip-
tions of species, morphology, distribution, behaviour, and has advertisement calls
available for many species. It has been recently updated to a cell phone app. “Frogs of
Southern Africa” and has relevant information about species that also occur in Angola.
In 2012 and 2013, studies of the lower catchments of the Cubango, Cuito and
Cuando rivers in southeastern Angola were organised by the Southern Africa
Regional Environmental Program (SAREP), funded by the USAID, and included
herpetological surveys. Preliminary results have been published (Brooks 2012,
2013), as well as an annotated checklist of the herpetofauna of the region (Conradie
et al. 2016).
In 2013, a partnership between the Kimpa Vita University in Uíge, the Technical
University Dresden and Senckenberg Natural History Collections, Dresden, pro-
moted herpetological surveys in the extremely poorly known Serra do Pingano eco-
system and surrounding forest fragments in Uíge Province. Two forest species,
Trichobatrachus robustus and Xenopus andrei, typical of the Congo Basin, were
added to the country’s list (Ernst et al. 2014, 2015). Both these observations repre-
sented southern range extensions of hundreds of kilometers. Additional important
discoveries from this survey await formal publication, and will certainly increase
current knowledge of the taxonomy and biogeography of Angolan amphibians, as
well as highlight the exceptional biodiversity of northern Angola (Ernst pers. comm.).
Since 2013, a project of the California Academy of Sciences in collaboration
with the National Institute of Biodiversity and Conservation Areas (INBAC),
Angola, initiated a study of the Angolan herpetofauna, including the development of
an atlas of the Angolan amphibians and reptiles, based on literature, analysis of
museum collections from several countries, and new findings (Marques et al. 2014,
2018). The Angolan type material deposited in the Porto Museum was studied, and
the nomenclature and taxonomy of hyperoliids, Leptopelis and Arthroleptis
described by Ferreira were discussed (Ceríaco et al. 2014a). Analysis of amphibians
collected in the Capanda Dam surroundings in Malanje (Ceríaco et al. 2014a)
included a possible record of Kassina maculosa, which if confirmed would be the
first for the country. In a study of the Namibe Province herpetofauna, Tomopterna
damarensis was recorded for the first time for Angola (Ceríaco et al. 2016a; Heinicke
et al. 2017), and a new species of pygmy toad has been described from Serra da
Neve (Ceríaco et al. 2018a). A booklet on the herpetofauna of the Cangandala
National Park in Malanje (Ceríaco et al. 2016c) was also released, followed by a
scientific publication on the same subject (Ceríaco et al. 2018b). Research on the
project’s findings and surveys to additional regions in Angola are ongoing.
In 2015 the Wild Bird Trust, supported by the National Geographic Society,
organised Angolan expeditions associated with the Okavango Wilderness Project.
Herpetological surveys took place in the headwaters of the Cuito, Cuanavale,
Cubango and Cuando rivers and other river sources in the region in both wet and dry
seasons. Whilst some of these results have been published (Conradie et al. 2016),
the project is ongoing but already two new country records (Kassinula wittei and
Leptopelis cf. parvus), numerous range extensions for Angolan herpetofauna, and a
number of candidate new species of amphibians have been identified.
Within the Southern African Science Service Centre for Climate Change and
Adaptive Land Management (SASSCAL) project, research on herpetology is being
undertaken by the Instituto Superior de Ciências da Educação (ISCED)-Huíla.
Observatories have been implemented in Tundavala, Bicuar National Park, Cameia
National Park, Iona National Park, Candelela and Cusseque (Jürgens et al. 2018).
Opportunistic surveys of herpetofauna are made at all observatories (SASSCAL
ObservationNet 2017), herpetofauna monitoring has been carried out at the
Tundavala observatory since 2016 (Baptista et al. 2018), and a checklist of Bicuar
National Park herpetofauna compiled (Baptista et al. in press). Additionally, in col-
laboration with Fundação Kissama, herpetological surveys have been made at sev-
eral sites in Huíla Province, and throughout Angola, with emphasis along the
Angolan escarpment: Cuanza-Norte, Cuanza-Sul (Cumbira) and Huíla Provinces.
An Angolan herpetofauna archive is being developed at ISCED Huíla, and research
undertaken in conjunction with these projects.
International and National Resources
Given the scarcity and the difficulties in obtaining information about Angolan
amphibians, the compilation and listing of existing information sources is relevant.
Table 12.1 lists generalist on-line platforms with relevant information about amphib-
ians that include Angolan species, as well as a list of institutions known to have
significant Angolan material in their assets.
The Current State of Knowledge on Angolan Amphibians
Despite some progress made during the last decade, the Angolan herpetofauna
remains one of the most poorly known in Africa (Conradie et al. 2016). This lack of
information becomes more evident when contrasted with the comprehensive infor-
mation compilations regarding adjacent Namibia, which include updated lists of
species (Herrmann and Branch 2013) and analysis of habitat availability, species
richness and conservation (Curtis et al. 1998). For Angola, even basic information,
such as accurate species checklists for the country, is absent. Existing information
is scattered in recent and historical publications, many of which are not easily acces-
sible. The recent Atlas of Angolan herpetofauna (Marques et al. 2018) contributes
to filling this gap. Figure 12.1 shows the localities where amphibians have been
collected before and after independence. Although recent surveys have filled some
gaps, many areas remain unsurveyed. Figure 12.2 depicts some of the amphibian
diversity present in Angola.
Checklist of Angolan Amphibians
Currently only 111 species are recorded from Angola (Appendix). Marques et al.
(2018) list 117 species for the country. This discrepancy results from the use of dif-
ferent criteria for synonymies, and of a conservative approach of the present authors
not incuding unconfirmed records, which are discussed elsewhere in this chapter.
Both these totals are considered to be underestimates, given the country’s size and
habitat richness, including the southern desert, the tropical northern forests, the
unique escarpment and the extensive plateau, many areas of which remain unsur-
veyed. This becomes more evident when compared with a country of similar size
such as South Africa, whose herpetofauna is the best studied in Africa and which is
considerably drier and cooler (and therefore less suitable for amphibians) than
Angola, and yet it has 128 species (Frost 2018), and new species continue to be
discovered (Turner and Channing 2017; Minter et al. 2017).
Table 12.1 List of relevant websites with information regarding Angolan amphibians, and
collections where Angolan amphibian specimens are deposited, according to available literature
On-line platforms and mobile phone apps
Amphibian Species of the World: http://research.amnh.org/vz/herpetology/amphibia/
AmphibiaWeb http://amphibiaweb.org/
IUCN Red List http://www.iucnredlist.org/initiatives/amphibians
Frogs of Southern Africa https://play.google.com/store/apps/details?id=com.coolideas.
eproducts.safrogs
Collections where amphibians from Angola are deposited
Angola Instituto Nacional para a Biodiversidade e áreas de Conservação, Ministério
do Ambiente (INBAC/MINAMB)a
Museu do Dundo (MD)
Museu Nacional de História Natural (Luanda)a
Southern African Science Service Centre for Climate Change and Adaptive
Land Management (SASSCAL) / Instituto Superior de Ciências da Educação
da Huíla (ISCED-Huíla)a
Austria Imperial Natural History Museum (K.K. Museum) / Natural History Museum
of Vienna (NHMW)
France National Museum of Natural History (Paris) (MNHNP)
Germany Berlin Zoological Museum (ZMB – Zoologisches Museum)a
Forschungsinstitut und Naturmuseum Senckenberg (SMF)
Hamburg Museum (ZMH – Zoologisches Museum für Hamburg)
Senckenberg Natural History Collections Dresden (MTD – Museum für
Tierkunde Dresden)a
Portugal Centro de Zoologia do Instituto de Investigação Científica Tropical, Lisbon
(IICT)
Museu de História Natural na Universidade do Porto (MUP)
Museu Nacional de História Natural e da Ciência, formerly Museu Bocage,
Lisbon (MBL) – collections destroyed on the 1978 fire
South Africa Ditsong National Museum of Natural History (formerly Transvaal Museum)
(TMP), Pretoria
Port Elizabeth Museum at Bayworld (PEM)a
South African Institute for Aquatic Biodiversity (SAIAB)a, Grahamstown
Spain Estación Biológica de Doñana (EBD-CSIC), Sevilla
Switzerland Musée de la Chaux-de-Fonds (LCFM)
Museum d’histoire naturelle de la Ville de Genève (MHNG – Geneva Natural
History Museum)
United Kingdom Natural History Museum, London (NHMUK, formerly British Museum)
Natural History Museum at Tring
United States of Carnegie Museum of Natural History (CM), Pittsburgh
America California Academy of Sciences (CAS), San Franciscoa
American Museum of Natural History (AMNH), New Yorka
Academy of Natural Sciences of Philadelphia (ANSP), Philadelphia
Field Museum of Natural History (FMNH), Chicago
Museum of Comparative Zoology (MCZ), Harvard University, Cambridge,
Massachusets
National Museum of Natural History, Smithsonian Institution (NMNH),
Washington, D.C.
indicates the institutions containing specimens from recent (post-1975) surveys
a
Fig. 12.1 Map with collecting localities for amphibians. Blue circles represent surveys before
1975 (based on literature records), and yellow triangles represent surveys after 1975 (literature
records, localities from the 2009 and 2011 expeditions, SAREP and NGOWP trips to southeastern
Angola, surveys in the scope of the SASSCAL Project and Fundação Kissama work, and
Senckenberg Technical University, Dresden)
Records That Require Confirmation
A number of unconfirmed records for Angola require further investigation. These

include Leptopelis notatus (Laurent 1964), Ptychadena schillukorum (Channing
2001), and Kassina maculosa (Ceríaco et al. 2014a). Monard (1937) noted one
specimen of Aubria subsigillata from Caquindo that Perret (1996) could not confi-
dently associate with any known genus, but that Channing (2001) considered to be
A. masako. However, the latter is a closed-canopy forest species that is not expected
to occur in southern Angola. The stated locality is either in error or the specimen
deserves further investigation. Phrynobatrachus dispar was recorded from Cabinda
by Peters (1877, as Arthroleptis dispar), but this species originates from São Tomé
and Príncipe islands (Uyeda et al. 2007; Frost et al. 2018), and it is therefore likely
that the Angolan record refers to another species. Hyperolius nitidulus was also
recorded from Angola (Peters 1877), but was described from Nigeria and is cur-
rently considered to extend south only to Cameroon (Amiet 2012). Hyperolius ocel-
latus has been described both from Angola and Fernando Pó, but the type locality
was later restricted to Fernando Pó (Perret 1975) which leaves Angolan specimens
Fig. 12.2 Representative of some of the families of frogs present in Angola. 1 Congulo Forest
Tree Frog (Leptopelis jordani) from Congulo. 2 Dombe Pigmy Toad (Poyntonophrynus domben-
sis) from Meva. 3 Kuvango Kassina (Kassina kuvangensis) from Cuanavale River Source. 4 Spot-
bellied Grass Frog (Ptychadena subpunctata) from Cameia National Park. 5 Marbled Rubber Frog
(Phrynomantis annectens) from Meva. 6 Marbled Snout-Burrower (Hemisus marmoratus) from
Bicuar National Park. 7 Angolan Reed Frog (Hyperolius cf. parallelus) from Quilengues. 8 Rain
frog (Breviceps sp. nov.) from Cuando River Source. (Photo credits – N Baptista: 4,6,7; P Vaz
Pinto: 1,2,5; W Conradie: 3,8)
with no appliable name. Phrynobatrachus auritus was recorded from Cabinda by

Peters (1877) as Arthroleptis plicatus, but the validity of this synonymy for Cabinda
requires further study. A number of species recorded from Angola were presumably
misidentified as the currently known species’ range does not include Angola, includ-
ing: Phrynobatrachus minutus recorded by Ruas (1996), but which is now restricted
to Ethiopia; Hyperolius microps recorded by Bocage (1895) and Monard (1937),
now restricted to Eastern Africa; Hyperolius multifasciatus Ahl 1931 which was
included provisionally by Monard (1937), but placed in the synonymy of H. kivuen-
sis Ahl 1931, by Pickersgill (2007); and Xenopus calcaratus recorded by Peters
(1877), but now restricted to western Africa. Specimens of Ptychadena cf. aequipli-
cata, which occurs approximately 50 km from the Cabinda Enclave (Nagy et al.
2013), exist in the AMNH collection, but their identity requires confirmation (Ernst
pers. comm.).
Species Likely to Occur in Angola But Not Yet Recorded
The ranges of many species occurring in adjacent countries (Namibia, Zambia and
the Democratic Republic of the Congo, DRC) are likely to extend into Angola and
are discussed below. A significant example is that of caecilians (Order Gymnophiona),
which are known from the Congo Basin but have not been recorded in Angola,
including Cabinda. Species that have been recorded close to the Angolan border and
that are likely to occur in the country are listed below.
Caecilians (Gymnophiona)
The Gaboon Caecilian (Geotrypetes seraphini (Duméril, 1859)) and the Congo
Caecilian (Herpele squalostoma (Stutchbury, 1836)) have both been recorded from
the extreme western DRC, in Mayombe, River Minkala, Vemba-Minionzi, around
Kidima, around 40 km from the Angolan border (Scheinberg and Fong 2017), and
are likely to occur in this poorly known region.
Frogs and Toads (Anura)
Arthroleptidae
Cryptic Tree Frog (Leptopelis parbocagii Poynton and Broadley, 1987). This tree
frog occurs in northern Mwinilunga district, northwest Zambia, less than 50 km
from Cazombo, eastern Angola (Schiøtz and Van Daele 2003), and may occur on
the Angolan side of the border.
Breviciptidae
Power’s Rain Frog (Breviceps poweri Parker, 1934). This rainfrog was found in
southwestern Zambia, less than 100 km from the Angolan border (Pietersen et al.
2017), and can be expected in Angola.
Bufonidae
Beira Pygmy Toad (Poyntonophrynus beiranus (Loveridge, 1932)). Recorded from

southwestern Zambia near the Angolan border (Poynton and Broadley 1991) and
may occur in Angola.
Northern Pygmy Toad (Poyntonophrynus fenoulheti (Hewitt and Methuen, 1913)).
This pigmy toad is recorded from Caprivi Strip in northeastern Namibia
(Channing and Griffin 1993) and southwestern Zambia (Pietersen et al. 2017),
less than 100 km from the Angolan border, and its presence is expected in south-
eastern Angola.
Hemisotidae
Perret’s Snout-burrower (Hemisus perreti Laurent, 1972). Recorded in Singa

Mbamba, Mayumbe (Royal Museum for Central Africa 2017) and in the region
of Kipanzu, Tshela (MHNG 2017) both in the Bas-Congo Province, DRC, in
close proximity to the Cabinda enclave, and it is therefore expected to occur in
Cabinda.
Barotse Snout-burrower (Hemisus barotseensis Channing and Broadley 2002).
Described from the Barotse floodplain near Mongu, in southwestern Zambia,
120 km east of the Angolan border, but may occur in suitable floodplain habitat
along the eastern Zambezi drainage.
Hyperoliidae
Foulassi Spiny Reed Frog (Afrixalus paradorsalis (Perret, 1960)). This hyperoliid
was found in Luango-Nzambi, DRC, around 50 km from the Cabinda Enclave
(Nagy et al. 2013) and is likely to occur in Angola.
Rainforest Reed Frog (Hyperolius tuberculatus (Mocquard, 1897)). Also found in
Luango-Nzambi, DRC (Nagy et al. 2013) and likely to occur at least in Cabinda.
Kachalola Reed Frog (Hyperolius kachalolae Schiøtz, 1975). Known from
Mwinilunga district, in northwestern Zambia (Schiøtz and Van Daele 2003), less
than 50 km from the eastern Angolan border.
Hyperolius major Laurent, 1957. This reed frog occurs in Mwinilunga district, in
northwestern Zambia, less than 50 km of Cazombo (Poynton and Broadley 1991;
Schiøtz and Van Daele 2003), eastern Angola.
Phrynobatrachidae
Golden Puddle Frog (Phrynobatrachus auritus Boulenger, 1900). This species of

puddle frog was found in Luki, DRC, only 20 km north of Angola (Nagy et al.
2013) and may occur in the country.
Horned Puddle Frog (Phrynobatrachus sp. aff. cornutus (Boulenger, 1906)), also
found in Luki, DRC (Nagy et al. 2013) and likely to occur in Angola.
Pipidae
Gaboon Dwarf Clawed Frog (Hymenochirus sp. aff. feae Boulenger 1906), and
Xenopus (Silurana) sp. This Dwarf Clawed Frog and an unidentified species of
clawed frog were found in Luki, DRC, 20 km north of the Angolan border (Nagy
et al. 2013) and are expected in Angolan territory.
Fraser’s Clawed Frog (Xenopus cf. fraseri Boulenger, 1905). This clawed frog has
been found in Luki, DRC, 20 km north of the Angolan border (Nagy et al. 2013)
and is expected in Angola, although the records of these species are considered
to need a critical revision (Ernst et al. 2015).
Common Platanna (Xenopus laevis (Daudin, 1802)). Recorded in Luki, DRC,
20 km north of the Angolan border, and in Tsumba-Kituti (Nagy et al. 2013) and
might occur in Angola.
Ptychadenidae
Dark Grass Frog (Ptychadena obscura (Schmidt and Inger, 1959)). This species has
been recorded in the Ikelenge pedicle, northern Mwinilunga district, northwest-
ern Zambia, in close proximity to the Angolan eastern border (Poynton and
Broadley 1991).
Mapacha Grass Frog (Ptychadena cf. mapacha Channing, 1993). This Grass Frog is
described from the Caprivi Strip in Namibia, near southeastern Angola (Channing
1993). It has also been recorded about 80 km east of Rundu (Haacke 1999), near
Vicota, around 30 km south of the Angolan border (Ceríaco et al. 2016a), and in
southwestern Zambia (Pietersen et al. 2017). Conradie et al. (2016) collected a
series of Ptychadena at Jamba provisionally assigned to P. cf. mossambica, but
mentioned that the specimens might be referable to P. mapacha. All these records
suggest that this species may occur in southeastern Angola.
Perret’s Grass Frog (Ptychadena cf. perreti Guibé and Lamotte, 1958). This grass
frog was found in Nkamuna, in the Bas-Congo province of DRC, near Angola
(Nagy et al. 2013).
Pyxicephalidae
Boettger’s Dainty Frog (Cacosternum boettgeri (Boulenger, 1882)). This species

has been recorded near the Angolan border in northern Namibia in Caprivi Strip
and in Omusati province (Channing and Griffin 1993), and Southern Province in
Zambia (Broadley 1971) and may occur in Angolan territory.
Knocking Sand Frog (Tomopterna krugerensis Passmore and Carruthers, 1975).
This frog has been recorded in northern Namibia close to the Angolan border
(Channing and Griffin 1993).
Tandy’s Sand Frog (Tomopterna tandyi Channing and Bogart, 1996). Recorded
from northern Namibia near the Angolan border (Coetzer 2017), and may occur
in southwestern Angola. A recent fing of Tomopterna has been made in Bicuar
National Park and its identification as T. tandyi is under discussion (Baptista
et al. in press).
Rhacophoridae
Southern Foam Nest Frog (Chiromantis xerampelina Peters, 1854). Recorded from
Caprivi Strip in northern Namibia (Channing and Griffin 1993) and from south-
eastern Zambia (Broadley 1971; Pietersen et al. 2017), and therefore expected in
southeastern Angola. It is recorded from southwestern Angola (Schiøtz 1999),
but the original source of this record is unknown. This odd distribution record
requires confirmation.
Western Foam-nest Tree Frog (Chiromantis rufescens (Günther, 1869)). This spe-
cies is known from near Boma, close to the northern bank of the Congo River
(Royal Belgian Institute of Natural Sciences 2017) and may occur in Angola.
According to Frost (2018), based on distribution and species’ habitat affinities,
around 20 additional species are expected in the country, mostly from the northern
forests and expected in the Cabinda enclave in northern Angola. These are general-
ist assumptions that do not necessarily take into account actual proximity to the
Angolan border. These include arthroleptids Silver Long-fingered Frog
(Cardioglossa leucomystax (Boulenger, 1903)), Kala Forest Treefrog (Leptopelis
aubryioides (Andersson, 1907)), Victoria Forest Treefrog (Leptopelis boulengeri
(Werner, 1898)), Red Treefrog (Leptopelis rufus Reichenow, 1874)); bufonids [High
Tropical Forest Toad (Sclerophrys latifrons (Boulenger, 1900))], hyperoliids
[African Wart Frog (Acanthixalus spinosus (Buchholz and Peters, 1875)); Greshoff’s
Wax Frog (Cryptothylax greshoffii (Schilthuis 1889) with an unconfirmed record
from northwestern Angola (Schiøtz 1999)), Olive Striped Frog (Phlyctimantis leon-
ardi (Boulenger, 1906), ptychadenids [Savanna Grass Frog (Ptychadena supercili-
aris (Günther, 1858)], and pipids [Western Dwarf Clawed Frog (Hymenochirus
curtipes Noble, 1924), False Fraser’s Clawed Frog (Xenopus allofraseri Evans,
Carter, Greenbaum, et al., 2015)].
Hidden Among the Unknown – Angolan Tadpoles
An important and often neglected component of studying amphibians is knowledge

of their larvae. Unlike adult frogs, whose activity is quite dependent on appropriate
weather conditions, breeding season, and nocturnal activity for most species, tad-
poles can be easily found in water bodies, during the day, and throughout the year
in some cases. The study of tadpoles includes not only morphology, but also micro-
habitat requirements, ecology, behaviour, feeding habits, predator-prey interactions,
etc. Although they look similar at first glance, tadpole morphology usually allows
the identification to genus, and a more precise analysis can often lead to species
identification.
Early literature regarding southern African tadpoles often covers species occur-
ring in Angola (Van Dijk 1966, 1971). Channing et al. (2012) provide a comprehen-
sive review of the knowledge about African tadpoles with keys to the identification
of genera and detailed description of species. Given the poorly known status of
Angolan amphibians, it is not surprising that very little is known about Angolan
tadpoles. Out of the 99 Angolan species that have tadpoles (i. e., Breviceps and
Arthroleptis genera not included), the tadpoles of only 44 species have been
described, and only those of Ptychadena porosissima (Channing et al. 2012),
Amietia angolensis (Channing et al. 2016) and of the recent discoveries of the
endemic Hyperolius chelaensis (Conradie et al. 2012), H. cinereus and H. raymondi
(Conradie et al. 2013) are based on Angolan material. A recent description of
Leptopelis anchietae tadpoles is also based on Angolan material (Channing et al.
2012), but it was not encountered with adult specimens, and was based on the asso-
ciation with the first description of that tadpole (Lamotte and Perret 1961), which
was based on a specimen from Cameroon that may involve another species. A list of
the Angolan frogs with undescribed tadpoles (Table 12.2) includes some of the
more common local species.
Comments on Selected Groups
As a consequence of the current poor knowledge of Angola’s amphibians, the taxo-

nomic status of many species in the checklist remains unresolved. Some of these are
discussed in this section, as well as recent discoveries from ongoing studies.
Species Complexes and Species with Unclear Boundaries
Some morphologically similar species display variation in calls or habitat and are
considered to form a complex of closely-related species, and the resolution of their
taxonomic status and distribution requires comprehensive investigation. This is
exemplified by the Hyperolius marmoratus/viridiflavus complex in Africa, in

which 15 names from Angola have been synonymised (Hyperolius cinctiventris,
H. decoratus, H. huillensis, H. insignis, H. marungaensis, H. microstictus, H. plic-
iferus, H. vermiculatus, Rappia cinctiventris, R. marmorata marginata, R. m.
paralella, R. m. variegata, R. plicifera, R. toulsonii, H. m. alborufus). Hyperolius
parallelus is closely related to this complex, and has several Angolan taxa in its
synonymy (H. angolensis, H. marmoratus var. angolensis, H. erythromelanus, H.
toulsonii, Rappia marmorata huillensis, R. m. insignis, R. m. taeniolata). Other
difficult groups are the Hyperolius platyceps complex, with four names currently
subsumed within it (Hyperolius angolanus, Rappia platyceps var. angolensis,
Table 12.2 Angolan frog species with undescribed tadpoles

Leptopelis bocagii (Günther, 1865) Hyperolius platyceps (Boulenger, 1900)
Leptopelis cynnamomeus (Bocage, 1893) Hyperolius polli (Laurent, 1943)
Leptopelis jordani (Parker, 1936) Hyperolius protchei (Rochebrune, 1885)
Leptopelis marginatus (Bocage, 1895) Hyperolius quinquevittatus (Bocage, 1866)
Leptopelis parvus (Schmidt and Inger, 1959) Hyperolius rhizophilus (Rochebrune, 1885)
Mertensophryne melanopleura (Schmidt and Hyperolius steindachneri (Bocage, 1866)
Inger, 1959) Hyperolius vilhenai (Laurent, 1964)
Mertensophryne mocquardi (Angel, 1924) Kassinula wittei (Laurent, 1940)
Poyntonophrynus grandisonae (Poynton and Phrynomantis affinis (Boulenger, 1901)
Haacke, 1993) Phrynobatrachus brevipalmatus (Ahl, 1925)
Poyntonophrynus kavangensis (Poynton and Phrynobatrachus cryptotis (Schmidt and
Broadley, 1988) Inger, 1959)
Poyntonophrynus pachnodes. (Ceríaco, Phrynobatrachus parvulus (Boulenger, 1905)
Marques, Bandeira et al. 2018a) Xenopus andrei (Loumont, 1983)
Sclerophrys buchneri (Peters, 1882) Xenopus petersii (Bocage, 1895)
Afrixalus osorioi (Ferreira, 1906) Xenopus epitropicalis (Fischberg, Colombelli,
Afrixalus fulvovittatus (Cope, 1861) and Picard, 1982)
Afrixalus wittei (Laurent, 1941). Hildebrandtia ornatissima (Bocage, 1879)
Hyperolius adspersus (Peters, 1877) Ptychadena ansorgii (Boulenger, 1905)
Hyperolius benguellensis (Bocage, 1893) Ptychadena bunoderma (Boulenger, 1907)
Hyperolius bicolor (Ahl, 1931) Ptychadena grandisonae (Laurent, 1954)
Hyperolius bocagei (Steindachner, 1867) Ptychadena guibei (Laurent, 1954)
Hyperolius cinnamomeoventris (Bocage, 1866) Ptychadena keilingi (Monard, 1937)
Hyperolius fasciatus (Ferreira, 1906) Ptychadena perplicata (Laurent, 1964)
Hyperolius ferreirai (Noble, 1924) Ptychadena taenioscelis (Laurent, 1954)
Hyperolius fuscigula (Bocage, 1866) Ptychadena upembae (Schmidt and Inger,
1959)
Hyperolius gularis Ahl, 1931 Ptychadena uzungwensis (Loveridge, 1932)
Hyperolius langi (Noble, 1924) Tomopterna damarensis (Dawood and
Hyperolius lucani (Rochebrune, 1885) Channing, 2002)
Hyperolius maestus (Rochebrune, 1885) Tomopterna tuberculosa (Boulenger, 1882)
Hyperolius nobrei (Ferreira, 1906) Amnirana parkeriana (Mertens, 1938)
Hyperolius parallelus (Günther, 1858)
Hyperolius fasciatus, Hyperolius ferreirai (originally Rappia bivittata)), and the

super-cryptic Hyperolius nasutus complex. Currently this is represented in Angola
by at least four species (H. adspersus, H. benguellensis, H. dartevellei, H. nasutus)
(Channing et al. 2013) and additional names that have been synonymised (H. punct-
ulatus, Rappia punctulata) (Channing et al. 2013) or not assigned to any known
species occurring in Angola (H. microps).
Typical toads are another problematic group. Formerly known as Bufo, which
was cosmopolitan in distribution and included the majority of bufonids, the genus
was partitioned with African typical toads transferred to Amietophrynus (Frost et al.
2006), and more recently renamed in the reinstated genus Sclerophrys (Poynton
et al. 2016). Seven species of typical toad occur in Angola (see Table 12.2). The
mysterious S. buchneri, known only from the holotype from northeastern Angola, is
considered as a valid species (Frost 2018), but synonymy with S. funerea has been
suggested and requires further studies (Tandy and Keith 1972). Apart from S. lemai-
rii which is easily distinguishable morphologically from the remaining species, dis-
tinction between the other Sclerophrys is difficult, even between the most common
species. Hybridisation between Sclerophrys species has been documented and dis-
cussed (Guttman 1967; Passmore 1972; Cunningham and Cherry 2004) and may
further complicate identification. The red coloration of the interior thigh and parotid
gland development are features commonly used to distinguish the often sympatric
S. pusilla, S. gutturalis and S. regularis (Du Preez and Carruthers 2017), but do not
distinguish these species in Angola. It is likely that cryptic diversity exists, and
understanding of the genus and delimitation of species boundaries requires an inte-
grative approach with comprehensive surveys, analysis of advertisement calls and
genetic studies.
Grass frogs, Ptychadena spp., are a challenging genus. At least 15 species of this
specious genus are represented in Angola (Appendix). P. mascareniensis, a large
species complex widespread in Africa and Madagascar, has been recently parti-
tioned (Dehling and Sinsch 2013b) with Ptychadena nilotica in much of continental
Africa, including Angola (Zimkus et al. 2017). Difficulties in distinguishing
Ptychadena species have been discussed (Poynton and Broadley 1985b; Dehling
and Sinsch 2013a, b), although coloration features such as triangular patch on the
head, pattern of the interior thigh (Poynton 1970) and several morphometric and
morphological features enable species identification (Dehling and Sinsch 2013a, b).
Species distinction in Angola is not clear, and in a recent study as many as six dif-
ferent species of Ptychadena were found in the same region (Conradie et al. 2016).
Rainfrogs in Angola are known from a single species, Breviceps adspersus.
However, analysis of material from Angola and adjacent regions has revealed that
the Angolan form has features of B. mossambicus and may indicate an undescribed
Angolan species (Poynton and Broadley 1985a, 1991).
Groups that remain not fully understood such as Phrynobatrachus (Zimkus et al.
2010), Xenopus (Furman et al. 2015) and Amnirana (Jongsma et al. 2018), all have
species widespread in Africa with type localities from Angola, and the resolution of
their taxonomy depends on detailed studies in Angola.
Species Synonymised with No Clear Justification
A number of putative Angolan species currently placed in synonymy require reas-

sessment as they may represent hidden diversity currently placed under a different
name. Cases are mentioned in the previous section, especially in the Hyperolius
genus. Other examples of this include the placement of Hylambates (=Leptopelis)
angolensis in the synonymy of Leptopelis bocagii. This resulted from comparison
between adult and juvenile specimens (Perret 1976) that may not be comparable.
Hylambates bocagei var. leucopunctata Ferreira 1904, has also been placed in the
synonymy of Leptopelis bocagii (Ceríaco et al. 2014b) and this also requires further
investigation as the well developed finger pads in the type specimen of H. b. leuco-
punctata suggests an arboreal habit, very different from the ground-dwelling habits
of L. bocagii, which lacks pads on fingers or toes.
Species with Questionable Distributions
Some species described from Angola have widespread distributions throughout

Africa and inhabit diverse habitats, suggesting that cryptic diversity may be involved
(see examples in Endemism section, below). The classic example of this is the
Common River Frog, Amietia angolensis, which was considered widespread in the
continent, but which was discovered to be in fact a complex of cryptic species, with
true A. angolensis being restricted to Angola (Channing and Baptista 2013). Another
potential example is Afrixalus osorioi, which was described from western Angola
and remains known in the country only from the type locality, whereas the closest
other records are in DRC, nearly 1000 km away from the type locality. Other exam-
ples include Ptychadena porosissima, Leptopelis cynnamomeus, L. bocagii,
Hyperolius bocagei, and highlight the earlier comments that study of Angolan
amphibians is crucial for solving many problems in African amphibian taxonomy.
Recent Discoveries and Ongoing Studies
The endemic Anchieta’s Treefrog, Leptopelis anchietae and Congulo Forest

Treefrog, Leptopelis jordani have been rediscovered in the Angolan escarpment
(Baptista et al. 2017), and together with other frogs belonging to the genus Kassina,
Arthroleptis and Amnirana found in the region, their conservation and taxonomic
status are being investigated (Baptista et al. in prep.). Further to this, ongoing stud-
ies (Baptista et al. in prep.) are assessing: a candidate new species of Schismaderma;
the taxonomic status of Hildebrandtia ornatissima from the Angolan central pla-
teau, previously discussed by Boulenger (1919); the status of Hyperolius punctula-
tus from the Cuanza River (currently in the synonymy of Hyperolius nasutus); and
the status of various populations of morphologically distinct pygmy toads that can-
not be assigned to known Poyntonophrynus species. During the 2011 expedition to
Lagoa Carumbo, a large white-lipped frog was morphologically assigned to the
Amnirana lepus group (Branch and Conradie 2015). This assigment has been con-
firmed in a phylogeny of the genus (Jongsma et al. 2018), and further studies are
underway to address the taxonomical status of the Angolan population (Conradie
pers. comm.). On the SAREP (2012/3) and the NGOWP (2016/7) expeditions to
southeastern Angola, numerous candidate new species were discovered, in the gen-
era Phrynobatrachus, Breviceps and Amnirana, and are currently under investiga-
tion (Conradie pers. comm.). The new country records of Kassinula wittei and
Leptopelis cf. parvus are being studied to determine if they conform to the nominal
forms from northern Zambia and southern DRC, respectively (Conradie pers.
comm.). During recent independent surveys conducted in the northern Angolan
provinces of Uíge (Ernst et al. 2014, 2015) and Zaire (Vaz Pinto and Baptista
Unpublished Data), two different Alexteroon spp. were discovered. The taxonomic
status of these, the first Angolan records for this poorly-known hyperoliid genus, are
under investigation with the Uíge species tentatively assigned to the nominal spe-
cies A. hypsiphonus, whilst the Zaire discovery has affinities to A. obstetricans. The
material awaits formal taxonomic assignment pending analysis of type material.
Biogeography
Angola is one of the most biogeographically rich countries in Africa (Huntley 1974,
2019). Geomorphologically, the country can be divided into various regions, includ-
ing the western lowlands of the Coast Belt, the Transition Zone which includes the
escarpment, the Marginal Mountain Chain, the Old (Highland) Plateau, also known
as central plateau, which progressively decreases in altitude to the east, where the
Congo Basin in the north and the Zambezi-Cubango Basin in the south are located
(Huntley 1974). Each of these regions have several biome associations, with habi-
tats ranging from the tropical rainforests on the Maiombe region in the north, to the
Namib Desert in the south, one of the oldest deserts in the world (Huntley 1974,
2019). This complexity is reflected in the country’s diverse fauna.
The difficulties in establishing clear biogeographic regions for amphibians is
demonstrated by Poynton and Broadley (1991), in their thorough analysis of the
biogeography of the Zambezian amphibians. For Angolan amphibians, which are
much more poorly known, this difficulty is immensely increased. The biogeography
of Angolan amphibians can only be assessed after major taxonomic issues are
resolved, which in some cases requires the revision of entire genera (Cei 1977;
Blanc and Frétey 2000). In early studies of the Angolan herpetofauna, several
attempts were made to group species according to the distributions known at the
time, and these will be summarised below.
Bocage (1895a) made the first grouping, distinguishing a northern and southern
region, each divided into coastal, intermediate and high-altitude zones, and listing
species occurring in each block. Monard (1937) used humidity to explain the higher
diversity of amphibian species in central Angola (a high-rainfall region), compared
to the south. He divided Angolan amphibians into four groups: (i) pan-African spe-
cies (4% of the country’s species; such as Rana mascareniensis (=Ptychadena nilot-
ica), and Bufo (=Sclerophrys) regularis; (ii) southern species (10%) which reached
their northern limit in Angola, such as Pyxicephalus adspersus; (iii) tropical species
(40%), from western, central and eastern Africa, highlighting the central African
tropics as the most significant influence, and including Rana (=Amnirana) albola-
bris and Rana (=Hoplobatrachus) occipitalis; and (iv) endemic species (46%),
most of which are no longer considered endemic (see Endemism section).
Based on the species known from Angola at the time, Cei (1977) organised
Angolan amphibians in three questionable groups, each with affinities to different
habitats and regions: (i) the northern and northeastern forests and savannas, (ii) the
plateau, and (iii) the arid and semi-arid regions of the coast and of the south, provid-
ing a map to delineate those areas. The first area is wide, with northern and north-
eastern limits in the Congo, Cuanza and Cassai rivers (in Zaire, Uíge, Malanje and
Lunda-Norte), and extending to the southeast through Moxico and Cuando Cubango.
Examples of species within this group are Arthroleptis carquejai and Hyperolius
steindachneri. The second region corresponds to the south of Congo and Cuango
rivers and comprises the southern tropics: Cuanza-Norte, Cuanza-Sul, Huambo,
Bié, Malanje and Huíla provinces. Characteristic species in this group include
Hildebrandtia ornatissima, Hyperolius cinereus, Hyperolius quinquevittatus,
Leptopelis anchietae. The third and southernmost region comprises the arid sections
of Benguela, Namibe, and Cunene provinces. The fauna on this group is related to
that of the Namib, Kalahari, and Namaqualand regions, and can be exemplified by
Pyxicephalus adspersus and Poyntonophrynus dombensis.
Surprisingly, the Great Escarpment of Angola has not been considered in any of
these studies. This escarpment is part of a much larger geomorphological unit that
dominates the African subcontinent and extends into western Angola, where it acts
as a barrier between the dry coastline and the inland plateau. Due to its climatic and
topographic peculiarities, it promotes isolation and thus speciation (Huntley 1974).
It is a well-documented center of endemism for birds (Hall 1960), and although the
escarpment herpetofauna is poorly understood, its endemism potential for herpeto-
fauna has been highlighted (Laurent 1964, Clark et al. 2011, Baptista et al. 2018,
Branch et al. 2019), and endemic amphibian species are known from the region
(Leptopelis jordani, L. marginatus, Amnirana parkeriana and Hyperolius che-
laensis). Bordering the Angolan escarpment to the east, the highlands of the ancient
massif include patches of Afromontane forest. These consist of islands of relic cool
moist Afromontane forest with great biogeographic interest (Huntley 1974), and
also potential for endemism.
Inland to the escarpment zone, the plateau is broadly dominated by miombo
woodlands, and its fauna often has influences from adjacent regions. Boundaries
between regions are not always clear or well understood. Some of these uncertain-
ties have been mentioned in early studies and still require explanation. Hellmich
(1957b) referred to the difficulty in establishing geographical limits between the
moist forests of the north and the central plateau. An example of this is the penetra-
tion of forest species in association with riverine habitats along the northern rivers
of Angola. He also noted that faunal boundaries between the slopes of the eastern
plateau and the flatlands between Cassai and Cuando were not clear, with the pres-
ence of ‘pockets’ of herpetological elements typical of the south on the central
plateau. Laurent (1964) referred to the known affinities between the species of
Katanga, in southeastern DRC, with the species of the Lundas and Moxico in
Angola.
All of these early biogeographic regions and the species assigned to them need
to be re-evaluated with updated taxonomy, accurate species distributions, and in
association with the study of phylogenetic relationships among the various amphib-
ian families and genera occurring in Angola. The confirmation of ancestral relation-
ships within these groups is a prerequisite for testing hypotheses about the timing
and environmental correlates of amphibian movement and speciation across the
Angolan landscape.
Endemism
The originality of the Angolan amphibians due to the richness of endemic species
has been highlighted (Blanc and Frétey 2000). Angola’s more unique amphibians
are also the most poorly known. There are 21 species of amphibians endemic to
Angola, of which about 75% are only known from the type locality or type speci-
mens (Table 12.3). Many have not been found for decades, and in some cases for
over 100 years. Most of these species are classified as Data Deficient in the IUCN
Red List (IUCN 2017).
A number of endemic taxa have been mentioned in the literature but still await
formal description: Hyperolius sp. I, Hyperolius sp. II, Hyperolius sp. III (Monard
1937), possibly unknown genus (Perret 1996), and as the taxonomic studies on
Angolan amphibians progress, more endemic species will very likely be discovered.
In contrast, many early species that were considered endemic have now been rele-
gated to the synonymy of wide-ranging species. Monard (1937), for instance, con-
sidered nearly half (46%) of the 80 Angolan species he considered to occur in the
country as endemic. However, of the 37 endemic species he identified, only eight
are still recognised. Sixteen of these former ‘endemics’ have been synonymised
with other species; e.g. Leptopelis angolensis (= L. bocagii), Rana buneli (=
Ptychadena bunoderma), Hyperolius seabrai (= H. bocagei), Hyperolius angola-
nus, H. ferreirai, H. fasciatus (all =Hyperolius platyceps), H. pliciferus, H. ver-
miculatus, H. marungaensis (all =Hyperolus marmoratus), H. angolensis, H.
erythromelanus, H. toulsonii (all =Hyperolius parallelus), H. punctulatus
(=Hyperolius nasutus), Rana myotympanum (=Hildebrandtia ornatissima), Rana
cacondana and R. signata (=Tomopterna tuberculosa). Many of these synonymies
have poor justification, and whilst some names may reflect regional variation, others
Table 12.3 List of amphibian species endemic to Angola, with IUCN Red List Category (LC least
concern, DD data deficient, N/A not assessed), and marked (X) when known only from the type
locality
Common name Scientific name IUCN TYPE
Angola River frog Amietia angolensis (Bocage, 1866) LC
Parker’s white-lipped Amnirana parkeriana (Mertens, 1938) DD X
frog
Cambondo squeaker Arthroleptis carquejai (Ferreira, 1906) DD X
Angola ornate frog Hildebrandtia ornatissima (Bocage, 1879) DD
Two-colored reed frog Hyperolius bicolor (Ahl, 1931) DD X
Chela Mountain Reed Hyperolius chelaensis (Conradie et al., 2012) N/A X
Frog
Monard’s Reed Frog Hyperolius cinereus (Monard, 1937) LC
Brown-throated Reed Hyperolius fuscigula (Bocage, 1866) DD X
Frog
Loanda Reed Frog Hyperolius gularis (Ahl, 1931) DD X
Landana Reed Frog Hyperolius lucani (Rochebrune, 1885) DD X
Cabinda Reed Frog Hyperolius maestus (Rochebrune, 1885) DD X
Nobre’s Reed Frog Hyperolius nobrei (Ferreira, 1906) N/A X
Rochebrune’s Reed Hyperolius protchei (Rochebrune, 1885) DD X
Frog
Raymond’s Reed Frog Hyperolius raymondi (Conradie et al., 2013) N/A
African Reed Frog Hyperolius rhizophilus (Rochebrune, 1885) DD X
Luita River Reed Frog Hyperolius vilhenai (Laurent, 1964) DD X
Congulo Forest Leptopelis jordani (Parker, 1936) DD X
Treefrog
Quissange Forest Leptopelis marginatus (Bocage, 1895) DD X
Treefrog
Ahl’s Puddle Frog Phrynobatrachus brevipalmatus (Ahl, 1925) DD X
Grandison’s Pygmy Poyntonophrynus grandisonae (Poynton and Haacke, DD X
Toad 1993)
Serra da Neve Pygmy Poyntonophrynus pachnodes (Ceríaco, Marques, N/A X
Toad Bandeira et al., 2018a)
Taxonomy follows Frost (2018)
referred to species found in other countries may not be conspecific (see Comments
on selected groups). All deserve careful re-examination.
At least four species (Leptopelis marginatus, L. jordani, Amnirana parkeriana,
and Hyperolius chelaensis) are escarpment-endemics, and others are plateau-
endemics (Hildebrandtia ornatissima, H. cinereus). However, in order to effectively
protect Angolan endemic amphibians and their habitats, further studies are needed
to reveal the relations between endemic amphibians and particular habitat, and also
the importance of other potential areas of endemism (e.g. relic Afromontane forest
patches, isolated mountains such as Serra da Neve, the Angolan escarpment).
Directions for Future Research in Angola
Detailed species lists for a country are an essential baseline tool for understanding
biodiversity, its distribution and conservation status. The confusing status of
Angolan amphibian taxonomy has been discussed in previous sections and demon-
strates how studying taxonomy forms the bedrock for resolving the many pressing
questions regarding Angolan amphibian conservation and biology.
A critical first taxonomic step is to revisit the type localities of all the species
described from the country to obtain new topotypical material. This is particularly
important for the 15 species described by Bocage (Amietia angolensis, Hyperolius
benguellensis, H. cinnamomeoventris, H. fuscigula, H. quinquevittatus, H. stein-
dachneri, Ptychadena anchietae, P. subpunctata, Sclerophrys funerea, Leptopelis
anchietae, L. cynnamomeus, L. marginatus, Hildebrandtia ornatissima,
Poyntonophrynus dombensis, Xenopus petersii), for which many of the type speci-
mens were lost in the fire that destroyed the collections of the Natural History
Museum of Lisbon, and for which the original descriptions are the only available
source of information. Possibly also lost are the type specimens of several Angolan
endemics described by Rochebrune (Hyperolius lucani, H. maestus, H. protchei, H.
rhizophilus) (Frost 2018), which have very vague descriptions. For many species,
neotypes may need to be designated in order to stabilise their taxonomy. Integrative
taxonomic studies, including analysis of genetic material, advertisement calls, adult
and larval morphology, habitat associations and natural history are crucial to bring
Angolan studies into the new millennium.
Many regions of Angola have never been surveyed for amphibians (see Fig. 12.1).
Surveying these areas would greatly improve understanding of amphibian distribu-
tions, habitat associations and relative abundance, but are also critical for making
assessments on their conservation status in terms of IUCN criteria. Priority areas
include the northwestern provinces (Uíge and Zaire), the extensive wetlands of
Moxico, the escarpment and the adjacent Afromontane forest patches that are rich
in endemic birds (Hall 1960), other vertebrates (Crawford-Cabral 1966; Clark et al.
2011) and also probably amphibians, and for which the urgent need of studies has
been highlighted (Laurent 1964; Clark et al. 2011).
The controversial frog from Caquindo (Perret 1996) for which genus assignment
lacks consensus (see Records that require confirmation), still has to be recollected
and its true affinities resolved. This could enrich Angolan herpetology possibly with
a new endemic genus. This begs the question – how much remains to be discovered
about Angolan amphibians? It also shows how the analysis of extant collections can
contribute significantly to the knowledge of the country’s fauna. Collections that
remain to be studied include those from the Rohan-Chabot Mission, the Vernay
Angola Expedition, and the Leptopelinae and Hyperoliidae from the Portuguese
Mission of Apiarian Studies of the Overseas.
Another important step to furthering amphibian knowledge is studying the biol-
ogy of individual species. Some studies are available for iconic species such as the
Dombe Pigmy toad Poyntonophrynus dombensis (Channing and Vences 1999),
based on individuals from Namibia, and Lemaire’s toad Sclerophrys lemairii, the
first study of this kind made in Angolan territory (Conradie and Bills 2017).
However, this is still missing for many species, and understanding their natural his-
tory, reproduction strategies, breeding sites, breeding seasons, behaviour, habitat
and microhabitat requirements, both for adults and tadpole stages, are key for an
effective planning of species conservation. All of this is even more relevant for the
extremely poorly known Angolan endemics.
Conservation-driven studies about Angolan amphibians require awareness of
potential threats to biodiversity, particularly those resulting from habitat loss and
climate change. Habitat degradation as a result of exploitation of natural resources
and associated with industrialisation have increased dramatically in Angola in
recent decades and will affect amphibians. The implementation of monitoring pro-
grammes are crucial for documenting and understanding this relation. Research
about the appearance and effect of global amphibian diseases such as the chytrid
fungus (Batrachochytrium dendrobatidis), viruses (Ranavirus spp.), and other
pathogens, are lacking in Angola, even though they are threatening amphibians
around the world and are reported from neighbouring countries (Greenbaum et al.
2014).
The study of Angolan amphibians is a broad and important component of biodi-
versity studies, for which many baseline questions remain unanswered, and exciting
discoveries are still to be made. This becomes more evident when confronted with
the fact that Angolan fauna is among the least studied in Africa. Increasing public
awareness about amphibians and their importance is necessary for their conserva-
tion, and requires developing local knowledge and expertise, as well as constructing
functional amphibian collections in national archives. These are essential steps for
understanding and protecting this rich, diverse and ecologically important group.
This is even more urgent in an era where an “amphibian decline crisis” is happening
around the world (Beebee and Griffiths 2005), and where this decline is known to
have major consequences in ecosystem function (Whiles et al. 2006).
Acknowledgements The writing of this chapter was made possible through a convergence of
efforts and projects. SASSCAL Project (sponsored by the German Federal Ministry of Education
and Research (BMBF) under promotion number 01LG1201M); Conservation Leadership
Programme (Project CLP ID: F01245015: Conserving Angolan Scarp Forests: a Holistic Approach
for Kumbira Forest); National Geographic/Okavango Wilderness Project (NGOWP); South
Africa’s National Research Foundation (2009–2017, WRB), National Geographic Society
(Explorer Grant 2011, WRB); Fundação para a Ciência e Tecnologia (contract SFRH/PD/
BD/140810/2018, NB); and Wild Bird Trust 2015–2018. Particular thanks go to Fernanda Lages
(ISCED Huíla), Brian Huntley (South Africa), John Hilton and Rainer Von Brandis (Wild Bird
Trust) for logistical and administrative support.
Appendix 1
Checklist of the amphibians recorded in Angola, based on historical records and on

confirmed records from recent surveys. Taxonomy follows Frost (2018).
Unconfirmed records are not included. To avoid redundancy, records included in
existing compilations (e.g. Monard 1937; Ruas 1996) are mentioned under the com-
pilation reference, and the original reference(s) is not included in the list
Common name Species References

Family Arthroleptidae
Carqueja’s Arthroleptis carquejai Ferreira (1906)
Squeaker (Ferreira, 1906)
Lameer’s Arthroleptis lameerei (De Laurent (1964) and Ruas (1996)
Squeaker Witte, 1921)
Tanganyika Arthroleptis spinalis Laurent (1950)
Screeching Frog (Boulenger, 1919)
Common Arthroleptis stenodactylus Laurent (1964), Ruas (1996) and Conradie et
Squeaker (Pfeffer, 1893) al. (unpub. data)
Variable Arthroleptis variabilis Baptista and Vaz Pinto (unpub. data)
Squeaker (Matschie, 1893)
Plain Squeaker Arthroleptis xenochirus Monard (1937), Laurent (1964), Ruas
(Boulenger, 1905) (1996), Ceríaco et al. (2018b), Conradie et
al. (unpub. data), Baptista and Vaz Pinto
(unpub. data), and Ernst (unpub. data)
Anchieta’s Leptopelis anchietae Bocage (1895), Boulenger (1905), Schmidt
Treefrog (Bocage, 1873) (1936), Monard (1937), Laurent (1964),
Conradie et al. (2016), Baptista et al. (2018),
(in prep.) and Ernst (unpub. data)
Gaboon Forest Leptopelis aubryi (Duméril, Peters (1887) and Laurent (1954)
Treefrog 1856)
Bocage’s Leptopelis bocagii (Günther, Bocage (1895), Monard (1937), Hellmich
Burrowing 1865) (1957b), Laurent (1954, 1964), Ceríaco et al.
Treefrog (2018b), Baptista et al (2018, in prep),
Baptista and Vaz Pinto (unpub. data) and
Conradie et al. (unpub. data)
Efulen Forest Leptopelis calcaratus Baptista and Vaz Pinto (unpub. data)
Treefrog (Boulenger, 1906)
Cinnamon Leptopelis cynnamomeus Bocage (1895), Monard (1937) and Laurent
Treefrog (Bocage, 1893) (1964)
Congulo Forest Leptopelis jordani (Parker, Parker (1936) and Baptista et al. (2017)
Treefrog 1936)
Quissange Forest Leptopelis marginatus Bocage (1895)
Treefrog (Bocage, 1895)
Kanole Forest Leptopelis cf. parvus Conradie et al. (unpub. data)
Treefrog (Schmidt and Inger, 1959)
Rusty Forest Leptopelis viridis (Günther, Boulenger (1882) and Bocage (1895)
Treefrog 1869)
Hairy Frog Trichobatrachus robustus Ernst et al. (2014)
(Boulenger, 1900)
(continued)

Family Brevicipitidae
Common Rain Breviceps cf. adspersus Bocage (1895), Monard (1937), Hellmich
Frog (Peters, 1882) (1957b), Laurent (1964), Ruas (1996) and
Family Bufonidae
Dark-sided Forest Mertensophryne Ruas (1996)
Toad melanopleura (Schmidt and
Inger, 1959)
Mocquard’s Mertensophryne mocquardi Monard (1937)
Forest Toad (Angel, 1924)
Dombe Pygmy Poyntonophrynus dombensis Bocage (1895), Poynton and Haake (1993),
Toad (Bocage, 1895) Ceríaco et al. (2018a) and Vaz Pinto and
Branch (unpub. data)
Grandison’s Poyntonophrynus Poynton and Haacke (1993) and Ceríaco et
Pygmy Toad grandisonae (Poynton and al. (2018a)
Haacke, 1993)
Kavango Pygmy Poyntonophrynus Poynton and Haacke (1993), Ruas (1996)
Toad Toad kavangensis (Poynton and and Vaz Pinto (unpub. data)
Broadley, 1988)
Serra da Neve Poyntonophrynus pachnodes Ceríaco et al. (2018a)
Pygmy Toad (Ceríaco et al. in press.)
Red Toad Schismaderma carens (Smith, Monard (1937), Ruas (1996) and Baptista
1848) and Vaz Pinto (unpub. data)
Buchner’s Toad Sclerophrys buchneri (Peters, Peters (1882)
1882)
Somber Toad Sclerophrys funerea (Bocage, Bocage (1895), Monard (1937), Laurent
1866) (1954, 1964), Ruas (1996) and Conradie et
al. (2016)
Guttural Toad Sclerophrys gutturalis Ruas (1996, 2002), Conradie et al. (2016,
(Power, 1927) unpub. data), Baptista et al. (2018), Baptista
and Vaz Pinto (unpub. data)
Lemaire’s Toad Sclerophrys lemairii Laurent (1950, 1964), Ruas (1996) and
(Boulenger, 1901) Conradie et al. (2016)
Western Olive Sclerophrys poweri (Hewitt, Conradie et al. (2016) and Baptista et al. (in
Toad 1935) press)
Merten’s Striped Sclerophrys pusilla (Mertens, Ruas (1996, 2002), Conradie et al. (2016),
Toad 1937) Poynton et al. (2016), Ceríaco et al. (2018b),
Baptista et al. (2018) and Baptista and Vaz
Pinto (unpub. data)
Common Toad Sclerophrys regularis (Reuss, Bocage (1895), Monard (1937), Laurent
1833) (1964), Ruas (1996), Ceríaco et al. (2014b)
and Vaz Pinto and Baptista (unpub. data)
Family Dicroglossidae
Crowned Hoplobatrachus occipitalis Bocage (1895), Monard (1937), Hellmich
Bullfrog (Günther, 1858) (1957b) and Ruas (1996), Baptista (unpub.
data)
(continued)

Family Hemisotidae
Guinea Hemisus guineensis (Cope, Laurent (1964), Ceríaco et al. (2018b),
Snout-burrower 1865) Conradie et al. (unpub. data) and Baptista
and Vaz Pinto (unpub. data)
Marbled Hemisus marmoratus (Peters, Bocage (1895), Monard (1937), Hellmich
Snout-burrower 1854) (1957b), Ruas (1996) and Baptista et al. (in
prep.)
Family Hyperoliidae
Striped Spiny Afrixalus dorsalis (Peters, Laurent (1964) and Baptista and Vaz Pinto
Reed Frog 1875) (unpub. data)
Four Lined Reed Afrixalus fulvovittatus (Cope, Bocage (1866a) and Ferreira (1904)
Frog 1861)
Osorio’s Spiny Afrixalus osorioi (Ferreira, Ferreira (1906), Baptista and Vaz Pinto
Reed Frog 1906) (unpub. data) and Ernst (unpub. data)
Four-Lined Spiny Afrixalus quadrivittatus Peters (1887) and Perret (1976)
Reed Frog (Werner, 1908)
De Witte’s Spiny Afrixalus wittei (Laurent, Ceríaco et al. (2018b), Baptista and Vaz
Reed Frog 1941) Pinto (unpub. data) and Ernst (unpub. data)
Sprinkled Long Hyperolius adspersus (Peters, Laurent (1964)
Reed Frog 1877)
Benguela Long Hyperolius benguellensis Bocage (1895), Ferreira (1906), Monard
Reed Frog (Bocage, 1893) (1937), Laurent (1950), Channing et al.
(2013), Conradie et al. (2016) and Baptista et
al. (2018)
Two-colored Hyperolius bicolor (Ahl, Ahl (1931)
Reed Frog 1931)
Bocage’s Reed Hyperolius bocagei Monard (1937), Laurent (1950, 1954, 1964),
Frog (Steindachner, 1867) Ceríaco et al. (2014b), Conradie (unpub.
data), Baptista and Vaz Pinto (unpub. data)
and Ernst (unpub. data)
Chela Mountain Hyperolius chelaensis Conradie et al. (2012)
Reed Frog (Conradie et al. 2012)
Monard’s Reed Hyperolius cinereus Monard (1937), Conradie et al. (2016),
Frog (Monard, 1937) Baptista et al. (2018), Baptista and Vaz Pinto
(unpub. data) and Conradie et al. (unpub.
data)
Dimorphic Reed Hyperolius Monard (1937), Laurent (1950, 1954, 1964),
Frog cinnamomeoventris (Bocage, Ceríaco et al. (2016c), (2018b) and Baptista
1866) and Vaz Pinto (unpub. data)
Variable Reed Hyperolius concolor Monard (1937)
Frog (Hallowell, 1844)
Dartevelle’s Long Hyperolius dartevellei Laurent (1964) and Channing et al. (2013)
Reed frog (Laurent, 1943)
Brown-throated Hyperolius fuscigula Bocage (1866)
Reed Frog (Bocage, 1866)
Family Hemisotidae (cont.)
Loanda Reed Hyperolius gularis (Ahl, Ahl (1931)
Frog 1931)
Kivu Reed Frog Hyperolius kivuensis (Ahl, Laurent (1950, 1954)
1931)
(continued)

Lang’s Reed Frog Hyperolius langi (Noble, Monard (1937)
1924)
Landana Reed Hyperolius lucani Rochebrune (1885)
Frog (Rochebrune, 1885)
Cabinda Reed Hyperolius maestus Rochebrune (1885)
Marbled Reed Hyperolius marmoratus Boulenger (1882), Bocage (1895) and
Frog (Rapp, 1842) Monard (1937)
Large-nosed Hyperolius nasutus (Günther, Bocage (1895), Monard (1937), Laurent
Long Reed Frog 1865) (1950, 1954, 1964), Hellmich (1957b),
Baptista and Vaz Pinto (unpub. data) and
Ceríaco et al. (2018b)
Nobre’s Reed Hyperolius nobrei (Ferreira, Ferreira (1906)
Frog 1906)
Angolan Reed Hyperolius parallelus Monard (1937), Laurent (1950, 1954, 1964),
Frog (Günther, 1858) Ceríaco et al. (2018b), Conradie et al.
(unpub. data), Baptista et al. (2018) and
Baptista and Vaz Pinto (unpub. data)
Leopard Reed Hyperolius pardalis (Laurent, Conradie (unpub. data)
Frog 1948)
Rio Luinha Reed Hyperolius platyceps Monard (1937), Laurent (1950, 1954) and
Frog (Boulenger, 1900) Baptista and Vaz Pinto (unpub. data)
Tshimbulu Reed Hyperolius polli (Laurent, Laurent (1954)
Frog 1943)
Rochebrune’s Hyperolius protchei Rochebrune (1885)
Reed Frog (Rochebrune, 1885)
Five-striped Reed Hyperolius quinquevittatus Bocage (1895), Laurent (1950, 1954) and
Frog (Bocage, 1866) Baptista and Vaz Pinto (unpub. data)
Raymond’s Reed Hyperolius raymondi Conradie et al. (2013)
Frog (Conradie et al. 2013)
African Reed Hyperolius rhizophilus Rochebrune (1885)
Steindachner’s Hyperolius steindachneri Bocage (1895), Monard (1937), Laurent
Reed Frog (Bocage, 1866) (1950, 1954, 1964), Poynton and Haacke
(1993) and Channing and Vaz Pinto (unpub.
data)
Luita River Reed Hyperolius vilhenai (Laurent, Laurent (1964)
Frog 1964)
Kuvangu Kassina Kassina kuvangensis Monard (1937) and Conradie et al. (2016,
(Monard, 1937) unpub. data)
Family Hemisotidae (cont.)
Bubbling Kassina Kassina senegalensis Monard (1937), Laurent (1954, 1964),
(Duméril and Bibron, 1841) Poynton and Haacke (1993), Conradie et al.
(2016), Baptista et al. (2018), Baptista and
Vaz Pinto (unpub. data), Conradie et al.
(unpub. data) and Ernst (unpub. data)
De Witte’s Kassinula wittei (Laurent, Conradie et. al (unpub. data)
Clicking Frog 1940)
(continued)

Family Microhylidae
Spotted Rubber Phrynomantis affinis Laurent (1964)
Frog (Boulenger, 1901)
Marbled Rubber Phrynomantis annectens Ruas (1996) and Vaz Pinto and Branch
Frog (Werner, 1910) (unpub. data)
Banded Rubber Phrynomantis bifasciatus Boulenger (1882), Monard (1937), Ruas
Frog (Smith, 1847) (1996), Channing (unpub. data) and Baptista
et al. (unpub. data)
Family Phrynobatrachidae
Ahl’s Puddle Phrynobatrachus Ahl (1925)
Frog brevipalmatus (Ahl, 1925)
Cryptic Puddle Phrynobatrachus cryptotis Laurent (1964)
Frog (Schmidt and Inger, 1959)
Mababe Puddle Phrynobatrachus Poynton and Haacke (1993), Conradie et al.
Frog mababiensis (FitzSimons, (2016, unpub. data) and Baptista and Vaz
1932) Pinto (unpub. data)
Snoring Puddle Phrynobatrachus natalensis Bocage (1895), Monard (1937), Hellmich
Frog (Smith, 1849) (1957b), Ruas (1996), Conradie et al. (2016,
unpub. data), Ceríaco et al. (2018b), Baptista
et al. (2018, in press) and Baptista and Vaz
Pinto (unpub. data)
Loanda River Phrynobatrachus parvulus Ruas (1996), Baptista and Vaz Pinto (unpub.
Frog (Boulenger, 1905) data) and Conradie et al. (unpub. data)
Family Pipidae
Andre’s Clawed Xenopus andrei (Loumont, Ernst et al. (2015)
Frog 1983)
Southern Tropical Xenopus epitropicalis Laurent (1950, 1954) and Klein (unpub.
Clawed Frog (Fischberg et al., 1982) data)
Müller’s Clawed Xenopus muelleri (Peters, Conradie et al. (2016)
Frog 1844)
Peters’ Clawed Xenopus petersii (Bocage, Bocage (1895), Monard (1937), Hellmich
Frog 1895) (1957b), Ruas (1996), Baptista et al. (2018),
Baptista and Vaz Pinto (unpub. data), Ceríaco
et al. (2018b) and Ernst (unpub. data)
Power’s Clawed Xenopus poweri (Hewitt, Conradie et al. (2016)
Frog 1927)
Family Pipidae (cont.)
Clawed Frog Xenopus sp. Laurent (1950)
Family Ptychadenidae
Common Ornate Hildebrandtia ornata (Peters, Poynton and Haacke (1993)
Frog 1878)
Angola Ornate Hildebrandtia ornatissima Bocage (1895), Monard (1937) and Baptista
Frog (Bocage, 1879) and Vaz Pinto (unpub. data)
Anchieta’s Grass Ptychadena anchietae Ruas (1996), Ceríaco et al. (In press.),
Frog (Bocage, 1868) Baptista et al (2018) and Baptista and Vaz
Pinto (unpub. data)
Ansorge’s Grass Ptychadena ansorgii Monard (1937) and Ruas (1996)
Frog (Boulenger, 1905)
(continued)

Rough Grass Ptychadena bunoderma Monard (1937), Ruas (1996) and Conradie et
Frog (Boulenger, 1907) al. (unpub. data)
Grandison’s Ptychadena grandisonae Ruas (1996)
Grass Frog (Laurent, 1954)
Guibe’s Grass Ptychadena guibei (Laurent, Ruas (1996), Ceríaco et al. (in press),
Frog 1954) Conradie et al. (2016) and Baptista and Vaz
Pinto (unpub. data)
Keiling’s Grass Ptychadena keilingi (Monard, Ruas (1996) and Conradie et al. (unpub.
Frog 1937) data)
Mozambique Ptychadena cf. mossambica Conradie et al. (2016) and Conradie (unpub.
Grass Frog (Peters, 1854) data)
Nile Grass Frog Ptychadena nilotica (Seetzen, Monard (1937), Schmidt and Inger (1959),
1855) Ruas (1996), Conradie et al. (2016), Dehling
and Sinsch (2013b) and Zimkus et al. (2017)
Sharp-nosed Ptychadena oxyrhynchus Monard (1937), Hellmich (1957b), Ruas
Grass Frog (Smith, 1849) (1996), Ceríaco et al. (2018b), Conradie et
al. (2016) and Baptista (unpub. data)
Many-Grass Frog Ptychadena perplicata Laurent (1964)
(Laurent, 1964)
Striped Grass Ptychadena porosissima Ruas (1996), Conradie et al. (unpub. data)
Frog (Steindachner, 1867) and Channing et al. (2012)
Spot-bellied Ptychadena subpunctata Ruas (1996), Conradie et al. (2016) and
Grass Frog (Bocage, 1866) Baptista (unpub. data)
Small Grass Frog Ptychadena taenioscelis Ruas (1996) and Conradie et al. (2016)
(Laurent, 1954)
Upemba Grass Ptychadena upembae Ruas (1996)
Frog (Schmidt and Inger, 1959)
Udzungwa Grass Ptychadena uzungwensis Ruas (1996) and Conradie et al. (2016,
Frog (Loveridge, 1932) unpub. data)
Family Pyxicephalidae
Angola River Amietia angolensis (Bocage, Bocage (1895), Monard (1937), Ruas (1996),
Frog 1866) Channing and Baptista (2013), Ceríaco et al.
(2016b), Channing et al. (2016), Baptista et
al. (2018) and Baptista and Vaz Pinto (unpub.
data)
African Bullfrog Pyxicephalus adspersus Monard (1937) and Ruas (1996)
(Tschudi, 1838)
Cryptic Sand Tomopterna cryptotis Monard (1937), Ruas (1996), Conradie et al.
Frog (Boulenger, 1907) (2016) and Baptista et al. (in press)
Damaraland Sand Tomopterna damarensis Ceríaco et al. (2016a) and Heinicke et al.
Frog (Dawood and Channing, (2017)
2002)
Rough Sand Frog Tomopterna tuberculosa Bocage (1895), Monard (1937), Ruas (1996),
(Boulenger, 1882) Baptista et al (2018, unpub. data) and
Family Ranidae
Forest White- Amnirana albolabris Bocage (1895), Monard (1937), Ruas (1996)
lipped Frog (Hallowell, 1856) and Jongsma et al. (2018)
(continued)

Darling’s Amnirana darlingi Monard (1937), Laurent (1964), Ruas
White-lipped (Boulenger, 1902) (1996), Ceríaco et al. (2018b)
Frog Branch and Conradie (2015) and Conradie et
al. (unpub. data)
Lemaire’s Amnirana lemairei (De Witte, Laurent (1964), Ruas (1996) and Baptista
White-lipped 1921) and Vaz Pinto (unpub. data)
Frog
Andersson’s Amnirana cf. lepus Branch and Conradie (2015)
White-lipped (Andersson, 1903)
Frog
Parker’s Amnirana parkeriana Mertens (1938)
White-lipped (Mertens, 1938)
Frog
References
Ahl E (1925 “1923”) Ueber neue afrikanische Frösche der Familie Ranidae. Sitzungsberichte der
Gesellschaft Naturforschender Freunde zu Berlin 1923:96–106
Ahl E (1931) Amphibia, Anura III, Polypedatidae. Das Tierreich 55: xvi + 477
Amiet JL (2012) Les Rainettes du Cameroun (Amphibiens Anoures). La Nef des Livres, Saint-
Nazaire, 591 pp
Baptista N, António T, Branch WR The herpetofauna of Bicuar National Park and surrounds,
southwestern Angola: a first description and preliminary checklist. Amphibian & Reptile
Conservation. (in press).
Baptista N, Vaz Pinto P, Ernst R et al (2017) Cryptic diversity in treefrogs (Leptopelis) of the
Angolan escarpment – fitting the pieces together. 13th conference of the Herpetological
Association of Africa, Bonamanzi, South Africa
Baptista N, António T, Branch WR (2018) Amphibians and reptiles of the Tundavala region of
the Angolan Escarpment. In: Revermann R, Krewenka KM, Schmiedel U et al (eds) Climate
change and adaptive land management in Southern Africa – assessments, changes, challenges,
and solutions, Biodiversity & ecology, vol 6, pp 397–403
Beard KH, Vogt KA, Kulmatiski A (2002) Top-down effects of a terrestrial frog on forest nutrient
dynamics. Oecologia 133(4):583–593
Beebee TJ, Griffiths RA (2005) The amphibian decline crisis: a watershed for conservation biol-
ogy? Biol Conserv 125(3):271–285
Blanc CP, Frétey T (2000) Biogeographie des Amphibiens d’Afrique Centrale et d’Angola.
Biogeographica 76(3):107–118
Bocage JVB (1866a) Lista dos reptis das possessões portuguesas d’ Africa occidental que existem
no Museu de Lisboa. Jornal de Sciências Mathemáticas, Physicas e Naturaes. Lisboa 1:37–56
Bocage JVB (1866b) Reptiles nouveaux ou peu connus recueillis dans les possessions por-
tugaises de l’Afrique occidentale, qui se trouvent au Muséum de Lisbonne. Jornal de Sciencias
Mathematicas, Physicas e Naturaes. Lisboa I(1):57–78
Bocage JVB (1867) Batraciens nouveaux de I’Afrique occidentale (Loanda et Benguella). Proc
Zool Soc London 35:843–846
Bocage JVB (1873) Mélanges erpétologiques. Sur quelques Reptiles et Batraciens nouveux,
rares ou peu connues de I’Afrique occidentale. Jornal de Sciências Mathemáticas, Physicas e
Naturaes. Lisboa 4(1):209–227
Bocage JVB (1879a) Reptiles et batraciens nouveaux d’ Angola. Jornal de Sciências Mathemáticas,
Physicas e Naturaes. Lisboa 7(26):97–99
Bocage JVB (1879b) Subsidio para a fauna das possessões portuguezas d’África occidental. Jornal
de Sciências Mathemáticas, Physicas e Naturaes. Lisboa (1) 7:85–95
Bocage JVB (1882) Reptiles rares ou nouveaux d’Angola. Jornal de Sciencias, Mathemáticas,
Physicas e Naturaes. Lisboa (1) 8:299–304
Bocage JVB (1893) Diagnose de quelques nouvelles espéces de reptiles et batraciens d’ Angola.
Jornal de Sciências Mathemáticas, Physicas e Naturaes. Lisboa (2) 10:115–121
Bocage JVB (1895a) Herpétologie d’Angola et du Congo. Lisbonne, Imprimerie Nationale, 203
pp, 19 pls
Bocage JVB (1895b) Sur une espêce de Crapaud à ajouter à la faune herpétologique d’Angola.
Jornal de Sciências Mathemáticas, Physicas e Naturaes. Lisboa (2) 4:51–53
Bocage JVB (1896a) Mamíferos, aves e réptis da Hanha, no sertão de Benguella. Jornal de
Sciências Mathemáticas, Physicas e Naturaes. Lisboa (2) 14:105–114
Bocage JVB (1896b) Répteis de algumas possessões portuguesas de Africa que existem no Museu
de Lisboa. Jornal de Sciências Mathemáticas, Physicas e Naturaes 2:65–104
Bocage JVB (1897a) Mamíferos, aves e reptis da Hanha, no sertão de BengueIla. Segunda lista.
Jornal de Sciências Mathemáticas, Physicas e Naturaes. Lisboa (2):207–211
Bocage JVB (1897b) Mamíferos, réptis e batrachios d’África de que existem exemplares típi-
cos no Museu de Lisboa. Jornal de Sciências Mathemáticas, Physicas e Naturaes. Lisboa (2)
4:187–206
Boulenger GA (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the
British museum, 2nd edn. Taylor and Francis, London
Boulenger GA (1905) A list of the batrachians and reptiles collected by Dr. W. J. Ansorge in
Angola, with descriptions of new species. Ann Mag Nat Hist Ser 7 16(92):8–115
Boulenger GA (1907a) Descriptions of three new lizards and a new frog, discovered by Dr. W
J. Ansorge in Angola. Ann Mag Nat Hist Ser 7 19:212–214
Boulenger GA (1907b) Description of a new frog discovered by Dr. W. J. Ansorge in Mossamedes,
Angola. Ann Mag Nat Hist Ser 7 20:109
Boulenger GA (1919) On Rana ornatissima, Bocage, and R. ruddi, Blgr. Trans Royal Soc S Afr
8:33–37
Branch WR, Bauer AM (2005) The life and herpetological contributions of Andrew Smith.
pp. 1-19 in Smith, A. The Herpetological Contributions of Sir Andrew Smith. Society for the
Study of Amphibians and Reptiles, Villanova, PA. iv + 84 pp
Branch WR, Conradie WC (2015) Herpetofauna da região da Lagoa Carumbo (Herpetofauna of
the Carumba Lagoon Area). In: Huntley BJ (ed), Relatório sobre a Expedição Avaliação rápida
da Biodiversidade de região da Lagoa Carumbo, Lunda-Norte – Angola, República de Angola.
Ministério do Ambiente, pp 194–209, 219p
Branch WR, Baptista N, Keates CW, et al (2019) Rediscovery, taxonomic status, and phylogenetic
relationships of two rare and endemic snakes (Serpentes: Psammophinae) from the Angolan
Escarpment. Zootaxa, in press
Brito D (2010) Overcoming the Linnean shortfall: data deficiency and biological survey priorities.
Basic Appl Ecol 11(8):709–713
Broadley DG (1971) The reptiles and amphibians of Zambia. The Puku, Occ Pap Dept Game Fish
Zambia 7:1–143
Brooks C (2012) Biodiversity survey of the upper Angolan Catchment of the Cubango-Okavango
River Basin. USAid-Southern Africa. 151 pp
Brooks C (2013) Trip report: aquatic biodiversity survey of the lower Cuito and Cuando river
systems in Angola. USAid-Southern Africa. 43 pp
Cei JM (1977) Chaves para uma identificação preliminar dos batráquios anuros da R. P. de Angola.
Boletim da Sociedade Portuguesa de Ciências Naturais 17:5–26
Ceríaco LMP, Bauer AM, Blackburn et al (2014a) The herpetofauna of the Capanda Dam region,
Malanje, Angola. Herpetol Rev 45(4):667–674
Ceríaco LMP, Blackburn DC, Marques MP et al (2014b) Catalogue of the amphibian and reptile
type specimens of the Museu de História Natural da Universidade do Porto in Portugal, with
some comments on problematic taxa. Alytes 31(1):13–36
Ceríaco LMP, Bauer AM, Heinicke MP et al (2016a) Geographical distributions:

Ptychadenidae, Ptychadena mapacha Channing, 1993 – Mapacha ridged frog in Namibia.
Afr Herp News 63:19–20
Ceríaco LMP, de Sá SAC, Bandeira SA et al (2016b) Herpetological survey of Iona National Park
and Namibe regional natural park, with a synoptic list of the amphibians and reptiles of Namibe
Province, Southwestern Angola. Proc Calif Acad Sci 63(2):15–61
Ceríaco LMP, Marques MP, Bandeira SA et al (2016c) Anfíbios e répteis do Parque Nacional da
Cangandala. Instituto Nacional da Biodiversidade e Áreas de Conservação & Museu Nacional
de História Natural e da Ciência, 96 pp
Ceríaco LMP, Marques MP, Bandeira S et al (2018a) A new earless species of Poyntonophrynus
(Anura: Bufonidae) from the Serra da Neve Inselberg, Namibe Province, Angola. Zookeys
780:109–136
Ceríaco LMP, Marques MP, Bandeira S et al (2018b) Herpetological survey of Cangandala
National Park, with a synoptic list of the amphibians and reptiles of Malanje Province, Central
Angola. Herpetological Review 49(3):408–431
Channing A (1993) A new grass frog from Namibia. S Afr J Zool 28:142–145
Channing A (1999) Historical overview of amphibian systematics in Southern Africa. Trans Royal
Soc S Afr 54(1):121–135
Channing A (2001) Amphibians of central and southern Africa. Cornell University Press,
New York, 470 pp
Channing A, Baptista N (2013) Amietia angolensis and A. fuscigula (Anura: Pyxicephalidae) in
southern Africa: a cold case reheated. Zootaxa 3640(4):501–520
Channing A, Griffin M (1993) An annotated checklist of the frogs of Namibia. Modoqua
18:101–116
Channing A, Vences M (1999) The advertisement call, breeding biology, description of the tadpole
and taxonomic status of Bufo dombensis, a little-known dwarf toad from southern Africa. S Afr
J Zool 34:74–79
Channing, A., and D. G. Broadley. 2002. A new snout-burrower from the Barotse Floodplain
(Anura: Hemisotidae: Hemisus). Journal of Herpetology 36: 367–372.
Channing A, Rödel MO, Channing J (2012) Tadpoles of Africa: the biology and identification of
all known tadpoles in sub-Saharan Africa. Edition Chimaira, Frankfurt, 401pp
Channing A, Hillers A, Lötters S et al (2013) Taxonomy of the super-cryptic Hyperolius nasutus
group of long reed frogs of Africa (Anura: Hyperoliidae), with descriptions of six new species.
Zootaxa 3620(3):301–350
Channing A, Dehling JM, Lötters S et al (2016) Species boundaries and taxonomy of the African
river frogs (Amphibia: Pyxicephalidae: Amietia). Zootaxa 4155(1):1–76
Clark VR, Barker NP, Mucina L (2011) The great escarpment of southern Africa: a new frontier for
biodiversity exploration. Biodivers Conserv 20(12):2543–2561
Coetzer W (2017) Occurrence records of southern African aquatic biodiversity. Version 1.10. The
south African Institute for Aquatic Biodiversity. Occurrence dataset https://doi.org/10.15468/
pv7vds. Accessed via GBIF.org
Conradie W, Bills R (2017) Wannabe Ranid: notes on the morphology and natural history of the
Lemaire’s toad (Bufonidae: Sclerophrys lemairii). Salamandra 53(3):439–444
Conradie W, Branch WR, Measey GJ et al (2012) A new species of Hyperolius Rapp, 1842
(Anura: Hyperoliidae) from the Serra da Chela mountains, South-Western Angola. Zootaxa
3269(1):1–17
Conradie W, Branch WR, Tolley KA (2013) Fifty Shades of Grey: giving colour to the poorly
known Angolan Ashy reed frog (Hyperoliidae: Hyperolius cinereus), with the description of a
new species. Zootaxa 2636(3):201–223
Conradie W, Bills R, Branch WR (2016) The herpetofauna of the Cubango, Cuito, and lower
Cuando river catchments of South-Eastern Angola. Amphibian Reptile Conserv 10(2):6–36
Crawford-Cabral JC (1966) Some new data on Angolan Muridae. Zool Afr 2:193–203
Crawford-Cabral J, Mesquitela LM (1989) Índice toponímico de colheitas zoológicas em Angola.

Instituto de Investigação Cientifica Tropical, Centro de Zoologia, Lisbon, 206
Cunningham M, Cherry MI (2004) Molecular systematics of African 20-chromosome toads
(Anura: Bufonidae). Mol Phylogenet Evol 32(3):671–685
Curtis B, Roberts KS, Griffin M et al (1998) Species richness and conservation of Namibian fresh-
water macro-invertebrates, fish and amphibians. Biodivers Conserv 7(4):447–466
Davic RD, Welsh JHH (2004) On the ecological roles of salamanders. Annu Rev Ecol Evol Syst
35:405–434
Dehling JM, Sinsch U (2013a) Diversity of Ptychadena in Rwanda and taxonomic status of P.
chrysogaster Laurent, 1954 (Amphibia, Anura, Ptychadenidae). Zoo Keys 356:69–102
Dehling JM, Sinsch U (2013b) Diversity of Ridged Frogs (Anura: Ptychadenidae: Ptychadena
spp.) in wetlands of the upper Nile in Rwanda: morphological, bioacoustic, and molecular
evidence. Zoologischer Anzeiger 253(2):143–157
Du Preez L, Carruthers V (2009) A complete guide to the frogs of Southern Africa. Struik
Publishers, Cape Town, 488 pp
Du Preez L, Carruthers V (2017) Frogs of Southern Africa: a complete guide. Struik Publishers,
Cape Town, 520 pp
Ernst R, Nienguesso ABT, Lautenschlaeger T et al (2014) Relicts of a forested past: southernmost
distribution of the hairy frog genus Trichobatrachus Boulenger, 1900 (Anura: Arthroleptidae)
in the Serra do Pingano region of Angola with comments on its taxonomic status. Zootaxa
3779(2):297–300
Ernst R, Schmitz A, Wagner P, Branquima MF et al (2015) A window to Central African forest
history: distribution of the Xenopus fraseri subgroup south of the Congo Basin, including a first
country record of Xenopus andrei from Angola. Salamandra 52(1):147–155
Ferreira JB (1897) Lista dos reptis e amphibios que fazem parte da última remessa de J. d’Anchieta.
Jornal de Sciências Mathemáticas, Physicas e Naturaes 5(2):240–246
Ferreira JB (1900) Sobre alguns exemplares pertencentes à fauna do norte de Angola (Reptis,
Batrachios, Aves e Mammiferos). Jornal de Sciências Mathemáticas, Physicas e Naturaes,
Lisboa 2(6):48–54
Ferreira JB (1904) Reptis e amphibios de Angola da região ao norte do Quanza (Collecção Newton –
1903). Jornal de Sciências Mathemáticas, Physicas e Naturaes, Segunda Série 7(26):111–117
Ferreira JB (1906) Algumas espécies novas ou pouco conhecidas de amphibios e reptis de Angola
(Collecção Newton – 1903). Jornal de Sciências Mathemáticas, Physicas e Naturaes, Segunda
Série 7(26):159–171
Frétey T, Dewynter M, Blanc CP (2011) Amphibiens d’Afrique central et d’Angola. Clé de
détermination ilustrée desamphibiens du Gabo et du Mbini/Illustrated identification key of
the amphibians from Gabon and Mbini. Biotope, Mèze/Muséum national d’Histoire naturelle,
Paris, 232 pp
Frost DR (2018) Amphibian species of the world: an online reference. Version 6.0. Electronic
database accessible at http://research.amnh.org/herpetology/amphibia/index.html. American
Museum of Natural History, New York
Frost DR, Grant T, Faivovich J et al (2006) The amphibian tree of life. Bull Am Mus Nat Hist
297:1–370
Furman BL, Bewick AJ, Harrison TL et al (2015) Pan-African phylogeography of a model organ-
ism, the African clawed frog ‘Xenopus laevis’. Mol Ecol 24(4):909–925
Greenbaum E, Meece J, Reed KD et al (2014) Amphibian chytrid infections in non-forested habi-
tats of Katanga, Democratic Republic of the Congo. Herpetol Rev 45:610–614
Günther ACLG (1865 ‘1864’) Descriptions of new species of batrachians from West Africa. Proc
Zool Soc London 3:479–482
Guttman SI (1967) Transferrin and hemoglobin polymorphism, hybridization and introgression
in two African toads, Bufo regularis and Bufo rangeri. Comp Biochem Physiol 23(3):871–877
Haacke WD (1999) Geographical distribution: Ptychadena mapacha Channing, 1993 – Mapacha
Grass Frog. Afr Herp News 30:35
Hall BP (1960) The faunistic importance of the scarp of Angola. Ibis 102(3):420–442
Heinicke MP, Ceríaco LM, Moore IM et al (2017) Tomopterna damarensis (Anura: Pixicephalidae)
is broadly distributed in Namibia and Angola. Salamandra 53(3):461–465
Hellmich W (1957a) Die reptilienausbeute der Hamburgischen Angola Expedition. Mitteilungen
aus dem Hamburgischen Zoologischen Museum und Institut 55:39–80
Hellmich W (1957b) Herpetologische Ergebnisse einer Forschungsreise in Angola.
Herrmann HW, Branch WR (2013) Fifty years of herpetological research in the Namib Desert and
Namibia with an updated and annotated species checklist. J Arid Environ 93:94–115
Huntley BJ (1974) Outlines of wildlife conservation in Angola. J S Afr Wildl Manag Assoc
4:157–166
tion: a modern synthesis. Springer, Cham
IUCN Red List of Threatened Species Version 2017-2. www.iucnredlist.org
IUCN Red List of Threatened Species. Version 2017-3. www.iucnredlist.org
Jongsma CF, Barej MF, Barratt CD et al (2018) Diversity and biogeography of frogs in the genus
Amnirana (Anura: Ranidae) across sub-Saharan Africa. Mol Phylogenet Evol 120:274–285
Jordan K (1936) Dr Karl Jordan’s expedition to South-West Africa and Angola. Narrative.
Novitates Zooligicae 40:17–62, 2 maps, 5 pls
Jürgens N, Strohbach B, Lages F et al (2018) Biodiversity observation – an overview of the cur-
rent state and first results of biodiversity monitoring studies. In: Revermann R, Krewenka KM,
Schmiedel U et al (eds) Climate change and adaptive land management in southern Africa –
assessments, changes, challenges, and solutions, Biodiversity & ecology, vol 6, pp 382–396
Köhler J, Vieites DR, Bonett RM et al (2005) New amphibians and global conservation: a boost in
species discoveries in a highly endangered vertebrate group. AIBS Bull 55(8):693–696
Lamotte M, Perret JL (1961) Les formes larvaires de quelques espèces de Leptopelis: L. aubryi, L.
viridis, L. anchietae, L. ocellatus et L. calcaratus. Bulletin de l’Institute fondamental d’Afrique
noire, Sér. A 23:855–885
Laurent RF (1950) Reptiles et Batraciens de la region de Dundo (Angola du Nord-Est). Publicações
culturais da Companhia de Diamantes de Angola 6:126–136
Laurent RF (1954) Reptiles et Batraciens de la région de Dundo (Angola) (Deuxième Note).
Publicações culturais da Companhia de Diamantes de Angola 23:35–84
Laurent RF (1964) Reptiles et Amphibiens de l’Angola (Troisième contribution). Publicações cul-
turais da Companhia de Diamantes de Angola 67:11–165
Marques MP, Ceríaco LMP, Bauer AM et al (2014) Geographic distribution of amphibians &
reptiles of Angola: towards an Atlas of the Angolan Herpetofauna. 12th conference of the
Herpetological Association of Africa, Gobabeb, Namibia
Marques MP, Ceríaco LMP, Blackburn DC et al (2018) Diversity and distribution of the amphibians
and terrestrial reptiles of Angola atlas of historical and bibliographic records (1840¬–2017).
Proceedings of the California academy of sciences, Series 4, Volume 65, Supplement II: 1-501
Mertens R (1938a) Amphibien und Reptilien aus Angola, gesammelt von W. Schack.
Senckenbergiana 20:425–443
Mertens R (1938b) Herpetologische Ergebnisse einer Reise nach Kamerun. Abhandlungen der
Senckenbergischen Naturforschenden Gesellschaft, Frankfurt am Main 442:1–52
MHNG – Muséum d’histoire naturelle de la Ville de Genève (2017) Partial Amphibians Collection.
Occurrence Dataset https://doi.org/10.15468/iftvxc. Accessed via GBIF.org
Minter LR, Netherlands EC, Du Preez LH (2017) Uncovering a hidden diversity: two new species
of Breviceps (Anura: Brevicipitidae) from northern KwaZulu-Natal, South Africa. Zootaxa
4300:195–216
Monard A (1937) Contribuition à la Batrachologie d’Angola. Bulletin de la Société Neuchâteloise
des Sciences Naturelles 62:1–59
Nagy ZT, Kusamba C, Collet M et al (2013) Notes on the herpetofauna of Western Bas-Congo,
Democratic Republic of the Congo. Herpetology Notes 6:413–419
Ohler A (1996) Systematics, morphometrics and biogeography of the genus Aubria (Ranidae,
Pyxicephalinae). Alytes 13:141–166
Parker HW (1936) Dr. Karl Jordan’s expedition to South West Africa and Angola: herpetological
collection. Novitates Zoologicae 40:115–146
Passmore NI (1972) Intergrading between members of the “regularis group” of toads in South
Africa. J Zool 167(2):143–151
Perret JL (1975) Les sous-espèces d’Hyperolius ocellatus Günther (Amphibia, Salientia). Annales
de la Faculté des Sciences du Cameroun 20:23–31
Perret JL (1976) Révision des amphibiens africains et principalement des types, conservés au
Musée Bocage de Lisbonne. Arquivos do Museu Bocage, Segunda Série 6(2):15–34
Perret JL (1977) Les Hylarana (Amphibiens, Ranidés) du Cameroun. Rev Suisse Zool 84:841–868
Perret JL (1996) Sur un énigmatique batracien d’Angola. Societé Neuchâteloise des Sciences
Naturelles 119:95–100
Peters WCH (1877) Übersicht der Amphibien aus Chinchoxo (Westafrika), welche von
der Afrikanischen Gesellschaft dem Berliner zoologischen Museum übergeben sind.
Monatsberichte der Königlichen Preussische Akademie des Wissenschaften zu Berlin
1877:611–621
Peters WCH (1882) Neue Batrachier (Amblystoma Krausei, Nyctibatrachus sinensis, Bufo buch-
neri). Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 1882:145–148
Pickersgill M (2007) Frog search. Results of expeditions to Southern and Eastern Africa from
1993–1999. Frankfurt contributions to natural history 28. Edition Chimaira, Frankfurt
Pietersen DW, Pietersen EW, Conradie W (2017) Preliminary herpetological survey of Ngonye
Falls and surrounding regions in southwestern Zambia. Amphibian Reptile Conserv 11(1)
[Special Section]:24–43 (e148)
Poynton JC (1964) The Amphibia of Southern Africa: a faunal study. Ann. Natal Mus 17:1–334
Poynton JC (1970) Guide to the Ptychadena (Amphibia: Ranidae) of the southern third of Africa.
Ann. Natal Mus 20(2):365–375
Poynton JC, Broadley DG (1985a) Amphibia Zambesiaca 1. Scolecomorphidae, Pipidae,
Microhylidae, Hemisidae, Arthroleptidae. Ann. Natal Mus 26(2):503–553
Poynton JC, Broadley DG (1985b) Amphibia Zambesiaca 2. Ranidae. Ann. Natal Mus
27(1):115–181
Poynton JC, Broadley DG (1987) Amphibia Zambesiaca 3. Rhacophoridae and Hyperoliidae. Ann.
Natal Mus 28(1):161–229
Poynton JC, Broadley DG (1988) Amphibia Zambesiaca, 4. Bufonidae. Ann. Natal Mus
29(2):447–490
Poynton JC, Broadley DG (1991) Amphibia Zambesiaca 5. Zoogeography. Ann. Natal Mus
32:221–277
Poynton JC, Haacke WD (1993) On a collection of amphibians from Angola, including a new spe-
cies of Bufo Laurenti. Ann Transv Mus 36(2):9–16
Poynton JC, Loader SP, Conradie W et al (2016) Designation and description of a neotype of
Sclerophrys maculata (Hallowell, 1854), and reinstatement of S. pusilla (Mertens, 1937)
(Amphibia: Anura: Bufonidae). Zootaxa 4098(1):73–94
Regester KJ, Lips KR, Whiles MR (2006) Energy flow and subsidies associated with the com-
plex life cycle of ambystomatid salamanders in ponds and adjacent forest in southern Illinois.
Oecologia 147(2):303–314
Rochebrune AT (1885) Vertebratorum novorum vel minus cognitorum orae Africae occidentalis
incolarum. Diagnoses (1). Bulletin de la Société Philomathique de Paris 7(9):86–99
Royal Belgian Institute of Natural Sciences (2017). RBINS DaRWIN. Occurrence Dataset https://
doi.org/10.15468/qxy4mc. Accessed via GBIF.org
Royal Museum for Central Africa, Belgium (2017). RMCA-HERP. Occurrence Dataset https://doi.
org/10.15468/0inmlf. Accessed via GBIF.org
Ruas C (1996) Contribuição para o conhecimento da fauna de batráquios de Angola. Garcia de

Orta, Série de Zoologia 21(1):19–41
Ruas C (2002) Batráquios de Angola em colecção no Centro de Zoologia. Garcia de Orta, Série de
Zoologia 24(1–2):139–154
SASSCAL ObservationNet (2017) http://www.sasscalobservationnet.org/ consulted on March 6th
2018
Scheinberg L, Fong J (2017) CAS herpetology (HERP). Version 33.10. California Academy of
Sciences. Occurrence dataset https://doi.org/10.15468/bvoyqy. Accessed via GBIF.org
Schick S, Kielgast J, Rödder D et al (2010) New species of reed frog from the Congo basin with
discussion of paraphyly in Cinnamon-belly reed frogs. Zootaxa 2501:23–36
Schiøtz A (1999) Treefrogs of Africa. Editions Chimaira, Frankfurt am Main, 350 pp
Schiøtz A, Van Daele P (2003) Notes on the treefrogs (Hyperoliidae) of North-Western Province,
Zambia. Alytes 20:137–149
Schmidt KP (1936) The amphibians of the Pulitzer-Angola expedition. Ann Carnegie Mus
25:127–133
Steindachner F (1867) Reise der österreichischen Fregatte Novara um die Erde in den Jahren 1857,
1858, 1859 unter den Bafehlen des Commodore B. von Wüllerstorf-Urbair. Zologischer Theil.
1. Amphibien. Wien: K. K. Hof- und Staatsdruckerei. 70 p + 5pls
Tandy M, Keith R (1972) Bufo of Africa. In: Blair WF (ed) Evolution in the Genus Bufo. University
of Texas Press, Austin/London, pp 119–170
Turner AA, Channing A (2017) Three new species of Arthroleptella Hewitt, 1926 (Anura:
Pyxicephalidae) from the Cape Fold Mountains, South Africa. Afr J Herpetol 66:53–78
Uyeda JC, Drewes RC, Zimkus BM (2007) The California Academy of Sciences Gulf of Guinea
expeditions (2001, 2006) VI. Proc Calif Acad Sci 58(13–22):367
Van Dijk DE (1966) Systematic and field keys to the families, genera and described species of
southern African anuran tadpoles. Ann Natal Mus 18:231–286
Van Dijk DE (1971) A further contribution to the systematics of Southern African anuran tad-
poles—the genus Bufo. Ann Natal Mus 21:71–76
Waddle JH (2006) Use of amphibians as ecosystem indicator species. PhD Thesis. University of
Florida, Gainesville
Whiles MR, Lips KR, Pringle CM et al (2006) The effects of amphibian population declines on
the structure and function of Neotropical stream ecosystems. Front Ecol Environ 4(1):27–34
Zimkus BM, Rödel MO, Hillers A (2010) Complex patterns of continental speciation: molec-
ular phylogenetics and biogeography of sub-Saharan puddle frogs (Phrynobatrachus). Mol
Phylogenet Evol 55(3):883–900
Zimkus BM, Lawson LP, Barej MF et al (2017) Leapfrogging into new territory: how Mascarene
ridged frogs diversified across Africa and Madagascar to maintain their ecological niche. Mol
Chapter 13
The Reptiles of Angola: History, Diversity,
Endemism and Hotspots
William R. Branch, Pedro Vaz Pinto, Ninda Baptista, and Werner Conradie
Abstract This review summarises the current status of our knowledge of Angolan
reptile diversity, and places it into a historical context of understanding and growth.
It is compared and contrasted with known diversity in adjacent regions to allow
insight into taxonomic status and biogeographic patterns. Over 67% of Angolan
reptiles were described by the end of the nineteenth century. Studies stagnated dur-
ing the twentieth century but have increased in the last decade. At least 278 reptiles
are currently known, but numerous new discoveries have been made during recent
surveys, and many novelties await description. Although lizard and snake diversity
is currently almost equal, most new discoveries occur in lizards, particularly geckos
and lacertids. Poorly known Angolan reptiles and others from adjacent regions that
W. R. Branch (deceased)
P. Vaz Pinto
N. Baptista
Instituto Superior de Ciências da Educaҫão da Huíla, Rua Sarmento Rodrigues,
Lubango, Angola
e-mail: nindabaptista@gmail.com
W. Conradie (*)
School of Natural Resource Management, Nelson Mandela University, George, South Africa
e-mail: werner@bayworld.co.za

https://doi.org/10.1007/978-3-030-03083-4_13
284 W. R. Branch et al.
may occur in the country are highlighted. Most endemic Angolan reptiles are lizards
and are associated with the escarpment and southwest arid region. Identification of
reptile diversity hotspots are resolving but require targeted surveys for their
delimitation and to enable protection. These include the Kaokoveld Centre of
Endemism, Angolan Escarpment and the Congo forests of the north. The fauna of
Angola remains poorly known and under-appreciated, but it is already evident that
it forms an important centre of African reptile diversity and endemism.
Keywords Herpetofauna · History · Priority studies · Reptilia · Review
Introduction
Systematic biology is the backbone of biology in that it describes the taxa and their relation-
ships which then serve as the objects of research. (Uetz and Stylianou 2018)
The extensive territory of Angola is for herpetologists one of the least known parts of
Africa. This is particularly unfortunate because there are indications that it may be one of
the most interesting areas of the continent. (Gans 1976)
The need to classify things is a basic human need. Initially it was simply utilitarian,
driven by the necessity for rural people to know what was edible, venomous, poi-
sonous, or useful. As the world’s diverse civilizations developed all were faced with
the need to refine this knowledge, and as they migrated, came together, experienced
new habitats and new life forms, the need for classification became essential. Only
when new technological innovations in such things as sailing and weaponry allowed
the reach of various nations to become global, did universal categorisation and clas-
sification really need to be become standardised. Driven by the Enlightenment and
during the rise of critical thinking and the scientific revolution there began the first
steps in developing a universally recognised system for classifying Life. The current
classification system was initiated by Carl Linnaeus (1707–1778), a Swedish bota-
nist, physician, and zoologist who formalised the modern system of naming organ-
isms, now known as binomial nomenclature. During the 250+ years since the
Linnaean revolution, the rules used to name this diversity have been refined and
modified, and increasingly sophisticated methods have been developed to gain
insight into the relationships of its components.
Since the equally important Darwinian Revolution and awareness of the evolution-
ary relatedness of Life, modern systematics places emphasis on revealing patterns of
relationship among groups, often figuratively represented as trees or cladograms. The
branches of these ‘trees’ are monophyletic when they include only the descendants of
a common ancestor. All modern classifications comprise hypotheses represented as
phylogenies of nested groups (clades) exhibiting monophyly. Biochemical adjuncts to
traditional taxonomy have proliferated since the middle of the last century, but detailed
multi-taxon genomic analysis, linked with increasingly sophisticated computer pro-
cessing of sequence data, is a phenomenon of the twenty-first century. These recent
technological advances have allowed objective assessments of hypotheses of phyloge-
netic relationships. It is important to emphasise that the assignment of any individual
13 The Reptiles of Angola: History, Diversity, Endemism and Hotspots 285
specimen, first to a species and then to any higher taxonomic group, tests hypotheses
of relationships. The placement of a specimen at any level in the nomenclatural hier-
archy, from species to phylum, must conform to the definitions of those groups. It
should be stressed, moreover, that taxonomy is a dynamic discipline and that every
assignment of a specimen to a species or higher taxonomic group is a hypothesis of
relationship. It is always subject to revision in the light of new knowledge.
It has become increasingly obvious that species may result from different mecha-
nisms and histories, and there is increasing use of evolutionary and phylogenetic
species concepts to reflect hypotheses about the boundaries of past and present gene
transfer within evolutionary lineages of Life’s diversity. Many phylogenies based on
molecular/genetic data conflict with historical ideas of relationships previously
based solely or largely on morphological analysis. It is evident that morphology is
often conservative (maintained by selective pressures) that may mask underlying
cryptic genetic diversity. This awareness has led to the burgeoning description of
new species, genera, and higher categories.
History: Early Research on Angolan Reptiles
Early studies on the Angolan herpetofauna have been summarised by Baptista et al.
(2019) in this volume, also see the recently published Angola Reptile and Amphibian
Atlas (Marques et al. 2018). Other recent summaries can be found in introductions
to regional herpetofaunas, e.g. Ceríaco et al. (2014a, 2016a) and Conradie et al.
(2016). To avoid duplication much will not be repeated here, where instead empha-
sis is placed on the main publications discussing Angolan reptiles during and after
the colonial period, and particularly the periodic attempts to overview its diversity
(Bocage 1895; Monard 1937).
José Vincente Barbosa du Bocage is rightly known as the ‘Father of Angolan
Herpetology’. He was, however, more than just a scientist and for much of his life
held multiple positions in government, administration and science, often simultane-
ously. De Almeida (2011) reviews diverse aspects of his multifaceted life; Madruga
(2013) discusses the development of his scientific career; and Gamito-Marques
(2017) explains Bocage’s role in the foundation of National Museum of Lisbon and
his importance, via his contacts with collectors in various Portuguese colonies as
well as other zoologists at major European museums, in developing the collections
and status of the museum. A list of his scientific publications is available at
TRIPLOV (2018). In his first attempt to review the known Angolan herpetofauna he
listed 26 reptile species from the Congo and 57 reptiles and amphibians from Angola
in the Museu de Lisboa collection (Bocage 1866). Part of this material came from
José de Anchieta collected during a zoological expedition in 1864 to Rio Quilo,
Cabinda, and the coast of Loango, and other material was collected by Bayão
Pinheiro when a military commander in the Duque de Bragança (Calandula) dis-
trict. After nearly 30 years of subsequent study, during which Bocage published at
least a paper a year on the herpetofauna of Portugal’s African colonies (see full list
in TRIPLOV 2018), he again summarised the Angolan herpetofauna in his mono-
graphic Herpetology d’Angola et du Congo (Bocage 1895). In this review he more

than trebled his previous summary, recording 143 reptile and 39 amphibian species
from Angola. Of these Bocage himself described 40 reptile species that are still
considered valid, and of which no less that 26 (65%) had been collected by José de
Anchieta, seven of which are still named in his honour. Following Bocage’s retire-
ment, herpetological studies at Lisbon were continued, but with less intensity, by
Bethencourt Ferreira (1897, 1900, 1903, 1904, 1906), who added a number of addi-
tional species to the country list but described only one new snake, Typhlops bocagei
(Ferreira 1904), that was later placed in the synonymy of Afrotyphlops lineolatus
(Jan, 1864).
George A Boulenger of the British Natural History Museum remains the most
prolific herpetologist of all time, and described a remarkable 659 reptiles still rec-
ognised today – no less than 5% of the world’s 13,000+ known reptiles (Uetz and
Stylianou 2018). In 1905 Boulenger published a note on the amphibians and reptiles
collected by WJ Ansorge during a prolonged visit (1903–1905) to Angola (Boulenger
1905). The material included three new species of frogs and a snake, of which two,
Rana ansorgii (= Ptychadena ansorgii) and Psammophis ansorgii, were named in
the collector’s honour. Two years later Boulenger described another three lizards
and a frog collected by Ansorge, including the gecko Phyllodactylus ansorgii, the
skinks Mabuia ansorgii (Trachylepis sulcata ansorgii) and Mabuia laevis
(Trachylepis laevis), and the frog Rana bunoderma (= Ptychadena bunoderma)
(Boulenger 1907a), soon followed (Boulenger 1907b) by another new frog from
‘Mossâmedes’ (actually Catequero, Cunene), Rana cryptotis (= Tomopterna crypto-
tis). Boulenger (1915) also prepared a List of the snakes of Belgian and Portuguese
Congo, Northern Rhodesia and Angola in which he documented 139 snakes for the
region, of which only 57 were considered to occur in Angola. This was lower than
Bocage’s (1895) assessment, but by this time Boulenger had completed his mono-
graphic, three volume ‘Catalogue’ of the World’s snakes (Boulenger 1893, 1894,
1896), in which he had synonymised many of Bocage’s taxa or shown some to be
synonyms. This included Angola’s most iconic snake, Vipera heraldica, which
Boulenger (1896) surprisingly and incorrectly synonymised with Bitis peringueyi,
and Bogert (1940) continued the confusion. There it languished until revalidated by
Mertens (1958).
The dramatic discovery of the Giant Sable in Angola early in the twentieth cen-
tury led to numerous expeditions to collect them as hunting trophies or taxidermy
specimens for European and American museum dioramas. Some expeditions col-
lected additional fauna, although the herpetological collections of the Vernay
Angola Expedition (VAE 1925) seem to have been plagued by poor documentation.
Although various specimens were incorporated in diverse taxonomic reviews, no
publication dedicated to the full herpetological results was produced. Bogert (1940)
incorporated 202 VAE snakes into his review of African snakes, but 10 species and
42 (21%) specimens lacked detailed locality information and were simply listed as
from ‘Angola’. Three new snakes were described from Angolan material: Vernay’s
File Snake (Mehelya vernayi, = Limaformosa vernayi Broadley et al. 2018) from
Hanha, and Cowles’ Shield Cobra (Aspidelaps lubricus cowlesi) and the Western
Banded Spitting Cobra (Naja nigricollis nigricincta) from Munhino. The first two
have subsequently been discovered at numerous localities in northern Namibia
(Haacke 1981; Broadley and Baldwin 2006), but remain known in Angola only
from the type or a few other specimens, respectively. Loveridge (1944) described
two new geckos (Afroedura bogerti and Pachydactylus scutatus angolensis) on VAE
material, and Stanley et al. (2016) discussed VAE Cordylus material labelled simply
as ‘Angola’ and that they assigned morphologically to a new species, Cordylus
namakuiyus, discovered in the Namibe region. The description of at least one other
new species from old VAE material is also in preparation (Ichnotropis sp. Branch in
prep.).
The main targets of the Pulitzer Angola Expedition (1930–1931) were birds and
mammals, but Rudyerd Boulton, who had earlier accompanied the VAE, also col-
lected reptiles and amphibians. Karl Schmidt (1933, 1936) documented the reptiles
and amphibians, respectively, collected during the expedition from diverse sites in
the centre and south of the country. The reptiles included two new species, but
Lygodactylus laurae was quickly synonymised when Schmidt realised he had over-
looked Bocage’s (1896) earlier description of L. angolensis. His description of
Rhoptropus boultoni not only honoured Rudyerd’s contribution to the collection of
Angolan reptiles, but was also the first record of this interesting diurnal rupicolous
gecko genus for the country. Two other new subspecies were also described, of
which Pachydactylus bibronii pulizerae was latter transferred to Chondrodactylus
(Bauer and Lamb 2005), and has also been recently validated as a full species, C.
pulizerae, that is mainly restricted to Angola but also extends into northern Namibia
(Heinz 2011; Ceríaco et al. 2014a). The Angolan Savanna Monitor (Varanus albigu-
laris angolensis) was described by Schmidt (1933) from ‘Gaúca, Bihe’ (= Zaúca
River, Malanje; Crawford-Cabral and Mesquitela 1989). Although additional mate-
rial had been collected the validity of the morphological diagnosis (small nuchal
scales and larger body scales) has not been reassessed, and neither has its genetic
affinities.
In 1933–1934 Karl Jordan, an entomologist, collected through northern Namibia
and Angola and made well documented collections and published a detailed itiner-
ary of his trip (Jordan 1936). Among these was an important herpetological collec-
tion, particularly from scarp forest habitats at Congulo and Quirimbo. These were
studied by Parker (1936), who recorded diverse Congo Basin snakes previously
unknown from Angola; e.g. Philothamnus heterodermus, Thelotornis kirtlandii,
Toxicodryas blandingii, T. pulverulenta, Pseudohaje goldii, Chamaelycus parkeri
and Hormonotus modestus, as well as the new wolf snake Lycophidion ornatum.
Swiss zoologist Albert Monard explored Angola during two extensive trips (July
1928–February 1929 and April 1932–October 1933) that resulted in extensive
reviews of Angolan birds (Monard 1934), mammals (Monard 1935), reptiles
(Monard 1937), and amphibians (Monard 1938). Ceríaco et al. (2016a, b) note that
Monard was so inspired by Angolan biodiversity that he unsuccessfully champi-
oned for the development of a local Natural History Museum that he offered to
direct and manage. His detailed reptile ‘Contribution’ (Monard 1937) was the first
overview of Angolan reptiles subsequent to Bocage’s (1895) monograph, and within
it Monard presented taxonomic updates and also initiated the first attempts to
generate a biogeographic overview of the herpetofauna (see section below). He
noted that only 19 lizards, 10 snakes and a single terrapin had been added to the
reptile fauna of Angola, and he even envisaged (incorrectly!) that most of the knowl-
edge on the subject was complete, and he therefore concentrated on understanding
biogeographic patterns. However, eight of the 19 additional lizards and three of the
10 snakes he included had already been synonymised (Boulenger 1915) or subse-
quently were. Certainly Monard seemed little interested in taxonomy and described
relatively few novelties, only one of which may remain valid. The rejected taxa
include: the worm lizards Amphisbaenia ambuellensis (= Zygaspis quadrifrons),
Monopeltis granti kuanyamarum (= Dalophia pistillum), M. devisi (= Monopeltis
anchietae), and M. okavangensis (= M. anchietae); the serpent plated lizard
Tetradactylus lundensis (= T. ellenbergeri); and the skink Mabuia striata angolen-
sis. The latter, however, remains problematic and under investigation (Conradie
et al. 2016). Marques et al. (2018) provided a replacement name, Trachylepis
monardi nom. nov., to stabilise taxonomy.
Throughout the early part of the twentieth century various other publications
discussed small collections made by explorers (e.g. Angel 1921, 1923; Mertens
1938). All added new locality records within and for the country, and also described
a number of new species (some no longer valid, e.g. Psammophis rohani, Angel
1921). Some reptile discoveries were especially serendipitous, e.g. the discovery
of the new lizard species Ichnotropis microlepidota (Marx 1956) based on three
specimens all found in the crop of a Dark Chanting Goshawk (Melierax metaba-
tes) collected during a bird survey of Mount Moco, that still awaits discovery in
the wild!
In 1952–1954 the Hamburg Museum made an expedition to various locations in
western Angola to collect mammals and herpetofauna, the most important being
numerous additional snake records from forested habitats at Piri-Dembos (= Piri,
Cuanza-Norte) (Hellmich 1957a, b). These confirmed, and sometimes added to
Parker’s (1936) records of snakes from the scarp forest isolates of Congulo and
Quirimbo. These included (e.g.): Philothamnus heterodermus, Thrasops flavigula-
ris, Toxicodryas blandingii, T. pulverulenta, Gonionotophis poensis, Pseudohaje
goldii, Atheris squamigera and Bitis nasicornis. All are Congo Basin species and
form an important biogeographic component of Angolan reptile diversity. Hellmich
undertook a follow up expedition in 1954–1955, but his expedition suffered delays
in obtaining permits and he missed the wet season activity and therefore shifted his
survey to more open habitats in the southern provinces. There he undertook some of
the first ecological studies on Angolan reptiles that were briefly discussed in a six-
part series of popular articles on his Angolan travels (Hellmich 1954–1955). On his
return he studied the reptile collections of the combined Hamburg expeditions
(Hellmich and Schmelcher 1956; Hellmich 1957a, b), but the amphibians were only
studied much later (Ceríaco et al. 2014b). As with Monard, Hellmich discovered
relatively few taxonomic novelties, i.e. the lizards Gerrhosaurus nigrolineatus ahle-
feldti (currently not considered valid) and Agama agama mucosoensis (now a full
species; Wagner et al. 2012).
Between 1950 and 1960 the Belgian herpetologist Raymond F Laurent lived in
Rwanda and Katanga (then Belgian colonies) and undertook detailed studies on
varied herpetological groups in the Congo Basin, describing numerous new species
and subspecies. During this period he studied the herpetological collections of the
Museu do Dundo in northeast Angola, made by António de Barros Machado, the
museum’s director. Summaries of the museum’s snake collections appeared first
(Laurent 1950, 1954), followed by another report on Dundo material including
Machado’s extensive herpetological collection from the arid southwest region of the
country (Laurent 1964). This was completed after Laurent had moved to the United
States (1961) and the report described a number of new Angolan species, including
Bogert’s Speckled Western Burrowing Skink (Typhlacontias bogerti), two Namib
Day geckos (Rhoptropus taeniostictus and R. boultoni montanus), and finally
Hellmich’s Wolf Snake (Lycophidion hellmichi), based (in part) on material col-
lected during the Hamburg expeditions. It also included additional information of
many previously poorly known species, as well as making ecological observations.
It set a new standard for herpetological research in the area, but sadly was the final
major Angolan herpetological work of the colonial period. Laurent did not study the
historical collections in the Museu de Lisboa, and therefore did not re-assign
Bocage’s original material to his new taxa or identify significant new distribution
records. This was unfortunate as he was one of the last herpetologists working on
the Angolan herpetofauna before the disastrous fire that destroyed (1978) the collec-
tions that Bocage had studied, as well as much of his correspondence with collec-
tors and fellow researchers. Manaças reported on collections of lizards (Manaças
1963), snakes (Manaças 1973), and venomous snakes (Manaças 1981) from Angola.
Bringing Knowledge of Angolan Reptiles into the Modern Era
Awareness of the interesting reptiles of the Angolan Namib region started inciden-
tally following expeditions in the 1950s by the enthusiastic entomologist Charles
Koch of the Transvaal Museum (TMP, now the Ditsong Natural History Museum,
Pretoria, South Africa). Koch did much to inventory the amazing diversity of the
tenebrionid beetle fauna of the Namib Desert, and much of his collecting involved
walking at night in the desert with a pressure lamp. In addition to his numerous
beetle discoveries Koch also collected many nocturnal and terrestrial reptiles, par-
ticularly geckos. These he gave to his colleague at the TMP, the Curator of Lower
Vertebrates Vivian FitzSimons. Koch visited the northern Namib in Angola on four
occasions (1951–1957), accompanied on the last trip by the Swedish zoologists
Lundholm and Rudebeck. The herpetological collections during these trips were
significant, and FitzSimons (1953, 1959) described a new genus of plated lizard
Angolosaurus (now subsumed within Gerrhosaurus) as well two new species,
Pachydactylus caraculicus and Prosymna visseri. Unreported, however, were many
of Koch’s numerous other reptile discoveries, including new records of the iconic
Namib Web-footed Gecko (Palmatogecko rangei, now included in Pachydactylus)
in 1951 and 1954, then unknown from the Angolan Namib region. Laurent (1964)
described the new Angolan Namib Day Gecko Rhoptropus taeniostictus from
Angola, although nine specimens had already been collected by Koch during his
trips, but remained undescribed. Also unrecorded were nine specimens of
Pachydactylus scutatus angolensis from Lungo, Lucira and São Nicolau, the first
collected since the description of the species by Loveridge (1944), and also 13 spec-
imens of Chondrodactylus fitzsimonsi, at the time known from only one Angolan
specimen (Pico Azevedo, Schmidt 1933).
In 1964 Wulf Haacke, born in Namibia, became the then Transvaal Museum’s
herpetologist with a special interest in the arid western areas of southern Africa. In
March–April 1971 he undertook his first trip to Angola, concentrating on the north-
ern extension of the Namib Desert into southwestern Angola. A follow up trip in
April–June 1974 targeted specific genera to confirm the northern range limits and
taxonomic status of Cordylus, Cordylosaurus, Gerrhosaurus, Pachydactylus,
Afroedura and Rhoptropus. Both trips were exceptionally successful resulting in
over 2000 specimens, the largest herpetological collections ever assembled by one
researcher in Angola. Although the amphibian collections made during these trips
were reviewed by Poynton and Haacke (1993), the vast majority of the numerous
new and rare reptiles contained in these collections were never formally published.
Haacke’s second trip in 1974 was designed in particular to collect new material for
his proposed thesis and revision of Rhoptropus. Prior to this trip, and excluding
Koch’s undescribed material, less than 30 specimens of Rhoptropus were known
from Angola (Bocage 1895; Parker 1936; Mertens 1938; Laurent 1964). At the end
of Haacke’s surveys the Transvaal Museum held 650 specimens of the genus,
included nearly 250 specimens of R. barnardi from over 25 localities, nearly 50
specimens of R. biporosus, and seven of R. afer. At the time R. barnardi in Angola
was known from very few specimens (Bocage 1895; Schmidt 1933; Parker 1936;
Laurent 1964) and R. biporosus was unknown in Angola and restricted to northern
Namibia. The status of R. afer in Angola was particularly confused. Bocage’s (1895)
knowledge of Namib Day geckos (Rhoptropus) seems to have been limited, and he
considered specimens from diverse localities from coastal Moçâmedes and the inte-
rior to Capangombe to all be applicable to R. afer Peters 1869. However, he noted
that his material had 6–8 preanal pores whilst R. afer, as Peters (1869) had correctly
recorded, had none. Schmidt also recorded R. barnardi for the first time from
Angola, noting numerous other specimens from the Moçâmedes region in the
British and Zoologisches museums that agreed with R. barnardi. These may have
been the source of Bocage’s confusion. The few recent records of Angolan R. afer
have all been restricted to the vicinity of the Cunene River mouth, and it is evident
that Bocage’s material from further north was not based on true R. afer.
Due to the limited access to Angola during the protracted civil war the TM expedi-
tions to Angola were curtailed, and for the next 34 years studies on Angolan reptiles
were based on museum material collected earlier. Details of some of the geckos col-
lected by the TM expeditions were incorporated into Haacke’s studies on the burrow-
ing geckos of southern Africa, which included the first records for Angola (Haacke
1976a) for the Palmatogecko rangei and Kaokogecko vanzyli (both now included in
Pachydactylus), and by implication Colopus (= Pachydactylus) wahlbergii, known to

Haacke (1976b) from three specimens (TM 38526—28) from the Angola-SWA bor-
der, 18 degrees E). The bizarre and iconic Plume-tailed Gecko Afrogecko plumicau-
dus, collected during Haacke’s 1971 trip, was of problematic taxonomic affinities
and not described until much later (Haacke 2008). Its true affinities, however, were
finally resolved later when fresh material was available for genetic analysis and it was
placed in the monotypic genus Kolekanus (Heinicke et al. 2014). The 64 burrowing
skinks of the genus Typhlacontias collected during these expeditions also formed an
integral part of Haacke’s (1997) revision of the genus, and which led to the recogni-
tion of a long overlooked Namib species, T. johnstonii, previously confused (Bocage
1895; Monard 1937) with T. punctatissimus, and described from Porto Alexandre
(=Tômbwa) at the northern limit of its range. A new Angolan species, T. rudebecki,
was also described, based on a single specimen collected from São Nicolau during
Koch’s 1956 expedition. Laurent’s (1964) species T. bogerti was treated as a northern
subspecies of the Speckled Burrowing Skink, T. punctatissimus bogerti (Haacke
1997). A number of other new Angola records were discovered by Haacke during his
trips. The rupicolous gecko Pachydactylus oreophilus, described from northern
Namibia (McLachlan and Spence 1967), was known only from the types until Haacke
discovered similar material from numerous localities in southwest Angola. No addi-
tional Angolan material was collected until the Huntley expedition in the region in
2009 (see below), when it was realised that Angolan material was not conspecific
with Namibian P. oreophilus. The status of the Angolan material is currently being
investigated (Branch et al. in prep.). Haacke also collected the first records of true P.
scutatus from Angola, as well as additional P. angolensis (Branch et al. 2017). Finally,
Broadley (1975) referred some small skinks collected by Haacke to Trachylepis lac-
ertiformis, creating a zoogeographic enigma as the nominotypic population of this
small skink is restricted to the lower Zambezi River valley. Fuller details of the Koch
and Haacke collections and other recent material will be incorporated into a full
review of the herpetofauna of the Angolan Namib region (Branch in prep.).
Legless burrowing worm lizards (Amphisbaenidae) are rarely found due to their
ability to burrow deep underground. Carl Gans (1976) described three new species
from Angola, including Monopeltis luandae based on fresh (1971) and historical
(1892) material from the Luanda region, and M. perplexus on material from the
Vernay-Angola expedition collected from ‘Hanha or Capelongo’ in 1925. Similarly,
Gans (1976) re-assessed old and new material when describing Dalophia angolen-
sis from Calombe near Vila Luso (= Luena), and reassigned specimens identified as
M. ellenbergeri, and then as M. granti transvaalensis (Monard 1937), to D. angolen-
sis. None of Gans’ three new species had been rediscovered in the intervening 40+
years until recently, when M. luandae was rediscovered from near the type locality
(Branch et al. 2018). In a companion paper that radically changed understanding of
the taxonomy of worm lizards in the southern half of Africa, Broadley et al. (1976)
revised the genera Monopeltis and Dalophia. The revision of Broadley et al. (1976)
affected most of the early names applied to Angolan worm lizards. The first large
worm lizard described from Angola was Lepidostsernon (Phractogonus) anchietae
Bocage 1873, from ‘Humbe, dans l’intérior de Mossamedes’, later transferred to the
genus Monopeltis (Boulenger 1885). Broadley et al. (1976) relegated Monard’s
(1937) species M. okavangensis and M. devisi to M. anchietae, which is now known
to have a wider range in northern Namibia and adjacent Botswana. Monopeltis
vanderysti vilhenai, described by Laurent (1954) from Dundo, Angola, was not rec-
ognised by Broadley et al. (1976) and returned to M. vanderysti, which is widely
distributed in the Congo region. The Dundo specimen remains the only Angolan
record of the species. Monopeltis capensis in Angola was first recorded by Bocage
(1873), and later by Monard (1937). Although provisionally placed in the M. capen-
sis capensis Group B (Broadley et al. 1976), with a wide range through the Kalahari
region (Northern Cape, South Africa, through Botswana to southern Angola), it was
later treated as a separate species, M. infuscata Broadley 1997a. Monard’s (1937)
species Monopeltis granti kuanyamarum, described from a single specimen from
Mupanda, was transferred to Dalophia pistillum (Broadley et al. 1976). The only
Angolan specimen of Dalophia ellenbergeri was collected whilst trench digging
during hostilities at Cuito Cuanavale (Branch and McCartney 1992; Broadley
1997b). Gray (1865) described Dalophia welwitschii from Pungo Andongo, and this
has not been rediscovered. It is the type species for the genus Dalophia, and Gans
(2005) was obviously in error when treating it as M. welwitschii and yet still con-
tinuing to recognise Dalophia. A phylogeny of African amphisbaenids (Measey and
Tolley 2013), albeit based on poor taxon sampling, recovered Monopeltis and
Dalophia as monophyletic clades, supporting the use of Dalophia welwitschii.
Two species of round-headed worm lizards of the genus Zygaspis are now known
to occur in southeast Angola, but the genus was unknown to Bocage from Angola,
and the first record from the country was Monard’s (1931) description of
Amphisbaena ambuellensis from ‘Chimporo’ (= Tchimpolo). This was subsequently
synonymised with A. quadrifrons by Loveridge (1941) with some misgivings, and
subsequently transferred to Zygaspis by Alexander and Gans (1966). It remained
known only from Monard’s material for many years, but has recently been collected
in southern Angola (Conradie et al. 2016, Baptista et al. in prep.), and the availabil-
ity of Monard’s ambuellensis for this material is being reassessed. More recently,
Laurent (1964) recorded A. q. capensis from Alto Chicapa in northeast Angola,
which was shown to be the new species Zygaspis nigra by Broadley and Gans
(1969). This small black worm lizard is near endemic to eastern Angola, with
records from adjacent Zambia (Kalobo, Broadley and Gans 1969; Ngonya Falls,
Pietersen et al. 2017) and Namibia (Katima Mulilo, Broadley and Gans 1969).
Recent material is known from the Okavango catchment (Conradie et al. 2016).
Following the cessation of hostilities, modern biodiversity surveys were initiated
by Brian Huntley with the multinational SANBI/ISCED/UAN Angolan Biodiversity
Assessment and Capacity Building Project (Huntley 2009). Surveys were under-
taken by botanists and zoologists in various habitats between Lubango and the
Cunene River, and 15 Angolan students were involved in fieldwork and training
sessions. The immediate reptile highlights of the survey involved the discovery of
two new species of the lacertid Pedioplanis (Conradie et al. 2012a), two specimens
of the rare Shovel-snout Snake Prosymna visseri were collected at Espinheira, Iona
National Park, only the 5th and 6th Angolan specimens since its description
(FitzSimons 1959); the 1st record of the Namib Wolf Snake (Lycophidion namibia-
num) from Angola, again at Espinheira; the southernmost records of the newly-
described Plume-tailed Gecko (Afrogecko plumicaudatus Haacke 2008) that allowed
its generic assignment to later be readjusted; and topotypic material of the rare
chameleon (Chamaeleo anchietae) were collected around Estação Zootécnica. This
chameleon has an unusual, disjunct distribution with scattered populations (treated
as separate subspecies by Laurent 1951) from the Upemba region, Democratic
Republic of the Congo (DRC) and Udzungwa Mountains, Tanzania. The status of
these disjunct C. anchietae populations is currently under investigation (Branch
et al. in prep.). A new species of reed frog, Hyperolius chelaensis, completed the
new discoveries (Conradie et al. 2012b). Following the success of the 2009 survey,
another expedition was organised in 2011 to Lagoa Carumbo, the second-largest
Angolan freshwater lake situated in Lunda Norte Province (Huntley and Francisco
2015). The herpetological results were summarised by Branch and Conradie (2015).
Significant herpetological discoveries included the discovery of at least two new
species of frog, one described (Hyperolius raymondi Conradie et al. 2013), and the
description of the other (Amnirana sp.) is in preparation (Jongsma et al. 2018), and
also the first record for Angola (Branch and Conradie 2013) of the Banded Water
Cobra (Naja annulata). Other reports include new insights into the distributions of
venomous snakes, such as Jameson’s Green Mamba, Dendroaspis jamesoni (Vaz
Pinto and Branch 2015) and the Gaboon Adder, Bitis gabonica (Oliveira et al.
2016), as well as a recent summary of Angolan venomous snakes (Oliveira 2017).
The unique World Heritage site of the Okavango Delta is situated in Botswana,
but depends on the Okavango drainage, which arises and is almost entirely con-
tained within southeastern Angola. During the last six years a series of international
collaborative surveys have been undertaken to explore this poorly known region of
southeast Angola, and to understand the hydrology and biodiversity of the Okavango
drainage. The first surveys were organised by the Okavango River Basin Water
Commission (OKACOM), in accord with the Angolan National Action Plan for the
Sustainable Management of the Cubango/Okavango River Basin (OKACOM 2011),
and occurred in the lower catchments of the Cubango and Cuito rivers (Brooks
2012, 2013). More recent surveys (2015–2018) formed part of the ongoing National
Geographic funded Okavango Wilderness Project (NGOWP 2018), which have
intensely surveyed the source lakes of the major Okavango tributaries in an unex-
plored region where the headwaters of the Cuanza, Zambezi and Okavango basins
meet. The herpetological results of the OKACOM surveys (2012–2013) and first
phase of the NGOWP surveys were presented by Conradie et al. (2016), who also
reviewed the region’s herpetofauna. In total 67 reptiles species are now known from
the region, comprising 38 snakes, 32 lizards, five chelonians, and a single crocodile
(NGOWP 2018). Three reptiles new for Angola, including Causus rasmusseni
(although the specific status of this taxon still requires genetic confirmation),
Acontias kgalagadi kgalagadi and Panaspis maculicollis were discovered (Conradie
and Bourquin 2013; Medina et al. 2016). The results of more recent surveys (2016–
2018) were presented (Conradie et al. 2017) and fuller details are being prepared for
publication, and online species lists are planned for public access.
Contemporaneous with the above surveys a number of other Angolan biodiver-

sity initiatives began. In partnership, the Kimpa Vita University in Uíge and Dresden
University, Germany, undertook herpetological surveys of Serra do Pingano, Uíge
Province, discovering diverse tropical Congo Basin species (Ernst 2015), including
two frogs previously unrecorded for Angola (Ernst et al. 2014, 2015). In addition
the California Academy of Sciences in conjunction with the National Institute of
Biodiversity and Conservation Areas and the Ministry of the Environment of Angola
(MINAMB/INBAC) initiated an ongoing Atlas of Angolan amphibian and reptiles
(Marques et al. 2014, 2018). Various areas have been surveyed and preliminary
results published in the scientific and popular literature (Ceríaco et al. 2014a, b,
2016a, b, 2018b). As part of the Southern African Science Service Centre for
Climate Change and Adaptive Land Management program (SASSCAL) the Instituto
Superior de Ciências da Educaҫão da Huíla (ISCED), Lubango, has been undertak-
ing herpetofaunal monitoring at several areas in Huíla Province and elsewhere in
Angola (Baptista et al. 2018, 2019), with emphasis on the escarpment. A herpeto-
faunal archive is also being developed at ISCED.
Angolan citizen science is in its infancy, but the FaceBook site Angola Ambiente
is a public group where members post observations (https://www.facebook.com/
groups/1045499302182009/). It is designed “to raise awareness of the fantastic
fauna and flora of this magnificent country”, and requests observations with detailed
locality data. It includes irregular lists of sightings with locality details in support of
mapping initiatives.
Checklist of Angolan Reptiles
How Many Species?
The first attempt to summarise the herpetofauna of Angola was undertaken by JV

Barbosa du Bocage (1866), who listed 50 reptile species from Angola in the Museu
de Lisboa collection, including 23 snakes, 21 lizards, four chelonians, and one croc-
odilian. After nearly 30 years of study he again summarised the Angolan herpeto-
fauna in his monographic Herpetology d’Angola et du Congo (Bocage 1895) in
which he listed 143 reptile and 39 amphibian species from the country. Of these
Bocage had described 37 of the taxa (although not all are now recognised). During
two trips to Angola (1928–1929 and 1932–1933) the Swiss collector Albert Monard
made important collections of amphibians and reptiles. In his monograph (Monard
1937) he presented an updated checklist of Angolan reptiles, listing 169 reptile spe-
cies in 10 families and 28 genera. There has been no subsequent updated and check-
list of the country’s reptiles, although Blanc and Fretey (2002) noted a total of 257
Angolan reptile species and published a breakdown of its composition. However, no
species list was included and therefore it is impossible to assess the accuracy or the
validity of the species included. In contrast, the online Reptile Database (Uetz et al.
2018, as on 14 October 2018) currently generates a list of 255 reptiles for Angola,
but unfortunately although close to the existing count, it is inaccurate in numerous

respects. Some species have been included that are unrecorded from the country but
may occur in the country (e.g. a snakes from the Congo Basin Calabaria reinhardti,
Namibian Lygodactylus lawrencei, etc.). Moreover, many other species are dupli-
cated and listed under both their historical and current taxonomic assignments (e.g.
Agama hispida = A. aculeata, Chamaesaura macrolepis = Ch. miopropus, Cordylus
cordylus = C. namakuiyus, C. vittifer = C. machadoi, etc.) These inaccuracies have
been discussed in the recent Atlas and being rectified (see Marques et al. 2018
Appendix Table A2).
Currently (as of mid-2018), there are 278 reptile species recorded from Angola,
comprising 15 chelonian, three crocodilian, 132 lizard and 128 snake species.
Table 13.1 details the historical growth of knowledge of reptile diversity in Angola
based on summaries in Bocage (1866, 1895), Monard (1937), and this study.
Table 13.2 summarises the number of genera, species, and endemic taxa in the
major reptile groups in Angola. Updated checklists of the major reptile groups,
including details of common and scientific names, historical scientific names used
by Bocage (1895) for the current taxa, as well as their endemic and conservation
status are summarised in: Appendix 1 – chelonians; Appendix 2 – lizards; and
Appendix 3 – snakes. Contained within these checklists are 43 Angolan species
named by Bocage, i.e. 15.5% of the current reptile diversity. This is less than the
25.9% (37 of 143) in Bocage’s (1895) summary, but no other researcher has
described more from Angola.
Table 13.1 Historical development of reptile diversity in Angola based on summaries in Bocage
(1866, 1895), Monard (1931), and this study
Bocage Monard This study
Group 1866 1895 1937 2018
Snakes 23 71 81 128
Lizards 21 59 78 132
Chelonians 4 8 9 15
Crocodilians 2 1 1 3
Total a 50 139 169 278
a
This includes ‘species’ known at the time of Bocage and Monard, some of which may have later
been relegated to synonymy (see discussion on Monard’s list)
Table 13.2 Summary of the taxonomic diversity and endemicity of the reptiles of Angola
Group Genera Species Endemic
Chelonians 11 15 0
Crocodilians 3 3 0
Lizards 40 132 27
Snakes 49 128 6
Total 103 278 33
Recent Discoveries
In the last decade, and resulting from the burgeoning scientific interest in Angola,
biodiversity surveys have led to the description of numerous new species and the
validation of the specific status of others. Perhaps the most exciting was the long
delayed description of the beautiful and bizarre Plume-tailed Gecko (Afrogecko
pulumicaudus Haacke 2008) from the Angolan Namib region. Other novelties
included the description of the lacertids Pedioplanis haackei and P. huntleyi
(Conradie et al. 2012a and the cordylid Cordylus namakuiyus (Stanley et al. 2016).
Some subspecies were validated as full species, including the geckos R. boultoni
benguellensis and R. boultoni montanus (Ceríaco et al. 2016a) and the snake
Psammophylax rhombeatus ocellatus (Branch et al. 2019), as well as the revival of
the skink Trachylepis damarana from the synonymy of T. varia (Weinell and Bauer
2018). Some species, e.g. Philothamnus nitidus loveridgei, however, have been
shown to lack genetic support for recognition (Engelbrecht et al. 2018) and are now
not recognised.
In addition, preliminary studies have revealed numerous problematic specimens
and populations that demonstrate the existence of cryptic, previously synonymised
species or unnamed taxa awaiting description in numerous genera. Geckos –
Hemidactylus, Rhoptropus (Ceríaco et al. 2016a; Bauer and Kuhn 2017), Afroedura
(Branch et al. 2017), and various Pachydactylus groups (Branch et al. 2017; Ceríaco
et al. 2016a; Heinz 2011); within the lacertids Nucras, Pedioplanis and Heliobolus
(Branch and Tolley 2017); and a skink in the Trachylepis varia complex (Weinell
and Bauer 2018). The descriptions of at least a dozen new species in these genera
are in preparation. Ceríaco et al. (2016b, 2018b) signalled the presence of a new
skink (T. cf. megalura) from Cangandala National Park. Snake-eyed skinks
(Panaspis wahlbergii-maculilabris complex) have been shown to include numerous
cryptic species in southern and east Africa (Medina et al. 2016). Records of P. wahl-
bergii in Angola are therefore also likely to represent taxonomic novelties. It is also
likely that genetic studies will further validate a number of other lizard taxa cur-
rently treated as subspecies, e.g. Ichnotropis bivittata palida and Trachylepis bayoni
huilensis, as full species. Moreover, the rare gecko Afrogecko ansorgi, described by
Boulenger (1907a), as Phyllodactylus ansorgi and not collected again for nearly
100 years, was recently rediscovered and a re-assessment of its generic status is in
preparation. In addition, ongoing surveys of the Angolan Okavango Project con-
tinue to confirm new species records for Angola, including most recently
Pachydactylus wahlbergii (G Neef pers. comm. July 2018), previously assumed to
enter southern Angola based on material collected on the Angolan-Nambian border
in the 1970s (Haacke 1976b). Cryptic diversity in snakes is also being unravelled,
and African forest cobras have been shown to include five species (Wüster et al.
2018), of which two enter Angola, whilst Angolan house snakes (Boaedon) is
expected to comprise at least eight species, with four potential new country records,
i.e. B. fuliginosus, B. radfordi, B. virgatus and B. mentalis (the latter signalled as a
valid species by Kelly et al. 2011, and will be formally revived for western arid
populations from South Africa to southwest Angola), revival from synonymy of two
Bocage names for Angolan endemics, i.e. B. angolensis and B. variegatus, and two
additional taxonomic novelties (Hallermann et al. personal communication).
Overview of Reptile Diversity
Chelonians
This ancient lineage of reptiles has relatively little global diversity and includes the
greatest proportion of threatened reptiles, particularly in Asia. They are relatively
poorly known in Angola, and their diversity is discussed below. The first sea turtle
to swim in the early South Atlantic, after the separation of Africa and South America
90 Million years ago, was the extinct chelonian Angolachelys mbaxi, discovered in
Angola in 2009 near the village Iembe, Bengo Province (Mateus et al. 2009). Five
of the seven species of sea turtles in the world have been recorded from Angolan
waters (Carr and Carr 1991), although only four occur regularly. They include (in
decreasing order of abundance): Olive Ridley Sea Turtle (Lepidochelys olivacea),
Leatherback Sea Turtle (Dermochelys coriacea), Green Sea Turtle (Chelonia
mydas), Loggerhead Sea Turtle (Caretta caretta), and Hawksbill Sea Turtle
(Eretmochelys imbricata). Although early surveys (2000–2006) indicated the
absence of Hawksbill in Angolan waters (Weir et al. 2007), juveniles were recently
recorded in the Soyo and Cabinda region (Morais 2008, 2016). These may still be
vagrants in Angolan waters (TTWG 2017) as Hawksbills forage on coral reefs
which are absent in Angola. Nesting in Angolan waters has only been confirmed for
Green, Olive Ridley and Leatherback sea turtles, and occurs during September–
March, peaking in November–December in the north and a month later in the south
(Morais 2017). It is widespread for the Olive Ridley, but restricted mainly to the
south for the Green Sea Turtle. The latter remains common in the Cunene River
estuary where adults and juveniles feed and also escape the cold waters of the
Benguela Current (Elwen and Braby 2015). The giant Leatherback nests primarily
in the warmer north, with little activity south of Benguela. The Angolan population
(approximately 1000 in 2005–2016, Morais 2016) forms the southern part of the
major Gabon nesting grounds, where 6000–7000 females breed annually (Billes
et al. 2006). Sea turtles face numerous threats, including by-catch and drowning in
trawler fishing nets, the collection of nesting females and their eggs for food, and
disturbance of the nest sites by beach activities, etc. (Morais et al. 2005; Morais
2008; Weir et al. 2007). The Projecto Kitabanga of Universidade Agostinho Neto
(https://www.facebook.com/Kitabanga/) is involved in research and public educa-
tion of Angolan sea turtles.
Only three land tortoises are recorded for Angola. The Leopard Tortoise
(Stigmochelys pardalis) is restricted to the southwest, with all records occurring
below the escarpment south of Benguela and along the Cunene valley. Bell’s Hinge-
back Tortoises (Kinixys belliana) is considered widespread in Angola (TTWG
2017), and Kinixys material from central and eastern Angola were confirmed as this
species in a molecular phylogeny of the genus (Kindler et al. 2012). Although mate-
rial from Capanda Dam has been referred to Kinixys spekii (Ceríaco et al. 2014a),
this species is not currently considered to occur in the country (TTWG 2017), but is
known from the Zambezian region of Namibia and the Ikelenge pedicle of north-
west Zambia. The Forest Hinge-back Tortoise (Kinixys erosa) occupies moist for-
ests of the Congo basin and West Africa, but only enters Angola in the extreme
northeast (Dundo, Laurent 1964) and the Cabinda enclave (Bocage 1895).
The Nile Soft-shelled Terrapin (Trionyx triunguis) is restricted to the coastal
region, entering the estuaries and lower stretches of the major rivers. It tolerates sea
water and grows to over a metre in length. Populations in the eastern Mediterranean
and lower Nile River are threatened, and its status in Angola is poorly known, but is
known from the coastal region and with populations in the Cunene River mouth and
extending some distance upstream in the Cuanza River. Aubry’s Soft-shelled
Terrapin (Cycloderma aubryi) was recorded once from Cabinda (Peters 1869), but
there are no recent records. Trade in chelonians, particularly soft-shelled terrapins
for food in Asia, has pushed many species to the brink of extinction (TTWG 2011),
and involvement in Africa is confirmed by the discovery of a turtle butchery on Lake
Malawi (Face of Malawi 2013) and the recent confiscation of a large T. triunguis in
a shopping mall in Luanda (Arruda 2018). All other Angolan terrapins have hard
shells and are restricted to fresh water ecosystems. They withdraw the head into the
shell sideways and are represented by the Pelomedusidae in Africa, including the
genera Pelomedusa and Pelusios. Although Bocage (1866) listed the forest species
Pelusios gabonensis from Cabinda and Duque de Bragança in his first review of
Angolan reptiles, the species was subsequently omitted (Bocage 1895). However, it
was subsequently recorded from Dundo (Laurent 1964), and mapped to enter
extreme northwest Angola south of the Congo River (TTWG 2017), but no docu-
mentation supporting this is presented. It was not recorded at Soyo (W Klein pers.
comm.), although Western Hinged Terrapin (Pelusios castaneus) was common. The
most widespread Angolan terrapins are Pelusios nanus, P. bechuanicus and P. rho-
desianus in the extensive wetlands of eastern Angola (Conradie et al. 2016; TTWG
2017).
Crocodilians
Of the three crocodilians that occur in Angola, only the Nile crocodile (Crocodylus
niloticus) is widespread, being absent only from the southwest although occurring
in the lower Cunene River. The remaining two species are both denizens of the
Congo Basin and have only a peripheral presence in Angola. The Sharp-snouted
Crocodile (Mecistops cataphractus) in Angola had been discussed by Machado
(1952), who noted an unusual early record from Lunda and others from Dundo, later
confirmed by Laurent (1964). Recent studies (Shirley et al. 2014) have found sig-
nificant molecular and morphological support for two divergent taxa in Mecistops –
one distributed entirely in West Africa and the other in Central Africa. As the type
locality is Senegal, West African populations would keep the name and Congo
Basin and Angolan populations have been considered to represent an undescribed
species (Shirley et al. 2014), and was subsequently described as a new species,
Mecistops leptorhynchus (Shirley et al. 2018). The Dwarf Crocodile (Osteolaemus
tetraspis) is known from nineteenth century of records from the Cabinda enclave
(Bocage 1866; Peters 1877), but no confirmed records exist for the natural occur-
rence of the species south of the Congo River (Eaton 2010). Ceríaco et al. (2018a)
discuss a problematic specimen collected in the lagoon at Luanda that they consider
to be indicative of an unknown population in the Cuanza River drainage and also the
first record of O. osborni for Angola. However, the specimen’s identity was not
confirmed by genetic monophyly, and its presence in Angola may also result from
an escapee brought to Luanda for the bushmeat trade. As with Mecistops recent
genetic studies indicate the existence of at least three species within the Osteolaemus
tetraspis complex (Eaton et al. 2009), but the taxonomic identity of Cabinda and the
putative Cuanza Osteolaemus populations require further study.
Squamates
Scaled reptiles (Squamata) form the major component of reptile diversity (Pincheira-
Donoso et al. 2013), with over 10,000 species currently recognised, of which over
60% are lizards. Reflecting this, lizards are also the dominant component of the
Angolan reptile diversity and are the group in which most recent discoveries have
been made (see above).
The 132 species of Angolan lizards are currently contained in nine families, with
skinks (Scincidae) containing the greatest diversity. This contrasts with Namibia
(Herrmann and Branch 2013) and South Africa (Branch 2014) where geckos form
the greatest component of lizard diversity and endemicity (Table 13.3). It is likely
that the current dominance of skinks in Angola is an artefact of our present knowl-
edge. Most skinks are diurnal and active and therefore more easily discovered.
Cryptic diversity has already been identified in certain Angolan gecko genera (e.g.
Afroedura, Pachydactylus and Rhoptropus, see above), and the discovery of these
and others is predicted to also promote geckos to dominance in diversity and ende-
micity in the Angolan reptile diversity. The evolutionary centre for girdled lizards
(Cordylidae) occurs in southern African (Stanley et al. 2016), but the family is rela-
tively poorly represented in Angola. Although it is unlikely to reach the species or
generic diversity of even Namibia, there are indications that the diversity of rupico-
lous Cordylus in the escarpment and central uplands is under-represented (e.g.
Stanley et al. 2016), and that rediscoveries and further new species await discovery
and description.
The families Agamidae, Chamaeleonidae, Gerrhosauridae and Varanidae all
have limited diversity in Angola, as do the last two families throughout Africa.
Blue-headed tree agamas have been revised (Wagner et al. 2018), with populations
from northern Namibia, Angola and northwest Zambia now referred to the revived
Acanthocercus cyanocephalus. However, it is evident that current species boundaries
Table 13.3 Comparison of diversity and endemicity of Angolan and South African Squamates
(excluding Chelonia), by genera (Gen.), species (Spp.), subspecies (Sub.), and endemism (End.)
Angola South Africa
Family Gen. Spp. End. Gen. Sp. End.
Lizards
Gekkonidae 8 34 8 12 89 55
Agamidae 2 7 2 2 7 0
Chamaeleonidae 2 5 0 2 19 15
Gerrhosauridae 4 8 0 5 13 6
Cordylidae 2 5 2 10 53 38
Scincidae 12 45 6 7 62 32
Lacertidae 6 15 6 8 29 9
Amphisbaenidae 3 11 3 4 12 2
Varanidae 1 2 0 1 2 0
subtotal 40 132 27 51 286 157
(20.5%) (54.9%)
Snakes
Leptotyphlopidae 2 5 2 3 10 3
Typhlopidae 2 8 1 3 7 0
Pythonidae 1 3 0 1 2 0
Colubridae 14 31 0 9 16 0
Natricidae 2 4 0 1 2 0
Lamprophiidaea 15 39 2 17 42 3
Atractaspididae 6 11 0 6 16 2
Elapidae 5 14 0 6 18 1
Viperidae 3 13 1 2 14 4
subtotal 50 128 6 48 127 13
(4.7%) (10.2%)
TOTAL 95 260 33 109 413 170
(12.7%) (41.2%)
Angola/South 83% 63%
Africa
a
Excludes additional Boaedon species (Hallerman et al. in prep.)
in Agama and Acanthocercus do not full reflect Angolan agamid diversity. The
remaining families, Lacertidae and Amphisbaenidae, are relatively well represented
in Angola, with worm lizard diversity in Angola (three genera, 11 species) second
only to that in South Africa (12) for diversity in Africa. Most are associated with the
sands of the Kalahari Basin, or in secondary deposition in the coastal zone of South
Africa and southern Mozambique. The role of river capture and hydrological
changes associated with nascent rifting on fossorial reptiles awaits fuller study.
Lacertid diversity in Angola (13 species) is reduced relative to South Africa (29) and
Namibia (24), but is known to be under-represented and recently described
Pedioplanis species (Conradie et al. 2012a), and recently discovered cryptic diver-
sity in other lacertid genera (Branch and Tolley 2017; Conradie et al. 2016) will
increase species numbers in the family. A number of additional tropical lacertids
may also enter the northern regions of Angola (see below).
There are several aspects of Angolan reptile diversity that stimulate interest. The
first is the absence of an endemic radiation of chameleons within Angolan forest
refugia. African countries with the highest chameleon diversity (Tilbury 2018), i.e.
South Africa and Tanzania, have largely endemic radiations of chameleons
(Kinyongia and Rhampholeon in Tanzania, Bradypodion in South Africa). All three
genera are absent from Angola, where only Chamaeleo and Trioceros occurs.
Greater knowledge of the history of forest habitats in Angola may give insight as to
the absence of a forest chameleon radiation. Sandy habitats in arid southwest
Angolan include a radiation of skinks of the genera Sepsina and Typhlacontias that
have reduced limbs, serpentine locomotion and fossorial behaviour. The ranges of
some species within these genera extend south into adjacent Namibia and Botswana.
In arid habitats at the southern end of the Namib Desert, in the southern Dune Sea
and adjacent Succulent Karoo biome these Angolan fossorial skink radiations are
almost completely replaced by another suite of sepentiform skinks of the genera
Scelotes, Typhlosaurus and Acontias. Only one species, Typhlacontias brevipes, of
the Angolan radiation occurs in the northern parts of the southern Dune Sea.
Increased knowledge of the history of aridification and dune movements of the
Namib Desert may again give insight into these distributions.
That snake diversity in Angola is probably the most well known component of
the reptile fauna is unsurprising. However, their distribution, particularly of forest-
adapted species in the northern and scarp forest isolates, remains poorly-known.
The taxonomic status of these isolated forest populations calls for genetic studies on
their phylogenetic relationships to confirm their conspecificity with northern popu-
lations. The diversity and composition of snake families in Angola reflects that of
Africa, with relatively low diversity in primitive groups such as scolecophidians
(Typhlopidae and Leptotyphlopidae) and haenophidians (Pythonidae). Again, in
Angola as in southern Africa the venomous families Elapidae and Viperidae have
slightly greater species diversity, but with more tropical representatives (e.g. the
elapids Pseudohaje goldi, Naja annulata and N. melanoleuca, and viperids Causus
lichtensteini, C. maculatus, Atheris squamigera and Bitis nasicornis). The dominant
African snake family is the Lamprophiidae, of which the Atractaspididae is closely
related and sometimes treated as a subfamily. The group appears to have originated
in Africa and subsequently radiated into Arabia and Asia, and the subfamilies
Lamprophiinae, Prosyminae and Psammophinae form important radiations in Sub-
Saharan Africa. Lamprophids thus form the dominant component of the Angolan
snake fauna (39 species), but includes only two endemic psammophines. As with
elapids and viperids a number of Congo Basin species enter the northern forests,
including some currently known from very few Angolan specimens, e.g.
Lycodonomorphus subtaeniatus, Chamaelycus parkeri, Boaedon olivaceus,
Bothrophthalmus lineatus, etc. Perhaps the greatest difference between South Africa
and Angola is reflected in the greater diversity of colubrids (Colubridae) in Angola
(28 vs 14 species). These include numerous tropical Congo Basin snakes that enter
the northern and scarp forests, and of particular interest are the rare Congo Basin
species Toxicodryas blandingii, T. pulverulenta, Rhamnophis aethiopissa,
Philothamnus nitidus, Dasypelis palmarum, etc. The family is considered of Asian
origin and to have entered and subsequently radiated in Africa.
Species Recorded from Angola but Poorly Known
Some species are known from Angola from either a single or very few specimens
and their presence and taxonomic status requires confirmation. This summary does
not include wide ranging species that peripherally enter Angola, either from the
Congo Basin (e.g. Pelusios gabonensis, Feylinia grandisquamis, Hypoptophis wil-
soni, etc.), or from the southern Kalahari or Namib deserts (e.g. Rhoptropus afer,
Pachydactylus rangei, P. vanzyli, Chamaeleo namaquensis, Amblyodipsas ventri-
maculata, etc.).
Grass Lizard – Chamaesaura anguina oligopholis Laurent (1964). Described from
Calonda, Lunda, but with no recent material. It may deserve specific status.
Angola Girdled Lizard – Cordylus angolensis (Bocage, 1895). Known only from
the type description of a single male from Caconda, but a population that con-
forms to the species has recently been discovered (Vaz Pinto Unpublished Data).
Scaled Sandveld Lizard -Nucras scalaris Laurent 1964. Still known only from type
series of four specimens from Alto Chicapa and Alto Chilo.
Dewitte’s Five-toed Skink – Leptosiaphos dewittei (Loveridge, 1934). Recorded by
Parker (1936, as Lygosoma dewittei) from Congulo. However, the only known
Angolan specimen lacks the diagnostic compressed tail. The species occurs in
the eastern Congo Basin, a considerable disjunction from Congulo.
African Shovel-nosed Snake – Scaphiophis albopunctatus Peters, 1870. Only once
recorded from Angola (Laurent 1950, Muita River) in Guinea-Congo savannah
habitat.
The only other know record is a juvenile specimen collected from Capaia, Lunde
Norte (Branch and Conradie 2015).
Collared Snake-eater – Polemon collaris (Peters 1881). Recorded by Peters (1881,
Cuango), Ferreira (1904, Golungo Alto) and Hellmich 1957a, b, Bella Vista, as
Miodon gabonensis). Isolated populations of small fossorial snakes such as
Polemon often include cryptic diversity (Portillo et al. 2018), and fresh material
is required for taxonomic assessment.
Lined Water Snake – Lycodonomorphus (?) subtaeniatus Laurent 1954. Described
from Keseki (DRC), with four paratypes from Dundo the only Angolan records.
Greenbaum et al. (2015) transferred L. s. upembae to Boaedon. This is probably
where L. subtaeniatus belongs but fresh material is required for genetic
analysis.
Speckled Wolf Snake – Lycophidion meleagre Boulenger 1893. Described from
Angola and known from Cabinda to Luanda, but Broadley (1996) also includes
records from coastal Tanzania in the species’ range, creating a biogeographic
anomaly that requires genetic assessment.
Parker’s Banded Snake – Chamaelycus parkeri (Angel, 1934). Parker’s (1936)
Congulo specimen (as Oophilositum parkeri) remains the only known Angolan
material. Elsewhere the species is restricted to Kivu (DRC) and Congo
Brazzaville, and confirmation of the specific status of the Congulo population is
required.
Angolan Coral Snake – Aspidelaps lubricus cowlesi Bogert 1940. Described from
Munhino (101 km east of Moçamedes, via railroad), and known from Angola
from the type and one additional specimen (Branch 2018). Considered wide-
spread in northern Namibia, but genetic monophyly between Angolan and
Nambian populations is required for confirmation.
Angolan Garter Snake – Elapsoidea semiannulata moebiusi Werner, 1897. Listed
by Broadley (2006) from northern Angola, but with no specific localities given.
All Bocage localities (1866, 1895, 1897) were restricted to Bissau specimens. A
southern subspecies is now treated as a valid species (E. boulengeri). The status
of E. s. moebiusi requires a modern taxonomic assessment and also confirmation
for Angola.
Angolan Dwarf Adder – Bitis heraldica (Bocage, 1889). Angola’s most iconic snake
for which no new material was collected for over 50 years has recently been re-
discovered. It has a disjunct distribution in montane grasslands of the Angolan
inland plateau, and the fresh material will allow its subgeneric relationships
within Bitis to be assessed as well as its conservation status.
Species Likely to Occur in Angola but Currently Unconfirmed
A number of species are recorded in close proximity to the Angolan border and live
in habitats contiguous with those in Angola, and are therefore likely to occur in the
country. They include:
Lizards
Heenen’s Dwarf Day Gecko – Lygodactylus heeneni De Witte, 1933. This small
diurnal gecko was recorded from the Ikelenge Pedicle in northwest Zambia
(Broadley 1991; Haagner et al. 2000) within 25 km of the Angolan border.
Long-tailed Worm Lizard – Dalophia longicauda (Werner, 1915). This fossorial
species was described from northern Namibia and is known to extend through
the Caprivi region to western Zimbabwe (Broadley et al. 1976; Gans 2005) and
also to southwest Zambia (Pietersen et al. 2017). Populations occur to the east
and west of the Okavango River and are expected to occur in southeast Angola.
Maurice’s Worm Lizard – Monopeltis mauricei Parker, 1935. This fossorial species
was described from central Botswana and is known to extend through the
Kalahari to Katima Mulilo in the Caprivi region (Broadley et al. 1976; Gans
2005). Elevated to a full species by Broadley (2001).
West African Striped Lizard – Poromera fordii (Hallowell, 1857). An arboreal spe-
cies recorded during a survey in the Bas-Congo region (Nagy et al. 2013) within
30 km of the Angolan border but currently unknown from Angola.
Fine-scaled forest lizard – Adolfus africanus (Boulenger, 1906). A terrestrial spe-
cies recorded from the Ikelenge Pedicle in northwest Zambia (Broadley 1991),
within 25 km of the Angolan border.
Snakes
Western Thread Snake – Namibiana occidentalis (FitzSimons, 1962) occurs in

extreme Kaokoveld (Broadley and Broadley 1999) but has not yet been recorded
from southern Angola. The single record of the Damara Thread Snake (N. labia-
lis Sternfeld, 1908) from southern Angola demonstrates that these small snakes
can cross the Cunene River.
Leptotyphlops sp. An unidentified thread snake was recorded during a survey of the
Bas-Congo region (Nagy et al. 2013). Based on its forest habitat it is unlikely to
be referable to any known Angolan species.
Slender Quill-snouted Snake – Xenocalamus b. bicolor (Günther 1868). Although
Broadley (1971) records no Angolan material, the species occurs in the Caprivi
area and adjacent western Zambia, and it is usually associated with Kalahari
sands. It is therefore likely to occur in southeast Angola
Bark Snake – Hemirhaggheris nototaenia (Günther, 1864). This dwarf arboreal
snake is recorded from the western Caprivi and Okavango region, and extends
eastwards through Zambia to East Africa. Earlier records from southwest Angola
(Bocage, 1895) were later referred to H. viperina (Broadley and Hughes 2000).
It is a secretive snake and may still be found in the miombo woodlands of south-
east Angola.
Cunene Racer – Mopanveldophis zebrinus (Broadley and Schätti 2000). This enig-
matic colubrid snake remains known from only a handful of specimens. The type
locality is the Cunene River at Ruacana, western Owamboland, Namibia (17°
25’S, 14° 10′E), and it appears restricted to the Mopaneveld of northern Namibia
and can be expected to occur in similar habitat in southern Angola.
Endemism in Angolan Reptiles
Species that are fully endemic or near endemic to a country (i.e. those that have over
90% of known records included in that country), should be highlighted for national
conservation monitoring as their protection depends completely on the national
authorities. Only six species of snake are endemic to Angola, but no chelonians or
crocodilians. Endemic snakes include two species of primitive thread snake,
Namibiana latifrons and N. rostrata, that are the northern members of a small genus
(five species) endemic to the western arid region of southern Africa (Adalsteinsson
et al. 2009). Three rare snakes are also endemic to the high plateau region, including
the psammophines Psammophis ansorgi and Psammophylax ocellatus (Branch
et al. 2019), as well as the rare and iconic Bitis heraldica, which may be now of high
conservation concern. During the Hamburg Expedition 10 specimens were col-
lected from Bela Vista (Hellmich 1957a, b), but only one other specimen (Mount
Moco) has been recorded in last 60 years (FM Gonçalves, photo 2010). Extensive
clearing of natural habitat for agriculture, and increased fire risk in these montane
grasslands may threaten the species.
Lizards contain the greatest number of endemic and near-endemic Angolan rep-
tiles, particularly among cordylids (two endemic, 50%), lacertids (one near endemic,
six endemic, 53.8%), rupicolous geckos (eight endemic, 23.5%), amphisbaenians
(one near endemic, three endemic species; 36.4%), and diverse skinks (one near
endemic, six endemic, 16.3%). Agama planiceps schacki is certainly a full species
that is well-defined morphologically, but requires genetic assessment. It would also
be endemic to Angola. None of these endemic lizards are currently considered of
conservation concern. Only 12.7% of all Angolan reptiles are endemic as opposed
to 41.2% of those in South Africa. This number increases to nearly 20.5% when
only lizards are considered, but is still much less than the 54.9% of endemic lizards
in South Africa (Table 13.3). However, the number of endemic species in the coun-
try has increased with the description of new Angolan taxa (e.g. Kolekanus plumi-
caudus, Pedioplanis haackei, P. huntleyi and Cordylus namakuiyus), and will
increase further as new species in the genera Nucras, Heliobolus, Pedioplanis,
Afroedura, Rhoptropus, Pachydactylus, Trachylepis and Boaedon discovered during
recent surveys are described.
Reptile Hotspots
The existing global protected area network and conservation priorities are heavily
biased towards amphibian, avian and mammal faunas (Roll et al. 2017). Reptiles,
which represent a third of terrestrial vertebrate diversity, have been largely ignored,
in part, because their diversity and distribution was not globally assessed until 2017.
Both the global (Roll et al. 2017) and African (Lewin et al. 2016) assessments dem-
onstrated that whilst the distribution patterns of species richness of all reptiles com-
bined, as well as those of snakes, revealed similar patterns to those of the other three
tetrapod classes, the patterns displayed by hotspots of total and endemic lizards and
chelonian richness do not overlap significantly with those of other terrestrial tetra-
pods. A detailed analysis of reptile hotspots within Angola awaits fuller details of
species diversity and distributions, both of which are still in their formative period.
However, it is already evident that certain regions and their associated habitats and
reptile faunas, particularly for endemic or near endemic species, present unique
associations, some of which may be confirmed as regionally important reptile
hotspots.
Kaokoveld Centre of Endemism
Lizard diversity in southern Africa, particularly in the western arid regions, is the
highest in Africa, and the existence of similar habitat structure and diversity in
southwest Angola indicates that this African lizard hotspot may also extend into
Angola in association with arid and hyper-arid habitats. In association with desert
habitats a number of characteristic Namib reptiles cross the Cunene River and just
enter extreme southwest Angola, including: Gerrhosaurus skoogi, Pachydactylus
rangei, P. vanzyli, Chamaeleo namaquensis, Meroles anchietae, M. reticulata,
Trachylepis puncutula, and Bitis caudalis. Recent discoveries also suggest the exis-
tence of an endemic Angolan Namib reptile fauna, including the existing Angolan
Namib endemics Pedioplanis benguellensis, Typhlacontias rudebecki, and T. punc-
tatissimus bogerti, as well as a number newly described species in the region, e.g.
Kolekanus plumicaudus (Haacke 2008), Pedioplanis huntleyi, P. haackei (Conradie
et al. 2012a), and Cordylus namakuiyus (Stanley et al. 2016). Moreover, recent sur-
veys in the region have revealed numerous examples of cryptic diversity in some
lizard genera, where new species of Afroedura, Pachydactylus, Nucras, Pedioplanis
endemic to the Angolan Namib region have been identified and await description.
At its northern and southern limits, the Namib Desert transforms into semi-arid,
often succulent vegetation that may be loosely termed the ‘Pro-Namib’ region. In
the south this forms the Succulent Karoo, a botanical hotspot of regional endemism
and floral beauty (CEPF 2003). The Succulent Karoo has diverse and specialised
reptile endemics (Branch 1994; Bauer and Branch 2003), and the region has been
highlighted as a regional reptile hotspot, including numerous species of conserva-
tion concern (Branch 2014). As with the Succulent Karoo, the recognition of a
unique reptile fauna in southwest Angola supports a corresponding northern ‘Pro-
Namib’, in some ways analogous to the Succulent Karoo, and that has been identi-
fied as a distinctive phytogeographical region – the Kaokoveld Centre of Endemism,
which extends as a narrow strip north of Namibe to Lucira and is characterised by a
number of localised succulents (see Craven 2009 for fuller discussion).
Angolan Escarpment
Inland from the coastal arid herpetofauna is the Bié section (sensu Clark et al. 2011)
of the Angolan Escarpment and adjacent high plateau. The southern African Great
Escarpment (GE) forms a semi-continuous U-shaped mountain chain that runs for
5000 km from western Angola through Namibia and South Africa to the Zimbabwe-
Mozambique border. Clark et al. (2011) noted that the GE hosts more than half of
southern Africa’s centres of plant endemism and is a repository of palaeo- and neo-
endemics. It also has a rich endemic fauna and its fragmented sections serve as
refugia and as episodic corridors for biological continuity. However, many sections
of the Great Escarpment have been poorly studied, particularly in Angola where the
Bié Escarpment summit and adjacent highlands is one of the most isolated sections
of the Afromontane archipelago in Africa. With ca. 20 endemic bird species it forms
the core of the Western Angola Endemic Bird Area. Other faunal groups have not
been as extensively studied, but endemic reptiles associated with the Serra da Chela
grasslands and wetlands include two endemic snakes (Psammophylax ocellatus and
Psammophis ansorgi), the chameleon Chamaeleo anchietae, the serpentine skink
Eumecia anchietae, the skink Trachylepis bayoni huilensis, the gecko Rhoptropus
montanus, and the lacertid Ichnotropis bivittata pallida. A new reed frog, Hyperolius
chelaensis was also recently discovered (Conradie et al. 2012b). In the adjacent
highlands, including Mount Moco, at least two new species of the Afroedura bogerti
complex have also been signalled (Branch et al. 2017).
Northern Congo Forests
The Congo Basin has numerous forest specialists, particularly snakes. Many of
these are found in forests in Cabinda and along the northern border of Angola.
These forests have only been incidentally surveyed, particularly the numerous
important snake records listed in a series of papers based on the Museu do Dundo
collections (Laurent 1950, 1954, 1964; Tys van den Audenaerde 1967). Among
these collections are the only known Angolan records Gonionotophis brussauxi
Letheobia praeocularis, Xenocalamus bicolour machadoi, Hypoptophis wilsoni
katangae, Grayia tholloni, Philothamnus nitidus, Bothrophthalmus lineatus,
Boaedon olivaceus, Lycodonomorphus subtaeniatus, Prosymna ambigua brevis and
Causus lichtensteini. In addition, other Congo Basin reptiles only recorded from
Dundo include the terrapin Pelusios gabonensis, the worm lizard Monopeltis
vanderysti, and the skinks Lepidothyris hinkeli joei (as Mochlus fernandi, Laurent
1964) and Feylinia grandisquamis (as F. elegans, Laurent 1964). Parker (1936) pre-
sented the first survey of the central scarp forests of the Angolan Escarpment and
recorded numerous Congo Basin snakes. For many species these remain their south-
ern records, and they probably occur as disjunct, relictual populations. Some were
subsequently recorded further north in forest habitats from Dundo or during the
Hamburg Angola Expedition at Piri Dembos (see above). They include: Toxicodryas
blandingii, T. pulverulenta, Atractaspis reticulata heterochilus, Bitis nasicornis,
and Pseudohaje goldii. Others remain known only from Parkers’ records: i.e. the
skinks Panaspis breviceps, Leptosiaophis dewittei, and Trachylepis affinis; and the
snakes Lycophidion ornatum, Chamaelycus parkeri (as Oophilositum parkeri) and
Hormonotus modestus. The Congo Basin snake Rhamnophis aethiopissa is recorded
in Angola only from Piri Dembos (Hellmich 1957a, b). The taxonomic status of all
these isolates requires genetic confirmation as some may have undergone vicariant
speciation. A phylogenetic assessment may give insight towards dating the separa-
tion of these forest isolates and understanding their biogeographic importance.
The forests of Cabinda form part of the Congo Basin and a number of reptiles
occur there which have not been recorded in Angola south of the Congo River.
Currently Cabinda remains the southern limit of the African Dwarf Crocodile
(Osteolaemus tetraspis) and the Soft-shelled Terrapin (Cycloderma). The presence
of two other reptiles recorded from Cabinda by Peters (1876, 1877), e.g. Owen’s
Horned Chameleon (Triceros oweni) and the skink Euprepes perrotetii
(= Trachylepis perrotetii) are problematic. The latter is widespread in West Africa
but not known even from Gabon. Peters (1877) recorded Euprepes perrotetii from
Chinchoxo, Cabinda, and in a supplement to the same article noted a specimen from
Pungo Andongo, upon which he considered it to form part of the fauna of Angola.
However, no subsequent records of this distinctive species have been recorded from
Angola. Although it is possible that these specimens were confused with large fire
skinks (Lepidothrys sp.), Wagner et al. (2009) reviewed the genus and noted no
misidentifications among the material they examined. It is more likely that Peters’
specimens were simply accompanied by incorrect locality data. Forest chameleons
are difficult to locate unless specifically targeted during faunal surveys, and Owen’s
Horned Chameleon is known from Gabon. No recent collections of both these spe-
cies confirm their presence in Cabinda. Research underlying the proposed Mayombe
Transfrontier Reserve (MTR) to protect forests in Cabinda and adjacent countries
has concentrated on the large mammals, particular the Great Apes, and no detailed
herpetological surveys have been undertaken. Recent surveys of the forest herpeto-
fauna of the Serra do Pingano Ecosystem, Uíge Province (Ernst 2015) concentrated
on amphibians but did record an number of interesting reptiles, particularly the
arboreal lacertid Holaspis guentheri and water snake Grayia ornata, the former
being the second record for the country (Laurent 1964) and the latter one of the few
records for the country (Branch 2018). The northern tropical forests of Angola are
threatened by massive timber extraction, and desperately need to be scientifically
surveyed before their associated herpetofauna is lost.
Future Directions for Reptile Research in Angola
he Continued Need for Further Field Surveys and Taxonomic

T
Studies
The conservation status and threats for African reptiles were reviewed by Tolley
et al. (2016), who noted the large discrepancy between taxonomic sampling and
documentation between many countries. They presented a scatter-plot of measured
reptile species richness relative to log-transformed country area from African coun-
tries. This illustrated the great contrast between known reptile diversity in well sur-
veyed countries such as South Africa, Kenya and Tanzania, with that of the majority
of Africa. Angola is the seventh largest African country and has both habitat and
topographic diversity. Together these features should generate rich biological diver-
sity, but this is not reflected in our current knowledge of Angolan reptile diversity.
Branch (2016) presented species accumulation curves documenting the growth in
taxonomic knowledge of Angolan and southern African reptiles, noting that there
has been no decline in the rate of new species discovery in the subcontinent during
the last 150 years. This is in marked contrast to the relative stagnation of taxonomic
discovery in Angola since the early part of the twentieth century (see Fig. 13.1 and
Table 13.1 for comparison). As noted earlier, despite Angola and South Africa being
of comparable size and habitat diversity there is a difference of over 150 species of
Fig. 13.1 Species accumulation curves for reptile discoveries in Angola (blue) and southern
Africa (red) showing the relative stagnation of Angolan reptile species descriptions during the
twentieth century. By the end of the nineteenth century 67.4% of Angolan reptiles had already been
described, in contrast to less than half (47.8%) of those in southern Africa
lizards between the two countries (Angola 132, South Africa 286). This contrast is
even higher in terms of endemicity, where only 27 (20.5%) of 132 Angolan lizards
are endemic in contrast to 157 (54.9%) of 286 South African lizards. It would appear
that perhaps as many as 75+ new lizard species await discovery in Angola, and that
many of these will be endemic. Branch (2014) noted that endemicity in South
African lizards was particularly evident in rupicolous forms (many geckos, cordylids
and skinks) associated with rocky outcrops. Rock exposures may form an archi-
pelago of ‘sky islands’ on which isolation inhibits gene flow and thus leads to spe-
ciation. It is the lizard families containing large numbers of rupicolous species, i.e.
Gekkonidae, Cordylidae and Scincidae, already show the greatest levels of ende-
micity among Angolan reptiles, and in which recent surveys have already identified
numerous cryptic taxa (Stanley et al. 2016; Branch et al. 2017).
Field Surveys of Potential Biodiversity ‘Hotspots’
Many African protected areas underperform in their stated conservation goals

(Lindsey et al. 2014; Bowker et al. 2017). It is now generally accepted that modern
national and internationally co-ordinated networks of protected area should be
designed to cover important biodiversity hotspots and also protect habitats essential
for the maintenance of ecosystem services such as water flow and quality, nutrient
transfer, etc. (NPAES 2010). Such a revised Angolan network was proposed many
years ago (Huntley 1974), and initial biodiversity surveys to gain insight into biodi-
versity in potential sites were undertaken (Huntley 2009; Huntley and Francisco
2015). Recent studies have shown that protected reserves designed to protect mam-
mals, birds and amphibians are effective in protecting snakes, but fare badly in
protecting African lizard diversity (Lewin et al. 2016; Roll et al. 2017). Future sys-
tematic biodiversity surveys should be directed to unique habitats and landforms in
undersampled regions. Some of the interesting species recorded on recent field sur-
veys are illustrated in Fig. 13.2.
Biogeography of Angolan Reptiles
Huntley (2019) in the introduction to this volume has presented a biogeographic

outline, summarising various aspects of climate, geology and vegetation, etc., that
characterise Angola. He noted the complexity of the Angolan landscape, where
seven of the nine African biomes are represented in Angola as well as the second
largest representation of ecoregion diversity in Africa. Monard (1937) and Hellmich
(1957a, b) made preliminary attempts to assess biogeographic patterns among
Angolan reptiles. However, they did little more than look for coarse habitat associa-
tions within the Ethiopian region. These attempts were constrained by lack of
knowledge of the true reptile diversity in the region and, more importantly, by the
ignorance of reptile distributions as large tracts of the country were still unexplored.
Moreover, recent studies indicate that reptile distributions, particularly those of liz-
ards, are more influenced by substrate specificity and isolation than by vegetation
type (Bauer and Lamb 2005; Roll et al. 2017). Recent biogeographic studies lay
greater emphasis on evolutionary relationships within the group studied, and explore
correlations between genetic divergence (as a proxy for time) and known dates of
major events in landscape evolution. This approach searches for historical barriers
to, or corridors for migration and gene flow. These may be generated, for example,
by climatic changes associated with Ice Age cycles and the resultant contraction and
expansion of forest and savanna, changes in historic coastlines and/or island con-
nectivity, as well as the development of an ‘arid’ corridor at an Ice Age maxima, etc.
The biological consequences of nascent rifting on river capture and other hydrologi-
cal consequences on palaeolakes and wetlands have also been explored (Cotterill
and De Wit 2011). However, the application of such approaches requires more
detailed knowledge of reptile distribution within Angola, as well as the availability
of genetic material and adequate taxon sampling within a chosen group. These will
allow historic climatic and landform events to be meaningful correlated with specia-
tion and radiation within groups for testing phylogeographic hypotheses. Such stud-
ies depend on meaningful progress in the topics discussed earlier in this section.
Advances in all these areas are required to fully understand and conserve the diver-
sity and evolution of Angolan reptiles.
Fig. 13.2 Angolan reptiles. Top to bottom, left to right. Bogert’s Flat Gecko (Afroedura cf.
bogerti), Omahua Lodge, Namibe; Angolan Namib Day Gecko (Rhoptropus taeniostictus), Chapéu
Armado, Namibe; Ansorge’s Leaf-toed Gecko (Afrogecko ansorgii), Meve, Benguela; Anchieta’s
Chameleon (Chamaeleo anchietae), Humpata, Huíla; Kaokoveld Girdled Lizard (Cordylus namak-
uiyus), Rio Makonga, Namibe; Bayon’s Legless Skink (Sepsina bayoni), Quiçama National Park,
Luanda; Water Cobra (Naja annulata), Lagoa Carumbo, Lunde Norte; Angolan Skaapstekker
(Psammophylax ocellatus), Humpata, Huíla
Acknowledgements This review results from extensive collaboration by the authors and the
Editor’s of this volume during recent studies on the Angolan herpetofauna. It has been both syner-
gistic and rewarding. Funding for research in Angola has been supported by: South Africa’s
National Research Foundation (2009–2017, WRB), National Geographic Society (Explorer Grant
2011, WRB; NGOWP and Wild Bird Trust 2015–2018, all authors), Fundação para a Ciência e
Tecnologia (contract SFRH/PD/BD/140810/2018, NB). We are all particularly indebted to
Fernandas Lages (ISCED), Brian Huntley (South Africa), and John Hilton (Wild Bird Trust) for
their support with the documentation, logistics and permitting required for successful fieldwork in
Angola.
Appendices
Appendix 1
Checklist of Angolan Chelonians and Crocodilians. C: Cabinda; Status: CITES (I,

II = Appendix 1 or 2), IUCN Conservation Status1. Species listed under
Order|Family
Common Name Scientific Name C Bocage (1895) Status

Chelonia | Chelonidae
Loggerhead Sea Turtle Caretta caretta (Linnaeus, 1758)
Thalassochelys I, VU
caretta
Green Sea Turtle Chelonia mydas (Linnaeus, 1758) Y Chelonia mydas I, EN
Olive Ridley Sea Turtle Lepidochelys olivacea Y I, VU
(Eschscholtz, 1829)
Hawksbill Sea Turtle Eretmochelys imbricata I, CR
(Linnaeus, 1766)
Chelonia | Dermochelyidae
Leatherback Sea Turtle Dermochelys coriacea (Vandelli, Y I, VU
1761)
Chelonia | Testudinidae
Bell’s Hinge-back Kinixys belliana (Gray, 1831) Cinixys belliana II
Tortoise
Forest Hinge-back Kinixys erosa (Schweigger, 1812) Y Cinixys erosa II, ENa
Tortoise
Leopard Tortoise Stigmochelys pardalis (Bell, Testudo pardalis II
1828)
Chelonia | Pelomedusidae
Southern Marsh Terrapin Pelomedusa subrufa (Bonnaterre, Pelomedusa
1789) galeata
Okavango Hinged Pelusios bechuanicus
Terrapin (FitzSimons, 1932)
Gabon Hinged Terrapin Pelusios gabonensis (Duméril,
1856)
(continued)
Common Name Scientific Name C Bocage (1895) Status

Dwarf Hinged Terrapin Pelusios nanus (Laurent, 1956)
Variable Hinged Terrapin Pelusios rhodesianus (Hewitt, Sternothaerus
1927) sinuatus
Western Hinged Terrapin Pelusios castaneus (Schweigger, Y Sternothaerus
1812) Derbianus
Chelonia | Trionycidae
Nile Soft-shelled Trionyx triunguis (Forskȧl, 1775) Y Trionyx triunguis II, VUa
Terrapin
Aubrey’s Flap-shelled Cycloderma aubryi (Dumeri,, Y Cycloderma II, VUa
Terrapin 1856) Aubryi
Crocodylia | Crocodylidae
Nile Crocodile Crocodylus niloticus (Laurenti, Crocodilus II
1768) vulgaris
Central African Mecistops leptorhynchus (Shirley Crocodylus I, DD
slender-snouted et al. 2018) cataphractus
Crocodile
African Dwarf Crocodile Osteolaemus tetraspis (Cope, Y Ostelolaemus I, EN
1861) tetraspis
Turtle Working Group 2017, Draft Red List
a
IUCN Conservation Status categories. CR Critically Endangered, EN Endangered, VU Vulnerable,

b
DD Data Deficient
Appendix 2
Checklist of Angolan Lizards. C: Cabinda; Obs: Observations (E:endemic; NE:

near-endemic). Species listed under Order|Family|Subfamily
Common name Scientific name C Bocage (1895) Obs

Sauria | Agamidae
Angolan Tree Agama Acanthocercus Stellio angolensis Bocage
cyanocephalus (Falk, 1925) 1866 is a nomen nudum
Stellio atricollis
Ground Agama Agama aculeata (Merrem, Agama armata
1820)
Anchieta’s Agama Agama anchietae (Bocage,
1896)
Congo Agama Agama congica (Peters, Y A. colonorum
1877)
Mucoso Agama Agama mucosoensis E
(Hellmich, 1957)
Namib Rock Agama Agama planiceps planiceps Agama planiceps
(Peters, 1862)
Schack’s Rock Agama Agama p. schacki (Mertens, E
1938)
(continued)

Sauria | Amphisbaenidae
Angola Blunt-tailed Dalophia angolensis NE
Worm Lizard (Gans, 1976)
Ellenberger’s Blunt-tailed Dalophia ellenbergeri
Worm Lizard (Angel, 1920)
Zambezi Blunt-tailed Dalophia pistillum
Worm Lizard (Boettger, 1895)
Welwitch’s Blunt-tailed Dalophia welwitschii Monopeltis Welwitschii E
Worm Lizard (Gray, 1865)
Anchieta’s Spade-snouted Monopeltis anchietae Monopeltis anchietae
Worm Lizard (Bocage, 1873)
Infuscate Spade-snouted Monopeltis infuscata Monopeltis capensis
Worm Lizard (Broadley, 1997)
Luanda Spade-snouted Monopeltis luandae (Gans, E
Worm Lizard 1976)
Confusing Spade-snouted Monopeltis perplexus E
Worm Lizard (Gans, 1976)
Vanderyst’s Spade- Monopeltis vanderysti (De
snouted Worm Lizard Witte, 1922)
Balck Round-headed Zygaspis nigra (Broadley
Worm Lizard and Gans, 1969)
Kalahari Round-headed Zygaspis quadrifrons
Worm Lizard (Peters, 1862)
Sauria | Chamaeleonidae
Angolan Chameleon Chamaeleo anchietae Chamaeleo anchietae
(Bocage, 1872)
Flap-necked Chameleon Chamaeleo dilepis (Leach, Y Chamaeleon dilepis & C.
1819) quilensis
Etienne’s Chameleon Chamaeleo gracilis Y Chamaeleon gracilis
etiennei (Schmidt, 1919)
Namaqua Chameleon Chamaeleo namaquensis Chamaeleon
(Smith, 1831) namaquensis
Owen’s Three-horned Trioceros oweni (Gray,
Chameleon 1831)
Sauria | Cordylidae
Northern Grass Lizard Chamaesaura miopropus Chamaesaura macrolepis
(Boulenger, 1895)
Angola Grass Lizard Chamaesaura anguina E
oligopholis (Laurent, 1964)
Angola Girdled Lizard Cordylus angolensis Zonurus cordylus E
(Bocage, 1895)
Machado’s Girdled Cordylus machadoi
Lizard (Laurent, 1964)
Kaokoveld Girdled Cordylus namakuiyus E
Lizard (Stanley et al, 2016)
(continued)

Sauria | Gekkonidae
Bogert’s Flat Gecko Afroedura bogerti
(Loveridge, 1944)
Ansorge’s Leaf-toed Afrogecko ansorgii E
Gecko (Boulenger, 1907)
Button-scaled Gecko Chondrodactylus
fitzsimonsi (Loveridge,
1947)
Pulitzer’s Gecko Chondrodactylus pulitzerae Pachydactylus Bibronii
(Schmidt, 1933) (part)
Fisher’s Gecko Chondrodactylus Pachydactylus Bibronii
laevigatus (Fischer, 1888) (part)
Pulitzer’s Gecko Chondrodactylus pulitzerae
(Smith, 1933)
Bayão’s House Gecko Hemidactylus bayonii Hemidactylus bayonii E
(Bocage, 1893)
Benguela House Gecko Hemidactylus benguellensis Hemidactylus E
(Bocage, 1893) benguellensis
Western House Gecko Hemidactylus brooki
angulatus (Hallowell 1852)
Long-headed House Hemidactylus Hemidactylus bocagii
Gecko longicephalus (Bocage,
1873)
Tropical House Gecko Hemidactylus mabouia Y Hemidactylus mabouia &
(Moreau De Jonnès, 1818) H. benguellensis
Forest House Gecko Hemidactylus murecius Hemidactylus murecius
(Peters, 1870)
Plume-tailed Geco Kolekanos plumicaudus E
(Haacke, 2008)
Angola Dwarf Day Lygodactylus angolensis
Gecko (Bocage, 1896)
Bradfield’s Dwarf Day Lygodactylus bradfieldi
Gecko (Hewitt, 1932)
Cape Dwarf Day Gecko Lygodactylus capensis Lygodacylus capensis
(Smith, 1849)
Hewitt’s Punctate Gecko Pachydactylus amoenoides Pachydactylus ocellatus
(Hewitt, 1935)
Angola Thick-toed Gecko Pachydactylus angolensis E
(Loveridge, 1944)
Caricul Thick-toed Gecko Pachydactylus caraculicus
(FitzSimons, 1959)
Kaokoveld Thick-toed Pachydactylus cf.
Gecko oreophilus (McLachlan and
Spence, 1976)
(continued)

Punctate Thick-toed Pachydactylus punctatus
Gecko (Peters, 1854)
Web-footed Gecko Pachydactylus rangei
(Andersson, 1908)
Scherz’s Thick-toed Pachydactylus scherzi
Gecko (Mertens, 1954)
Rough-scaled Thick-toed Pachydactylus cf. rugosus NR
Gecko (Smith, 1849)
Large-scaled Thick-toed Pachydactylus scutatus
Kalahari Ground Gecko Pachydactylus wahlbergii
(Peters, 1869)
Van Zyl’s Web-footed Pachydactylus vanzyli
Gecko (Steyn and Haacke, 1966)
Common Namib Day Rhoptropus afer (Peters,
Gecko 1869)
Barnard’s Namib Day Rhoptropus barnardi Rhoptropus afer ?
Benguella Namib Day Rhoptropus benguellensis E
Gecko (Mertens 1938)
Two-pored Namib Day Rhoptropus biporosus
Gecko (FitzSimons, 1957)
Boulton’s Namib Day Rhoptropus boultoni
Gecko (Schmidt, 1933)
Montane Namib Day Rhoptropus montanus E
Gecko (Laurent, 1964)
Angolan Namib Day Rhoptropus taeniostictus E
Gecko (Laurent, 1964)
Sauria | Gerrhosauridae
Dwarf Plated Lizard Cordylosaurus Cordylosaurus trivittatus
subtessellatus (Smith,
1844)
Kalahari Plated Lizard Gerrhosaurus auritus
(Boettger, 1887)
Laurent’s Plated Lizard Gerrhosaurus bulsi
(Laurent, 1954)
Keeled Plated Lizard Gerrhosaurus multilineatus
(Bocage, 1866)
Black-lined Plated Lizard Gerrhosaurus nigrolineatus Y Gerrhosaurus
(Hallowell, 1857) nigrolineatus
Desert Plated Lizard Gerrhosaurus skoogi
(Andersson, 1916)
Western Giant Plated Matobosaurus maltzahni Gerrhosaurus validus
Lizard (De Grys, 1938)
Ellenberger’s Snake Tetradactylus ellenbergeri Caitia africana Gray
Lizard (Angel, 1922)
(continued)

Sauria | Lacertidae
Bushveld Lizard Heliobolus lugubris Eremias lugubris
(Smith, 1838)
Northern Blue-tailed Tree Holaspis guentheri (Gray,
Lizard 1863)
Western Rough-scaled Ichnotropis b. bivittata Ichnotropis capensis
Lizard (Bocage, 1866)
Pale Rough-scaled Lizard Ichnotropis b. pallida E
(Laurent, 1964)
Cape Rough-scaled Ichnotropis c. capensis
Lizard (Smith, 1838)
Overlaete’s Rough-scaled Ichnotropis c. overlaeti
Lizard (Witte and Laurent 1942)
Small-scaled Rough- Ichnotropis microlepidota E
scaled Lizard (Marx, 1956)
Shovel-snouted Lizard Meroles anchietae Pachyrhynchus Anchietae
(Bocage, 1867)
Reticulate Desert Lizard Meroles reticulatus Scaptira reticulata
(Bocage, 1867)
Rough-scaled Desert Meroles squamulosus
Lizard (Peters, 1854)
Laurent’s Sandveld Nucras scalaris (Laurent, E
Lizard 1964)
Western Sandveld Lizard Nucras aff. tesselata NE
(Smith, 1838)
Benguella Sand Lizard Pedioplanis benguellensis Eremias namaquensis E
(Bocage, 1867)
Haacke’s Sand Lizard Pedioplanis haackei E
Conradie et al. 2012
Huntley’s Sand Lizard Pedioplanis huntleyi E
(Conradie et al. 2012)
Sauria | Scincidae | Acontinae
Japp’s Burrowing Skink Acontias jappi (Broadley,
1968)
Kalahari Burrowing Acontias kgalagadi
Skink kgalagadi (Lamb et al.,
2010)
Western Burrowing Skink Acontias occidentalis
(FitzSimons, 1941)
Sauria | Scincidae | Eugongylinae
Shorted-headed Panaspis breviceps (Peters,
Snake-eyed Skink 1873)
Cabinda Snake-eyed Panaspis cabindae Y Ablepharus cabindae
Skink (Bocage, 1866)
Speckle-lipped Snake- Panaspis maculicollis
eyed Skink (Jacobsen and Broadley,
2000)
(continued)

Angolan Snake-eyed Panaspis aff. wahlbergii Ablepharus wahlbergii
Skink complex
De Witte’s Leaf-litter Leptosiaphos dewittei
Skink (Loveridge, 1934)
Sauria | Scincidae | Lygosominae
Hinkel’s Red-sided Skink Lepidothyris hinkeli
(Wagner et al., 2009)
Sundevall’s Writhing Mochlus sundevalli (Smith, Lygosoma Sundevallii
Skink 1849)
Sauria | Scincidae | Mabuyinae
Anchieta’s Snake Skink Eumecia anchietae Lygosoma Anchietae
anchietae (Bocage, 1870)
Lunda Western Snake Eumecia a. major (Laurent, E
Skink 1964)
Iven’s Water Skink Lubuya ivensii (Bocage, Lygosoma Ivensii
1879)
Wedge Snouted Skink Trachylepis acutilabris Y Mabuia acutilabris
(Peters, 1862)
Senegal Skink Trachylepis affinis (Gray, Y Mabuia Raddonii (not in
1838) Angola)
Monard’s Skink Trachylepis monardi E
(Marques et al. 2018)
Bayão’s Skink Trachylepis b. bayoni Mabuia Bayonii
(Bocage, 1872)
Huila Skink Trachylepis b. huilensis E
(Laurent, 1964)
Ovambo Stree Skink Trachylepis binotata Mabuia bionotata
(Bocage, 1867)
Bocage’s Skink Trachylepis bocagii Mabuia Petersi
(Boulenger, 1887)
Chimba Skink Trachylepis chimbana Mabuia chimbana
(Boulenger, 1887)
Damara Skink Trachylepis damarana Mabuia varia (part)
(Peters, 1870)
Hoesch’s Skink Trachylepis hoeschi
(Mertens, 1954)
Bronze Rock Skink Trachylepis cf.
lacertiformis (Peters, 1854)
Angolan Blue-tailed Trachylepis laevis
Skink (Boulenger, 1907)
Speckled-lipped Skink Trachylepis maculilabris Y Mabuia maculilabris
(Gray, 1845)
Grass Skink Trachylepis cf. megalura
(Peters, 1878)
Western Three Striped Trachylepis occidentalis Mabuia occidentalis
Skink (Peters, 1867)
(continued)

Speckled Skink Trachylepis punctulata Mabuia punctulata
(Bocage, 1872)
Kalahari Tree Skink Trachylepis spilogaster
(Peters, 1882)
Striped Skink Trachylepis striata (Peters, Mabuia striata
1844)
Ansorge’s Rock Skink Trachylepis sulcata Mabuia sulcata
ansorgii (Boulenger, 1907)
Angolan Variable Skink Trachylepis cf. Mabuia varia (part)
albopunctata (Bocage,
1867)
Wahlberg’s Skink Trachylepis wahlbergi
(Peters, 1869)
Sauria | Scincidae | Scincinae
Curror’s giant burrowing Feylinia currori (Gray, Y Feylinia Currori
Skink 1845)
Large-scaled burrowing Feylinia grandisquamis
Skink (Müller, 1910)
Western Limbless Skink Melanoseps occidentalis
(Peters, 1877)
Angolan burrowing Skink Sepsina angolensis Sepsina angolensis
(Bocage, 1866)
Bayão’s Burrowing Skink Sepsina bayoni (Bocage, Y Sepsina Bayonii NE
1866)
Cope’s Burrowing Skink Sepsina copei (Bocage, Sepsina Copei E
1873)
Johnson’s Western Typhlacontias johnsonii
Burrowing Skink (Andersson, 1916)
Speckled Western Typhlacontias p. Typhlacontias
Burrowing Skink punctatissimus (Bocage, punctatissimus
1873)
Bogert’s Western Typhlacontias p. bogerti E
Burrowing Skink (Laurent, 1964)
Rohan’s Western Typhlacontias rohani
Burrowing Skink (Angel, 1923)
Rudebeck’s Western Typhlacontias rudebecki E
Burrowing Skink (Haacke, 1997)
Sauria | Varanidae
Savanna Monitor Varanus albigularis Varanus albigularis
angolensis (Schmidt, 1933)
Water Monitor Varanus niloticus Y Varanus niloticus
(Linneaus, 1766)
Appendix 3
Checklist of Angolan Snakes. C: Cabinda; Obs: Observations (E:endemic; NE:

near-endemic; NR: new record for Angola; RC: requires confirmation). Species
listed under Order | Family | Subfamily

Scolecophidia | Leptotyphlopidae
Shaba Thread Snake Leptotyphlops kafubi
(Boulenger, 1919)
Peter’s Thread Snake Leptotyphlops scutifrons Stenosoma
(Peters, 1854) scutifrons
Damara Thread Snake Namibiana labialis (Sternfeld,
1908)
Benguela Thread Snake Namibiana latifrons (Sternfeld, E
1908)
Angolan Beaked Thread Namibiana rostrata (Bocage, Stenosoma E
Snake 1886) rostratum
Scolecophidia | Typhlopidae
Angolan Blind Snake Afrotyphlops angolensis Typhlops punctatus
(Bocage, 1866)
Angolan Giant Blind Afrotyphlops anomalus Typhlops anomalus
Snake (Bocage, 1873) & T. anchietae
Blotched Blian Snake Afrotyphlops congestus Y
(Duméril and Bibron, 1844)
Lined Blind Snake Afrotyphlops lineolatus (Jan, Y Typhlops punctatus
1864) var. lineolatus &
Typhlops boulengeri
Schmidt’s Blind Snake Afrotyphlops schmidti (Laurent,
1956)
Schlegel’s Blind Snake Afrotyphlops schlegelii Typhlops petersii,
(Bianconi, 1847) Typhlops humbo
& Typhlops
hottentotus
Giant Blind Snake Afrotyphlops mucruso (Peters, Typhlops mucruso
1854)
Leopoldville Beaked Letheobia praeocularis
Blind Snake (Stejneger, 1894)
Henophidia | Pythonidae
Namib Dwarf Python Python anchietae (Bocage, Python anchietae
1887)
Southern African Python Python natalensis (Smith, Python natalensis
1840)
Northern African Python Python sebae (Gmelin, Y
1789)
(continued)

Henophidia | Colubridae | Colubrinae
White-lipped Snake Crotaphopeltis hotamboeia Y Crotaphopeltis
(Laurenti, 1768) rufescens
Barotse Water Snake Crotaphopeltis barotseensis NR
(Broadley, 1968)
Confusing Egg-eater Dasylepis confusa (Trape and NR
Mané, 2006)
Palm Egg-Eater Dasypeltis palmarum (Leach, Y Dasypeltis scabra
1818) var. palmarum
Rhombic Edd-Eater Dasypeltis scabra (Linnaeus, Dasypeltis scabra
1758)
Shreve’s Tree Snake Dipsadoboa shrevei
(Loveridge, 1932)
Punctate Boomslang Dispholidus typus punctatus Bucephalus capensis
(Laurent, 1955) (part)
Green Boomslang Dispholidys t. viridis (Smith, Bucephalus capensis
1838) (part)
Emerald Snake Hapsidophrys smaragdinus Y Hapsidophrys
(Schlegel, 1837) smaragdina
Angolan Green Snake Philothamnus angolensis Y Philothamnus
(Bocage, 1882) irregularis
Thirteen-scaled Green Philothamnus carinatus
Snake (Andersson, 1901)
Striped Green Snake Philothamnus dorsalis Philothamnus
(Bocage, 1866) dorsalis
Emerald Green Snake Philothamnus heterodermus Philothamnus
(Hallowell, 1857) heterodermus
Slender Green Snake Philothamnus heterolepidotus Philothamnus
(Günther, 1863) heterolepidotus
Southeastern Green Snake Philothamnus hoplogaster
(Günther, 1863)
Loveridge’s Green Snake Philothamnus nitidus
loveridgei (Laurent, 1960)
Ornate Green Snake Philothamnus ornatus (Bocage, Philothamnus
1872) ornatus
Spotted Bush Snake Philothamnus semivariegatus Philothamnus
(Smith, 1840) semivariegatus
Large-eyed Green Rhamnophis aethiopissa
Treesnake (Günther, 1862)
Hook-nosed Snake Scaphiophis albopunctatus Scaphiophis
(Peters, 1870) albopunctatus
Damara Tiger Snake Telescopus finkeldeyi (Haacke,
2013)
Western Tiger Snake Telescopus polystictus Crotaphlopeltis
(Mertens, 1954) semiannulatus
Oates’ Vine Snake Thelotornis capensis oatesi Thelotornis kirtlandii
(Günther, 1881)
(continued)

Forest Vine Snake Thelotornis kirtlandii Thelotornis
(Hallowell, 1844) kirtlandii
Yellow-throated Treesnake Thrasops flavigularis Y Thrasops
(Hallowell, 1852) flavigularis
Jackson’s Treesnake Thrasops jacksoni (Günther,
1895)
Blanding’s Treesnake Toxicodryas blandingii Y Dipsas Blandingii
(Hallowell, 1844)
Powdered Treesnake Toxicodryas pulverulenta Y Dipsas pulverulenta
(Fischer, 1856)
Henophidia | Colubridae | Grayinae
Ornate Water Snake Grayia ornata (Bocage, 1866) Y Grayia ornata
Smith’s Water Snake Grayia smithii (Leach, 1818) Grayia triangularis
Thollon’s Water Snake Grayia tholloni (Mocquard,
1897)
Henophidia | Natricidae
Bangweulu Swamp Snake Limnophis bangweolicus
(Mertens, 1936)
Striped Swamp Snake Limnophis bicolor (Günther, Helocops bicolour
1865)
Broadley’s Marsh Snake Natriciteres bipostocularis
(Broadley, 1962)
Olive Marsh Snake Natriciteres olivacea (Peters, Y Mizodon olivaceus
1854)
Henophidia | Lamprophiidae | Atractaspidinae
Common Purple-glossed Amblyodipsas polylepis Calamelaps
Snake (Bocage, 1873) polylepis
Kalahari Purple-glossed Amblyodipsas ventrimaculata NR
Snake (Roux, 1907)
Cape Centipede Eater Aparallactus capensis (Smith, Uriechis capensis
1849)
Birbon’s Burrowing Asp Atractaspis bibronii (Smith, Atractaspis Bibronii
1849)
Congo Burrowing Asp Atractaspis congica (Peters, Y Atractaspis congica
1877)
Reticulate Burrowing Asp Atractaspis reticulata RC
heterochilus (Boulenger, 1901)
Wilson’s burrowing snake Hypoptophis wilsoni
(Boulenger, 1908)
Collared Snake-Eater Polemon collaris (Peters, 1881) Microsoma collare
Bi-coloured Quill-snouted Xenocalamus bicolor
Snake machadoi (Laurent, 1954)
Elongate Quill-snouted Xenocalamus mechowii
Snake mechowii (Peters, 1881)
Inorante Elongate Xenocalamus m. inorantus (de
Quill-snouted Snake Witte and Laurent, 1947)
(continued)

Henophidia | Lamprophiidae | Lamprophiinae
Angolan House Snake Boaedon angolensis (Bocage, Boodon lineatus var.
1895) angolensis, Bocage,
1895
Brown House Snakea Boaedon fuliginosus (Boie,
1827)
Olive House Snake Boaedon olivaceus (Dumeril, Y Boodon olivaceus
1856)
Red-Black Striped House Bothrophthalmus lineatus Bothrophthalmus
Snake (Peters, 1863) lineatus
Parker’s Banded Snake Chamaelycus parkeri (Angel,
1934)
Mocquard’s Dwarf File Gonionotophis brusseauxi
Snake (Mocquard, 1889)
Yellow Forest Snake Hormonotus modestus
(Duméril, Bibron and Duméril,
1854)
Western Forest File Snake Mehelya poensis (Smith, 1849)
Cape File Snake Limaformosa capensis (Smith, Heterolepis Guirali ?
1847)
Savorgan’s File Snake Limaformosa savorgani ? NR
(Moquard, 1887)
Vernay’s File Snake Limaformosa vernayi (Bogert,
1940)
White-bellied Water Snake Lycodonomorphus (?)
subtaeniatus (Laurent, 1954)
Hellmich’s Wolf Snake Lycophidion hellmichi
(Laurent, 1964)
Flat Wolf Snake Lycophidion laterale Lycophidium
(Hallowell, 1857) laterale
Speckled Wolf Snake Lycophidion meleagre Lycophidium
(Boulenger, 1893) meleagris
Spotted Wolf Snake Lycophidion multimaculatum Y Lycophium capense
(Boettger, 1888)
Namib Wolf Snake Lycophidion namibianum NR
(Broadley, 1991)
Ornate Wolf Snake Lycophidion ornatum (Parker, ?
1936)
Viperine Rock Snake Hemirhagerrhis viperina Psammophylax
(Bocage, 1873) nototaenia
Henophidia | Lamprophiidae | Psammophinae
Angolan Sand Snake Psammophis angolensis Amphiophis
(Bocage, 1872) angolensis
Ansorge’s Sand Snake Psammophis ansorgii E
(Boulenger, 1905)
Jalla’s Sand Snake Psammophis jallae (Peracca,
1896)
(continued)

Leopard Sand Snake Psammophis leopardinus
(Bocage, 1887)
Namib Sand Snake Psammophis namibensis
(Broadley, 1975)
Karoo Sand Snake Psammophis notostictus
(Peters, 1867)
Mozambique Grass Snake Psammophis mossambicus Y Psammophis
(Peters, 1882) sibilans (Linnaeus,
1758)
Strip-bellied Sand SnakePsammophis subtaeniatus
(Peters, 1882)
Western Sand Snake Psammophis trigrammus
(Günther, 1865)
Fork-marked Sand Snake Psammophis trinasalis
(Werner, 1902)
Zambezi Sand Snake Psammophis zambiensis
(Hughes and Wade, 2000)
Striped Beaked Psammophylax acutus Rhageheris acutus
Skaapstekker (Günther, 1888)
Huila Skaapstekker Psammophylax ocellatus Psammophylax E
(Bocage, 1873) rhombeatus
Striped Skaapstekker Psammophylax tritaeniatus Rhagerhis
(Günther, 1868) tritaeniata
Henophidia | Lamprophiidae | Prosymnidae
Zambezi Shovel-snout Prosymna ambigua (Bocage, Prosymna ambigua
Snake 1873)
Angola Shovel-snout Prosymna angolensis Prosymna frontalis
Snake (Boulenger, 1915)
South-west Shovel-snout Prosymna frontalis (Peters,
Snake 1867)
Visser’s Shivel-snout Prosymna visseri (FitzSimons,
Snake 1959)
Henophidia | Lamprophiidae | Pseudaspidae
Mole Snake Pseudaspis cana (Linnaeus, Pseudaspis cana
1758)
Western-keeled Snake Pythonodipsas carinata
(Günther, 1868)
Henophidia | Lamprophiidae | Elapidae
Cowles’ Shield Cobra Aspidelaps lubricus cowlesi
(Bogert, 1940)
Jameson’s Mamba Dendroaspis jamesoni (Traill, ? Dendroaspis
1843) neglectus
Black Mamba Dendroaspis polylepis ? Dendroaspis
(Günther, 1864) angusticeps
Günther’s Garter Snake Elapsoidea guentherii (Bocage, Elapsoidea
1866) Guentherii
(continued)

Angolan Garter Snake Elapsoidea s. semiannulata
(Bocage, 1882)
Western Garter Snake Elapsoidea s. moebiusi
(Werner, 1897)
Anchiete’s Cobra Naja (Ureaus) anchietae Naja anchietae &
(Bocage, 1879) Naja haje
Banded Water Cobra Naja (Boulengerina) annulata
(Peters, 1876)
Central African Forest Naja (Boulengerina)
Cobra melanoleauca (Hallowell,
1857)
Savanna Forest Cobra Naja (Boulengerina) subfulva ?
(Laurent, 1956)
Mozambique Cobra Naja (Afronaja) mossambica
(Peters, 1854)
Western Banded Spitting Naja (Afronaja) nigricincta
Cobra (Bogert, 1940)
Black Spitting Cobra Naja (Afronaja) nigricollis Naja nigricollis
(Reinhardt, 1843)
Gold’s Tree Cobra Pseudohaje goldii (Boulenger,
1895)
Henophidia | Viperidae
Variable Bush Viper Atheris squamigera (Hallowell, Atheris squamigera
1854)
Puff Adder Bitis arietans (Merrem, 1820) Vipera arietans
Horned Adder Bitis caudalis (Smith, 1839) Vipera caudalis
Gaboon Adder Bitis gabonica Duméril, ? Vipera rhinoceros
(Bibron and Duméril, 1854)
Angolan Adder Bitis heraldica (Bocage, 1889) Vipera heraldica E
Rhinoceros Viper Bitis nasicornis (Shaw, 1802) ?
Peringuey’s Adder Bitis peringueyi (Boulenger, ?
1888)
Two-lined Night Adder Causus bilineatus (Boulenger,
1905)
Lichtenstein’s Night Causus lichtensteini (Jan,
Adder 1859)
West African Night Adder Causus maculatus (Hallowell,
1842)
Rasmussen’s Night Adder Causus rasmusseni (Broadley,
2014)
Angola Green Night Causus resimus (Peters, 1862) Causus resimus
Adder
Rhombic Night Adder Causus rhombeatus Y Causus rhombeatus
(Lichtenstein, 1823)
a
Don’t include the additional Boaedon species (Hallerman et al. in prep.)
References
Adalsteinsson SA, Branch WR, Trapé S, Vitt LJ, Hedges SB (2009) Molecular phylogeny, clas-
sification, and biogeography of snakes of the family leptotyphlopidae (Reptilia, Squamata).
Zootaxa 2244:1–50
Alexander AA, Gans C (1966) The pattern of dermal-vertebral correlation in snakes and amphis-
baenians. Zool Med 41(11):171–190
Angel F (1921) Description d’un ophidien nouveau de l’Angola appartenant au genre Psammophis.
Bull Soc Zool Fr Paris 46(8–10):116–118
Angel MF (1923) Reptiles. In: Rohan-Chabot (ed) Mission Rohan-Chabot, Angola et Rhodesia
(1912–1914), Histoire Naturelle, Fascicule 1 (Mammifères – Oiseaux –Reptiles – Poissons),
vol IV. Imprimerie Nationale, Paris, pp 157–169, 1 pl
Arruda M (2018) Confiscation by police of illegal soft-shell terrapin trade in Luanda. https://www.
facebook.com/katimbala.ingles/videos/pcb.1576072472480736/1576072072480776/?type=3
&theater
Baptista N, António T, Branch WR (2018) Amphibians and reptiles of the Tundavala region of
the Angolan Escarpment. In: Revermann R, Krewenka KM, Schmiedel U et al (eds) Climate
change and adaptive land management in southern Africa – assessments, changes, challenges,
and solutions, Biodiversity & ecology, vol 6. Klaus Hess Publishers, Göttingen, pp 397–403
Baptista N, Conradie W, Vaz Pinto P et al (2019) The Amphibians of Angola: early studies and the
Bauer AM, Branch WR (2003) The herpetofauna of the Richterveld National Park and the adjacent
northern Richtersveld, Northern Cape Province, Republic of South Africa. Herpetol Nat Hist
8(2):111–160
Bauer AM, Kuhn AL (2017) Historical climate change and the evolution of the Namib Day Geckos
(Squamata: Gekkonidae: Rhoptropus) Oral Presentation (abst). Afr Herp News 66:9
Bauer AM, Lamb T (2005) Phylogenetic relationships of southern African geckos in the
Pachydactylus group (Squamata: Gekkonidae). Afr J Herpetol 54(2):105–129
Billes A, Fretey J, Verhage B et al (2006) First evidence of leatherback movement from Africa to
South America. Mar Turt Newsl 111:13–14
Blanc CP, Fretey J (2002) Analyse Zoogegraphique du peuplement reptilien de L’Afrique Centrale
et de L’Angola. Biogeographica 78:49–75
Bocage JVB (1866) Lista dos reptis das possessões portuguesas d’ Africa occidental que existem
no Museu de Lisboa. J Sci Math Phys Nat Lisboa 1:37–56
Bocage JVB (1873) Reptiles nouveaux de l’interieur de Mossamedes. J Sci Math Phy Nat Lisboa
4:247–253
Bocage JVB (1895) Herpétologie d’Angola et du Congo. Lisbonne, Imprimerie Nationale, 203
pp, 19 pls
Bocage JVB (1896) Mammiferos, aves e reptis da Hanha, no sertào de Benguella. J Sci Math Phys
Nat Lisboa 14(2):105–114
Bogert CM (1940) Herpetological results of the Vernay Angola Expedition. I. Snakes, including an
arrangement of the African Colubridae. Bull Am Mus Nat Hist 77:1–107
Boulenger GA (1885) Catalogue of the lizards in the British Museum (Natural History). Volume
II, Iguanidae, Xenosauridae, Zonuridae, Anguidae, Anniellidae, Helodermatidae, Varanidae,
Xantusiidae, Teiidae, Amphisbaenidae. London: British Museum of (Natural History), London,
xiv + 492 pp, 54 figs., 24 pls
Boulenger GA (1893) Catalogue of the snakes in the British Museum (Natural History). Volume
I, containing the families Typhlopidae, Glauconiidae, Boidae, Ilysiidae, Uropeltidae,
Xenopeltidae and Colubridae aglyphae, part. British Museum (Natural History), London, xiv
+ 448 pp, 26 figs., 28 pls
II, containing the conclusion of the Colubridae aglyphae. British Museum (Natural History),
London, xii + 382 pp, 25 figs., 20 pls
III, containing the Colubridae (Opisthoglyphae and Proteroglyphae), Amblycephalidae, and
Viperidae. British Museum (Natural History), London, xiv + 727 pp, 37 figs., 25 pls
Boulenger GA (1905) A list of the batrachians and reptiles collected by Dr. W. J. Ansorge in
Angola, with descriptions of new species. Ann Mag Nat Hist Ser 7 16:105–115
Boulenger GA (1907a) Descriptions of three new lizards and a frog, discovered by Dr. W. J.
Ansorge in Angola. Ann Mag Nat Hist Seventh Ser 19:212–214
Boulenger GA (1907b) Descriptions of a new frog discovered by Dr. W. J. Ansorge in Mossamedes,
Angola. Ann Mag Nat Hist Seventh Ser 20:109
Boulenger GA (1915) A list of the snakes of the Belgian and Portuguese Congo, northern Rhodesia,
and Angola. Proc Zool Soc London 1915:193–223
Bowker JN, De Vos A, Ament JM et al (2017) Effectiveness of Africa’s tropical protected areas for
maintaining forest cover. Conserv Biol 31(3):559–569
Branch WR (1994) Herpetofauna of the Sperrgebeit region of southern Namibia. Herpetol Nat
Hist 2(1):1–11
Branch WR (2014) Reptiles of South Africa, Lesotho and Swaziland: conservation status, diver-
sity, endemism, hotspots and threats. In: Bates MF, Branch WR, Bauer AM, Burger M, Marais
J, Alexander GJ, de Villiers MS (eds) Atlas and Red Data Book of the Reptiles of South Africa,
Lesotho and Swaziland, Suricata 1. South African National Biodiversity Institute, Pretoria,
pp 22–50
Branch WR (2016) Preface, amphibian & reptile conservation special Angola-Africa issue.
Amphib Rep Conserv 10(2):2. i-iii
Branch WR (2018) Snakes of Angola: an annotated checklist. Amphibian & Reptile Conservation
12(2) [General Section]: 41–82 (e159)
Branch WR, Conradie W (2013) Naja annulata annulata (Bucholtz & Peters, 1876). African Herp
News 59:50–53
Branch WR, Conradie WC (2015) Vl Herpetofauna da regiáo da Lagoa Carumbo (Herpetofauna
of the Carumba Lagoon Area). In: Huntley B & Francisco P (eds) Relatório sobre a Expedição
Avaliação rápida da Biodiversidade de regiáo da Lagoa Carumbo, Lunda-Norte – Angola,
pp 194–209. Republica de Angola Ministerio do Ambiente, 219p
Branch WR, McCartney CJ (1992) A report on a small collection of reptiles from southern Angola.
J Herpetol Assoc Afr 41:1–3
Branch WR, Tolley KA (2017) Oral presentation (abst). New Lacertids from Angola. Afr Herp
News 66:11
Branch WR, Haacke W, Vaz Pinto P et al (2017) Loveridge’s Angolan geckos, Afroedura karroica
bogerti and Pachydactylus scutatus angolensis (Sauria, Gekkonidae): new distribution records,
comments on type localities and taxonomic status. Zoosyst Evol 93(1):157–166
Branch WR, Baptista N, Vaz Pinto P (2018) Angolan amphisbaenians: rediscovery of Monopeltis
luandae Gans 1976, with comments on the type locality of Monopeltis perplexus Gans 1976
(Sauria: Amphisbaenidae). Herpetology Notes 11:603–606
Branch WR, Baptista N, Keates C et al (2019) Rediscovery, taxonomic status and phylogenetic
relationships of two enigmatic Psammophine snakes (Serpentes: Psammophinae) from the
southwestern Angola plateau. Zootaxa (in press)
Broadley DG (1971) A revision of the African snake genera Amblyodipsas and Xenocalamus. Occ
Pap Natl Mus Rhod B4(33):629–697
Broadley DG (1975) A review of the Mabuya lacertiformis complex in southern Africa (Sauria:
Scincidae). Arnoldia (Rhod) 7(18):1–16
Broadley DG (1991) The Herpetofauna of Northern Mwinilunga Distr., Northw. Zambia. Arnoldia
Zimbabwe 9(37):519–538
Broadley DG (1996) A review of the tribe Atherini (Serpentes: Viperidae), with the descriptions of
two new genera. Afr J Herpetol 45:40–48
Broadley DG (1997a) A review of the Monopeltis capensis complex in southern Africa (Reptilia:
Amphisbaenidae). Afr J Herpetol 46(1):1–12
Broadley DG (1997b) Amphibaenia. Dalophia ellenbergeri (Angel, 1920). African Herp News
26:34–35
Broadley DG (2001) Geographical distribution. Monopeltis sphenorhynchus. Afr Herp News
32:23–24
Broadley DG, Baldwin AS (2006) Taxonomy, natural history, and zoogeography of the
Southern African Shield Cobras, Genus Aspidelaps (Serpentes: Elapidae). Herpetol Nat Hist
9(2):163–176
Broadley DG, Broadley S (1999) A review of the African worm snakes from south of latitude 12°S
(Serpentes: Leptotyphlopidae). Syntarsus 5:1–36
Broadley DG, Gans C (1969) A new species of Zygaspis (Amphisbaenia: Reptilia) from Zambia
and Angola. Arnoldia (Rhod) 4(25):1–4
Broadley DG, Hughes B (2000) A revision of the African genus Hemirhagerrhis Boettger 1893
(Serpentes: Colubridae). Syntarsus 6:1–17
Broadley DG, Schätti B (2000) A new species of Coluber from northern Namibia (Reptilia:
Serpentes). Modoqua 19(2):171–174
Broadley DG, Gans C, Visser J (1976) Studies on Amphisbaenians (6). The Genera Monopeltis and
Dalophia in Southern Africa. Bull Am Mus Nat Hist 157(5):311–486
Broadley DG, Tolley KA, Conradie W et al (2018) A phylogeny and revision of the African File
Snakes Gonionotophis Boulenger (Squamata: Lamprophiidae). Afr J Herpetol. https://doi.org/
10.1080/21564574.2018.1423578
Brooks C (2012) Biodiversity survey of the upper Angolan Catchment of the Cubango-Okavango
River Basin. USAid-Southern Africa, 151 p
Brooks C (2013) Trip report: aquatic biodiversity survey of the lower Cuito and Cuando river
systems in Angola. USAid-Southern Africa. 43 p
Carr T, Carr P (1991) Surveys of the sea turtles of Angola. Biol Conserv 58(1):19–29
CEPF (2003) Ecosystem profile. The succulent Karoo hotspot, Namibia and South Africa. Critical
ecosystem partnership fund. South African National Biodiversity Institute. https://www.sanbi.
org/documents/ecosystem-profile-succulent-karoo-hotspot
Ceríaco LMP, Bauer AM, Blackburn DC et al (2014a) The herpetofauna of the Capanda Dam
Region, Malanje, Angola. Herpetol Rev 45(4):667–674
Ceríaco LMP, Blackburn DC, Marques MP et al (2014b) Catalogue of the amphibian and reptile
type specimens of the Museu de História Natural da Universidade do Porto in Portugal, with
some comments on problematic taxa. Alytes 31(1):13–36
Ceríaco LMP, de Sá SAC, Bandeira S, Valério H et al (2016a) Herpetological survey of Iona
National Park and Namibe Regional Natural Park, with a Synoptic list of the Amphibians and
reptiles of Namibe Province, Southwestern Angola. Proc Calif Acad Sci 63(2):15–61
Ceríaco LMP, Marques MP, Bandeira SA (2016b) Anfíbios e Répteis do Parque Nacional da
Cagandala. Publ. Instituto Nacional da Biodiversidade e Áreas de Conservação & Museu
Nacional de História Natural e da Ciência, 96 p
Ceríaco LMP, de Sá S, Bauer AM (2018a) The genus Osteolaemus (Crocodylidae) in Angola and
a new southernmost record for the genus. Herpetol Notes 11:337–341
Ceríaco LMP, Marques MP, Bandeira S et al (2018b) Herpetological survey of Cangandala
National Park, with a synoptic list of the amphibians and reptiles of Malanje Province, Central
Angola. Herpetol Rev 49(3):408–431
Clark VR, Barker NP, Mucina L (2011) The great escarpment of southern Africa: a new frontier for
biodiversity exploration. Biodivers Conserv 20:2543–2561
Conradie W, Bourquin S (2013) Geographical Distributions: Acontias kgalagadi kgalagadi (Lamb,
Biswas and Bauer, 2010). Afr Hep News 60:29–30
Conradie W, Branch WR, Measey GJ et al (2012a) Revised phylogeny of Sand lizards (Pedioplanis)
and the description of two new species from south-western Angola. Afr J Herpetol 60(2):91–112
Conradie W, Branch WR, Measey JG et al (2012b) A new species of Hyperolius Rapp, 1842 (Anura:
Hyperoliidae) from the Serra da Chela mountains, southwestern Angola. Zootaxa 3269:1–17
Conradie W, Branch WR, Tolley KA (2013) Fifty shades of grey: giving colour to the poorly
known Angolan Ash reed frog (Hyperoliidae: Hyperolius cinereus), with the description of a
new species. Zootaxa 3635(3):201–223
Conradie W, Bills R, and Branch WR (2016). The herpetofauna of the Cubango, Cuito, and lower
Cuando river catchments of south-eastern Angola. Amphibian Reptile Conserv 10(2) [Special
Section]:6–36
Conradie WC, Bills R, Baptista N et al (2017) Oral presentation (abst). Across river basins:
Exploring the unknown southeastern Angola. African Herp News 66:14–15
Cotterill F, De Wit M (2011) Geoecodynamics and the Kalahari Epeirogeny: linking its genomic
record, tree of life and palimpsest into a unified narrative of landscape evolution. S Afr J Geol
114:489–514
Craven P (2009) Phytogeographic study of the Kaokoveld Centre of Endemism. Unpublished
Ph.D. thesis, University of Stellenbosch, 234 p
Crawford-Cabral J, Mesquitela LM (1989) Índice toponímico de colheitas zoológicas em Angola.
Instituto de Investigação Cientifica Tropical. Centro de Zoologia, Lisbon 206 pp
De Almeida MAP (2011) José Vicente Barbosa du Bocage. In: Biographies of scientists and
engineers, Centro Interuniversitário de História da Ciência e Tecnologia. http://ciuhct.org/pt/
bocage-jose-vicente-barbosa-du
Eaton MJ (2010) Dwarf crocodile Osteolaemus tetraspis. In: Manolis SC, Stevenson C (eds)
Crocodiles. Status survey and conservation action plan, 3rd edn. Crocodile Specialist Group,
Darwin, pp 127–132
Eaton MJ, Martin A, Thorbjarnarson J, Amato G (2009) Species-level diversification of African
dwarf crocodiles (genus Osteolaemus): a geographic and phylogenetic perspective. Mol
Phylogenet Evol 50(3):496–506
Elwen S, Braby RJ (2015) Report on a turtle and cetacean assessment survey to the Kunene River
mouth, northern Namibia – January 2014. Afr Sea Turtle Newsl 4:22–27
Engelbrecht HM, Branch WR, Greenbaum E et al (2018) Diversifying into the branches: spe-
cies boundaries in African green and bush snakes, Philothamnus (Serpentes: Colubridae). Mol
Phylo Evol. https://doi.org/10.1016/j.ympev.2018.10.023
Ernst R (2015) A rapid assessment of the herpetofauna of the Serra do Pingano ecosystem in Uíge
Province, northern Angola. Unpubl. Report to Instituto Nacional da Biodiversidade e Áreas de
Conservação, Ministério do Ambiente, República de Angola, 11 p
Ernst R, Nienguesso ABT, Lautenschläger T et al (2014) Relicts of a forested past: southernmost
distribution of the hairy frog genus Trichobatrachus Boulenger, 1900 (Anura: Arthroleptidae)
in the Serra do Pingano region of Angola with comments on its taxonomic status. Zootaxa
3779(2):297–300
Ernst R, Schmitz A, Wagner P et al (2015) A window to Central African forest history: distribution
of the Xenopus fraseri subgroup south of the Congo Basin, including a first country record of
Xenopus andrei from Angola. Salamandra 52(1):147–155
Face of Malawi (2013) Chinese ‘managed’ Turtle butchery discovered on Lake Malawi. http://www.
faceofmalawi.com/2013/11/chinese-managed-turtle-butchery-discovered-on-lake-malawi/
Ferreira JB (1897) Lista dos reptis e amphibios que fazem parte da última remessa de J. d’Anchieta.
J Sci Math Phys Nat Lisboa 5(2):240–246
Ferreira JB (1900) Sobre alguns exemplares pertencentes á fauna do norte de Angola. J Sci Math
Phys Nat Lisboa 21:48–54
Ferreira JB (1903) Reptis de Angola da região norte do Quanza da collecção Pereira do Nascimento
(1902). J Sci Math Phys Nat Lisboa Segunda Série 7(25):9–16
Ferreira JB (1904) Reptis e amphibios de Angola da região ao norte do Quanza (Collecção
Newton – 1903). J Sci Math Phys Nat Lisboa Segunda Série 7(26):111–117
Ferreira JB (1906) Algumas espécies novas ou pouco conhecidas de amphibios e reptis de Angola
(Collecção Newton – 1903). J Sci Math Phys Nat Lisboa Segunda Série 7(26):159–171
FitzSimons VFM (1953) A new genus of gerrhosaurid from southern Angola. Ann Transv Mus
22(2):215–217
FitzSimons VFM (1959) Some new reptiles from southern Africa and southern Angola. Ann
Transv Mus 23(4):405–409
Gamito-Marques D (2017) A space of one’s own: Barbosa du Bocage, the foundation of the
National Museum of Lisbon, and the construction of a career in zoology (1851–1907). J Hist
Biol. https://doi.org/10.1007/s10739-017-9487-6
Gans C (1976) Three new spade-snoted amphisbaenians from Angola (Amphisbaenia, reptilia).
Am Mus Novit 2590:1–11
Gans C (2005) Checklist and bibliography of the Amphisbaenia of the World. Bull Am Mus Nat
Hist 289(1):130
Gray JE (1865) A revision of the genera and species of amphisbaenians with the descriptions
of some new species now in the collection of the British Museum. Proc Zool Soc London
1865:442–455
Greenbaum E, Portillo F, Jackson K et al (2015) A phylogeny of central African Boaedon
(Serpentes: Lamprophiidae), with the description of a new cryptic species from the Albertine
Rift. Afr J Herpetol 64(1):18–38
Haacke WD (1976a) The burrowing geckos of southern Africa, 2 (Reptilia: Gekkonidae). Ann
Transv Mus 30(2):13–29
Haacke WD (1976b) The burrowing geckos of southern Africa, 3 (Reptilia: Gekkonidae). Ann
Transv Mus 30(3):29–44
Haacke WD (1981) The file snakes of the genus Mehelya in Southern Africa with special reference
to South West Africa/Namibia. Modoqua 12(4):217–224
Haacke WD (1997) Systematics and biogeography of the southern African scincine genus
Typhlacontias (Reptilia: Scincidae). Bonner Zool Beiträge 47(1–2):139–163
Haacke WD (2008) A new leaf-toed gecko (Reptilia: Gekkonidae) from south-western Angola. Afr
J Herpetol 57:85–92
Haacke WD (2013) Description of a new Tiger Snake (Colubridae, Telescopus) from south-western
Africa. Zootaxa 3737(3):280–288
Haagner GV, Branch WR, Haagner AJF (2000) Notes on a collection of reptiles from Zambia and
adjacent areas of the Democratic Republic of the Congo. Ann East Cape Mus 1:1–25
Heinicke MP, Daza JD, Greenbaum E et al (2014) Phylogeny, taxonomy and biogeography of a
circum-Indian Ocean clade of leaf-toed geckos (Reptilia: Gekkota), with a description of two
new genera. Syst Biodivers 12(1):23–42
Heinz HM (2011) Comparative phylogeography of two widespread geckos from the typically
narrow-ranging Pachydactylus group in Southern Africa. Unpublished MSc thesis, Villanova
University, Villanova, Pennsylvania, USA, vii + 107 pp
Hellmich W (1954–1955) Auf herpetologischer Forschungsfahrt in Angola (Portugeisisch
Westafrika). Die Aquarien und Terrarien Zeitschrift. In six parts: 1954 – I, 7(11):
302–304; II, 7(12): 324–326; 1955 – III, 8(1): 23–26; IV, 8(2): 51–53; V, 8(3):78–81; VI,
8(4):103–107
Hellmich W (1957a) Die reptilienausbeute der Hamburgischen Angola Expedition. Mitteilungen
Hellmich W (1957b) Herpetologische Ergebnisse einer Forschungsreise in Angola.
Hellmich W, Schmelcher D (1956) Eine neue Rasse von Gerrhosaurus nigrolineatus Hallowell
(Gerrhosauridae). Zool Anz 156(7/8):202–205
Herrmann H-W, Branch WR (2013) Fifty years of herpetological research in the Namib Desert and
Namibia with an updated and annotated species checklist. J Arid Environ 93:94–115
Huntley BJ (1974) Outlines of wildlife conservation in Angola. J S Afr Wildl Manage Assoc
4:157–166

ect. Report on Pilot project. Unpublished report to ministry of environment, Luanda, 97 pp,
27 figures
Huntley BJ, Francisco P (eds) (2015) Relatório sobre a Expedição Avaliação rápida da
Biodiversidade de regiáo da Lagoa Carumbo, Lunda-Norte – Angola/Biodiversity Rapid
Assessment of the Carumbo Lagoon Area, Lunda-Norte. Angola Ministerio do Ambiente,
Luanda 219 pp
Jordan K (1936) Dr Karl Jordan’s expedition to South-West Africa and Angola. Narrat Nov Zool
40:17–62, 2 maps, 5 pls
Jongsma GFM, Barej MF, Barratt CD, Burger M, Conradie W, Ernst R, Greenbaum E, Hirschfeld
M, Leaché AD, Penner J, Portik DM, Zassi-Boulou A-G, Rödel M-O, Blackburn DC (2018)
Diversity and biogeography of frogs in the genus Amnirana (Anura: Ranidae) across sub-Saha-
ran Africa. Mol Phylogenet Evol 120:274–285
Kelly CMR, Branch WR, Broadley DG et al (2011) Molecular systematics of the African snake
family Lamprophiidae Fitzinger, 1843 (Serpentes: Elapoidea), with particular focus on the
genera Lamprophis Fitzinger 1843 and Mehelya Csiki 1903. Mol Phylogenet Evol 58:415–426
Kindler C, Branch WR, Hofmeyr MF et al (2012) Molecular phylogeny of African hinge-back
tortoises (Kinixys Bell, 1827): implications for phylogeography and taxonomy (Testudines:
Testudinidae). J Zool Syst Evol Res 50(3):192–201
Laurent RF (1950) Reptiles et Batraciens de la region de Dundo (Angola du Nord-Est). Publicações
culturais da Companhia de Diamantes de Angola 10:7–17
Laurent RF (1954) Reptiles et Batraciens de la région de Dundo (Angola) (Deuxième Note).
Publicações culturais da Companhia de Diamantes de Angola 23:35–84
Laurent RF (1964) Reptiles et Amphibiens de l’Angola (Troisième contribution). Publicações cul-
turais da Companhia de Diamantes de Angola 67:11–165
Lewin A, Feldman A, Bauer AM et al (2016) Patterns of species richness, endemism and environ-
mental gradients of African reptiles. J Biogeogr 43:2380–2390
Lindsey PA, Nyirenda VR, Barnes JI et al (2014) Underperformance of African protected area
networks and the case for new conservation models: insights from Zambia. PLoS One
9(5):e94109
Loveridge A (1941) Revision of the African lizards of the family Amphisbaenidae. Bull Mus Comp
Zool 87(5):353–451
Loveridge A (1944) New geckos of the genera Afroedura, new genus, and Pachydactylus from
Angola. Am Mus Novit 1254:1–4
Machado AdB (1952) Generalidades acerca da Lunda e da sua exploraçao biolôgica. Publ. Cult.
Comp. Diam. Angola, 12
Madruga CM (2013) José Vicente Barbosa du Bocage (1823–1907): a construção de uma persona
científica. Unpublished M.Sc. thesis, Universidade de Lisboa
Manaças S (1963) Sáurios de Angola. Memórias da Junta de Investigações do Ultramar, Lisboa,
43, segunda série. Estudos Zool:223–240
Manaças S (1973) Alguns ofídeos de Angola. Memórias da Junta de Investigações do Ultramar,
Lisboa, 58, segunda série. Estudos de Zoologia:187–200
Manaças S (1981) Ofídeos venenosos da guiné, S. Tomé, Angola e Moçambique. Garcia Orta Sér
de Zool 10(1/2):13–46
Marques MP, Ceríaco LMP, Bauer AM, et al. (2014) Geographic distribution of Amphibians and
reptiles of Angola: towards an Atlas of Angolan herpetofauna. Poster. 12th Herpetological
association of Africa conference, Gobabeb, Namibia, 20–22 November, 2014
Marques MP, Ceríaco LMP, Blackburn DC, Bauer AM (2018) Diveristy and distribution of the
amphibians and terrestrail reptiles of Angola atlas of historical and bibliographic records
(1840–2017). Proceedings of the California academy of sciences, Series 4, Volume 65,
Supplement II: 1–501
Marx H (1956) A new lacertid lizard from Angola. Fieldiana: Zoology 39:5–9
Mateus O, Jacobs L, Polcyn M (2009) The oldest African eucryptodiran turtle from the Cretaceous
of Angola. Acta Palaeontol Pol 54(4):581–588
McLachlan GR, Spence JM (1967) A new species of Pachydactylus (Pachydactylus oreophilus sp.
nov.) from Sesfontein, South West Africa. Cimbebasia (21):3–8
Measey J, Tolley KA (2013) A molecular phylogeny for sub-Saharan amphisbaenians. Afr
J Herpetol 62(2):100–108
Medina MF, Bauer AM, Branch WR et al (2016) Molecular phylogeny of Panaspis and
Afroablepharus skinks (Squamata: Scincidae) in the savannas of sub-Saharan Africa. Mol
Mertens R (1938) Amphibien und Reptilien aus Angola, gesammelt von W. Schack.
Senckenbergiana 20:425–443
Mertens R (1958) Bitis heraldica, eine oft verkannte Otter aus Angola. Senckenbergiana Biologica
Frankfurt-am-Main 39(3–4):145–148, 4 figs
Monard A (1931) Mission scientifique Suisse dans l’Angola. Résultats scientifiques. Reptiles. Bull
Soc Neuchâtel Sci Nat ser 2 40(55):89–111
Monard A (1934) Ornithologie de l’Angola. Arquivos do Museu Bocage 5:1–110
Arquivos do Museu Bocage 6:1–314 44 fig
Monard A (1937) Contribution á l’herpétologie d’Angola. Arquivos do Museu Bocage 8:19–154
Monard A (1938) Contribution à la batrachologie d’Angola. Bull Soc Neuch Se Nat 62:5–59, 19
fig
Morais M (2008) Tartarugas Marinhas na Costa de Cabinda. Plano de conservação e gestão para a
implementação do projecto de prospecção sísmica “on shore”. Holisticos/Chevron. 67 p
temporada 2015/2016. Universidade Agostinho Neto/Faculdade de Ciências, Luanda
temporada 2016/2017. Universidade Agostinho Neto / Faculdade de Ciências, Luanda
Morais M, Torres MOF, Martins MJ (2005) Análise da Biodiversidade Marinha e Costeira e
Identificação das Pressões de Origem Humana sobre os Ecossistemas Marinhos e Costeiros.
Ministerio do Urbanismo e Ambiente, Luanda, 140 pp
Nagy ZT, Kusamba C, Collet M et al (2013) Notes on the herpetofauna of western Bas-Congo,
democratic Republic of the Congo. Herpetology Notes 6:413–419
NGOWP (2018) National geographic Okavango wilderness project. Initial findings from explora-
tion of the upper catchments of the Cuito, Cuanavale, and Cuando Rivers, May 2015–December
2016
NPAES (2010) National protected area expansion strategy for South Africa 2008. Priorities for
expanding the protected area network for ecological sustainability and climate change adapta-
tion. Government of South Africa, Pretoria
OKACOM (2011) National action plan for the sustainable management of the Cubango/Okavango
River Basin, Angola – Draft 3. Available http://www.okacom.org/site-documents. Accessed 24
June 2016
Oliveira PRS d (2017) Sepentes em Angola. Uma visão toxinolŏgca e clínica dos envenenamentos.
Glaciar, Lisbon, p 159
Oliveira PRS d, Rocha MT, Castro AG et al (2016) New records of Gaboon viper (Bitis gabonica)
in Angola. Herpetol Bull 136:42–43
Parker HW (1936) Dr. Karl Jordan’s expedition to South West Africa and Angola: herpetological
collection. Nov Zool 40:115–146
Peters WCH (1869) Eine Mittheilung über neue Gattungen und Arten von Eidechsen. Monatsber
Königl Preuss Akad Wissensch Berlin 1869:57–66
Peters WCH (1876) Über die von Hrn. Professor Dr. R. Buchholz in Westafrika gesammelten
Amphibien. Auszug aus dem Monatsberich der Königl. Akademie der Wissenschafen zu
Berlin:117–124
Peters WCH (1877) Übersicht der Amphibien aus Chinchoxo (Westafrika), welche von
der Africanischen Gesellschaft dem Berliner zoologischen Museum übergeben sind.
Monatsberichte der königlich Akademie der Wissenschaften zu Berlin 10:611–620
Peters WCH (1881) Zwei neue von Herrn Major von Mechow während seiner letzten Expedition
nach West-Afrika entdeckte Schlangen und eine Übersicht der von ihm mitgebrachten herpe-
tologischen Sammlung. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin
9:147–150
Pietersen DW, Pietersen EW, Conradie W (2017) Preliminary herpetological survey of Ngonye
falls and surrounding regions in southwestern Zambia. Amphibian Reptile Conserv 11(1)
[Special Section]:24–43
Pincheira-Donoso D, Bauer AM, Meiri S, Uetz P (2013) Global taxonomic diversity of living
reptiles. PLoS One 8(3):e59741
Portillo F, Branch WR, Conradie W et al (2018) Phylogeny and biogeography of the African bur-
rowing snake subfamily Aparallactinae (Squamata: Lamprophiidae). Mol Phylogenet Evol
127:288–303
Poynton JC, Haacke WD (1993) On a collection of amphibians from Angola including a new spe-
cies of Bufo Laurenti. Ann Transv Mus 36(2):9–16
Roll U, Feldman A, Novosolov M et al (2017) The global distribution of tetrapods reveals a need
for targeted reptile conservation. Nat Ecol Conserv 1:1677–1682
Schmidt KP (1933) The reptiles of the Pulitzer-Angola expedition. Ann Carnegie Mus 22:1–15
Schmidt KP (1936) The amphibians of the Pulitzer-Angola expedition. Ann Carnegie Mus
25:127–133
Shirley MH, Vliet KA, Carr AN et al (2014) Rigorous approaches to species delimitation have
significant implications for African crocodilian systematics and conservation. Proc R Soc B
281:20132483
Shirley MH, Carr AN, Nestler JH et al (2018) Systematic revision of the living African slender-
snouted crocodiles (Mecistops Gray, 1844). Zootaxa 4504:151–193
Stanley EL, Ceríaco LMP, Bandeira S et al (2016) A review of Cordylus machadoi (Squamata:
Cordylidae) in southwestern Angola, with the description of a new species from the Pro-Namib
desert. Zootaxa 4061(3):201–226
Tilbury C (2018) Chameleons of Africa: an atlas, including the Chameleons of Europe, the Middle
East and Asia. Edition Chimaira, Frankfurt M, 831 pp
Tolley KA, Alexander GJ, Branch WR et al (2016) Conservation status and threats for African
reptiles. Biol Conserv 204:63–67
Trape J-F, Mediannikov O (2016) Cinq serpents nouveaux du genre Boaedon Duméril, Bibron
& Duméril, 1854 (Serpentes : Lamprophiidae) en Afrique centrale. Bull Soc Herp France
159:61–111
TRIPLOV (2018) Publications José Vincente Barbosa du Bocage. http://www.triplov.com/biblos/
bocage.htm
TTWG Turtle Taxonomy Working Group (2011) Turtle conservation Coalition. Turtles in trou-
ble: the world’s 25+ most endangered tortoises and freshwater turtles. Chelonian Research
Foundation, Conservation International, Wildlife Conservation Society, and San Diego Zoo
Global, 54 pp
TTWG Turtle Taxonomy Working Group (2017) Turtles of the world: annotated checklist and
atlas of taxonomy, synonymy, distribution, and conservation status (8th Ed.). Chelonian Res
Monogr 7:1–292
Tys van den Audenaerde DFE (1967) Les serpents des environs de Dundo (Angola) (Note complé-
mentaire). Publicações culturais da Companhia de Diamantes de Angola 76:31–37
Uetz P, Freed P, Hošek J (eds) (2018) The reptile database. Available http://www.reptile-database.
org. Accessed 11 Jan 2018
Vaz Pinto P, Branch WR (2015) Geographical distribution: Dendroaspis jamesoni (Thraill, 1843),
Jameson’s Mamba. African Herp News 62:45–47
Uetz P, Stylianou A (2018) The original descriptions of reptiles and their subspecies. Zootaxa
4375(2):257
Wagner P, Böhme W, Pauwels OSG, Schmitz A (2009) A review of the African red-flanked skinks
of the Lygosoma fernandi (Burton, 1836) species group (Squamata: Scincidae) and the role of
climate change in their speciation. Zootaxa 2050:1–30
Wagner P, Bauer AM, Wilms TM et al (2012) Miscellanea accrodontia: notes on nomenclature,
taxonomy and distribution. Russ J Herpetol 19:177–189
Wagner P, Greenbaum E, Bauer AM et al (2018) Lifting the blue-headed veil – integrative tax-
onomy of the Acanthocercus atricollis species complex (Squamata: Agamidae). J Nat Hist.
https://doi.org/10.1080/00222933.2018.1435833
Weinell JL, Bauer AM (2018) Systematics and phylogeography of the widely distributed African
skink Trachylepis varia species complex. Mol Phylogenet Evol 120:103–117
Weir CR, Ron T, Morais M et al (2007) Nesting and at-sea distribution of marine turtles in
Angola, West Africa, 2000–2006: occurrence, threats and conservation implications. Oryx
41(2):224–231
Wüster W, Chirio L, Trape J-F et al (2018) Integration of nuclear and mitochondrial gene sequences
and morphology reveals unexpected diversity in the forest cobra (Naja melanoleuca) species
complex in Central and West Africa (Serpentes: Elapidae). Zootaxa 4455:68–98
Chapter 14
The Avifauna of Angola: Richness,
Endemism and Rarity
W. Richard J. Dean, Martim Melo, and Michael S. L. Mills
Abstract Angola has a rich history of ornithological exploration going back to the
early 1800s. From the early-1970s to 2002, however, the civil war prevented access
to many areas, and very little work on birds was done. From about the early 2000s
information on birds in Angola has been gathered at an increasing rate, with new
species being added to the list and a steady rise in publications on biogeography and
biology of birds. With about 940 species, Angola has an impressive array of bird
species, including c. 29 endemic species, and several species that are rare and poorly
known. For the future, there are many areas of avian biology to attract studies, not
only to gather more data on the rare and endemic species, but also local surveys of
bird communities, the identification of major threats to the avifauna from landuse
changes (concomitantly with suggestions for remedial action) and more.
Understanding the role of birds in ecosystem processes, long term studies on the
biology and breeding of individual species, and inferring the evolutionary history of
the endemic species and of those species that occur in small isolated populations in
Angola are all areas for future research. The future of ornithological research and
conservation in Angola is dependent on it being carried out by Angolans – outreach,
W. R. J. Dean (*)
DST-NRF Centre of Excellence at the FitzPatrick Institute, University of Cape Town,
Rondebosch, South Africa
e-mail: lycium@telkomsa.net
M. Melo
DST-NRF Centre of Excellence at the FitzPatrick Institute, University of Cape Town,
Rondebosch, South Africa
e-mail: melo.martim@gmail.com
M. S. L. Mills
A. P. Leventis Ornithological Research Institute, University of Jos, Jos, Plateau State, Nigeria
e-mail: birdsangola@gmail.com

https://doi.org/10.1007/978-3-030-03083-4_14
336 W. R. J. Dean et al.
capacity building, and advanced training must all come together in order to find and
train the motivated ornithologists that such a biodiversity-rich country deserves.
Keywords Afromontane forests · Angolan escarpment · Conservation · Endemic

bird area · Ornithology · Rare birds
Early Ornithological History
The richness, endemism and rarity of the avifauna in Angola has attracted many
ornithologists, with the early studies during the late 1800s and first few years of the
1900s being almost entirely simple collections of birds. A chronology of bird col-
lections is given in Table 14.1. Publications and results of many of these collections
have been well covered by several authors, including Traylor (1963), Pinto (1983)
and Dean (2000). From the 1960s until the early 1970s extensive collections were
made at a number of localities in Angola by the Instituto de Investigação Científica
de Angola (IICA) (Fig. 14.1). Details of some of these collections, and records of
special interest, were published in a series of papers by Pinto (see references) pro-
viding much needed data on the biogeography and habitats of birds. The bird speci-
men collection assembled by the IICA, and now held by the Instituto Superior de
Ciências da Educação (ISCED) in Lubango was catalogued by Mills et al. (2010).
This was revised by Fernanda Lages and colleagues in 2016, aided by the discovery
of the field notebooks associated with the collections. This database, of what is
probably the third largest bird collection in Africa, will be available soon through
the Global Biodiversity Information Facility (GBIF) portal.
Although a war for independence in Angola had been going on since 1961,
armed conflict escalated with the start of a civil war in 1975, immediately after
Angola became independent from Portugal. The war went on to last almost three
decades, pre-empting any significant field-based biological research. Many of the
reports on the avifauna of Angola that were published during the 1960s, 1970s and
1980s were “desktop” studies of museum specimens, all using data that had been
collected before 1974. Despite issues with security and hazards imposed by the
localised patches of unexploded ordinances and the extensive use of landmines,
some avian studies were done in this period. Two East German biologists, Dr. Rainer
Günther and Dr. Alfred Feiler, based at the Museum für Naturkunde der Humboldt-
Universität zu Berlin and the Staatliches Museum für Tierkunde, Dresden, respec-
tively, were commissioned to survey biodiversity, including birds, in Angola
(Günther and Feiler 1986a, b). The ICBP (International Council for Bird
Preservation, now BirdLife International) attempted a project to gather data on the
status of (inter alia) the threatened endemic bird species on the Angolan “Scarp”.
The subsequent report adds little to what was known about the avifauna of this area,
but the report usefully outlines the major threats to the biodiversity in the southern-
most patches of Guinea-Congolian forest (Hawkins 1993).
It was only after the war ended definitely in 2002 that ornithologists returned to
the country with most expeditions targeting the regions that had been classified as
Important Bird Areas (IBAs, Dean 2001) and, in particular, the core habitats of the
14 The Avifauna of Angola: Richness, Endemism and Rarity 337
Table 14.1 A chronology of the collecting expeditions made in Angola, adapted from Traylor
(1963) and Dean (2000), and reproduced, in part, by courtesy of the British Ornithologists’ Union
1850– J. Anchieta collected birds mainly in central Angola. The specimens, many of which
1892 were Types, were described and reported on by Bocage in a number of separate
papers, summarised up to 1881 (Bocage 1877, 1881).
1858– J. J. Monteiro lived in Angola and collected birds. His collections, together with
1868 those made by Charles Hamilton (a visitor), were reported on by Hartlaub and
Monteiro (1860), Hartlaub (1865), Sharpe and Monteiro (1869) and Sharpe (1871).
1876– A. Lucan and L. A. Petit collected birds in Cabinda. Some of their specimens were
1877 deposited in the Natural History Museum, Tring. The collection was reported on by
Sharpe and Bouvier (1876a, b, 1877, 1878).
1880 and A. W. Eriksson collected in Cunene, between the Cunene and Cubango rivers. His
1887 specimens are in the Älvsborgs Länsmuseum, Vänersborg, Sweden (for details see
Rudebeck 1955 and Lundevall and Ängermark 1989), and in the Zoological
Museum, Uppsala, Sweden.
1884– P. J. van der Kellen collected for the Nationaal Natuurhistorisch Museum, Leiden,
1888 in Namibe and Huíla (Büttikofer 1888, 1889a, b).
Early Francisco Newton, a Portuguese naturalist, collected in southern Cuanza-Norte and
1900s along the coast (Seabra 1905–1907). Some of the material he collected is in the Museu
de História Natural – Zoologia, Oporto (Seabra 1905a, b, 1906a, b, c, d, 1907).
1901 C. H. Pemberton collected along the Cunene River and in the area between the
Cuanza River and Bailundo for the Rothschild’s Museum at Tring, UK.
1903– W. J. Ansorge collected extensively throughout western Angola for the Rothschild’s
1906 Museum at Tring, UK.
1908– W. J. Ansorge collected in Cuanza-Norte for the British Museum in London.
1909
1910– W. Lowe spent a few days collecting in December 1910 and March 1911 in the
1911 Luanda area (Bannerman 1912).
1912– Some birds were collected in Cuando Cubango, Cunene and Huíla by the Mission
1913 Rohan-Chabot (Ménégaux and Berlioz 1923).
1920s and R. Braun lived and studied birds mainly in Cuanza-Norte, northern Malanje and on
1930s the escarpment of Cuanza-Sul (Braun 1930, 1934; Sick 1934; Stresemann 1934,
1937).
1925 R. Boulton collected in Namibe, Huíla and Benguela for the American Museum of
Natural History.
1926– H. Lynes and B. B. Osmaston collected cisticolas on the Huambo highlands and
1927 along the Benguela coast.
1927 H. & C. Chapman collected on the central plateau for the American Museum of
Natural History.
1928 P. Koester collected in the highlands of Huambo and southern Cuanza-Sul and sent
the skins to O. Neumann. Some of these skins are now in the Museum of
Comparative Zoology, Harvard University.
1928– A. Monard (1932, 1934) collected in eastern and southern Huíla and in Lunda-
1929, Norte. His specimens are in the Musée D’Histoire Naturelle, La-Chaux-de-Fonds,
1932– Switzerland.
1933
1929– The gray African expedition of the Academy of Natural Sciences, Philadelphia
1930 made two collections in Bié and southern Malanje (Bowen 1931, 1932).
(continued)
1930 L. Fenaroli collected in the northwest and on the north-central plateau (Moltoni
1932).
1930– H. Lynes and J. Vincent collected cisticolas and other species on the plateau and
1931 from Dundo, Lunda-Norte, to Vila Luso (now Luena) in Moxico (Lynes and Sclater
1933, 1934).
1931 R. Boulton collected in central and southern Angola for the Carnegie Museum of
Natural History (Boulton 1931).
1931– Jean Bodaly made large collections in northern Bié and sent them to the Carnegie
1934 Museum and Chicago Natural History Museum (now Field Museum of Natural
History).
1931– H. K. Prior collected at Dondi in Huambo and sent skins to the Field Museum of
1934 Natural History.
1932– The Phipps-Bradley Expedition made a collection on the plateau for the American
1933 Museum of Natural History.
1933– H. Lynes and J. Vincent collected birds (mainly cisticolas) in Benguela, Huambo,
1934 southern Lunda-Sul and northern Lunda-Norte (Lynes 1938).
1944– C. M. N. White (1950) collected in areas in the extreme east of Moxico.
1949
1952 H. A. Beatty collected in the northwest and sent skins to the Field Museum of
Natural History, Chicago.
1954 W. Serle visited coastal areas for only 1 week but published interesting data (Serle
1955).
1954– G. Heinrich (1958a, b, c) collected extensively in the western half of Angola and
1955 sent some skins to the Field Museum of Natural History, Chicago, and the
Zoologisches Institut und Zoologisches Museum, Hamburg (Meise 1958).
1956 G. Rudebeck collected for the Visser-Transvaal Museum Expedition in
southwestern Angola, but only a few details have been published (Rudebeck 1958).
1957 B. P. Hall led an expedition to central and western Angola which resulted in two
major publications on zoogeography and taxonomy (Hall 1960a, b).
1957 R. Boulton collected in northwestern and northeastern Angola, and sent skins to the
Field Museum of Natural History, Chicago.
1957– G. Heinrich collected in Cuanza-Norte, Malanje and Lunda-Norte for the Peabody
1958 Museum of Natural History, Yale University, and the Smithsonian Institution.
1958– Staff at the Instituto de Investigação Científica de Angola (IICA) collected in
1973 Moxico, Bengo, Luanda, Malange, Cuanza-Sul, Bié, Benguela, Huíla, Namibe and
Cuando-Cubango for the IICA collection at Lubango.
1972 W.R.J. Dean collected in Huíla, Malange and Cabinda for the Peabody Museum,
New Haven (Dean 1974).
1972 M.E. Ferreira collected in Huíla for the Zoologishes Forschungsinstitut und
Museum Alexander Koenig, Bonn.
1973 W.R.J. Dean collected in Huíla, Cuanza-Norte, Cuanza-Sul and Malange for the
Peabody Museum, New Haven (Dean 1974).
1982– R. Günther and A. Feiler collected in Luanda, Bengo, Uige, Cuanza-Norte and
1983 Lunda-Norte for the Staatliches Museum für Tierkunde, Dresden and the Museum
für Naturkunde der Humboldt-Universität zu Berlin (Günther and Feiler 1986a, b)
Fig. 14.1 Localities where bird specimens were collected by the Instituto de Investigação
Científica de Angola (IICA) with duplicates held by the Instituto de Investigação Científica
Tropical (IICT), shown as the number of specimens collected in each 30-minute square. (Figure
extracted from Monteiro et al. (2014) and reproduced by permission of the authors)
endemic birds: the forests of the Western Escarpment and the highlands. Over the
last 15 years the knowledge of avian diversity, distribution and biogeography for the
country has steadily increased, with several species new to the Angola list being
recorded from areas that had never been previously explored.
ecent History and the Exponential Increase in Information

R
on Birds
One of the first “post war” studies was the publication of field notes on the Grey-
striped Francolin Francolinus griseostriatus that provided some information on the
biology of this endemic and rare species (Vaz Pinto 2002). Studies of individual
species and surveys of areas of particular interest gained momentum during the
early 2000s, with notes on the Gabela Akalat Sheppardia gabela (Mills et al. 2004),
and more general reports on the conservation status and vocalisations of endemic
and threatened bird species from the forests of the western escarpment (‘Scarp for-
ests’) of Angola (Ryan et al. 2004; Mills 2010) and a survey of the birds in Cumbira
Forest, Gabela (Sekercioğlu and Riley 2005). Cumbira Forest, a representative of

Central Scarp forests, is rich in endemic bird species, and has since been the focus
of one of the most detailed studies of birds in Angola (Cáceres et al. 2015, 2016,
2017). Most recent publications on birds in Angola, however, deal with extensions
of distributional range, vocalisations and lists of birds from specific areas, all pro-
viding data towards an atlas of Angolan birds (Table 14.2). Reports on the birds of
particular areas, such as Cangandala National Park (Mills et al. 2008) and the Soyo
area (Dean and le Maitre 2008; Stavrou and Mills 2013) are valuable in that there is
a large gap in time between when the places were last surveyed for birds and now.
The avifauna of many areas is known only from collections of specimens during the
1950s and there have been significant and rapid changes in landuse in many areas
after the war, such as with the conversion of old-growth miombo woodland for
charcoal making, or the replacement of secondary forest and shade-forest coffee
plantations by slash-and-burn agriculture in the Scarp (Leite et al. 2018), the most
important area of bird endemism (Cáceres et al. 2017). Reports on the avifauna of
Mount Moco (Mills et al. 2011a, b), the Namba Mountains (Mills et al. 2013a, b),
and Lagoa Carumbo (Mills and Dean 2013) have highlighted the bird species rich-
ness of these areas and, inter alia, noted threats to the local ecosystems and their
Table 14.2 New species on the Angolan bird list recorded since 1975
Northern Royal Albatross Diomedea (epomorpha) sanfordi Lambert (2001)
Spectacled Petrel Procellaria conspicillata Lambert (2001)
Red-billed Tropicbird Phaethon aethereus Lambert (2001)
White-tailed Tropicbird Phaethon lepturus Lambert (2001)
Cape Vulture Gyps coprotheres Bamford et al. (2007)a
Lesser Spotted Eagle Aquila pomarina Meyburg et al. (2001)
Booted Eagle Hieraaetus pennatus Sinclair (1981)
Red-necked Falcon Falco chicquera Mills et al. (2016)
Pacific Golden Plover Pluvialis fulva Mills (2015)
Red Phalarope Phalaropus fulicarius Lambert (2001)
European Oystercatcher Haematopus ostralegus Simmons et al. 2009
Black-headed Gull Chroicocephalus ridibundus Lambert (2001)
Little Tern Sternula albifrons Lambert (2001)
Greater Crested Tern Thalasseus bergii Dean et al. (2002)
Lemon Dove Columba larvata Mills and Dowd (2007)
Yellow-throated Cuckoo Chrysococcyx flavigularis Mills et al. (2013a, b)
Pink-billed Lark Spizocorys conirostris Mills (2006)
Red-tailed Leaflove Phyllastrephus scandens Mills et al. (2013a, b)
Forest Swallow Petrochelidon fuliginosa Mills and Tebb (2015)
South African Cliff Swallow Petrochelidon spilodera Mills et al. (2013a, b)
Singing Cisticola Cisticola cantans Dean et al. (2003)
White-collared Oliveback Nesocharis ansorgei Mills and Vaz Pinto (2015)
The list does not include unconfirmed records
a
Record based on satellite tracking
importance for conservation. Notes on rare and little known species, such as
Brazza’s Martin Phedina brazzae (Mills and Cohen 2007), Black-tailed Cisticola
Cisticola melanurus (Mills et al. 2011a, b), Bocage’s Sunbird Nectarinia bocagii
(Mills 2013), and the Red-necked Falcon Falco chicquera (Mills et al. 2016), have
provided some insights into the biology of these species. Breeding data for all spe-
cies in Angola are few, but MSLM and co-workers (see references) have recently
published several notes on the biology and first descriptions of nests and eggs and
some useful notes on brood parasitism and nests of a number of species.
The creation of the Internet group Angola Birders by MSLM in 2012 has greatly
facilitated and promoted the acquisition and sharing of data on bird distributions.
This has led to a large number of records of the occurrence of species, some of
which have cleared up distributional anomalies. For example, records of Black
Bishop Euplectes gierowii by Pedro Vaz Pinto (2 June 2015), together with earlier
records by MSLM, provide some evidence that the species is not as rare as previ-
ously thought (see Dean 2000). The Angola Birders Group has proved very useful,
not only for new distribution records, but also for new breeding records. More
importantly, it has generated interest in birds in many people, including diplomats
and executives of companies now trading in Angola, who now spend their spare
time “off the beaten track”. Their contributions towards a national database (at pres-
ent held by MSLM) of bird records are valuable. The facebook group “Angola
Ambiente” has completely overtaken Angola Birders and is currently providing a lot
of useful information.
One of the useful products from the national database of bird records is the bilin-
gual annotated checklist published by Mills and Melo (2013), based on the cata-
logue of the specimen collection at Lubango (Mills et al. 2010) and sight records.
This updates earlier lists, and includes some notes on unconfirmed and doubtful
records of species unlikely to occur in Angola. These records vary from (probably)
confusion with known species or sightings of species well outside their distribution
range.
The Richness of the Angolan Avifauna
About 940 species of birds have been recorded in Angola (Mills and Melo 2013).
This increases the number of species recorded for the country; a total of 915 species
were listed by Dean (2000), and 12 additional species were added by Mills and
Dean (2007), of which seven marine species, mostly pelagic, were recorded by
Lambert (2001). Some specimens thought to be of Red-faced Cisticola proved to be
those of an isolated population of Singing Cisticola Cisticola cantans (Dean et al.
2003), and the occurrence of the Lesser Spotted Eagle Aquila pomarina (Meyburg
et al. 2001) and the Swift Tern Sterna bergii (Dean et al. 2002) have been verified.
One additional species, Slaty Egret Egretta vinaceigula, has been added as a result
of cataloguing the bird collection at Lubango (Mills et al. 2010). Observations made
in poorly known areas of Angola by MSLM and co-workers have added additional
species to the list, including the Yellow-throated Cuckoo Chrysococcyx flavigularis,

[Red-tailed] Leaf-love Phyllastrephus scandens and South African Cliff Swallow
Petrochelidon spilodera (Mills et al. 2013a, b). Developments in assessing species
boundaries using molecular methods, have added another 16 species (Mills and
Dean 2007). One example of this is the Common Fiscal Lanius collaris that has now
been split into two species, the Northern Fiscal L. humeralis and Southern Fiscal L.
collaris (Fuchs et al. 2011). There have also been some corrections to the list, such
as specimens of the White-bellied Sunbird Cinnyrus talatala from Mount Moco and
Mount Soque listed by Dean (2000) that were found to be miscatalogued Oustalet’s
Sunbirds Cinnyrus oustaleti (Mills and Dean 2007). There are also a number of
avian taxa that require further investigation, either in the field or using molecular
analyses to establish species boundaries (Mills and Dean 2007).
The list for Angola does not include 88 species for which there are sight records
and that, in many cases, and for various reasons, are unlikely to occur in Angola
(Mills and Melo 2013). Some species are simply misidentifications of similar,
related species. Others, however, particularly migrant waders (shorebirds) from the
Northern Hemisphere, may have been correctly identified, but require more records,
photographs or specimens for verification. Unconfirmed species are dealt with by
Mills and Dean (2007) and more fully by Mills and Melo (2013) and will not be
listed here.
Endemism in the Angolan Avifauna
There are about 29 species of birds endemic to Angola, the number depending on
the taxonomic authority that is followed (Table 14.3). Most occur in the forests of
the Western Escarpment and in the last remnants of Afromontane forest of the high-
lands, the two core habitats of the Western Angola Endemic Bird Area (BirdLife
International 2017). Two species, Red-backed Mousebird Colius castanotus and
Bubbling Cisticola Cisticola bulliens, are widely distributed in Angola, including
the Western Escarpment and associated coastal plains and in a range of woodlands
and forest patches modified for the cultivation of coffee (Dean 2000).
The semi-evergreen humid forests of the Angolan Escarpment (‘the Scarp
Forests’) are impoverished outliers of the Congolian rainforest (Huntley and Matos
1994). They have been the major speciation hotspot for birds in Angola by: (i) creat-
ing a barrier between arid-adapted species of the coastal plains and of the miombo
woodlands of the plateau, (ii) creating a steep ecological gradient, and (iii) function-
ing as a refuge for moist forest specialists that were isolated here during the dry
periods of the glacial cycles (Hall 1960a) – 75% of the endemic bird species are
associated with this region.
The Afromontane forests of west-central Angola make the most isolated repre-
sentatives of all the Afromontane centres of endemism, separated by >2000 km
from other similar habitats. This isolation has allowed the development of plant and
animal communities that are quite distinct from those of other montane centres. The
Table 14.3 Provisional list of bird species endemic or near endemic to Angola, with their IUCN
Red List Category, and main area of occurrence
Scientific name English name I S M O N
Pternistis griseostriatus Grey-striped Francolin VU •
Pternistis swierstrai Swierstra’s Francolin LC •
Tauraco erythrolophus Red-crested Turaco LC
Colius castanotus Red-backed Mousebird LC •
Gymnobucco vernayi Angola Naked-faced Barbet LC • *
Lybius leucogaster Angola White-headed Barbet LC • *
Platysteira albifrons White-fronted Wattle-eye LC •
Prionops gabela Gabela Helmetshrike EN •
Malaconotus monteiri Monteiro’s Bushshrike LC • 1
Laniarius amboimensis Gabela Bushshrike EN •
Laniarius brauni Braun’s Bushshrike EN •
Phyllastrephus viridiceps Angola White-throated Greenbul LC • *
Phyllastrephus fulviventris Pale-olive Greenbul LC •
Macrosphenus pulitzeri Pulitzer’s Longbill LC •
Cisticola bulliens Bubbling Cisticola LC •
Cisticola bailunduensis Huambo Cisticola LC • *
Cisticola melanura Black-tailed Cisticola LC •
Sheppardia gabela Gabela Akalat LC •
Xenocopsychus ansorgei Angola Cave-Chat LC • • 2
Dioptrornis brunneus Angola Slaty Flycatcher LC •
Nectarinia bocagii Bocage’s Sunbird LC •
Cinnyris ludovicensis Ludwig’s Double-collared Sunbird LC • • 3
Ploceus temporalis Bocage’s Weaver LC •
Euplectes aureus Golden-backed Bishop LC • 4
Lagonosticta ansorgei Ansorge’s Firefinch LC •
Coccopygia bocagei Angolan Swee Waxbill LC •
Estrilda thomensis Cinderella Waxbill LC •
Macronyx grimwoodi Grimwood’s Longclaw LC •
Crithagra benguelensis Benguela Seedeater LC •
IUCN categories, LC Least Concern, NT Near Threatened, VU Vulnerable, EN Endangered, S
Forests of the Western Escarpment (‘Scarp forests’), M Afromontane forests, O Other habitats, N
Notes
1. Recent records from Cameroon are considered doubtful (Mills 2010)
2. Near-endemic as a marginal population was recently found in Namibia (Swanepoel 2013)
3. Isolated populations in Malawi and Tanzania sometimes treated as sub-species are better treated
as distinct species (Bowie et al. 2016)
4. Population on São Tomé Island was very likely introduced by humans as cage birds (Jones and
Tye 2006)
*indicates recent species splits proposed by HBW and BirdLife International (2017) following the
criteria in Tobias et al. (2010). In these cases only phenotypic data (morphology and song) was
used and it would be useful to measure the levels of genetic differentiation from sister taxa
total number of endemic bird species associated with these forests is small
(Table 14.3), but many endemic subspecies are present (Mills et al. 2011a, b) and
molecular studies are likely to support the treatment of several of these populations
as distinct species. It is likely that because of their small size, the Afromontane for-
ests of Angola were not included in the ‘Afromontane archipelago’ biome as defined
by White (1978; cf. Fig. 1). Current research has uncovered a key role of these for-
ests in the evolutionary history of the bird communities of the montane forests of
Africa. Genetic data, together with the reconstruction of past climates and associ-
ated habitats, have shown that the small Angolan Afromontane forests were areas of
high climatic stability throughout glacial cycles and constituted the link between the
montane bird communities of East Africa and the Cameroon mountains (Vaz da
Silva 2015). For species such as the African Hill Babbler Sylvia [Pseudalcippe]
abyssinica and Bocage’s Akalat Sheppardia bocagei, populations were isolated in
the Angola mountains from very early on and are likely to constitute distinct species
(Vaz da Silva 2015).
Apart from the Afromontane and Scarp forests, most other vegetation types and
bird habitats are all part of much larger areas that extend into Angola from (i) the
North: Guinea-Congolian forests; (ii) the East: miombo woodlands; and (iii) the
South: Namib Desert. The avifauna of these biomes is endemic to the habitat type
and thus not confined to Angola. An exception might be Bocage’s Sunbird Nectarinia
bocagii that is known only from Angola (Dean 2000; Mills 2013) and western DRC
(Dowsett et al. 2008), Black-tailed Cisticola Cisticola melanurus (Irwin 1991; Mills
et al. 2011a, b) and the White-headed Robin-Chat Cossypha heinrichi that show a
similar distribution. There are no data on the relative abundance of these species in
the DRC, but in Angola they are considered uncommon to locally common (Dean
2000; Mills and Melo 2013), and thus Angola is very likely to house most of the
population, giving them near endemic status. On the other hand, the formerly
endemic Angolan Cave Chat Xenocopsychus ansorgei is now treated as ‘near-
endemic’ after the discovery of an isolated population in northern Namibia
(Swanepoel 2013).
Commonness and Rarity
The relative abundance of birds in Angola is covered by Mills and Melo (2013).
Most bird species that are widespread in Angola are, if not common, then frequently
seen. About 170 species can be considered uncommon (134 species) or rare (35 spe-
cies). The status of many of the uncommon and rare species is uncertain – some
species are known from a few, or a single specimen, collected a long time ago and
not subsequently recorded. Examples are the Congo Serpent Eagle Dryotriorchis
spectabilis, collected in 1954 at Canzele, Cuanza-Norte, and another specimen col-
lected at Mwaoka, Lunda-Norte, in 1964, and not seen since. The status of the
Lemon-bellied Crombec Sylvietta denti of which a single specimen was collected at
Dundo, Lunda-Norte, in 1958, and a second specimen sound-recorded at Lago
Carumbo (Mills and Dean 2013) is uncertain. Similarly, the status of the Long-
tailed Hawk Urotriorchis macrourus is not known. A specimen was collected at
Cacongo (Lậndana) in Cabinda by L. Petit, probably in 1876, and not recorded
since, despite an extensive collecting trip to Cabinda by the IICA in 1969 (Pinto
1972). Species that have restricted ranges with small populations in Angola, are
generally not rare, and may be locally common within their particular habitat.
Anomalies in Bird Distribution Ranges and Recent Findings
A few species are known from isolated communities within certain areas, with the
nearest conspecifics many kilometres away. These patterns could be real, or they
could be the result of the geographical bias in surveys and collecting. Most collec-
tors favoured the western half of Angola. With the exception of Lunda-Norte, and
parts of Moxico, the coverage by collectors across the east-west gradient was poor
(e.g., Fig. 3 in Monteiro et al. 2014). For a few species we can be certain that the gap
in the distribution between western Angola and western Zambia is real, and is likely
to be the result of relict mountain chains that no longer exist.
The recent exploration of places such as Lagoa Carumbo in Lunda-Norte has
provided much new information on distributions thought to be disjunct (Mills and
Dean 2013). Only 67 species had been collected in the Lagoa Carumbo area during
the 1950s by Heinrich (1958a, b, c). Field surveys by MSLM in 2011 recorded 175
species, with 21 species that had been collected by Heinrich not seen. The data on
the species seen at Lagoa Carumbo included new records for the area, extensions of
ranges and two new records for Angola.
Some remarkable recent finds have been made. The presence of the White-
collared Oliveback Nesocharis ansorgei in Angola was unknown and not even sug-
gested until populations were found in 2011, 2012, and 2013 at Quibaxi and Quitexe,
Cuanza-Norte, and at Uíge (Mills and Vaz Pinto 2015). Before this the nearest
known populations were > 1500 km away in eastern DRC.
Ecotourism in Angola: Birding
Ecotourism is becoming a significant means of raising funds for the protection of

sites of high biodiversity value. With recent changes in the entry requirements for
visitors, improvements in road and hotel infrastructure, and its high biodiversity,
Angola stands to attract a large number of visitors for ecotourism purposes. Most of
the key bird watching sites are unprotected, thus making income from tourism even
more important (Cáceres 2011). For visiting birders the endemic and near-endemic
birds (Table 14.3) are a major drawcard, but the country also holds a variety of spe-
cials summarised in Mills (2018), including Finsch’s Francolin Scleroptila finschii,
Anchieta’s Barbet Stactolaema anchietae, Angola Batis Batis minulla,
Yellow-throated Nicator Nicator vireo, Angola Lark Mirafra angolensis, Brazza’s

Martin Phedina brazzae, Black-and-rufous Swallow Hirundo nigrorufa, Sharp-
tailed Starling Lamprotornis acuticaudus, White-headed Robin-Chat Cossypha
heinrichi, Forest Scrub Robin Erythropygia leucosticta, Oustalet’s Sunbird Cinnyris
oustaleti, Black-chinned Weaver Ploceus nigrimentus, Dusky Twinspot
Euschistospiza cinereovinacea and Black-faced Canary Crithagra capistrata, all
arguably seen more easily in Angola than any other country. Table 14.4 lists the key
sites for visiting birders, with main habitats and most sought-after birds – some of
which are depicted on Fig. 14.2. Most ‘Namibian specials’ are also easily found in
Angola.
here to from Here? Future Directions for Ornithological

W
Research in Angola
As noted in the introduction, it is clear that there have been major advances in the
knowledge of avian species diversity and distribution and in the relative abundance
of species during the last 15 years. This information is crucial to identify potential
conservation areas, although more local surveys of bird communities are needed to
paint a complete picture. We can run algorithms on the species distribution and
abundance data to identify precisely where conservation areas should be, and the
inclusion of other parameters such as endemism and/or threat levels can be used to
Table 14.4 Key sites for birdwatching in Angola

Site name Habitat Key birds
Northern Congo forest Braun’s Bushshrike, Congo basin birds
Escarpment
Calandula Gallery forest, White-headed Robin-Chat, Anchieta’s Barbet, Bannerman’s
Falls area miombo Sunbird
Quiçama NP Gallery forest, Grey-striped Francolin, White-fronted Wattle-eye,
thickets Monteiro’s Bushshrike, Gabela Helmetshrike, Bubbling
Cisticola, Red-backed Mousebird
Cumbira Forest Gabela Akalat, Gabela Bushshrike, Pulitzer’s Longbill,
Forest Red-crested Turaco, Hartert’s Camaroptera, Black-faced
Canary, Forest Scrub Robin
Mount Moco Montane forest, Swierstra’s Francolin, Finsch’s Francolin, Ludwig’s
grassland, Double-collared Sunbird, Bocage’s Sunbird, Black-and-
miombo rufous Swallow, Dusky Twinspot, Angola Lark
Benguela area Arid bushveld Hartlaub’s Francolin, White-tailed Shrike, Bare-cheeked
Babbler
Tundavala Montane forest, Angola Cave Chat, Swierstra’s Francolin, Angola Swee
grassland, rocks Waxbill, Angola Slaty Flycatcher, Oustalet’s Sunbird,
Ludwig’s Double-collared Sunbird
Lubango- Arid bushveld, Cinderella Waxbill, Benguela Long-billed Lark, Rüppell’s
Namibe desert Korhaan
Fig. 14.2 Some special birds of Angola. Top to bottom, left to right: Red-crested Turaco, the
endemic national bird of Angola. (Photo: Lars Petersson); Anchieta’s Barbet, a sought-after spe-
cies with a range extending to the DRC and Zambia but best seen in Angola. (Photo: Maans
Booysen); Braun’s Bushshrike, an endemic restricted to the forests of the northern escarpment.
(Photo: Fiona Tweedie); Monteiro’s Bushshrike, a difficult-to-see endemic associated primarily
with the central escarpment. (Photo: Tasso Leventis); Gabela Helmetshrike, an endemic that occurs
primarily at the base of the central escarpment, as in Quiçama NP. (Photo: Tasso Leventis);
Bocage’s Sunbird is only present in the highlands of Angola and the southwest of the DRC. (Photo:
Alexandre Vaz)
prioritise conservation efforts. Detailed data on patterns of bird diversity and con-
servation threats have been obtained for the core habitats of the only Endemic Bird
Area of Angola: the Afromontane forests at Mount Moco (Mills et al. 2011a, b) and
at the Namba Mountains (Mills et al. 2013a, b); and the Angolan Scarp forests, in
particular for the central scarp forests where most endemism is concentrated (Mills
2010), with special emphasis in Cumbira Forest (Cáceres et al. 2015, 2016, 2017).
So far, and understandably, almost all the research on the birds of Angola has
been on species diversity and distribution patterns, and not processes. Avian diver-
sity surveys – that would feed a permanently updated atlas for the breeding birds of
Angola – should continue, as many areas remain poorly explored or not visited at all
for decades. In parallel with such exploration efforts, research on ecological and
evolutionary processes must be promoted, as this will provide the information that
ultimately is essential for guiding conservation efforts.
Very few studies have been carried out on the biology of individual species, no
long-term studies of breeding have been done, and the nests and eggs of many spe-
cies have yet to be discovered (e.g., Mills and Vaz 2011). Seed dispersal and fru-
givory by birds in Angola is another field that needs investigation, particularly now
where so much habitat is being destroyed for slash-and-burn cultivation and char-
coal making. Birds can play a key role in the rehabilitation of damaged areas. Seeds
regurgitated by birds often germinate below roost sites, and the seedlings can be
collected and planted out. Rehabilitation initiatives are already underway, albeit at
a small scale, on Mount Moco and Cumbira. The Mount Moco reforestation project
has been running since 2010, with the community-run nursery holding over 1400
saplings grown from locally collected seeds, and with almost 950 trees planted back
in the wild (MSLM, unpublished). The Cumbira project is still at its early steps,
with the creation of a pilot-nursery (Aimy Cáceres & Ninda Baptista,
unpublished).
Research on the evolutionary history of the endemic species and subspecies of
Angola is likely to provide novel insights into bird diversification in Africa and on
the uniqueness of the Angolan avifauna (see Endemism section above). The use of
molecular tools will clarify the taxonomic status of species with small and isolated
populations in Angola such as the Orange Ground Thrush Geokichla gurneyi
restricted to Mount Namba or Margaret’s Batis Batis margaritae present only on
Mount Moco and Namba, and separated from its nearest conspecific in western
Zambia by about 800 km. Moreover, the two subspecies occupy rather different
habitats – in Angola Margaret’s Batis is present in patches of Afromontane forest,
whereas in Zambia the species is present in dissimilar evergreen Cryptosepalum
forest. It is highly likely that the two forms constitute well-separated evolutionary
lineages and could be considered different species. This situation, with one “subspe-
cies” present in western Angola and the nearest other “subspecies” present 800–
900 km to the east in Zambia or the Katanga area is repeated in many of the western
Angolan avian taxa, and raises many questions about whether the isolated popula-
tions are two recently diverged forms of one species, or two species. Similarly, the
identification of the complex of swamp-dwelling weavers along the eastern border
of Angola, western Zambia and Katanga is still something of a mystery. The q uestion
has been addressed by several authors (Louette and Benson 1982; Louette 1984;
Dean 1996) but remains unsettled. Molecular tools may be required to clarify the
situation.
The distinctive endemic subspecies of Horus Swift (Apus horus fuscobrunneus)
is known from a single series of specimens taken on the coastal plain of Namibe and
has not been recorded in Angola since the early 1970s. Likewise, the endemic sub-
species of White-headed Barbet (Lybius leucocephalus leucogaster), which was
fairly common around the southern escarpment, was only rediscovered in 2017, in
Tundavala, after almost 40 years of being undetected (Baptista and Mills 2018).
Both taxa have been proposed as endemic species, so finding extant populations in
the field is a high priority.
Future ornithological research will only succeed and grow with greater local
input. There is a great need to stimulate more interest within Angola for the study of
birds, both by engaging students more directly and producing relevant educational
materials for local students. To these ends, joint Portuguese-English language books
have already been produced on The Common Birds of Luanda (Mills and Melo
2015) and The Special Birds of Angola (Mills 2018), to raise interest and awareness.
A basic handbook on ornithology relevant to Angola and written in Portuguese
would be a welcome addition. There is also a need for field courses to provide train-
ing to Angolan students, working together with Angolan universities. Most impor-
tantly, finding ways to encourage the interest of local students in field studies is
greatly needed.
Appendix 14.1 Publications Post 1975 Not Cited in the Text
Beel C (1992) Species new to the Angolan list. Zambian Ornithological Society
Newsletter 22(1):2
Bowen PStJ (1979) Some notes on Margaret’s Batis (Batis margaritae) in Zambia.
Bulletin of the Zambian Ornithological Society 11:1–10
Bowen PStJ (1983) The Black-collared Bulbul Neolestes torquatus in Mwinilunga
District and the first Zambian breeding record. Bulletin of the Zambian
Ornithological Society 13–15:7–14
Bowen PStJ, Colebrook-Robjent JFR (1984) The nest and eggs of the Black-and-
rufous Swallow Hirundo nigrorufa. Bulletin of the British Ornithologists’ Club
104:146–147
Braine S (1990) Records of birds of the Cunene River estuary. Lanioturdus
25:38–44
Brooke RK (1981a) The Feral Pigeon - a ‘new’ bird for the South African list.
Bokmakierie 33:37–40
Brooke RK (1981b) The seabirds of the Moçâmedes Province, Angola. Gerfaut
71:209–225
Collar NJ (1998) Monotypy of Francolinus griseostriatus. Bulletin of the British
Ornithologists’ Club 118:124–126
Dean WRJ (1976) Breeding records of Crex egregia, Myrmecocichla nigra and
Cichladusa ruficauda from Angola. Bulletin of the British Ornithologists’ Club
96:48–49
Dean WRJ (1988) The avifauna of Angolan miombo woodlands. Tauraco
1:99–104
Dean WRJ (2001) The distribution of vultures in Angola. Vulture News 45:20–25
Dean WRJ (2006) Age structure of a Palm-nut Vulture Gypohierax angolensis pop-
ulation. Vulture News 55:8–9
Dean WRJ (2007) Type specimens of birds (Aves) in the Transvaal Museum collec-
tion. Annals of the Transvaal Museum 44:67–121
Dean WRJ, Milton SJ (2005) Stomach contents of birds (Aves) in The Natural
History Museum, Tring, U.K., collected in southern Africa, northern Mozambique
and Angola. Durban Museum Novitates 30:15–23
Dean WRJ, Milton SJ (2007) Some additional breeding records for birds in Angola.
Ostrich 78:645–648
Dean WRJ, Vernon CJ (1988) Notes on the White-winged Babbling Starling
Neocichla gutturalis in Angola. Ostrich 59:39–40
Dean WRJ, Sandwith M, Milton SJ (2006) The bird collections of C. J. Andersson
in southern Africa, 1850–1867. Archives of Natural History 33:159–171
Dean WRJ, Walters MP, Dowsett RJ (2003) Records of birds breeding collected by
Dr WJ Ansorge in Angola and Gabon. Bulletin of the British Ornithologists’ Club
123:239–250
Dean WRJ, Franke U, Joseph G, et al. (2012) Type specimens in the bird collection
at Lubango, Angola. Bulletin of the British Ornithologists’ Club 132:41–45
Dean WRJ, Franke U, Joseph G, et al. (2014) Further breeding records for birds
(Aves) in Angola. Durban Museum Novitates 36:1–36
Dean WRJ, Huntley MA, Huntley BJ, et al. (1988) Notes on some birds of Angola.
Durban Museum Novitates 14:43–92
Lambert K (2006) Seabirds sighted in the waters off Angola, 1966–1988. Marine
Ornithology 34:77–80
Louette M (2002) Relationship of the Red-thighed Sparrowhawk Accipiter erythro-
pus and the African Little Sparrowhawk A. minullus. Bulletin of the British
Ornithologists’ Club 122:218–222
Mendelsohn JM, Haraes L (2018) Aerial census of Cape Cormorants and Cape Fur
Seals at Baía dos Tigres, Angola. Namibian Journal of Environment 2A:1–6
Meyburg B-U, Mendelsohn JM, Ellis DH, et al. (1995) Year-round movements of a
Wahlberg’s Eagle Aquila wahlbergi tracked by satellite. Ostrich 66:35–140
Mills MSL (2007a) Swierstra’s Francolin Francolinus swierstrai: a bibliography
and summary of specimens. Bulletin of the African Bird Club 14:175–180
Mills MSL (2007b) Vocalisations of Angolan Birds. Vol. 1. CD. Birds Angola &
Birding Africa, Cape Town
Mills MSL (2009) Vocalisations of Angolan birds: new descriptions and other notes.
Bulletin of the African Bird Club 16:150–166
Mills MSL (2014a) Dusky Twinspot Euschistospiza cinereovinacea, a new host
species for indigobirds Vidua. Bulletin of the African Bird Club 21:193–199
Mills MSL (2014b) Observations of the rarely seen aerial display of Short-winged
Cisticola Cisticola brachypterus. Bulletin of the African Bird Club 21:200–201
Mills MSL, Oschadleus HD (2013). Black-chinned Weaver Ploceus nigrimentus in
Angola, and its nest. Bulletin of the African Bird Club 20:60–66
Mills MSL, Vaz Pinto P, Haber S (2012). Grey-striped Francolin Pternistis griseo-
striatus: specimens, distribution and morphometrics. Bulletin of the African Bird
Club 19:172–177
Mills MSL, Melo M, Borrow N, et al (2011) The Endangered Braun’s Bushshrike
Laniarius brauni: a summary. Bulletin of the African Bird Club 18:175–181
Morant PD (Compiler). (1996) Environmental Study of the Kunene River Mouth.
CSIR Report EMAS-C96023. CSIR, Stellenbosch
Oschadleus HD, Mills MSL, Monadjem A (2014) Roadside colony densities of
weavers in southern Angola. Lanioturdus 47:17–20
Paterson J, Boorman M, Glendenning J, et al. (2009) Vagrants, range extensions and
interesting bird records for Skeleton Coast Park Namibia and southern Angola.
Lanioturdus 42:4–10
Ripley SD, Bond GM (1979) A third set of additions to the avifauna of Angola.
Bulletin of the British Ornithologists’ Club 99:140–142
Ryan PG, Cooper J, Stutterheim CJ (1984) Waders (Charadrii) and other coastal
birds of the Skeleton Coast, South West Africa. Madoqua 14:71-78
Simmons RE (2010) First breeding records for Damara Terns and density of other
shorebirds along Angola’s Namib Desert coast. Ostrich 81:19–-23
Simmons RE, Braby R, Braby SJ (1993) Ecological studies of the Cunene River
mouth: avifauna, herpetofauna, water quality, flow rates, geomorphology and
implications of the Epupa Dam. Madoqua 18:163-180
Simmons RE, Sakko A, Paterson J, et al. (2010) Birds and conservation significance
of the Namib Desert’s least known coastal wetlands: Baia and Ilha dos Tigres,
Angola. African Journal of Marine Science 28:713-717
Sinclair I (2007) First record of Bob-tailed Weaver Brachycope anomala for Angola.
Bulletin of the African Bird Club 14:78-78
Sinclair I, Chamberlain D, Chamberlain M, et al. (2007) Observations of three little-
known bird species in northern Angola. Bulletin of the African Bird Club
14:55-56
Sinclair I, Spottiswoode CN, Cohen C, et al. (2004) Birding western Angola.
Bulletin of the African Bird Club 16:211-212
Steinheimer FD, Dean WRJ (2007) Avian type specimens and their type localities
from Otto Schütt’s and Friedrich von Mechow’s Angolan collections in the
Museum für Naturkunde of the Humboldt-University of Berlin. Zootaxa
1387:1-25
Stjernstedt R, Aspinwall DR (1979) The nest and eggs of the Bar-winged Weaver
Ploceus angolensis. Bulletin of the British Ornithologists’ Club 99:138-140
Tye A (1992) A new subspecies of Cisticola bulliens from northern Angola. Bulletin
of the British Ornithologists’ Club 112:55-56
References
Bamford AJ, Diekmann M, Monadjem A et al (2007) Ranging behaviour of Cape Vultures Gyps
coprotheres from an endangered population in Namibia. Bird Conserv Int 17:331–339
Bannerman DA (1912) On a collection of birds made by Mr Willoughby P. Lowe on the West
Coast of Africa and outlying islands; with field notes by the collectors. Ibis 54:219–229
Baptista NL, Mills MSL (2018) Angola White-headed Barbet Lybius [leucocephalus] leucogaster
rediscovered. Bull Afr Bird Club 25:225–229
BirdLife International (2017) Endemic bird areas factsheet: Western Angola. Downloaded from
http://www.birdlife.org on 19/04/2017
Bocage JVB du (1877) Ornithologie d’Angola. Part 1. Imprimerie Nationale, Lisbonne (Lisbon),
pp 1–256
Bocage JVB du (1881) Ornithologie d’Angola. Part 2. Imprimerie Nationale, Lisbonne (Lisbon),
pp 257–576
Boulton R (1931) New species and subspecies of African birds. Ann Carnegie Museum 21:43–56
Bowen WW (1931) Angolan birds collected during the Gray African expedition – 1929. Proc Acad
Natl Sci Phila 83:263–299
Bowen WW (1932) Angolan birds collected during the second Gray African expedition – 1930.
Proc Acad Natl Sci Phila 84:281–289
Bowie RCK, Fjeldsa J, Kiure J et al (2016) A new member of the greater double-collared sunbird
complex (Passeriformes: Nectariniidae) from the Eastern Arc Mountains of Africa. Zootaxa
4175:23–42
Braun R (1930) Beitrage zur Biologie der Vögel von Angola. J Ornithol 78:47–49
Braun R (1934) Biologische Notizen über einige Vögel Nord-Angolas. J Ornithol 82:553–560
Büttikofer J (1888) On birds from the Congo and South Western Africa. Notes Leyden Mus
10:209–244
Büttikofer J (1889a) On a new collection of birds from South Western Africa. Notes Leyden Mus
11:65–79
Büttikofer J (1889b) Third list of birds from South Western Africa. Notes Leyden Mus 11:193–200
Cáceres A (2011) Implementation of eco-tourism as a conservation tool to save the last remnants of
Afromontane Forest of Mount Moco, Angola. MSc thesis. University of Porto, Porto
Cáceres A, Melo M, Barlow J et al (2016) Radio telemetry reveals key data for the conservation
of Sheppardia gabela (Rand, 1957) in the Angolan Escarpment Forest. Afr J Ecol 54:317–327
Cáceres A, Melo M, Barlow J et al (2017) Drivers of bird diversity in an understudied African
centre of endemism: the Angolan Escarpment Forest. Bird Conserv Int 27:256–268
Dean WRJ (1974) Breeding and distributional notes on some birds of Angola. Durban Mus Novit
10:109–125
Dean WRJ (1996) The distribution of the Masked Weaver Ploceus velatus in Angola. Bull Br
Ornithol Club 116:254–256
Dean WRJ (2000) The birds of Angola: an annotated checklist. BOU Checklist No. 18. British
Ornithologists’ Union, Tring
Dean WRJ (2001) Angola. In: Evans MI (ed) Fishpool LDC. Important Bird Areas in Africa and
Associated Islands – Priority Sites for Conservation. BirdLife International, Cambridge &
Pisces Publications, Newbury
Dean WRJ, Le Maitre DC (2008) The birds of the Soyo area, Northwest Angola. Malimbus 30:1–18
Dean WRJ, Dowsett RJ, Sakko A et al (2002) New records and amendments to the birds of Angola.
Bull Br Ornithol Club 122:180–184
Dean WRJ, Irwin MPS, Pearson DJ (2003) An isolated population of Singing Cisticola, Cisticola
cantans, in Angola. Ostrich 74:231–232
Dowsett RJ, Aspinwall DR, Dowsett-Lemaire F (2008) The birds of Zambia. An atlas and hand-
book. Tauraco Press and Aves, Liège
Fuchs J, Crowe TM, Bowie RCK (2011) Phylogeography of the fiscal shrike (Lanius collaris):
a novel pattern of genetic structure across the arid zones and savannas of Africa. J Biogeogr
38:2210–2222
Günther R, Feiler A (1986a) Zur phänologie, ökologie und morphologie angolanischer Vögel
(Aves). Teil I: Non-Passeriformes. Faunistische Abhandlungen aus dem Staatlichen Museum
für Tierkunde in Dresden 13:189–227
Günther R, Feiler A (1986b) Zur phänologie, ökologie und morphologie angolanischer Vögel
(Aves). Teil II: Passeriformes. Faunistische Abhandlungen aus dem Staatlichen Museum für
Tierkunde in Dresden 14:1–29
Hall BP (1960a) The faunistic importance of the scarp of Angola. Ibis 102:420–442
Hall BP (1960b) The ecology and taxonomy of some Angolan birds. Bull Brit Mus (Nat Hist) Zool
6:367–463
Hartlaub G (1865) Descriptions of seven new species of birds discovered by Mr J. J. Monteiro in
the Province of Benguela, Angola, West Africa. Proc Zool Soc London 33:86–88
Hartlaub G, Monteiro JJ (1860) On some birds collected in Angola. Proc Zool Soc London
28:109–112
Angola Scarp Project. Proceedings of the VIII Pan-African Ornithological Congress: 279–284.
Kigali, Rwanda, 1992. Koninklijk Museum voor Midden-Afrika, Tervuren
HBW and BirdLife International (2017) Handbook of the Birds of the World and BirdLife
International digital checklist of the birds of the world. Version 2. Available at: http://datazone.
birdlife.org
Heinrich G (1958a) Zur Verbreitung und Lebensweise der Vögel von Angola. J Ornithol 99:121–141
Heinrich G (1958b) Zur Verbreitung und Lebensweise der Vögel von Angola. Systematischer Teil
I (Galli – Muscicapidae). J Ornithol 99:322–362
Heinrich G (1958c) Zur Verbreitung und Lebensweise der Vögel von Angola. Systematischer Teil
III (Hirundinidae – Fringillidae). Journal für Ornitholgie 99:399–421
1:53–74
Irwin MPS (1991) The specific characters of the Slender-tailed Cisticola Cisticola melanura
(Cabanis). Bull Br Ornithol Club 111:228–236
Jones PJ, Tye A (2006) The Birds of São Tomé and Príncipe, with Annobón: Islands of the Gulf of
Guinea. BOU Checklist No. 22. British Ornithologists’ Union & British Ornithologists’ Club,
Oxford
Lambert K (2001) Sightings of new and rarely reported seabirds in Southern African waters. Mar
Ornithol 29:115–118
Leite A, Cáceres A, Melo M, Mills MSL, Monteiro AT (2018) Reducing Emissions from
Deforestation and forest Degradation (REDD+) in Angola: insights from the Scarp Forest con-
servation hotspot. Land Degrad Dev 29:4291–4300. https://doi.org/10.1002/ldr.3178
Louette M (1984) The identity of swamp-dwelling weavers in North-East Angola. Bull Br Ornithol
Club 104:22–24
Louette M, Benson CW (1982) Swamp-dwelling weavers of the Ploceus velatus/vitellinus com-
plex, with the description of a new species. Bull Br Ornithol Club 102:24–31
Lundevall C-F, Ängermark W (1989) Fåglar från Namibia. Axel W. Erikssons fågelsamling från
Sydvästafrika på Vänersborg Museum. Älvsborgs Länsmuseum, Vänersborg
Lynes H (1938) Contribution to the ornithology of the Southern Congo Basin. Revue de Zoologie
et Botanique Africaines 31:3–128
Lynes H, Sclater WL (1933) Lynes-Vincent tour in Central and West Africa in 1930–1931. Part
I. Ibis 75:694–729
Lynes H, Sclater WL (1934) Lynes-Vincent tour in Central and West Africa in 1930–1931. Part
II. Ibis 76:1–51
Meise W (1958) Über neue Hühner-, Specht- und Singvögelrassen von Angola. Abhandlungen des
Naturwissenschaftlichen Vereins in Hamburg, N.F. 2:63–83
Ménégaux A, Berlioz J (1923) Oiseaux. In: Mission Rohan-Chabot: Angola et Rhodesia (1912–
1914). Tome IV: Histoire Naturelle. Fascicle 1: Mammifères (anatomie comparée, embryolo-
gie). Oiseaux. Reptiles. Poissons. Imprimerie Nationale, Paris, pp 107–155
Meyburg B-U, Ellis DH, Meyburg C et al (2001) Satellite tracking of two lesser spotted eagles,
Aquila pomarina, migrating from Namibia. Ostrich 72:35–40
Mills MSL (2006) First record of Pink-billed Lark Spizocorys conirostris for Angola. Bull Afr
Bird Club 13:212
Mills MSL (2013) Little-known African bird: Bocage’s Sunbird Nectarinia bocagii—an Angolan
near-endemic. Bull Afr Bird Club 20:80–88
Mills MSL (2015) First record of Pacific Golden Plover Pluvialis fulva for Angola. Bull Afr Bird
Club 22:223–224
Mills MSL (2018) The Special Birds of Angola/As Aves Especiais de Angola. Go-away-birding,
Cape Town & Kissama Foundation, Luanda
Mills MSL, Cohen C (2007) Brazza’s Martin Phedina brazzae: new information on range and
vocalisations. Ostrich 78:51–54
Mills MSL, Dean WRJ (2007) Notes on Angolan birds: new country records, range extensions and
taxonomic questions. Ostrich 78:55–63
Mills MSL, Dean WRJ (2013) The avifauna of the Lagoa Carumbo area, Northeast Angola.
Malimbus 35:77–92
Mills MSL, Dowd AD (2007) First records of Lemon Dove Aplopelia larvata for Angola. Bull Afr
Bird Club 14:77–78
Mills MSL, Melo M (2013) The checklist of the birds of Angola/A Lista das Aves de Angola.
Associação Angolana para Aves e Natureza & Birds Angola, Luanda
Mills MSL, Melo M (2015) As Aves Comuns de Luanda/The Common Birds of Luanda. Associação
Aves e Natureza Angola, Luanda
Mills MSL, Tebb G (2015) First record of forest swallow Petrochelidon fuliginosa for Angola. Bull
Afr Bird Club 22:221–222
Mills MSL, Vaz A (2011) The nest and eggs of Margaret’s Batis Batis margaritae. Bull Br Ornithol
Club 131:208–210
Mills MSL, Vaz Pinto P (2015) An overlooked population of White-collared Oliveback Nesocharis
ansorgei, in Angola. Bull Afr Bird Club 22:64–67
Mills MSL, Cohen C, Spottiswoode C (2004) Little-known African bird: Gabela Akalat, Angola’s
long-neglected Gabelatrix. Bull Afr Bird Club 11:149–151
Mills MSL, Vaz Pinto P, Dean WRJ (2008) The avifauna of Cangandala National Park, Angola.
Bull Afr Bird Club 15:113–116
Mills MSL, Franke U, Joseph G et al (2010) Cataloguing the Lubango Bird Skin Collection:
towards an atlas of Angolan bird distributions. Bull Afr Bird Club 17:43–53
Mills MSL, Melo M, Vaz A (2011a) Black-tailed Cisticola Cisticola melanurus in eastern
Angola: behavioural notes and the first photographs and sound recordings. Bull Afr Bird Club
18:193–198
Mills MSL, Olmos F, Melo M et al (2011b) Mount Moco: its importance to the conservation
of Swierstra’s Francolin Pternistis swierstrai and the Afromontane avifauna of Angola. Bird
Mills MSL, Melo M, Vaz A (2013a) The Namba mountains: new hope for Afromontane forest
birds in Angola. Bird Conserv Int 23:159–167
Mills MSL, Vaz Pinto P, Palmerim JM (2013b) First records for Angola of Yellow-throated Cuckoo
Chrysococcyx flavigularis, South African Cliff Swallow Petrochelidon spilodera and Red-
tailed Leaflove Phyllastrephus scandens. Bull Afr Bird Club 20:200–204
Mills MSL, Bennett B, Baptista N et al (2016) Red-necked Falcon Falco chicquera in Angola. Bull
Afr Bird Club 23:89–90
Moltoni E (1932) Uccelli d’Angola raccolti da L. Fenaroli durante la spedizione 1930 Baragioli-
Durini. Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale in
Milano 71:169–178
Monard A (1932) Matériaux de la mission scientifique suisse en Angola. Oiseaux. Bull Soc
Neuchâteloise Sci Nat 56:301–355
Monard A (1934) Ornithologie de l’Angola. Arquivos do Museu Bocage 5:1–110
Monteiro M, Reino L, Beja P et al (2014) The collection and database of birds of Angola hosted
at IICT (Instituto de Investigação Científica Tropical), Lisboa, Portugal. ZooKeys 387:89–99
Pinto AA da R (1972) Contribuição para o estudo da avifauna do Distrito de Cabinda (Angola).
Memórias e Trabalhos do Instituto de Investigação Científica de Angola 10:1–103
Pinto AA da R (1983) Ornitologia de Angola, vol 1. Instituto de Investigacão Cientifica Tropical,
Lisboa
Rudebeck G (1955) Aves I. S Afr Anim Life 2:426–576
Rudebeck G (1958) A new race of the Bunting Fringillaria capensis (L.) from Angola. Bull Br
Ornithol Club 78:129–132
Ryan PG, Sinclair I, Cohen C et al (2004) The conservation status and vocalizations of threat-
ened birds from the scarp forests of the Western Angola Endemic Bird Area. Bird Conserv Int
14:247–260
Seabra A (1905a) Aves de Angola da exploração de F. Newton. J Sci Math Phys Nat 7(26):118–128
Seabra A (1905b) Mammiferos e aves da exploração de F. Newton em Angola. J Sci Math Phys
Nat 7(26):103–110
Seabra A (1906a) Aves da exploração de Fr. Newton em Angola – Subsidios para o conhecimento
da destribuição geographica das aves d’Africa occidental. Ann Sci Nat 10:153–159
Seabra A (1906b) Aves de Porto Alexandre. J Sci Math Phys Nat 7(27):143–148
Seabra A (1906c) Nota sobre a existencia de “Diomedia imutabilis” nas costas occidentaes de
Africa. J Sci Math Phys Nat 7(27):141–142
Seabra A (1906d) Ribeirinhas e palmípedes das margens do Rio Cunene. Ann Sci Nat 10:83–90
Seabra A (1907) Sur quelques oiseaux d’Angola envoyés par Francisco Newton. Contribution
à l’étude de la distribution géographique des oiseaux de l’Afrique occidentale. Bull Soc
Portugaise Sci Naturelles 1:41–45
Sekercioğlu CH, Riley A (2005) A brief survey of the birds in Kumbira Forest, Gabela, Angola.
Ostrich 76:111–117
Serle W (1955) The bird life of the Angolan littoral. Ibis 97:425–431
Sharpe RB (1871) On the birds of Angola.–Part III. Proc Zool Soc London 39:130–135
Sharpe RB, Bouvier A (1876a) Catalogue d’une collection recueille à Lândana et Chinchoxo
(Congo), par M. Louis Petit, pendant les mois de janvier février, mars et avril 1876. Bull Soc
Zool Fr 1:36–53
Sharpe RB, Bouvier A (1876b) Sur les collections recueilles dans la région du Congo par MM. le
Dr A. Lucan et L. Petit, depuis le mois de mai jusqu’en septembre. Bull Soc Zool Fr 1:300–314
Sharpe RB, Bouvier A (1877) Nouvelle liste d’oiseaux recueillis dans la région du Congo par MM.
le Dr A. Lucan et L. Petit, de Septembre 1876 à Septembre 1877. Bull Soc Zool Fr 2:470–481
Sharpe RB, Bouvier A (1878) Nouvelle liste d’oiseaux recueillis dans la région du Congo par MM.
le Dr A. Lucan et L. Petit, de Septembre 1876 à Septembre 1877. Bull Soc Zool Fr 3:73–80
Sharpe RB, Monteiro JJ (1869) On the birds of Angola.–Part I. Proc Zool Soc London 37:563–571
Sick H (1934) Ueber einige Vogelbälge aus Nord-Angola, gesammelt von Herrn R. Braun. Ornithol
Monatsberichte 42:167–172
Simmons RE, Mills MSL, Dean WRJ (2009) Oystercatcher Haematopus records from Angola.
Bull Afr Bird Club 16:211–212
Sinclair JC (1981) First sight records of the Booted Eagle in Angola. Ostrich 52:57
Stavrou C, Mills MSL (2013) Observations of birds of the Soyo area, Northwest Angola. Malimbus
35:27–36
Stresemann E (1934) Apalis rufogularis brauni subsp. nov. Ornith Monatsber 62:156–157
Stresemann E (1937) Weitere Vogelbälge aus Nord-Angola, gesammelt von Herrn R. Braun.
Ornithol Monatsberichte 45:51–53
Swanepoel W (2013) Rock star. Angola cave chat: a new species for Namibia. Afr Birdlife 1:30–32
Tobias JA, Seddon N, Spottiswoode CN et al (2010) Quantitative criteria for species delimitation.
Ibis 152:724–746
Traylor MA (1963) Check-list of Angolan Birds. Publicaҫões Culturais 6. Companhia de
Diamantes de Angola (DIAMANG), Lisboa
Vaz da Silva B (2015) Evolutionary history of the birds of the Angolan Highlands – the missing
piece to understand biogeography of the Afromontane Forests. MSc thesis. University of Porto,
Porto
Vaz Pinto P (2002) Field notes on the Grey-striped Francolin (Francolinus griseostriatus) in w
Angola. Newsletter of the Partridge, Quail and Francolin Specialist Group 17:3–5
White CMN (1950) Some records from Eastern Angola. Bull Br Ornithol Club 70:35
White F (1978) The Afromontane region. In: Werger MJA (ed) Biogeography and ecology of
Southern Africa. Springer, Dordrecht, pp 463–513
Chapter 15
The Mammals of Angola
Pedro Beja, Pedro Vaz Pinto, Luís Veríssimo, Elena Bersacola,

Ezequiel Fabiano, Jorge M. Palmeirim, Ara Monadjem, Pedro Monterroso,
Magdalena S. Svensson, and Peter John Taylor
Abstract Scientific investigations on the mammals of Angola started over 150 years

ago, but information remains scarce and scattered, with only one recent published
account. Here we provide a synthesis of the mammals of Angola based on a thorough
survey of primary and grey literature, as well as recent unpublished records. We present
a short history of mammal research, and provide brief information on each species
known to occur in the country. Particular attention is given to endemic and near endemic
species. We also provide a zoogeographic outline and information on the conservation
of Angolan mammals. We found confirmed records for 291 native species, most of
which from the orders Rodentia (85), Chiroptera (73), Carnivora (39), and
Cetartiodactyla (33). There is a large number of endemic and near endemic species,
most of which are rodents or bats. The large diversity of species is favoured by the wide
P. Beja (*)
CEABN-InBio, Centro de Ecologia Aplicada “Professor Baeta Neves”, Instituto Superior de
Agronomia, Universidade de Lisboa, Lisboa, Portugal
e-mail: pbeja@cibio.up.pt
P. Vaz Pinto
do Porto, Campus de Vairão, Vairão, Portugal
L. Veríssimo
e-mail: lmnverissimo@gmail.com
E. Bersacola · M. S. Svensson
Nocturnal Primate Research Group, Faculty of Humanities and Social Sciences, Oxford
Brookes University, Oxford, UK
e-mail: hellenbers@gmail.com; m.svensson@brookes.ac.uk
E. Fabiano
Department of Wildlife Management and Ecotourism, Katima Mulilo Campus, Faculty of
Agriculture and Natural Resources, University of Namibia, Katima Mulilo, Namibia
e-mail: fabianoezekiel@gmail.com

https://doi.org/10.1007/978-3-030-03083-4_15
358 P. Beja et al.
range of habitats with contrasting environmental conditions, while endemism tends to

be associated with unique physiographic settings such as the Angolan Escarpment. The
mammal fauna of Angola includes 2 Critically Endangered, 2 Endangered, 11
Vulnerable, and 14 Near-Threatened species at the global scale. There are also 12 data
deficient species, most of which are endemics or near endemics to the country.
Keywords Africa · Angolan escarpment · Conservation · Endemism · History of

mammalogy · Threatened species · Zoogeography
Introduction
The mammals of Africa, particularly the great apes, large herbivores, and carnivores
are among the most iconic wild species in the world, catching the imagination of
scientists and the general public alike (Monsarrat and Kerley 2018). These species
provided the motivation in the late nineteenth and early twentieth century for some
of the first efforts in wildlife conservation and sustainable use, initially with the
establishment of game reserves and later with the creation of National Parks and
other protected areas (Adams 2013). Today, over one hundred years later, the inter-
est in these charismatic species has increased even further, attracting ever larger
numbers of visitors each year from around the world to protected areas in Africa,
and thus representing important sources of economic revenue in some African coun-
tries. This interest has also been fuelled by appreciation that many mammalian spe-
cies have critical influences on the structure and functioning of African natural
ecosystems (Keesing and Young 2014; Malhi et al. 2016), and that they may provide
important services such as biological pest control in human-dominated landscapes
(Kunz et al. 2011; Sirami et al. 2013; Taylor et al. 2018a). At the same time, how-
ever, African mammals have become involved in some of the most challenging and
J. M. Palmeirim
Departamento de Biologia Animal, Faculdade de Ciências, cE3c – Centre for Ecology,
Evolution and Environmental Changes, Universidade de Lisboa, Lisboa, Portugal
e-mail: jmpalmeirim@fc.ul.pt
A. Monadjem
Department of Biological Sciences, University of Swaziland, Kwaluseni, Swaziland
Mammal Research Institute, Department of Zoology and Entomology, University of Pretoria,
Pretoria, South Africa
e-mail: aramonadjem@gmail.com
P. Monterroso
e-mail: pmonterroso@cibio.up.pt
P. J. Taylor
School of Mathematical & Natural Sciences, University of Venda,
Thohoyandou, South Africa
e-mail: peter.taylor.univen@gmail.com
15 The Mammals of Angola 359
controversial conservation problems in the world, due in particular to the rapid

growth of human populations, agricultural and pastoralism expansion and the asso-
ciated loss of natural habitats (Laurance et al. 2014; Searchinger et al. 2015), defor-
estation (Hansen et al. 2013), conflicts due to crop raiding (Hoare 2015; Seiler and
Robbins 2016) and predation on people and livestock (Loveridge et al. 2017;
McNutt et al. 2017), and poaching for bushmeat (Wilkie et al. 2016; van Velden
et al. 2018) and international trade (Biggs et al. 2013; Wasser et al. 2015; Cerling
et al. 2016). Conservation of African mammals is thus at a crossroads, with a com-
bination of multiple threats and opportunities, demanding a good understanding of
species diversity and ecological requirements, and how they interact with humans in
the context of complex and ever changing social-ecological systems.
In Angola, mammals have long been the focus of research and conservation
efforts. Like elsewhere in Africa, albeit later than in some other countries, mammals
provided the main motivation for the creation of the first Angolan Game Reserves
and National Parks (NPs) in the 1930s, which were mostly located in areas with
particularly important populations of large herbivores (Huntley et al. 2019).
Scientific research started as early as the mid-nineteenth century, with collectors and
zoologists describing the mammalian diversity of the country, including many spe-
cies new to science. Research continued over the years and until the present, but it
was plagued by long periods of interruption, particularly during the civil war of
1975–2002, making Angola one of the least known African countries in terms of its
mammalian fauna. During this period of turmoil there were very few mammalogical
studies (but see, e.g., Anstey 1991, 1993), but efforts to assess the status of the most
charismatic and highly endangered species resumed soon after the situation improved
in the early years of the twenty-first century (Morais et al. 2006a, b; Veríssimo 2008;
Chase and Griffin 2011; Carmignani 2015; Chase and Schlossberg 2016; Fabiano
et al. 2017; Overton et al. 2017; Vaz Pinto 2018; NGOWP 2018; Schlossberg et al.
2018). Scientific interest in Angolan mammals is slowly mounting again, with recent
studies reporting the discovery of new species to science (Carleton et al. 2015;
Svensson et al. 2017), describing important aspects of species distribution and ecol-
ogy (Bersacola et al. 2015; Svensson 2017), and even using cutting-edge tools for
answering complex questions related to species biogeography, phylogeography and
evolution (Rodrigues et al. 2015; Vaz Pinto 2018). This renewed interest is timely, as
Angola is currently striving to expand, reorganise and improve the management of
its system of protected areas, in which there will once again be a strong focus on the
conservation and sustainable use of mammal populations. This endeavour needs to
be solidly rooted in scientific information, profiting from data that has been col-
lected for over 150 years in the country, and promoting new studies that will help
designing cost-effective conservation and management strategies.
This chapter provides a synthesis of what is known at present about the mammals
of Angola. We have considered all mammalian species except cetaceans, which are
treated in Weir (2019). Regarding the pinnipeds, we have only considered the Brown
Fur Seal (Arctocephalus pusillus), which is the sole species of this group breeding
in Angola. We start by presenting a short history of mammal research in the country,
beginning with the studies of the pioneer Portuguese naturalist José Vicente Barbosa
du Bocage, and finishing with the present-day efforts to resume mammological
360 P. Beja et al.
research and to clarify the status of many species that have virtually vanished during
and in the years following the civil war. We then present a brief description of the
mammalian species recorded in Angola, which accompanies the checklist presented
in Appendix. Poorly known endemics and near endemics for which Angola may be
particularly relevant at the global scale are highlighted, but special attention is also
given to iconic species of high conservation concern, though the charismatic Giant
Sable Antelope is dealt with in more detail by Vaz Pinto (2018, 2019). The next sec-
tion provides an overview on the biogeography of the mammalian fauna in Angola,
based primarily on the study of Linder et al. (2012) for sub-Saharan Africa, and the
study of Rodrigues et al. (2015) dealing specifically with the mammals of Angola.
Finally, we provide a summary of the conservation status of Angolan mammals,
largely based on the global assessments by the IUCN (IUCN 2018). We also make
a brief assessment of threats and conservation opportunities for mammals, but leave
the details to Huntley et al. (2019), which deals specifically with the challenges of
biodiversity conservation in the country.
History of Mammal Collecting in Angola
The first truly scientific studies on the vertebrate fauna of Angola, which included
the classification and characterisation of several species of mammals, date back to
the end of the nineteenth century. They are mainly due to José Vicente Barbosa du
Bocage (1869, 1878, 1889a, b, 1890, 1897, 1902), professor of zoology at the
Polytechnic School of Lisbon, to whom the explorer José Alberto de Oliveira
Anchieta regularly sent specimens he collected in various parts of western Angola
(‘Sertão de Loanda’; ‘Sertão de Benguella’, ‘Sertão de Mossâmedes’), and three
other scientists of the time, who exchanged correspondence and opinions with
Bocage. The latter included the German WCH Peters, who published on Angolan
mammals based on the observations of the botanist Frederich MJ Welwitsch (Peters
1865) and the collections made by the German Expedition to Loango-Cabinda
(Peters 1879), and the British WL Sclater and JE Gray, who studied the specimens
sent to the British Museum of Natural History (Gray 1868, 1869) by geologist
Joachim José Monteiro, who lived in Angola at the time. Other collectors, espe-
cially at the end of the nineteenth century and the first decade of the twentieth cen-
tury, also sent specimens to the Polytechnic School’s Zoological Museum. Initially
founded on the collections studied by Bocage, this museum later came to be known
as the Bocage Museum which officially constituted the Zoology Section of the
National Museum of Natural History. Unfortunately, the collections deposited in it
were lost in their entirety, due to a fire that broke out in 1978. Further material was
also sent to other Portuguese museums and universities, such as the specimens
offered to the museum of the University of Coimbra, by Lieutenant Colonel Teodoro
da Cruz, and much later studied by A.A. THEMIDO (THEMIDO, 1931, 1946).
Besides museums in Portugal and the British Museum, other institutions receiv-
ing material from Angola at that time included: the Berlin Museum, which included
material obtained by the German expeditions to Loango and the northeast of Angola,
as well as the Kunene-Sambesi Expedition where the zoologist A Sokolowski col-

lected mammals (Sokolowski 1903); the Leyden Museum, where P Van Der Kellen
collected in southern Angola, and his material was studied by FA Jentink (1887,
1893, 1900, 1901); and the Tring Museum, which, along with the British Museum,
received material collected by J Ansorge in various parts of Angola, largely to be
studied by Thomas Oldfield (Thomas 1892, 1900, 1926, Thomas and Wroughton
1905). In 1916, Thomas would leave his name linked to the classification of
Hippotragus niger variani, the Giant Sable Antelope (Thomas 1916), whose discov-
ery and dispatch of specimens to the British Museum was due to the chief engineer
of the Benguela Railway HF Varian. The amassing of ever larger collections of
mammals from Angola and deposited at the British Museum at this time led to fur-
ther publications of catalogues and other papers on Angola’s mammalian fauna
(Lydekker 1899, 1903, 1904; Lydekker and Blaine 1913–1916; Blaine 1922, 1925)
The 1920s and 1930s witnessed a resurgence of zoological holdings collected in
Angola. Two Swiss missions by Albert Monard, curator of the Museum of Natural
History of La Chaux-de-Fonds, published important contributions to the mamma-
lian fauna of Angola (Monard 1930, 1931, 1933, 1935). Various American expedi-
tions also carried out work in Angola during this period, including the Vernay
Angola Expedition, organised by Arthur Vernay in 1925 to obtain material for the
American Museum of Natural History, New York; the Gray African Expeditions,
led by Prentiss Gray, who in 1929 obtained material, including specimens of
Hippotragus niger variani, for the Philadelphia Academy of Natural Sciences; the
Pulitzer Angola Expedition (1930–1931), organised by the Carnegie Museum and
directed by Rudyard Boulton, who, despite being an ornithologist, collected mam-
malian material; and the Phipps-Bradley Expedition in 1932–1933, organised by
John H Phipps, whose material was donated to the American Museum of Natural
History. It was especially the material in these museums which served as the basis
for the classic work of Hill and Carter (1941), The Mammals of Angola, Africa,
published in 1941, as well as other papers (Hill 1941). Other minor expeditions
included the Karl Jordan Expedition in 1934 whose material was deposited at the
Tring Museum, and reported by St. Leger (1936); and the Percy Sladen and
Kaffrarian Museum Expedition in 1934, organised by the Kaffrarian Museum and
directed by Capt. GC Shortridge. Shortridge collected mainly in Namibia and was
limited in Angola to the banks of the Cunene River. The increase in mammalian
specimens collected in Angola and held in museums around the world up to this
time allowed for the description of new subspecies by Hinton (1921), Matschie
(1900, 1906), Zukowsky (1964) and Zukowsky and Haltenorth (1957).
Following World War II, Portuguese participation in zoological surveys became
predominant in Angola. In fact, a board of overseas research was founded in Portugal
as a branch office of the Ministry of the Colonies – the Junta das Missões Geográficas
e de Investigações Coloniais. From the mid-1950s, the then-established Overseas
Research Board, based in Lisbon, later becoming the Institute of Tropical Scientific
Research (IICT), was the official institution in Portugal to oversee scientific mis-
sions to the Overseas Provinces at the time and, in fact, between 1957 and 1959, a
zoological mission to Angola was conducted, directed by F Frade, the materials of
362 P. Beja et al.
which were deposited at the then IICT Zoology Center. This researcher, later direc-
tor of the Zoology Center, was a prolific contributor of scientific papers on Angolan
mammals on topics including anatomy, taxonomy and conservation (Frade 1933,
1936, 1955, 1956, 1958, 1959a, b, 1960, 1963; Frade and Sieiro 1960). Nonetheless,
the majority of these scientific initiatives were mainly from institutions that were
effectively based in Angola, which, during the 1950s and 1960s, promoted zoologi-
cal explorations and collections in Angola.
Of the greatest importance was the Laboratory of Biology at the Dundo Museum,
in Lunda-Norte, in the extreme northeast of Angola. This museum had two sections,
one for ethnographic and the other for biological studies. Directed by António de
Barros Machado, it became world-renowned for its invaluable collections, as well
as for its prestigious magazine, Cultural Publications of the Diamond Company of
Angola. Barros Machado, in spite of his specialisation as an entomologist, made an
important contribution to the mammalogy of Angola (Machado 1952, 1968, 1969).
RW Hayman of the British Museum studied the mammal material housed in the
Dundo Museum (Hayman 1951, 1963).
The other Angolan institution of importance to mammalogy was the former
Institute of Scientific Research of Angola (IICA), specifically its Sections of
Ornithology and Mammalogy, based in Lubango, Huíla. The first section was
directed by AA Rosa Pinto and the second by J Crawford-Cabral. As a result of
several years of fieldwork and the collaborative work of various personnel, includ-
ing collectors and taxidermists, it was possible to organise, in both these Sections,
an excellent repository of zoological material from Angola. Both Sections still
remain in Lubango, where they are currently part of the Higher Institute of Sciences
and Education (Instituto Superior de Ciências da Educaҫão – ISCED). The study of
the material of the Mammalogy Section has been partly published mainly by
Crawford-Cabral in an extensive number of articles, initially in the Bulletin of the
Institute of Scientific Research of Angola and, more recently, in the Zoology Series
of the magazine Garcia de Orta, and elsewhere (Crawford-Cabral 1961, 1966a,b,
1967, 1968, 1969a, b, 1970a, b, 1971, 1982, 1986, 1987, 1992, 1996, 1997, 1998;
Crawford-Cabral and Fernandes 2001; Crawford-Cabral and Simões 1987, 1988;
Crawford-Cabral and Veríssimo 2005).
However, the interest of foreign countries in the Angolan fauna had not dimin-
ished. During the 1950s and mid-1960s important collections were made by the
German explorer Gerd Heinrich, mostly deposited in the Field Museum of Natural
History, Chicago; Werner Trense, who undertook a collecting expedition in Angola
between 1952–1954, which were deposited at the Hamburg Institute and Zoological
Museum, and studied by him (Trense 1959); and, a decade later, another expedition
from this last museum, which included the museum’s anatomist H Oboussier, whose
collections in Angola were related with her studies on the hypophysis of antelopes
(Oboussier 1962, 1963, 1964, 1965, 1966, 1972, 1976; Oboussier and Von Tyszka
1964).
In the late 1960s and until the mid-1970s the interest of South African zoologists
in Angolan mammalogy was also felt. In 1969, the State Museum of Namibia organ-
ised an expedition to southwestern Angola (mainly within the Namibe Provice),
under the direction of its director, CG Coetzee, which was repeated in 1974; and, in
June and July of the same year, the University of Cape Town and the Wildlife
Society, undertook an expedition to the same regions (Broom et al. 1974). Worthy
of reference, in this period just before independence, are the scientists who per-
formed field work in Angola, such as Richard Estes, with his studies on the Giant
Sable (Estes and Estes 1974) and the ecologist and conservationist Brian Huntley
(1972a, b, 1973a, b, c, d, e, f, 1974).
Following the independence of Angola in 1975, the political situation deterio-
rated rapidly and soon after a civil war raged on until 2002. During this period very
little was added to the knowledge of the Angolan mammalian fauna. However one
should highlight the contributions of Alfred Feiler, assistant to AG Marques at the
University Agostinho Neto, in Luanda, who undertook studies on mammal fauna
(Feiler 1986, 1989, 1990); as well as by a short mammal survey conducted in some
conservation areas (Juste and Carballo 1992); and a rapid assessment of the environ-
mental conditions and fauna in some of the protected areas conducted by Huntley
and Matos (1992).
With the end of the civil war, in 2002, the return of field work conditions and
initiatives was severely hampered by the unknown status of the war legacy such as
land mine fields, and the overall disruption of infrastructure and government institu-
tions. However, the first aerial survey for large mammals in Iona NP was conducted
in 2003 by a joint initiative between the government of Angola and the Namibia
Ministry of Environment (Kolberg and Kilian 2003). At this same time, a concerted
effort was ongoing to assess the status of the Giant Sable Antelope. This later cul-
minated in the establishment of the Giant Sable Project with the assistance of the
Kissama Foundation, which has since been in the forefront of the protection and
recovery of this species Vaz Pinto (2019). The first complete historical review of the
distribution of the ungulate fauna of Angola was published in 2005 (Crawford-
Cabral and Verissimo 2005). Further wildlife monitoring initiatives have been
developed in the southeast of the country. The first aerial surveys conducted in
Cuando Cubango province were undertaken by the organisation Elephants Without
Borders to assess the status of elephant populations, within the Luiana Partial
Reserve, in 2004, 2005 and 2006, and extended in 2015 (Chase and Griffin 2011,
Chase and Schlossberg 2016; Schlossberg et al. 2018). In 2007, the first systematic
ground mammal survey was developed in the former Mucusso Game Reserve
(Veríssimo 2008), in an effort to assist the Angolan Ministry of Environment to
review the protected areas status of southeast Angola. Recent and ongoing initia-
tives, including a large carnivores assessment developed by the organisation
Panthera in Cuando Cubango (see Funston et al. 2017); Huntley et al. 2019), as well
as other initiatives of mammal surveys in Mupa, Bicuar and Iona NPs (Overton
et al. 2017; Fabiano et al. 2017), and elsewhere (INBAC 2016), will continue to
improve the knowledge of the unique mammalian fauna of Angola, and hopefully,
its long term recovery and conservation. Despite these recent efforts, only a single
recent publication has provided a checklist of mammals of Angola (Taylor et al.
2018c).
364 P. Beja et al.
The Mammal Fauna
In this section we provide an overview of the mammalian fauna of Angola, giving at

least a brief comment on each species recorded until now, all of which are presented
in the checklist of Appendix. We also refer to some species that have never been
collected in the country, but that occur very close to the border in neighbouring
countries and thus are likely to occur in Angola. We have also reviewed cases of
species that were once judged to occur in Angola, usually based on old records, but
that have been probably misidentified and thus are no longer considered in the
checklist. The section is based on a wide range of sources, including for example
previous reviews focusing specifically on Angolan mammals (e.g., Hill and Carter
1941; Crawford-Cabral 1998; Crawford-Cabral and Simões 1987,1988; Crawford-
Cabral and Veríssimo 2005), monographs on the mammals of Africa (e.g., Happold
2013; Happold and Happold 2013; Monadjem et al. 2010a, 2015), data from muse-
ums and historical observations available through GBIF (e.g. Bohm and Jonsson
2017; Conroy 2018; Grant and Ferguson 2018; MNHN 2018; MHNG 2018;
Rodrigues et al. 2018; Taylor et al. 2018c), and unpublished data from the co-
authors, among others. These sources reflect a highly uneven survey effort across
Angola, as illustrated by the distribution of records in the GBIF database, and so it
is likely that new mammal species for Angola are still to be discovered, particularly
in less explored regions (Fig. 15.1).
In this review the higher taxonomy (i.e., family level and above) follows Kingdon
et al. (2013), and the taxa are presented in alphabetical order, following the hierar-
chy of orders and families. The taxonomy at species and infraspecific levels is based
largely on that adopted by the Red List of the IUCN (IUCN 2018), which in turn
mostly follows the 3rd edition of Mammal Species of the World – A Taxonomic and
Geographic Reference (Wilson and Reeder 2005). This option was chosen because
this is a generally recognised taxonomy, and because information on global conser-
vation status is available for each of these species. In a few cases we have not fol-
lowed this taxonomy, mainly when there were recent splitting of taxa treated as
conspecific by the IUCN. Although they are not treated systematically, we have
provided information on some particular subspecies, mainly in cases of type locali-
ties or restricted ranges in Angola, distinctive morphologies or ecologies, high con-
servation value, or that may warrant species status upon taxonomic revision.
Afrosoricida (Otter-shrews, Golden Moles)
The two species of Afrosoricida recorded in Angola are Congo Golden Mole (Huetia
leucorhina) and Giant Otter-shrew (Potamogale velox). Little has been published on
the Congo Golden Mole in the country, and the species is only known from a hand-
ful of records from northern Angola where it seems to occur in mosaics of grassland
and moist forests (Hayman 1963; Crawford-Cabral and Veríssimo, unpublished
Fig. 15.1 Distribution of the number of records of species occurrences in Angola registered in the
GBIF database.
366 P. Beja et al.
data). Notably, the first record of Golden Mole in Angola was collected by Von
Mechow in the Cuango river and initially attributed to the Hottentot Golden Mole
(Amblysomus hottentotus), as Chrysochloris albirostrus, but was later assumed to
be a mistake and has since been provisionally synonymised with H. leucorhina (Hill
and Carter 1941; Crawford-Cabral and Veríssimo, unpublished data). The Giant
Otter-shrew is known from relatively few records mostly dating from the nineteenth
century and studied by Barbosa du Bocage (du Bocage 1865, 1882, 1890), or from
the first half of the twentieth century (Seabra 1905; Hill and Carter 1941). This spe-
cies was mainly found in small forest-lined streams in northern Angola, but a couple
of records were obtained in the provinces of Bié and Huíla (Crawford-Cabral and
Veríssimo, unpublished data), likely corresponding to the southernmost distribution
of the species in Angola.
Carnivora (Carnivores)
The Carnivora in Angola are represented by at least 38 species within seven fami-
lies, most of which belong to the family Herpestidae. Although this group is one of
the most studied in Angola, there are still uncertainties regarding the occurrence of
some species. For most species there is very little data on current distribution and
abundance.
Family Canidae
There are at least five canid species in Angola, the most charismatic of which is the
African Wild Dog (Lycaon pictus). This species appears to have once been wide-
spread across Angola occurring from northeast in the province of Lunda-Norte to
southwest in the Namibe province and southeast in the Cunene and Cuando Cubango
province (Crawford-Cabral and Simões 1988; Huntley 1974). Although no esti-
mates of abundance are available, some populations were probably abundant in the
south, along the border with Namibia (Fabiano, unpublished data). Recent surveys
indicate that the species is still resident in southern Angola, with confirmed popula-
tions at Bicuar, Luengue-Luiana and Mavinga NPs (Veríssimo 2008; Overton et al.
2017; Fabiano et al. 2017; Funston et al. 2017; Monterroso et al., unpublished data).
There may also be other populations in the Angolan section of the greater Kavango-
Zambezi (KAZA) region, westwards to the Mupa NP and northwards to the Cameia
NP, where wild dogs were recently confirmed as resident (INBAC 2016; Fabiano
et al. 2017). Preliminary surveys recently estimated wild dog densities at 0.65 indi-
viduals/100 km2, which is comparable to other populations in southern Africa
(Overton et al. 2017). Based on spoor counts and camera trapping, Overton et al.
(2017) estimated a resident population size of 40–50 individuals in Bicuar NP, dis-
tributed through numerous small packs. In the same area, a camera trapping study
by Fabiano et al. (2017) suggested a minimum population size of ca. 60 individuals
in 10 different packs (2–38 individuals each), and confirmed reproduction in one

wild dog pack. At the Luengue-Luiana and Mavinga NPs, Funston et al. (2017)
estimated densities of 0.7 individuals/100 km2 and a population size of 599 ± 260
individuals using camera trapping and spoor tracking.
The four species of small canids occurring in Angola are Bat-eared Fox (Otocyon
megalotis), Cape Fox (Vulpes chama), Black-backed Jackal (Canis mesomelas), and
Side-stripped Jackal (Canis adustus). Bat-eared Fox range based on few historical
records seems to be confined to the Kaokoveld Desert, Namibian Savanna Woodlands
and Angolan Mopane Woodlands, in southern Angola (Crawford-Cabral and Simões
1987). Recent surveys have confirmed its presence at Iona, Bicuar, Mupa and
Luengue-Luiana and Mavinga NPs (Veríssimo 2008; Fabiano et al. 2017; Overton
et al. 2017; Funston et al. 2017). The few historical records of Cape Fox also sug-
gest that it is confined to the Kaokoveld Desert and Namibian Savanna Woodlands
in southwest Angola (Crawford-Cabral and Simões 1987). Recent surveys indicate
that the species still persist at Iona and Bicuar (Overton et al. 2017; Fabiano et al.,
unpublished data), but it was not detected at Mupa (Overton et al. 2017). The spe-
cies is probably absent from the southeast, where it was not detected in surveys
carried out at Luengue-Luiana and Mavinga NPs (Veríssimo 2008; Overton et al.
2017; Funston et al. 2017).
The historical records of the Black-backed Jackal suggest a distribution mostly
restricted to the arid coastal belt within the Kaokoveld Desert, Namibian Savanna
Woodlands and Angolan Mopane Woodlands (Crawford-Cabral and Simões 1987),
but has been recorded recently as far north as the outskirts of Luanda and above
2000 m at the Humpata plateau (Vaz Pinto, unpublished data). Contemporary
records indicate its presence in Iona NP (Fabiano et al., unpublished data), but also
in Bicuar, Mupa, Luengue-Luiana and Mavinga NPs, which are dominated by the
Zambezian Baikiaea Woodlands and Angolan Mopane Woodlands (Fabiano et al.
2017; Overton et al. 2017; Funston et al. 2017). The Side-stripped Jackal appears to
have had a wider historical range, ranging from a few records in the Southern
Congolian Forest-Savanna Mosaic in northeast Angola (Lunda-Norte province),
and more records falling within the Angolan Miombo Woodlands from central
northwest to central-southwest and highlands (Crawford-Cabral and Simões 1987).
Recent surveys confirmed its presence in the Luengue-Luiana and Mavinga NPs
(Veríssimo 2008; Funston et al. 2017), and in Cangandala NP (Vaz Pinto, unpub-
lished data). Surprisingly, it has also been recorded recently in dry coastal savanna
in Quiçama NP (Groom et al. 2018). It may also occur in Bicuar and Mupa NPs,
though it was probably overlooked in recent surveys more directed to endangered
species (Overton et al. 2017; Fabiano et al. 2017).
Family Felidae
The Felidae are represented in Angola by at least seven species, including iconic
and threatened species such as African Lion (Panthera leo), Leopard (Panthera par-
dus), and Cheetah (Acinonyx jubatus). Historically, lions were probably widespread,
368 P. Beja et al.
inhabiting the Western Congolian Forest-Savanna Mosaic in the northeast, the

Miombo Woodlands in central Angola, and the Savanna, Mopane and Baikiaea
Woodlands in the south along the border with Namibia (Crawford-Cabral and
Simões 1988; Veríssimo 2008; Huntley 1973c, 1974). The lion subspecies occurring
in the country are poorly known, but recent phylogeographic studies suggest that
Angola may represent a contact zone between the Central (P. l. leo) and Southern
African (P. l. melanochaita) lineages (Barnett et al. 2014), and thus the genetic
composition of Angolan lions could help elucidating the evolutionary history of this
species in the African continent. Recent surveys indicate that lions still exist in the
Luengue-Luiana and Mavinga NPs, while two recent records indicate their possible
presence in the surroundings of the Cameia NP (iNaturalist.org 2018a, b). Funston
et al. (2017) estimated the lion population of the Luengue-Luiana and Mavinga NPs
to be about 10–30 individuals. They concluded that the very low biomass of pre-
ferred prey species was the main limiting factor for lions locally, as observed else-
where (Bauer et al. 2015; Lindsey et al. 2017; Wolf and Ripple 2016). Recent
surveys have failed to detect the species in Bicuar, Mupa and Quiçama NPs (Overton
et al. 2017; Fabiano et al. 2017; Groom et al. 2018; Monterroso et al., unpublished
data). However, park rangers and farmers in the vicinity of Bicuar NP have reported
recent sightings of vagrant animals, suggesting that they may still occur in low num-
bers in the region (Fabiano, unpublished data). Recent observations of lone indi-
viduals have been obtained in Luando Reserve (Vaz Pinto, unpublished data). Other
surveys, mostly relying on desktop surveys and interviews, indicate that lions might
still occur in Cameia NP, and in the former Coutada do Mucusso (Veríssimo 2008;
Purchase et al. 2007a, b), but these are unlikely to hold resident populations.
Cheetah historically appear to have occurred across Angola, inhabiting various
habitats/ecoregions such as the Miombo Woodlands in the northeast and south-
central Angola, the Angolan Scarp Savanna and Woodlands in the northwest, and in
the Namibian Savanna and Mopane Woodlands as well as Zambezian Baikiaea
Woodlands along the border with Namibia (Crawford-Cabral and Simões 1988;
Veríssimo 2008). The subspecies represented in the country is the Southern African
Cheetah (A. j. jubatus) (Kitchener et al. 2017). The current status of the Cheetah in
Angola is poorly known, but it still occurs in some NPs (Funston et al. 2017; Kolberg
and Kilian 2003; Purchase et al. 2007a, b; Fabiano, unpublished data; Álvaro
Batista, personal communication). Funston et al. (2017) estimated that Cheetah
occupy roughly 8% of the Luengue-Luiana and Mavinga NPs, occurring at a density
of 0.2 individuals/100 km2. Fabiano et al. (unpublished data) also using spoor counts
estimated that cheetahs occupy approximately 28% of Iona’s plains, at a density of
0.61 individuals/100 km2 (0.17–1.98). This population is considered resident given
the frequent report of sightings, including that of females with cubs (Bruce Bennett
and Álvaro Batista, personal communication). The Iona cheetah population appears
to be genetically similar to the Namibian counterpart based on a limited scat sample
size (n = 22) genotyped at 8 loci (Fabiano et al., unpublished data). Other occasional
sightings suggest cheetahs may be present in Cameia NP. Cheetahs were not
detected in Bicuar and Mupa NPs (Overton et al. 2017; Fabiano et al. 2017), and
Overton et al. (2017) even suggests that they have been absent from the region for
at least a decade. Recent camera-trapping detected the species in western Cuando
Cubango (Stefan van Wyk, personal communication) and southern Moxico
(NGOWP 2018).
African Leopard historically had a wide distribution across Angola, ranging from
the northwest and the northeast, through central Angola to the southern border with
Namibia. However, most historical records were from National Parks. The subspe-
cies occurring in Angola is the African Leopard (Panthera pardus pardus) (Kitchener
et al. 2017). The distribution range appears not to have reduced since the 1970s, as
they still appear to be thriving throughout most of the country, including NPs and
their surrounding areas (INBAC 2016). Using spoor counts Overton et al. (2017)
estimated a density of 4.37–14.7 leopards/100 km2 in the Bicuar NP. Camera-
trapping also detected leopards in Mupa NP, though apparently at lower densities
than in Bicuar (Overton et al. 2017). Based on spoor counts, Funston et al. (2017)
found leopards to be widely distributed throughout Luengue-Luiana and Mavinga
NPs, with an estimated population size of 518 ± 190 individuals. Their camera trap-
ping efforts along the Cuando, Luiana and Luengue rivers allowed the detection of
120 different individuals, and estimated a density of 1.5 ± 0.14 leopards/100 km2.
Leopards are also found in the Iona NP (INBAC 2016), at a density of 1.02 (0.41–
2.39) leopards/100 km2 (Fabiano et al., unpublished data), as well as in Luando
reserve, and Quiçama and Cangandala NPs (INBAC 2016; Groom et al. 2018;
Fabiano, unpublished data; Vaz Pinto, unpublished data).
The other Felidae known to occur in Angola are Caracal (Caracal caracal),
African Golden Cat (Caracal aurata), African Wildcat (Felis silvestris) and Serval
(Leptailurus serval). Historical records indicate the presence of Caracal in the arid
zone of southwestern Angola and the Miombo Woodlands of Cunene (Crawford-
Cabral and Simões 1987). Recent surveys have confirmed its presence at Iona NP
(Fabiano et al., unpublished data) and in Bicuar and Mupa NPs (Fabiano et al. 2017;
Overton et al. 2017), and in the Luengue-Luiana and Mavinga NPs (Veríssimo
2008; Funston et al. 2017). The occurrence in Angola of the closely related African
Golden Cat (Caracal aurata) was only confirmed very recently from an animal
photographed on a local bushmeat market in northwestern Angola (Errol de Beer,
personal communication). This cat is endemic to the forests of Equatorial Africa,
particularly associated to areas of riverine forests with low human disturbance, and
even penetrating savanna regions (Sunquist and Sunquist 2009; Bahaa-el-din et al.
2015). From the two recognised subspecies of Golden Cat, the one probably occur-
ring in Angola is C. a. aurata (Sunquist and Sunquist 2009; Bahaa-el-din et al.
2015; Kitchener et al. 2017). African Wild Cat was historically widespread, occur-
ring across most of the country (Crawford-Cabral and Simões 1987; Crawford-
Cabral and Veríssimo, unpublished data). The species was associated with Miombo
Woodlands, the Scarp Savanna and Woodlands, and the Kaokoveld Desert. Recent
surveys have confirmed its presence in Bicuar and Mupa NPs (Fabiano et al. 2017;
Overton et al. 2017), Quiçama NP (Groom et al. 2018), and in the Luengue-Luiana
and Mavinga NP (Veríssimo 2008; Funston et al. 2017). Serval has a similar histori-
cal widespread distribution, occurring particularly across the western part of the
370 P. Beja et al.
country (Crawford-Cabral and Simões 1987). Within its range, Serval was associ-
ated with Miombo Woodlands, the Scarp Savanna and Woodlands, and the
Kaokoveld Desert (Crawford-Cabral and Simões 1987). Two historical records
were retrieved from northeastern Angola. Recent surveys have confirmed the pres-
ence of Serval in Bicuar and Mupa NPs (Fabiano et al. 2017; Overton et al. 2017),
Quiçama NP (Groom et al. 2018), as well as in Mucusso area in Cuando Cubango
(Veríssimo 2008). Additionally, it regularly surfaces as bushmeat near Luanda (Vaz
Pinto, unpublished data). Black-Footed Cat (Felis nigripes) may occur marginally
in southern Angola, but there seems to be no confirmed records (Sliwa 2013).
Family Herpestidae
The Family Herpestidae in Angola is represented by at least 12 species. The most

well-known and charismatic is certainly the Meerkat (Suricata suricatta), a social
mongoose widespread in the western parts of southern Africa (Jordan and Do Linh
San 2015). Historical records suggest that in Angola it is confined to the arid south-
west, including the Iona NP (Crawford-Cabral and Simões 1987), corresponding to
the northwest edge of the species’ range. Recent surveys suggest that meerkats still
occur in this area (Fabiano et al., unpublished data; Monterroso et al., unpublished
data). Crawford-Cabral (1971), proposed that the population occuring in Angola is
a distinct subspecies, Suricata suricatta iona.
Another interesting group of species is that including the Herpestes mongooses,
for which there are considerable taxonomic uncertainties (Taylor and Goldman
1993; Crawford-Cabral 1996; Gilchrist et al. 2009; Rapson et al. 2012; Veron et al.
2018). The three species known to occur in Angola are Kaokoveld Slender Mongoose
(Herpestes flavescens), Egyptian Mongoose (H. ichneumon) and Common Slender
Mongoose (H. sanguineus) (Crawford-Cabral 1996). Kaokoveld Slender Mongoose
was described by Barbosa du Bocage from specimens collected in Angola, and is
endemic to southwestern Angola and northwestern Namibia. However, there are
marked variations between two subpopulations, which have been assigned to differ-
ent subspecies or even species (Rathbun and Cowley 2008; Rapson et al. 2012).
Individuals with tan or yellowish pelage confined to southwestern Angola are
assigned to the subspecies H. f. flavescens (or H. flavescens sensu stricto), while
those with very dark pelage and with a distinctive rufus tinge that occur in north-
western and northcentral Namibia have been assigned to H. f. nigrata (or H. nigra-
tus) (Crawford-Cabral 1996; Tromp 2011; Taylor 2013). Very little is known about
this species, particularly in Angola, where most assumptions on their distribution
derive from habitat-interpreted satellite imagery (Rapson and Rathbun 2015). In
contrast to the previous species, the Egyptian Mongoose and the Common Slender
Mongoose are thought to be widespread (Crawford-Cabral and Simões 1987).
A group of five species of Herpestidae are thought to have wide distributions in
Angola, though their current range and abundances are poorly known. Possibly the
most widespread of these are the White-tailed Mongoose (Ichneumia albicauda),
the Banded Mongoose (Mungos mungo) and the Dwarf Mongoose (Helogale par-
vula), as historical records report their presence throughout the country (Crawford-
Cabral and Simões 1987; Trombone 2016; Figueira 2017; Grant and Ferguson
2018; Rodrigues et al. 2018). The White-tailed Mongoose has recently been con-
firmed in Cameia and Cangandala NPs, and the Banded Mongoose appears to be
abundant in Luando Reserve and Cangandala NP (Vaz Pinto, unpublished data).
The Marsh Mongoose (Atilax paludinosus) is associated with riparian habitats, such
as swamps and streambed areas, though occurring also in estuarine and marine hab-
itats (Gilchrist et al. 2009). Historical records of these species have mainly been
from western Angola (Trombone 2016; Rodrigues et al. 2018), though it may be
more widepread and has recently been confirmed in Cangandala NP (Vaz Pinto,
unpublished data). The Selous Mongoose (Paracynictis selousi) has the northwest-
ern edge of its range in Angola, with historical records from southern provinces
(Crawford-Cabral and Simões 1987; Trombone 2016; Conroy 2018; Grant and
Ferguson 2018; Rodrigues et al. 2018). Recent surveys confirmed its presence at
Luengue-Luiana and Mavinga NPs (Funston et al. 2017) and Bicuar NP (Overton
et al. 2017), albeit at low densities.
Finally, another three Herpestidae have very restricted or probably underesti-
mated ranges in Angola. Ansorge’s Cusimanse (Crossarchus ansorgei) may be more
widespread than usually believed because it has a relatively wide distribution in
rainforests of neighbouring regions of DRC (Angelici and Do Linh San 2015).
Although in Angola it was known from a single specimen collected in 1908 north of
the Cuanza River (Crawford-Cabral and Simões 1987), recent records suggest it
may extend its distribution along the escarpment to Cuanza-Sul (Michael Mills,
personal communication). The population in Angola is assigned to the endemic
subspecies C. a. ansorgei. Black-legged Mongoose (Bdeogale nigripes) seems to be
restricted in Angola to the enclave of Cabinda (Crawford-Cabral and Simões 1987).
The Yellow Mongoose (Cynictis penicillata) only occurs in a narrow fringe in the
extreme southwest of Angola (Crawford-Cabral and Simões 1987), though it is
widespread elsewhere in southern Africa. We are unaware of recent records of any
of these species in Angola.
Family Hyaenidae
There are three species of the family Hyaenidae recorded in Angola (Crawford-
Cabral and Simões 1988). Historical and contemporary records indicate that the
Brown Hyaena (Parahyaena brunnea) is largely confined to the arid parts of south-
western Angola, in Kaokoveld Desert and Namibian Savanna Woodlands (Huntley
1974; Fabiano et al., unpublished data). This range encompasses the Skeleton Coast
Transfrontier Park between Angola and Namibia. Recent surveys indicate that it is
widespread in the Iona NP (Fabiano et al., unpublished data), but it was not detected
in Luengue-Luiana NP, despite its presence in the nearby Bwabwata NP of Namibia
(Funston et al. 2017). Spotted hyaenas (Crocuta crocuta) historically were widely
distributed across Angola with main populations occurring in Zambezian Baikiaea
Woodlands in the south of the country, though it also appeared to have been
372 P. Beja et al.
widespread in the west. The Spotted Hyaena is one of the few large carnivores spe-
cies that shows no evidence of recent population declines throughout its African
range (Bohm and Höner 2015), although the situation may not necessarily be identi-
cal in Angola. Recently, populations were estimated at 10.8–18.0 individu-
als/100 km2 in Bicuar NP (Overton et al. 2017), 1.4 individuals/100 km2 in Mupa
NP (Overton et al. 2017), and 0.9 individuals/100 km2 in Luengue-Luiana and
Mavinga NPs (Funston et al. 2017). The presence of Spotted Hyaena has not been
confirmed in surveys of various protected areas, and they appear to have been extir-
pated from Luando Reserve and Cangandala NP (Vaz Pinto, unpublished data). In
Quiçama NP an active den was known to be present on the Cuanza floodplain as
recently as 2013 (Vaz Pinto, unpublished data), but a recent large mammal survey
could not find evidence of the species (Groom et al. 2018). Overall, the species is
expected to remain widely distributed in Angola (INBAC 2016). Aardwolf (Proteles
cristata) is the least known of the three Hyaenidae of Angola, occurring only in the
south of the country (Crawford-Cabral and Veríssimo, unpublished data). Recent
records from direct observations and roadkills suggest the species to be relatively
common along the arid coastal plain to as far north as Benguela, and on the high-
lands of Humpata plateau (Vaz Pinto, unpublished data).
Family Mustelidae
The Mustelidae are represented in Angola by Honey Badger (Mellivora capensis),

Striped Polecat (Ictonyx striatus), African Striped Weasel (Poecilogale albinucha),
and three species of otters (Crawford-Cabral and Simões 1987). Historical data on
Honey Badger suggest that it was more frequent in the southwest and east of the
country, within the Angolan Miombo Woodlands and the Zambezian Baikiaea
Woodlands (Crawford-Cabral and Simões 1987), but also occurring in the prov-
inces of Malanje and Moxico (Trombone 2016; MHNG 2018; Rodrigues et al.
2018). Recent surveys have confirmed its presence at Iona, Bicuar, Mupa, Quiçama,
Cangandala, Luengue-Luiana and Mavinga NPs (Veríssimo 2008; Fabiano et al.
2017; Fabiano, unpublished data; Overton et al. 2017; Funston et al. 2017; Groom
et al. 2018; Monterroso et al., unpublished data; Vaz Pinto, unpublished data).
The Striped Polecat is a generalist that occurs in most habitats except dense rain-
forests. Historically this species was recorded throughout the country (Trombone
2016; Figueira 2017; Grant and Ferguson 2018; Rodrigues et al. 2018), and is
expected to maintain a wide distribution. It was detected in the recent surveys car-
ried out in NPs of southern Angola (Veríssimo 2008; Funston et al. 2017; Monterroso
et al., unpublished data). The African Striped Weasel is also a widespread habitat
generalist, though it is often overlooked due to its secretive habits. Probably because
of this it has a relatively small number of records in Angola, though widely distrib-
uted throughout the country (Crawford-Cabral and Simões 1987)
Little is known about the three otter species occurring in Angola, with no system-
atic surveys available to describe their current status (Crawford-Cabral and Simões
1987). The Congo Clawless Otter (Aonyx congicus) is associated with the rainfor-
ests of the Congo Basin (Jacques et al. 2015a), and so in Angola should be restricted
to Cabinda and Lunda-Norte. This species is sometimes treated as conspecifc of the

African Clawless Otter (Aonyx capensis) (Wozencraft 2005), which in spite of the
paucity of historical records is thought to have a wide distribution in the east and
south of Angola (Veríssimo 2008; Jacques et al. 2015b), and has recently been
recorded at the Humpata plateau (Vaz Pinto, unpublished data). The Spotted-necked
Otter (Hydrictis maculicollis) is likely to be the most widespread otter in freshwater
systems throughout Angola (Reed-Smith et al. 2015), though the historical records
for the country are relatively few and scattered.
Family Nandiniidae
The African Palm Civet (Nandinia bionotata) is the sole representative of the family
Nandiniidae (Crawford-Cabral and Simões 1987). Historical records of the species
have been mainly made in the provinces of Uíge, Cuanza-Norte, and Lunda-Norte,
suggesting its occurrence throughout the north of Angola. There have been no recent
published records of this species in Angola, though it probably is still widespread
within its former known range in the northern half of the country.
Family Otariidae
The Brown Fur Seal (Artocephalus pusillus) is the only pinniped breeding in
Angola, with several large colonies in the island of Tigres (Meÿer 2007). This is the
northern limit of the species distribution, which extends along the coast of Namibia
to Algoa Bay in South Africa (Hofmeyr 2015). Other species occur occasionally
along the coast of Angola, with records of for instance Sub-Antarctic Fur Seal
(Arctocephalus tropicalis) (Carr et al. 1985) and South-Atlantic Elephant Seal
(Mirounga leonina) (França 1967).
Family Viverridae
The Viverridae in Angola include the African Civet (Civettictis civetta), and three
species of the genus Genetta. The civet was reported to occur in northern (Bengo,
Cuanza-Norte and Malanje), central-west (Benguela), and southwest Angola
(Namibe and Huíla province) (Crawford-Cabral and Simões 1987), mainly associ-
ated with the Western Congolian Forest-Savanna Mosaic, Angolan Scarp Savanna
and Woodlands, Angolan Miombo Woodlands. Recent surveys have confirmed its
presence at the Mupa, Quiçama, Cangandala, Mavinga and Luengue-Luiana NPs
(Veríssimo 2008; Overton et al. 2017; Funston et al. 2017; Groom et al. 2018; Vaz
Pinto, unpublished data). The three genets currently recognised in Angola are the
Small Spotted Genet (Genetta genetta), the Large Spotted Genet (Genetta macu-
lata), and the Miombo or Angola Genet (Genetta angolensis) (Crawford-Cabral and
Simões 1987). Small Spotted Genet was identified as G. g. felina by Crawford-
Cabral and Simões (1987). Historical records suggest that the species occurred
374 P. Beja et al.
predominantly in southwestern Angola within the Kaokoveld Desert (Namibe) and

the Angolan Mopane and Miombo Woodlands (Huíla and Cunene) (Crawford-
Cabral and Simões 1987). Recent surveys have confirmed its presence in southeast
Angola in Luengue-Luiana and Mavinga NPs (Funston et al. 2017). Small Spotted
Genets may still occur in Bicuar and Mupa NPs, and their surrounding areas, as
spotted genets (Genetta spp.) have been detected on cameras and roadkills, but not
identified to the species level (Overton et al. 2017; Fabiano et al. 2017; Monterroso
et al., unpublished data; Vaz Pinto, unpublished data). The taxonomy of the Large
Spotted Genet is still to be resolved (Angelici et al. 2016), and it appears to be a
‘superspecies’ comprising several valid species. Large Spotted Genet was recorded
as G. m. rubiginosa by Crawford-Cabral and Simões (1987). It appears to have had
widespread distribution occurring across the western part of the country (Crawford-
Cabral and Simões 1987). Recent surveys have confirmed its presence in southeast
Angola in the Luengue-Luiana and Mavinga NPs (Funston et al. 2017), and in
Quiçama NP (Groom et al. 2018). The Miombo Genet is considered a near-endemic
of the miombo ecoregion (Timberlake and Chidumayo 2011). According to Gaubert
et al. (2016), Miombo Genets’ westernmost distribution range should be restricted
to central Angola’s miombo woodlands, as it was mainly present in central, south-
west and south-central Angola (Crawford-Cabral and Simões 1987; Trombone
2016; Bohm and Jonsson 2017; Rodrigues et al. 2018). However, Huntley and
Francisco (2015) suggest that the species could be widespread across Lunda-Norte
province, suggesting its distribution in Angola may be underestimated. Recent sur-
veys have confirmed its presence in Cangandala and Bicuar NPs (Overton et al.
2017; Vaz Pinto, unpublished data).
etartiodactyla (Pigs, Hippopotamuses, Chevrotain, Giraffes,

C
Deer, Bovids)
The Cetartiodactyla includes 33 native species of 5 families in Angola. Most species

belong to the Bovidae, which is represented by at least 27 species (Crawford-Cabral
and Veríssimo 2005). Although this is one of the best-known animal groups in the
country, there are many uncertainties regarding the current status and distribution of
most species. Given the large number of species, information on this group is pro-
vided per family, while the bovidae are presented per tribe.
Family Bovidae
Tribe Aepycerotini
The tribe Aepycerotini comprises only one genus and one species, the Impala
(Aepyceros melampus). Although up to six subspecies of impala have been listed,
their validity was problematic and limits poorly defined (Ansell 1972; Fritz and
Bourgarel 2013). Most often only two races are recognised, the Common Impala (A.
m. melampus) and the Black-faced Impala (A. m. petersi), which is also supported
by molecular data (Lorenzen et al. 2006). The Black-faced Impala was described as
a new species from a specimen collected at Humbe, Cunene Province (du Bocage
1879), and some authors maintained a dual-species classification (Shortridge 1934;
Groves and Grubb 2011). Both taxa naturally occurred in Angola, in two disjunct
and well demarcated populations (Crawford-Cabral and Veríssimo 2005). The
Common Impala occurs in the southeast of the country between the Cubango and
the Cuando rivers, with most historical records obtained along the former
(Sokolowski 1903; Wilhelm 1933; Hill and Carter 1941; Huntley 1973c; Crawford-
Cabral and Veríssimo 2005). The Common Impala was given as extirpated along the
Cubango River by Veríssimo (2008), but subsequently relatively small numbers
have been confirmed in the Luengue-Luiana NP (Chase and Schlossberg 2016;
Funston et al. 2017). The Black-faced Impala is endemic to Kaokoland in north-
western Namibia and southwestern Angola (Fritz and Bourgarel 2013), but the bulk
of its distribution used to be in Angola where it extended as far north as Benguela
and mostly west of the Cunene River (Hill and Carter 1941; Swart 1967; Crawford-
Cabral and Veríssimo 2005). Before 1975 the Black-faced Impala was represented
in protected areas such as Iona, Bicuar and Mupa NPs (Huntley 1972c, 1973c;
Crawford-Cabral and Veríssimo 2005). Currently it is likely on the verge of extinc-
tion in Angola, as recent surveys have failed to record it in Iona, Bicuar and Mupa
NPs (Kolberg and Kilian 2003; Overton et al. 2017; van der Westhuizen et al. 2017).
Very small numbers may still linger in eastern Iona (Álvaro Baptista, personal com-
munication), and one single specimen was observed in 2016 near Serra da Neve in
northern Namibe province (Vaz Pinto, unpublished data).
Tribe Alcelaphini
The Alcelaphini are represented in Angola by the genera Damaliscus, Alcephalus

and Connochaetes. The earlier references mentioning the presence of Tsessebe
(Damaliscus lunatus) in Angola are scattered (Sokolowski 1903; Monard 1935; Hill
and Carter 1941; Varian 1953), but at some point the species was likely to be rela-
tively common and widely distributed across southeastern Angola to the east of the
Cunene river, and along the eastern border as far north as the upper Zambezi drain-
age (Newton da Silva 1970; Huntley 1973c; Crawford-Cabral and Veríssimo 2005).
It was once considered common in the plains of Cameia NP and in the areas now
included in Luengue-Luiana NP (Crawford-Cabral and Veríssimo 2005). Very little
is known in terms of current status of the species in Angola, but it appears to have
been much reduced in numbers. Veríssimo (2008) suggested their persistence in
northern Luengue and Luiana based only on witness reports, but an extensive aerial
survey did not record any Tsessebe in Luengue-Luiana and Mavinga NPs (Chase
and Schlossberg 2016). However, the species was confirmed in the area through
camera-trapping (Funston et al. 2017).
376 P. Beja et al.
The taxonomy of the hartebeest, genus Alcephalus, is controversial, with some

authors considering it a monospecific genus with up to eight subspecies (Flagstad
et al. 2001; Gosling and Capellini 2013), while others recognise different species
(e.g. Ansell 1972; Groves and Grubb 2011). Two morphologically very distinct taxa
occur in Angola in disjunct populations, Red Hartebeest (A. b. caama) and
Lichtenstein’s Hartebeest (A. b. lichtensteini), which are often treated as full species
(e.g. Huntley 1973c; Crawford-Cabral and Veríssimo 2005). Red Hartebeest used to
be widely distributed in southern Angola in the region between the Cunene and
Cubango rivers (Sokolowski 1903; Monard 1935; Hill and Carter 1941; Huntley
1973c; Crawford-Cabral and Veríssimo 2005). Red Hartebeest could be found in
only one protected area, Mupa NP (Huntley 1973c; Crawford-Cabral and Veríssimo
2005), and by the 1970s the Angolan population was already much reduced and
endangered (Huntley 1973c), and was feared extinct in the 1990s (Huntley and
Matos 1992). Recent surveys found no evidence of Red Hartebeest in Mupa
(Overton et al. 2017) and this taxon is possibly currently extinct in Angola.
Lichtenstein’s Hartebeest was found in eastern Angola where it was generally
uncommon to rare (Huntley 1973c; Crawford-Cabral and Veríssimo 2005). Most
records were obtained in Lundas and Moxico (Machado 1969), although some ear-
lier authors reported the species in Luiana along the Cuando River, even if not fully
supported (Statham 1926; Crawford-Cabral and Veríssimo 2005). Lichtenstein’s
Hartebeest was once present in Cameia NP, but their current status in Angola is
unknown and likely extinct.
Blue Wildebeest (Connochaetes taurinus taurinus) used to be widespread across
southern and eastern Angola, in Huíla, Cunene, Moxico and Cuando Cubango prov-
inces (Hill and Carter 1941; Newton da Silva 1970; Crawford-Cabral and Veríssimo
2005). It was once common in the protected areas of Bicuar and Cameia NPs, and
the region currently ascribed to Mavinga and to Luengue-Luiana NPs (Huntley
1973c; Crawford-Cabral and Veríssimo 2005). Wildebeest numbers in Angola must
have plummeted during the war (Huntley and Matos 1992), while recent surveys
and anecdotal records suggest that the species is likely extinct in the western areas
of their Angolan range, such as Bicuar NP (Overton et al. 2017). Nevertheless small
numbers have recently been recorded in the southeastern corner, both from camera-
trapping (Funston et al. 2017) and aerial counts (Chase and Schlossberg 2016).
Tribe Antilopini
Springbok (Antidorcas marsupialis) is the only species of gazelle sensu lato occur-
ring in Angola and southern Africa. Based on specimens collected south of Benguela,
and comparing these with Springbok from South Africa, Blaine (1922) claimed
enough differences to justify the description of a new species, Angolan Springbok
(A. angolensis). It was subsequently considered as one of three subspecies of spring-
bok, A. m. angolensis, and extending into Namibia (Ansell 1972; Hill and Carter
1941). However, the distinction among geographical boundaries and intergradation
has led to questioning the validity of these races (Skinner 2013). In Angola,
Springbok is strongly associated with the arid coastal belt, and present in the pro-
tected areas of Chimalavera, Namibe and Iona (Huntley 1973c, 1974; Crawford-
Cabral and Veríssimo 2005). Nevertheless, a few old records are also known from
the region of Naulila (Monard 1935; Galvão and Montês 1943–1945; Crawford-
Cabral and Veríssimo 2005). Currently it is still present along the coastal plain south
of Benguela, albeit in much reduced numbers (Vaz Pinto, unpublished data). Recent
aerial surveys in Iona NP have allowed population estimations, suggesting a decreas-
ing trend with an estimated 21% reduction in total numbers between 2003 and 2017
(Kolberg and Kilian 2003; van der Westhuizen et al. 2017).
Tribe Bovini
Buffalo (Syncerus caffer) is the sole representative of the Bovini in Africa.

Nevertheless its taxonomy remains controversial due to a marked geographical vari-
ation and the existence of intermediate forms, which has led to the recognition of
several species, subspecies or variants (Grubb 1972; Prins and Sinclair 2013). In
Angola at least two forms are known, the typical Cape Buffalo (S. c. caffer) of larger
body size, dark coloration and large hook-shaped horns, and the Forest Buffalo (S.
c. nanus) of smaller body size, reddish colour and smaller backward-pointing horns.
Notwithstanding, quite a lot of variation has been observed particularly among
Angolan forest buffalos, as specimens from northern Angola tend to be larger and
darker than those in Cabinda (Crawford-Cabral and Veríssimo 2005). Even though
Matschie (1906) has described an additional subspecies (S. c. mayi) based on a
specimen from Luanda, it was subsequently synonymised with S. c. nanus. Forest
Buffalo used to have a wide distribution across Cabinda and northern Angola,
including in protected areas such as Quiçama NP and Luando Strict Reserve
(Huntley 1973c; Crawford-Cabral and Veríssimo 2005). Cape Buffalo was mostly
present in the southeast, even though an additional small population was known
from Bicuar NP (Huntley 1973c) and a few other isolated pockets in Benguela,
Cuanza-Sul and Lunda provinces (Crawford-Cabral and Veríssimo 2005). The buf-
falo populations were severely reduced during the war (Huntley and Matos 1992),
but very small numbers of Forest Buffalo are still present in Quiçama NP (Groom
et al. 2018), and Cangandala NP and Luando Reserve (Vaz Pinto, unpublished data).
Cape Buffalo is not uncommon in Mucusso region and Luengue-Luiana NP
(Veríssimo 2008; Chase and Schlossberg 2016; Funston et al. 2017). On the other
hand, smaller and isolated populations of Cape Buffalo may have been extirpated,
such as that which used to occur in Bicuar NP (Overton et al. 2017).
Tribe Cephalophini
The tribe Cephalophini is represented by three genera and six species in Angola.
The genus Cephalophus corresponds to typical forest duikers, of which four species
are known for Angola: White-bellied Duiker (C. leucogaster), Bay Duiker (C.
378 P. Beja et al.
dorsalis), Black-fronted Duiker (C. nigrifrons), and Yellow-backed Duiker (C. silvi-
cultor). The White-bellied Duiker was reported from Cabinda based on a witness
account obtained in the 1970s (Brian Huntley, personal communication), and subse-
quently added to the Angolan list (Crawford-Cabral and Veríssimo 2005). The spe-
cies is known to occur in Maiombe forest across the border in Congo (Malbrant and
Maclatchy 1949; East 1999), but further evidence of its presence in Angola is still
lacking. Both Bay Duiker and Black-fronted Duiker have been recorded from moist
forest habitats in the northern half of the country, including Cabinda (Huntley
1973c; Huntley and Matos 1992; Crawford-Cabral and Veríssimo 2005). Although
no recent studies are available, the regular presence of both species in bushmeat
markets in northwestern Angola (Vaz Pinto, unpublished data) suggest they may
still be relatively common in spite of the poaching pressure. Yellow-backed Duiker
is the largest of duikers and is also widely, yet discontinuously, distributed across
the northern half of the country and including Cabinda (Machado 1969; Huntley
1973c; Crawford-Cabral and Veríssimo 2005). Unlike the previous species, Yellow-
backed Duiker is less dependent on moist forest habitats, being mostly an ecotone
species (Kingdon and Lahm 2013). In spite of the scarcity of records, it appears well
adapted to the riverine forests and thickets of central Angola, venturing into nearby
well-developed miombo woodlands (Vaz Pinto and Veríssimo 2016) and present
even on the Angolan highlands (Statham 1922; Hill and Carter 1941), possibly in
transition to Afromontane patches. Recent observations suggest the species to be
relatively common in Luando Strict Reserve, and venturing into the upper catch-
ments of the Okavango and Zambezi drainages (Vaz Pinto and Veríssimo 2016;
NGOWP 2018). Angolan Yellow-backed Duikers are assigned to the subspecies C.
s. ruficrista, of which the type locality given is Luanda (du Bocage 1869).
Blue Duiker (Philantomba monticola) is especially common along the escarp-
ment and in various types of forests and thickets along the coastal plain north of 15°
latitude, and including Cabinda (Crawford-Cabral and Veríssimo 2005). On the pla-
teau it is present north of 13° latitude but it is here less common, patchily distrib-
uted, and associated with riverine forests (Crawford-Cabral and Veríssimo 2005;
Vaz Pinto, unpublished data). Blue Duiker is still abundant in Quiçama NP (Groom
et al. 2018), while present in Cangandala NP (Vaz Pinto unpublished data), and has
recently been photographed on the upper catchments of the Cuito River (NGOWP
2018). Three subspecies have been tentatively ascribed to Blue Duiker from Angola,
but their validity remains unclear (Ansell 1972; Crawford-Cabral and Veríssimo
2005; Hart and Kingdon 2013). Under this classification, the population in Cabinda
is assigned to P. m. congicus, and those on the plateau to P. m. defriesi, while the
blue duikers from the escarpment and western Angola correspond to an endemic
race, P. m. anchietae (Ansell 1972; Crawford-Cabral and Veríssimo 2005; Hart and
Kingdon 2013).
Grey or Common Duiker (Sylvicapra grimmia) is likely the most widespread and
common of all Angolan antelopes (Statham 1922; Crawford-Cabral and Veríssimo
2005). It probably still occurs in all Angolan protected areas, except Iona and
Maiombe NPs, and throughout the country (Crawford-Cabral and Veríssimo 2005;
Veríssimo 2008; Funston et al. 2017; Overton et al. 2017; Groom et al. 2018;
NGOWP 2018; Vaz Pinto, unpublished data). Although many subspecies of Grey
Duiker have been suggested, the continuous distribution of the species in sub-
Saharan Africa and the existence of local intergrading variants prevents clear delin-
eation of boundaries (Ansell 1972; Wilson 2013). In Angola most are assigned to S.
g. splendidula, and the race is thought to intergrade with S. g. steinhardti in south-
western Angola (Hill and Carter 1941; Crawford-Cabral and Veríssimo 2005;
Wilson 2013).
Tribe Hippotragini
The tribe Hippotragini contains seven extant species, of which three can be found in
Angola, including two representatives of the genus Hippotragus and one of Oryx.
The Roan Antelope (Hippotragus equinus) is the most common and widely distrib-
uted large antelope in Angola, historically being absent only from Cabinda and the
arid southwest (Huntley 1973c; Newton da Silva 1970; Crawford-Cabral and
Veríssimo 2005). It used to be present in all existing protected areas except Iona NP
and Namibe Partial Reserve (Crawford-Cabral and Veríssimo 2005), and was once
considered abundant in Quiçama and Bicuar NPs (Huntley 1973c; Huntley and
Matos 1992). As a result of the civil war the species has been extirpated from
Quiçama NP (Huntley and Matos 1992; Groom et al. 2018; Vaz Pinto, unpublished
data), but it remains relatively common in Bicuar NP (Overton et al. 2017) and
Luando Strict Nature Reserve (Vaz Pinto, unpublished data), while small numbers
still linger in Mupa (Overton et al. 2017) and Cangandala NPs (Vaz Pinto et al.
2016). In addition the species has also been confirmed recently in various surveys
conducted across central and eastern Angola (Veríssimo 2008; Chase and
Schlossberg 2016; Funston et al. 2017; NGOWP 2018), and likely remains wide-
spread even if in reduced numbers throughout most of the country except on the
coastal plain. Roan intraspecific taxonomy is still unresolved, but the Angolan race
is usually ascribed to the Zambezian region subspecies H. e. cottoni (Ansell 1972;
Chardonnet and Crosmary 2013; Vaz Pinto 2018).
Sable Antelope (Hippotragus niger) had a highly fragmented distribution in
Angola, with three disjunct populations corresponding to three different subspecies
(Crawford-Cabral and Veríssimo 2005; Estes 2013; Vaz Pinto 2018, 2019). Giant
Sable (H. n. variani) is an endemic and critically endangered taxon, confined to the
Cuanza drainage, and being the most famous Angolan mammal is dealt with in a
separate dedicated chapter (see Vaz Pinto 2019). The occurrence of Kirk’s Sable (H.
n. kirkii) in eastern Angola was confirmed by a few scattered records from Cazombo,
east of the Zambezi River, and in the Lundas, on the western banks of the Cassai
River (Huntley 1973c; Crawford-Cabral and Veríssimo 2005). No records for this
taxon have been obtained in Angola for over 40 years and we are unaware of any
witness reports, thus suggesting the possibility of local extinction. The typical race
H. n. niger is known from southeastern Angola to the east of the Cuito river (Hill
and Carter 1941; Huntley 1973c; Crawford-Cabral and Veríssimo 2005; Vaz Pinto
2018), a region that broadly corresponds to the newly proclaimed Mavinga and
Luengue-Luiana NPs. Recent surveys have confirmed typical sable to be still rela-
tively common in the region and clearly outnumbering congeneric roan antelope
(Veríssimo 2008; Chase and Schlossberg 2016; Funston et al. 2017), and very
380 P. Beja et al.
recently a dispersing male has been recorded as far north as southern Moxico
(Kerllen Costa, personal communication)
Gemsbok (Oryx gazelle) in Angola is mostly associated with the Namib desert in
the southwestern corner, but its distribution used to extend along the semi-arid
coastal plain as far north as near Benguela (Blaine 1922; Statham 1922; Hill and
Carter 1941; Crawford-Cabral and Veríssimo 2005), and in Cunene province at least
as far inland as Cuamato and Chimporo (Monard 1935; Crawford-Cabral and
Veríssimo 2005). Specimens from southwestern Angola have led to the description
of a local endemic subspecies of gemsbok O. g. blainei based on facial mask differ-
ences (Blaine 1922; Hill and Carter 1941; Newton da Silva 1970). However the
species is currently assumed to be monotypic (Knight 2013). Gemsbok may have
been extirpated from most of its Angolan range in the second half of the twentieth
century (Newton da Silva 1970; Crawford-Cabral and Veríssimo 2005), while
remaining abundant in Iona NP (Huntley 1973c). The numbers were then likely
much reduced during the civil war (Huntley and Matos 1992), and although they
may still be relatively common in Iona, recent surveys suggest the general trend has
remained negative (Kolberg and Kilian 2003; van der Westhuizen et al. 2017).
Tribe Madoquini
The sole Madoquini in Angola is Kirk’s Dik-dik (Madoqua kirkii). The species was
first collected in Angola by Anchieta in 1878, and specimens obtained near Lobito
led to the description of a new subspecies, M. k. variani, which was later syn-
onymised with M. k. damarensis (Drake-Brockman 1909, 1930; Hill and Carter
1941; Newton da Silva 1970; Kingswood and Kumamoto 1997). Angolan dik-diks
are part of a southwest African population that extends well into Namibia, ascribed
to the subspecies M. k. damarensis, although the huge geographical gap that sepa-
rates these from the populations in the horn of Africa added by some morphological
characters and genetic evidence, suggest they should be best treated as full species
(Kumamoto et al. 1994; Zhang and Ryder 1995; Brotherton 2013). In Angola dik-
diks are associated with the semi-arid environments and particularly with Mopane
(Colophospermum mopane) woodlands, but also extending into the southern pla-
teau west of the Cunene River (Crawford-Cabral and Veríssimo 2005). The species
is well represented in protected areas such as Chimalavera and Namibe Reserves,
and Iona and Bicuar NPs (Huntley 1973c; Crawford-Cabral and Veríssimo 2005;
Vaz Pinto, unpublished data).
Tribe Oreotragini
Klipspringer (Oreotragus oreotragus) is the only representative of the Oreotragini,

but its taxonomy is one of the most hotly debated within African bovids, both in
terms of the relationships with other clades and among various populations. Ansell
(1972) recognised 11 subspecies, while Groves and Grubb (2011) distinguished up
to 20 taxa and elevated them to full species, but the latter still lacks molecular sup-
port and the monospecific classification is still the most widely accepted. Angolan
klipspringers have been attributed to the subspecies O. o. tyleri described from a

specimen obtained in Equimina, Benguela (Hinton 1921). They likely form part of
a metapopulation that extends into Namibia (Crawford-Cabral and Veríssimo 2005;
Roberts 2013), where a second subspecies O. o. cunenensis (Zukowsky 1924)
described near Ruacana falls has been synonymised with the former (Hill and Carter
1941; Ansell 1972). Klipspringer occurs in Angola in rocky mountainous habitats,
particularly in southern Angola and along the escarpment, with the northernmost
population present at Pungo Andongo (Crawford-Cabral and Veríssimo 2005), and
the easternmost population was reported from the region of Cassinga (Monard
1935; Newton da Silva 1970; Crawford-Cabral and Veríssimo 2005). The species
used to be relatively common in Iona NP, and Chimalavera and Namibe Reserves
(Huntley 1973c; Juste and Carballo 1992; Crawford-Cabral and Veríssimo 2005),
and in spite of absence of recent data it is still often observed in southwestern
Angola and along the southern escarpment (Vaz Pinto, unpublished data).
Tribe Ourebiini
This tribe is monospecific, comprising only the Oribi (Ourebia ourebi), which has a
wide distribution in Africa and across southcentral and eastern Angola (Crawford-
Cabral and Veríssimo 2005; Brashares and Arcese 2013). Up to 13 subspecies of
Oribi have been described, but their validity remains problematic (Brashares and
Arcese 2013). Two subspecies were described from specimens collected in Angola,
namely O. o. rutila from Luando Reserve (Statham 1922), and O. o. leucopus
(Monard 1930). The latter was subsequently synonymised with the former, and con-
sidered to extend into Caprivi, Botswana and west Zambia (Ansell 1972; Crawford-
Cabral and Veríssimo 2005; Brashares and Arcese 2013). In Angola the species
occurs in open savanna habitats above 1000 m, and used to be present in protected
areas such as Luando Reserve, Bicuar, Mupa and Cameia NPs, and across south-
eastern Angola (Huntley 1973c; Crawford-Cabral and Veríssimo 2005). Two refer-
ences reporting the presence of oribi on the southern coastal plain (Statham 1922;
Fenykovi 1953) are dubious and may result from misidentification of Steenbok
(Crawford-Cabral and Veríssimo 2005). Recent surveys have failed to record the
species in Mupa and Bicuar (Overton et al. 2017) and in Mucusso (Veríssimo 2008).
Anecdotal evidence suggestes its presence in the upper catchments of the Okavango
(NGOWP 2018), and they were recorded by camera traps in Luengue-Luiana NP
(Funston et al. 2017). Small numbers are still present and have been recently
observed and photographed in Luando Reserve and Cameia NP (Vaz Pinto, unpub-
lished data).
Tribe Raphicerini
Only one species of Raphicerini has been confirmed in Angola, the Steenbok
(Raphicerus campestris), occurring south of 12° latitude, being most common in
semi-arid habitats in the coastal plain but also present inland from Huíla to Cuando
Cubango provinces (Crawford-Cabral and Veríssimo 2005). The species used to be
382 P. Beja et al.
common and is still present in Iona, Bicuar and Mupa NPs, and in the recently des-
ignated Luengue-Luiana and Mavinga NPs (Huntley 1973c; Veríssimo 2008;
Funston et al. 2017; Overton et al. 2017). A remarkable record was one specimen
collected south of Namibe at Lagoa S. João do Sul, with very long hooves, suggest-
ing an isolated population and local adaptation to muddy terrain (Simões and
Crawford-Cabral 1988; Crawford-Cabral and Veríssimo 2005). Recent surveys have
extended the species distribution northwards into the upper catchment of the
Okavango (NGOWP 2018), adding to records from the Cuito River source and from
near Cuemba (Vaz Pinto, unpublished data). Interestingly, witness accounts reported
unusual behaviour displayed by the latter Steenbok, also suggesting isolation and
local adaptation (Vaz Pinto, unpublished data). Several subspecies have been pro-
posed and Ansell (1972) recognised eight races, but these remain unclear and often
only two are accepted, with the nominate subspecies R. c. campestris being assigned
to all populations in southern Africa (du Toit 2013). A congeneric species, Sharpe’s
Grysbok (Raphicerus sharpei), has never been recorded in Angola but it may well
be present in the regions of Cazombo or Luiana, as it is known to occur in western
Zambia and east Caprivi, very close to the Angolan border (Ansell 1972; Hoffman
and Wilson 2013).
Tribe Reduncini
The tribe is represented in Angola by the genera Redunca and Kobus, comprising
four species in total. Southern Reedbuck (Redunca arudinum) had a wide, albeit
discontinuous distribution associated with grassy patches near drainage lines, and
was present throughout the country except in Cabinda and the arid southwest
(Newton da Silva 1970; Huntley 1973c; Crawford-Cabral and Veríssimo 2005). The
species was especially common in protected areas such as Quiçama, Cangandala,
Cameia and Bicuar NPs, Luando Reserve, and the southeast regions (Huntley
1973c, 1974; Crawford-Cabral and Veríssimo 2005). Although a recent camera-trap
survey has failed to record the species in Bicuar and Mupa (Overton et al. 2017),
they may still be present, and have been recorded on surveys conducted in Mucusso,
Luengue-Luiana and Mavinga (Veríssimo 2008; Chase and Schlossberg 2016;
Funston et al. 2017). Even if in much reduced numbers, Reedbuck is also still found
in Quiçama NP (Groom et al. 2018), and in Cangandala NP and Luando Strict
Reserve (Vaz Pinto, unpublished data). The intraspecific taxonomy of Reedbuck is
still unresolved, but often two subspecies are recognised and separated by the
Zambezi River (Ansell 1972; Kingdon and Hoffmann 2013), with the Angolan pop-
ulations corresponding to the typical race R. a. arundinum.
The genus Kobus comprises the remaining three species of Reduncini present in
Angola. Puku (Kobus vardonii), is a relatively rare antelope that used to have its
westernmost populations in Angola (Jenkins 2013). The species was mostly
recorded in northeastern Angola, and a lot of what is known is due to the studies of
Machado (1969). Puku used to occur, albeit in low numbers, in Luando Strict
Reserve (Statham 1922; Huntley 1973c), possibly corresponding to an isolated and

westernmost subpopulation, but appears now to be absent (Vaz Pinto, unpublished
data). An old record reported to have been obtained in Huíla province (du Bocage
1902), is generally dismissed as mistaken (Crawford-Cabral and Veríssimo 2005),
and the southernmost record was obtained by Wilhelm Trense in Luiana (Crawford-
Cabral and Veríssimo 2005). The species has not been recorded in Angola in over
40 years and it is thus possibly extinct. Southern Lechwe (Kobus leche) used to be
widely distributed along river drainages in central and eastern Angola, only margin-
ally overlapping with Puku in Luando Reserve and possibly Luiana (Crawford-
Cabral and Veríssimo 2005). Angolan Lechwe was suggested by Sokolowski (1903)
to constitute a separate species Adenota (=Kobus) amboellensis, but it was subse-
quently synonymised with the typical race also known as Red Lechwe K. l. lechwe.
Red Lechwe seems to have been extirpated from a large portion of their former
range in Angola, when only a relic population survived in Luando Reserve (Vaz
Pinto, unpublished data), but larger populations have recently been recorded in the
upper catchments of the Okavango (NGOWP 2018), and in the Luengue-Luiana and
Mavinga NPs (Veríssimo 2008; Chase and Schlossberg 2016; Funston et al. 2017).
Waterbuck (Kobus ellipsiprymnus) comprises two very distinct subspecies that
often are considered to warrant full species status, and both have been recorded in
Angola. Common Waterbuck (K. e. ellipsiprymnus) is very localised in the country,
and only known to occur along the lower Cuando River in Luengue-Luiana NP
(Huntley 1973c; Crawford-Cabral and Veríssimo 2005). Current status is unknown,
as various surveys failed to record the species (Veríssimo 2008; Chase and
Schlossberg 2016), though a recent survey reported finding carcasses (Funston et al.
2017). The other taxon present in Angola is currently ascribed to Defassa Waterbuck
(K. e. defassa), as previous attempts to recognise geographical variants attributed to
Angolan waterbuck such as K. e. penricei (Hill and Carter 1941; Ansell 1972) have
been abandoned in favour of synonymy with K. e. defassa. Defassa Waterbuck used
to be widely distributed in Angola but were usually uncommon (Huntley 1973c;
Crawford-Cabral and Veríssimo 2005), and in protected areas were only represented
in Cangandala, Luando and Bicuar NPs (Statham 1922; Huntley 1973c). A recent
survey failed to record the species in Bicuar NP where it is feared extinct (Overton
et al. 2017). Nevertheless small numbers are still present across northeasten Angola,
including Cangandala and Luando (Vaz Pinto, unpublished data).
Tribe Tragelaphini
Four species of Tragelaphini are known from Angola (Crawford-Cabral and

Veríssimo 2005), all currently lumped within the genus Tragelaphus: Kudu (T.
strepsiceros), Bushbuck (T. scriptus), Sitatunga (T. spekii) and Common Eland (T.
oryx). Kudus used to be widely distributed in southern Angola, and along the semi-
arid coastal plain (Huntley 1973c; Crawford-Cabral and Veríssimo 2005). Although
reported previously (Huntley 1972c, 1973c), recent surveys have not recorded Kudu
in Iona NP (Kolberg and Kilian 2003; van der Westhuizen et al. 2017), but they are
384 P. Beja et al.
still relatively common in Bicuar and Mupa NPs (Overton et al. 2017) and small
numbers have been confirmed in the southeastern Cuando Cubango (Veríssimo
2008; Funston et al. 2017). The distinction among Kudu subspecies has remained
dubious due to variability of morphological characteristics and intergradation
(Owen-Smith 2013).
Bushbuck in Angola have been ascribed to the subspecies T. scriptus ornatus
(Crawford-Cabral and Veríssimo 2005), but the species taxonomy is highly prob-
lematic. Genetic studies have revealed two highly divergent and non-monophyletic
mitochondrial clades which provided support for two species: Tragelaphus scriptus
and T. sylvaticus (Moodley and Bruford 2007; Moodley et al. 2009). This however
still lacks confirmation with nuclear and morphological data (Hassanin et al. 2012).
Angolan animals could be important to disentangle the phylogenetic relationships
within Bushbuck, as they are relatively common and widespread across central and
northern Angola, where the boundary between both clades may be found. Bushbuck
has remained common in northern Angola, including in protected areas such as
Quiçama, Cangandala and Luando (Huntley 1973c; Groom et al. 2018; Vaz Pinto,
unpublished data), and features prominently in bushmeat markets (Bersacola et al.
2014; Groom et al. 2018; Vaz Pinto, unpublished data). It was generally uncommon
in the southern regions, and recent surveys have failed to detect the species in Mupa
and Bicuar NPs (Overton et al. 2017). Small numbers were recorded in the Luengue-
Luiana region (Veríssimo 2008; Funston et al. 2017).
Sitatunga had a relatively wide distribution in Angola, following the main river
systems but excluding the coastal plain (Huntley 1973c; Crawford-Cabral and
Veríssimo 2005). The species is still present in protected areas such as Cangandala
and Luando Reserve (Vaz Pinto, unpublished data) or in Luengue-Luiana (Veríssimo
2008; Funston et al. 2017). Additional records have been obtained from northern
Malanje and Lunda-Norte (Huntley and Francisco 2015; Vaz Pinto, unpublished
data). Traditionally two subspecies of Sitatunga have been recognised in Angola, T.
s. gratus in northern and central Angola, and T. s. selousi within the Zambezi and
Okavango drainages (Crawford-Cabral and Veríssimo 2005).
Common Eland used to be relatively widespread in Angola, except in the forest
biomes of the northwest (Crawford-Cabral and Veríssimo 2005). It was once abun-
dant in Quiçama NP and in the hunting concessions of southeastern Angola (Newton
da Silva 1970; Huntley 1973c; Crawford-Cabral and Veríssimo 2005), while present
in low numbers in Iona, Bicuar, Mupa and Cangandala NPs and Luando Reserve
(Huntley 1973c; Crawford-Cabral and Veríssimo 2005). The species is now extinct
in Quiçama NP (Groom et al. 2018) and Cangandala NP (Vaz Pinto, unpublished
data), and has not been recorded in mammal surveys conducted in Iona and Mupa
NPs (Kolberg and Kilian 2003; Overton et al. 2017; van der Westhuizen et al. 2017).
A small population may still linger in Luando (Vaz Pinto, unpublished data) and
Bicuar NP (Overton et al. 2017), and relatively larger numbers in Luengue-Luiana
NP (Veríssimo 2008; Funston et al. 2017). Angolan Eland have often been attributed
to the subspecies T. oryx livingstoni (Crawford-Cabral and Veríssimo 2005; Thouless
2013), but intergrading with the nominate race T. o. oryx in southern Angola (Ansell
1972; Crawford-Cabral and Veríssimo 2005; Thouless 2013). It has been argued
that Eland of the semi-arid biomes along the coastal plain could be ascribed to the
nominate race, while the remaining could be ascribed to T. o. livingstoni (Crawford-
Cabral and Veríssimo 2005).
Family Giraffidae
Angolan giraffes used to be ascribed to Giraffa camelopardalis angolensis (Lydekker

1904; Ciofolo and Le Pendu 2013), although recent molecular studies have pro-
posed a four species classification, under which they would correspond to G. giraffa
angolensis (Fennessy et al. 2016). Two disjunct populations were known in Angola,
one in Cunene and southern Huíla provinces, and the other in Cuando Cubango east
of Cuito River (Newton da Silva 1970; Crawford-Cabral and Veríssimo 2005). Once
the symbol of Mupa NP, in the early 1970s the giraffe was already on the brink of
extinction (Huntley 1973c), and was presumed extinct in Angola by the 1990s
(Huntley and Matos 1992; Juste and Carballo 1992). Recent surveys are consistent
with the species extinction in Mupa (Overton et al. 2017), but records obtained from
spoor and from aerial surveys have demonstrated the persistence of pocket popula-
tions in the areas corresponding to the current Luengue-Luiana NP (Veríssimo
2008; Chase and Schlossberg 2016).
Family Hippopotamidae
The sole hippopotamus species in Angola is the Common Hippopotamus

(Hippopotamus amphibius). This species is relatively widespread, and used to be
found across most large rivers and drainage basins in the country (Newton da Silva
1970; Crawford-Cabral and Veríssimo 2005). The hippopotamus in Angola have
been ascribed to H. a. constrictus, although subspecific taxonomy remains contro-
versial for this species and is often ignored (Klingel 2013). Although once common,
the hippopotamus has suffered from human direct persecution. In the 1970s it had
become rare in the country and reduced to small pockets in larger rivers, and this
situation may have deteriorated since (Huntley 1973c; Huntley and Matos 1992;
Juste and Carballo 1992). Although no study has addressed specifically the hippo-
potamus populations in Angola, recent general surveys have reported its presence in
isolated pockets along rivers such as Cuanza, Queve and Luando (Vaz Pinto, unpub-
lished data), and in the middle to lower sections of various rivers within the
Okavango and Zambezi basins (Veríssimo 2008; Chase and Schlossberg 2016;
Funston et al. 2017; NGOWP 2018). Small populations are still present in Quiçama
NP and Luando Strict Reserve (Groom et al. 2018; Vaz Pinto, unpublished data), but
based on recent surveys they appear to have disappeared from the southern Cunene
system, namely within Iona, Mupa and Bicuar NPs (Kolberg and Kilian 2003;
Overton et al. 2017; van der Westhuizen et al. 2017).
386 P. Beja et al.
Family Suidae
Three species of wild suids occur in Angola, but no studies have yet focused on
Angolan pigs. The Bushpig (Potamochoerus larvatus) is widespread, and possibly
only naturally absent from Cabinda and the arid southwest (Crawford-Cabral and
Veríssimo 2005). Data on Bushpig distribution have been reported by several
authors (e.g. Statham 1922; Monard 1935; Hill and Carter 1941; Newton da Silva
1970; Huntley 1973c), but mostly refer to scattered and localised records. The spe-
cies has recently been recorded in various surveys in protected areas (Veríssimo
2008; Chase and Schlossberg 2016; Funston et al. 2017; Overton et al. 2017; Groom
et al. 2018; NGOWP 2018; Vaz Pinto, unpublished data). Two subspecies have been
recognised in the past in Angola, P. l. johnstoni and P. l. cottoni, but the species
displays a large amount of individual variation and both have been subsequently
synonymised with P. l. koiropotamus (Grubb 1993; Crawford-Cabral and Veríssimo
2005; Seydack 2013). Formerly treated as conspecific with bushpigs, the Red River
Hog (Potomochoerus porcus) in Angola is confirmed only in Cabinda, and little is
known about its possible occurrence in Lunda’s provinces or possible intergradation
with the previous species (Crawford-Cabral and Veríssimo 2005; Leus and
Vercammen 2013; Seydack 2013). Common Warthog (Phacochoerus africanus),
were relatively widespread in Angola, and used to be locally common, even if irreg-
ularly distributed (Crawford-Cabral and Veríssimo 2005; Huntley 1973c). Recently,
it has been regularly recorded in various protected areas (e.g. Funston et al. 2017;
Overton et al. 2017; Groom et al. 2018; NGOWP 2018; Vaz Pinto, unpublished
data). Angolan warthogs may correspond to the subspecies P. a. sundervalli
(Cumming 2013), but geographic variation and transition among different forms
remains unresolved. Some early authors (Statham 1922; Monard 1935; Varian
1953) have reported the existence of a giant pig in central Angola, based on local
witness accounts, which they tentatively ascribed to the giant forest hog (Hylochoerus
meinertzhageni), but these reports are generally dismissed, and were likely based on
tales or misidentification of bushpigs (Newton da Silva 1970; Crawford-Cabral and
Veríssimo 2005).
Family Tragulidae
Water Chevrotain (Hyemoschus aquaticus) was first confirmed for Angola in

Cabinda (Huntley 1973c), although it was also reported to occur in Lunda-Norte
along the Cassai River (Crawford-Cabral and Veríssimo 2005). More recent witness
accounts suggest the occurrence of the species in forest streams and nearby flooded
areas in Uíge Province (Michael Mills, personal communication). It is possible that
the Water Chevrotain has a wider distribution across northern Angola in forest habi-
tats, but the elusive nature of the species and lack of studies prevent any conclusions
at this stage.
Chiroptera (Bats)
With 73 species recorded, bats (Chiroptera) are the second most speciose order of
mammals in Angola after the rodents. This represents about one third of the bat spe-
cies known to occur in mainland Africa (Happold and Happold 2013), and almost
two thirds of the species reported for the southern Africa region (Monadjem et al.
2010a). Angola is therefore a particulary rich country for bats in southern Africa,
compared with other species-rich countries such as Mozambique (67 species),
Zambia (65), Malawi (62) and Zimbabwe (62) (Monadjem et al. 2010b). Considering
that Angola is one of the least known southern African countries with respect to
bats, more species are likely to be found as new surveys are carried out throughout
the country. For instance, three of the 73 species reported here have been found
recently during the course of just a few weeks of fieldwork (Taylor et al. 2018c).
Even new (and possibly endemic) species may be found, particularly on the western
escarpment region and Afromontane forests that are high in bird endemism, and in
species with restricted and fragmented distributions (Mills et al. 2011, 2013).
The inventory of Angolan bats started in the nineteenth century (e.g. Peters 1870;
du Bocage 1889a, 1898; Seabra 1898a, b). A substantial number of records was
added later by various authors including Thomas (1904), Hill and Carter (1941),
Sanborn (1950), Hayman (1963) and finally by Crawford-Cabral (1986), who also
reviewed all the information then available. Little research on bats has been done in
Angola in the past few decades, and to our knowledge no studies of bat ecology
have been carried out in the country. There are important taxonomic issues to be
resolved in most families, and the existing records for the majority of species are old
and scarce, so the descriptions of bat ranges in the following sections should be
considered as provisional. Also, no recent bat surveys have been conducted in any
of the remaining Afromontane forests and adjacent grasslands along the escarp-
ment, and this must remain a critical zone for future surveys.
Family Emballonuridae
There are three species of Emballonuridae in Angola (Monadjem et al. 2010a;
Happold and Happold 2013). The Mauritian Tomb Bat (Taphozous mauritianus)
occurs widely in a variety of habitats and often roosts in buildings inhabited by
people, but in Angola it is only known from scattered records. African Sheath-tailed
Bat (Coleura afra) is a rare species in southern Africa, which in Angola is known
from just two records in the coastal area near Benguela, but this species may have
been overlooked. Giant Pouched Bat (Saccolaimus peli) is a species mainly found
in the Rainforest Belt Zone of Africa, with an isolated record in eastern Angola, but
it may have been overlooked in much of the northern region of the country.
388 P. Beja et al.
Family Hipposideridae
There are five species of Hipposideridae recorded in Angola (Monadjem et al.
2010a; Happold and Happold 2013). One of these is the African Trident Bat
(Triaenops afer), which is now included in the separate family Rhinonycteridae
(Foley et al. 2015). This is a tree-roosting species with a widespread, albeit patchy
distribution, with the isolated Congolese population encompassing Cabinda and the
extreme northwest of the country corresponding to the subspecies T. a. majusculus
Allen et Brosset, 1968, though this is not widely recognised (Benda and Vallo 2009).
The Sundevall’s Leaf-nosed Bat (Hipposideros caffer) is also wide ranging, but
represents a complex of species and requires urgent taxonomic revision (Vallo et al.
2008). Most existing records are from western Angola, where the populations pre-
sumably belong to the form H. caffer angolensis Seabra, 1898 (which is currently
placed in the species H. ruber - see below), but the species also seems to be present
in the extreme east of the country, representing an edge of a much larger range in
Central and East Africa (Kock et al. 2008). The Giant Leaf-nosed Bat (Macronycteris
gigas) is mostly a lowland forest and moist savanna species, which in Angola occurs
in Cabinda and penetrates southwards along the northern section of the escarpment
(Monadjem et al. 2010a). The type locality of this species is in Benguela, in semi-
arid savanna that is atypical regarding its habitat requirements. This species is dif-
ficult to separate from Striped Leaf-nosed Bat (M. vittatus), making it problematic
to ascertain its actual distribution. Noack’s Leaf-nosed Bat (H. ruber) has similar
habitat affinities but may be widespread in the country. However, due to previous
confusion of this species with H. caffer means that the distribution of H. ruber
remains mostly unknown in Angola, which itself represents a complex of multiple
species (Monadjem et al. 2013a). Finally, Striped Leaf-nosed Bat is a species form-
ing very large cave roosting colonies, which is judged to occur mainly in southern
Angola.
Family Miniopteridae
The Natal Clinging Bat (Miniopterus natalensis) is the only Miniopteridae judged
at present to occur in Angola (Monadjem et al. 2010a; Happold and Happold 2013).
However, this is a species distributed mostly in East and southern Africa, with a
handful of records in western and southern Angola (e.g., Grant and Ferguson 2018;
MHNG 2018). The population of Namibia and Angola appears to be largely isolated
from eastern populations, but they do not appear to be phylogenetically distinct
(Monadjem et al. 2013b). Some specimens collected in 1954 in the Huíla province
have been identified as Black Clinging Bat (Miniopterus fraterculus), but this is a
great distance from the known distribution of the species in the eastern parts of
South Africa, suggesting that they are probably misidentifications (Monadjem et al.
2013b). The Greater Long-fingered Bat (Miniopterus inflatus) may occur in Angola
and has probably been overlooked there (Monadjem et al. 2010a).
Family Molossidae
The Molossidae is represented by ten species in Angola. Many of these are known
from only a few and scattered records in the country (Monadjem et al. 2010a), mak-
ing it difficult to recognise distribution patterns. The White-bellied Free-tailed Bat
(Mops niveiventer) seems to be one of the most widespread species. It is mostly
associated with mature miombo woodland and occurs widely in Central Africa. The
similar Angolan Free-tailed Bat (M. condylurus) is widespread in sub-Saharan
Africa, but its distribution in Angola is poorly known. Some Molossidae seem to
have rather isolated populations in Angola (Monadjem et al. 2010a), which may
prove to be phylogenetically distinctive. Ansorge’s Free-tailed Bat (Chaerephon
ansorgei) is a species of dry woodland savanna, with a restricted population in west-
ern Angola that is isolated from the remaining known range of the species in the
eastern side of southern Africa. Little Free-tailed Bat (C. pumilus) occurs widely in
sub-Saharan Africa and is widespread and abundant in eastern parts of southern
Africa, but in the west seems to be restricted to an isolated population in northwest-
ern Angola and neighbouring DRC. Pale Free-tailed Bat (C. chapini) is sparsely
distributed in southern Africa, occurring mainly in northern Botswana and north-
eastern Zimbabwe and Zambia, with records in northern Namibia, western Angola
and DRC representing an isolated population.
The other Molossidae occur marginally or have only scattered records in Angola,
which may reflect environmental constraints but may also be due to poor survey
efforts. Roberts’s Flat-headed Bat (Sauromys petrophilus) is an arid zone specialist
for which there is an old record from Moçamedes (Crawford-Cabral 1986). Egyptian
Free-tailed Bat (Tadarida aegyptiaca) is widespread in southern Africa, but it occurs
only marginally in the south of Angola. The range of Midas Free-tailed Bat (M.
midas) also seems to reach the south of Angola, where it was tentatively detected
accoustically in a recent survey (Taylor et al. 2018c). In southern Africa the Nigerian
Free-tailed Bat (C. nigeriae) occurs in northwestern Namibia, northern Botswana,
Zimbabwe, Zambia and marginally in Angola and the DRC, though modelling of
the environmental niche suggests that the species may have been overlooked in the
country (Monadjem et al. 2010a). The Large-eared Free-tailed Bat (Otomops mar-
tiensseni) has a localised distribution in southern Africa, with scattered records from
Angola, Zimbabwe, Zambia, Malawi and the DRC.
Family Nycteridae
All but one of the six Nycteridae species recorded in Angola have distributions cen-
tered in Central and/or Western Africa and the country is at the southern edge of
their distribution (Monadjem et al. 2010a; Happold and Happold 2013). Dwarf Slit-
faced Bat (Nycteris nana), Intermediate Slit-faced Bat (N. intermedia), and Bate’s
Slit-faced Bat (N. arge) are mostly associated with lowland rainforests with very
few records in Angola, where they are probably restricted to the northern regions.
Large-eared Slit-faced Bat (N. macrotis) is thought to prefer savannas, and although
390 P. Beja et al.
the known records are all in northern Angola its range in Zambia and Botswana sug-
gests that it is also present further south. Hairy Slit-faced Bat (N. hispida) is a spe-
cies widespread in sub-Saharan Africa that uses a variety of habitats, which occurs
throughout central and northern Angola. Egyptian Slit-faced Bat (Nycteris theba-
ica) is a savanna species with wide habitat tolerance and very widespread in south-
ern Africa, though most records in Angola were obtained in the southwest
(Monadjem et al. 2010a).
Family Pteropodidae
A total of 15 species of pteropid fruit bats has been reported in Angola (Monadjem
et al. 2010a; Happold and Happold 2013), although the actual number of species
present is likely to be greater than this. Further work may reveal the presence of
additional species, particularly in the poorly surveyed Maiombe rainforest and
savanna-forest mosaics of Cabinda. It is interesting to note that several of the cur-
rently recognised pteropid species have a type locality in the country. This is the
case in the Angolan Soft-furred Fruit Bat (Myonycteris angolensis), Anchieta’s
Fruit Bat (Plerotes anchietae), Dobson’s Epauletted Fruit Bat (Epomops dobsonii),
Lesser Angolan Epauletted Fruit Bat (Epomophorus grandis), Hayman’s Dwarf
Epauletted Fruit Bat (Micropteropus intermedius), and Angolan Epauletted Fruit
Bat (E. angolensis). The latter species is a near endemic, present only in Angola and
northern Namibia, while Hayman’s Dwarf and Lesser Angolan Epauletted Fruit
Bats are only known from Angola and Congo.
The information available on the range of pteropids in Angola is insufficient to
make a definitive identification of distribution patterns, but some can be provision-
ally suggested. Overall, species diversity declines from north to south, because most
species of pteropids are dependent on well-wooded habitats and abundant fruit
resources. In fact, a few rainforest species that have the centre of their range in
Congo are known only in Cabinda and in some of the large pockets of moist forests
in the northern provinces. Species displaying this pattern include the Woermann’s
Long-tongued Fruit Bat (Megaloglossus woermanni), Franquet’s Epauletted Fruit
Bat (E. franqueti), Angolan Soft-furred Fruit Bat, Hammer-headed Fruit Bat
(Hypsignathus monstrosus), and Little Collared Fruit Bat (Myonycteris torquata).
Existing data suggest that at least some of these species extend their range south-
ward along the narrow band of forests of the Angolan Escarpment. Other species
associated with moist tropical forests and savannas occur more widely, extending
their range into northern Angola, as it is the case with Little Collared Fruit Bat and
Peter’s Dwarf Epauletted Fruit Bat (Micropteropus pusillus).
The most abundant pteropid in the species-poor south of Angola seems to be the
Angolan Epauletted Fruit Bat. It has a broad latitudinal range, although all the
records are in the western half of the country. In the drier regions it may be mostly
dependent on riverine woodlands. Peters’s Epauletted Fruit Bat (E. crypturus) has
been captured in one locality in the south of Angola but its range in the neighbour-
ing countries suggests that it may have a broad distribution in the south and east of
Angola, where only limited surveys have been conducted (Crawford-Cabral 1986).
Wahlberg’s Epauletted Fruit Bat (E. wahlbergi) and Straw-coloured Fruit Bat
(Eidolon helvum) have not been recorded in the drier areas of southern Angola, but
are present in the rest of the country. The latter occurs in Namibia and is known to
make long migrations tracking fruiting patterns, so it may also occur in much of
southern Angola, albeit only seasonally. Two species are only known from the cen-
tral part of the country and may be associated with its highlands. This is the case of
Dobson’s Epauletted Fruit Bat and Anchieta’s Fruit Bat, a rare or localised species.
Finally, Egyptian Rousette (Rousettus aegyptiacus) is so far only known in the
northwest of the country, which may be explained by the need to satisfy its require-
ments for abundant fruit resources and caves (Crawford-Cabral 1986).
Family Rhinolophidae
There are five species of Rhinolophidae currently recognised to occur in Angola

(Monadjem et al. 2010a; Happold and Happold 2013). Rüppell’s Horseshoe Bat
(Rhinolophus fumigatus) is a savanna species with separate populations in the west
and east of southern Africa, with the western population occupying southwest
Angola and central and northern Namibia. Lander’s Horseshoe Bat (R. landeri
sensu lato) was thought until recently to have an isolated population in Angola.
However, recent taxonomic work recognised populations from southern Africa as a
separate species, Peters’s Horseshoe Bat (R. lobatus), though no material from
Angola was examined (Taylor et al. 2018b). Given the proximity to populations
analysed it may be provisionally accepted that Peters’s Horseshoe Bat is the species
occurring in Angola, though it is possible that the country hosts a different species,
described by Seabra (1898b) as Angola’s Horseshoe Bat (R. angolensis) (Monadjem
et al. 2010a; Taylor et al. 2018b). Damara Horseshoe Bat (R. damarensis) was previ-
ously treated as a subspecies of Darling’s Horseshoe Bat (R. darlingi) (Monadjem
et al. 2010a), but has been given full species status in the IUCN Red List (Monadjem
et al. 2017). It is a species from arid habitats that is restricted to southwest Angola
and western Namibia. Dent’s Horseshoe Bat (Rhinolophus denti) is only known
from the Ruacaná falls, on the Angola-Namibia border (Crawford-Cabral 1986),
though it is widespread in Namibia and western Botswana. Eloquent Horseshoe Bat
(Rhinolophus eloquens) has been collected from Jau, Huíla Province (Grant and
Ferguson 2018), a location that is over 2000 km away from the closest records in the
eastern DRC and Rwanda. These specimens from the American Museum of Natural
History are worth re-examining as they may refer to a new species within the R.
eloquens/R. hildebrandtii group. Further surveys are likely to record more species
of this genus that are known to occur in the neighbouring countries close to the
border with Angola.
392 P. Beja et al.
Family Vespertilionidae
There are 28 species of Vespertilionidae recorded in Angola. This is a poorly known

group in the country, with many species represented by just one or a few scattered
and old records (Monadjem et al. 2010a). This is for instance the case of Beatrix’s
Butterfly Bat (Glauconycteris beatrix), Lesser Woolly Bat (Kerivoula lanosa),
Dobson’s Pipistrelle (Neoromicia grandidieri), White-bellied House Bat
(Scotophilus leucogaster), Green House Bat (S. viridis), and Moloney’s Flat-headed
Bat (Mimetillus moloneyi), which are known in Angola from just one or two records.
Seemingly, Silvered Woolly Bat (K. argentata) and Dusky Pipistrelle (Pipistrellus
hesperidus) have been tentatively recorded at single locations based on acoustic
identifications during recent surveys (Taylor et al. 2018c). Even species that are
likely to be widespread and abundant in the region, and that are known to roost in
houses in relatively large numbers, such as the Cape Serotine (Neoromicia capen-
sis), have been scarcely recorded in the country possibly due to under-sampling.
Despite the paucity of studies, it is noteworthy that five currently recognised
Vespertilionidae species have their type locality in Angola. This is the case of
Anchieta’s Pipistrelle (Hypsugo anchietae), which was collected in western Angola
(Cahata) in atypical environmental conditions compared to the main range of the
species, though it may be widespread in the southeast. Angolan Long-eared Bat
(Laephotis angolensis) is an endemic to southern Africa that is known from just
four specimens collected at one locality in DRC and two in central Angola, includ-
ing the type locality (Tyihumbwe). Bocage’s Mouse-eared Bat (Myotis bocagii) is
widespread in tropical Africa, with just a few records in northern Angola, including
the type locality. Welwitsch’s Mouse-eared Bat (M. welwitschii) was described
from northern Angola, although the great majority of the species’ records are in
East Africa. Angolan Hairy Bat (Cistugo seabrae) was described from specimens
collected in Moçamedes. It is endemic to southern Africa, where it occurs from the
extreme southwest of Angola through western Namibia to the extreme northwest of
South Africa. Finally, it is worth mentioning Machado’s Butterfly Bat
(Glauconycteris machadoi), a species described from eastern Angola that is not
retained here because it has been suggested that it may correspond to a colour phase
of G. variegata, although this issue needs clarification (Crawford-Cabral 1986).
For several Vespertilionidae Angola is at the edge of their distribution, and so
they occur marginally in the country, which may justify at least partly the scarcity
of records. This is the case of species associated with rainforests and other forests
of tropical Africa, which are largely restricted to Cabinda and northern Angola.
Besides the above mentioned Bocage’s Mouse-eared Bat, the species showing this
pattern are Common Butterfly Bat (G. argentata), Beatrix’s Butterfly Bat, White-
winged Serotine (Neoromicia tenuipinnis), and Broad-headed Pipistrelle (Hypsugo
crassulus). Other species have their core range in eastern and southern Africa and
have isolate records in Angola, including Lesser Woolly Bat, Green House Bat,
Damara Woolly Bat, and Dusky Pipistrelle. It is possible that these patterns are
shaped to at least some extent by under-sampling in Angola. Finally, Long-tailed
Serotine (Eptesicus hottentotus) is only known from southwest Angola, but it has a
much wider, albeit sparse distribution in southern Africa (Monadjem et al. 2010a).
Most Vespertilionidae from woodlands and savannas have also been scarcely
recorded in Angola, though climatic niche modelling suggests that many savanna
species may occur more widely in the south of the country (Monadjem et al. 2010a).
In contrast, only Thomas’s Flat-headed Bat (M. thomasi) has been associated with
the moist miombo belt of southcentral Africa, which occupies a large part of central
Angola. Banana Bat (Neoromicia nana) is associated with well-wooded habitats
such as riparian vegetation and forest patches, and it has been recorded in western
and central Angola. Savanna and open woodland species, many of which are associ-
ated with aquatic and riparian habitats, include Variegated Butterfly Bat (G. varie-
gata), Botswana Long-eared Bat (Laephotis botswanae), Zulu Serotine (Neoromicia
zuluensis), Schlieffen’s Twilight Bat (Nycticeinops schlieffeni), Rüppell’s Pipistrelle
(Pipistrellus rueppellii), Rusty Pipistrelle (P. rusticus), Thomas’s House Bat
(Scotoecus hindei), Yellow-bellied House Bat (Scotophilus dinganii), and White-
bellied House Bat. The inclusion of Thomas’s House Bat is tentative, because the
taxonomy of this genus is still unresolved and confusion with White-bellied Lesser
House Bat (S. albigula) is possible (Monadjem et al. 2010a). Specimens of Yellow-
bellied House Bat from northern Angola, the southern DRC and northern Zambia
were collected in environmental conditions considered unsuitable, and may repre-
sent a distinct species.
Erinaceopmorpha (Hedgehogs)
The Southern African Hedgehog (Atelerix frontalis) is the sole representative of the
order Erinaceopmorpha in Angola, with a distribution restricted to the southwest of
the country, in the Namibe and Huíla provinces, which extends into central Namibia
(Cassola 2016b). This population is disjunct from another occurring in Zimbabwe,
Botswana and South Africa (Cassola 2016b).
Hyracoidea (Hyraxes)
Four species of Hyracoidea are known from Angola. The Angolan Bush Hyrax
(Heterohyrax brucei bocagei) is widely distributed in rocky habitats across the
western half of the country, and is mostly a plateau species absent below 500 m of
altitude (Gray 1869; du Bocage 1889b; Crawford-Cabral and Veríssimo 2005).
Nevertheless, the Bush Hyrax may be sympatric just below the southern escarpment
with the Kaokoveld Rock Hyrax (Procavia capensis welwitschii), the latter being
mostly associated with rock outcrops in the southwest arid coastal plain (du Bocage
1889b; Hill and Carter 1941; Crawford-Cabral and Veríssimo 2005). In addition,
two species of Tree Hyrax, the Southern Tree Hyrax (Dendrohyrax arboreus cf
braueri) and the Western Tree Hyrax (Dendrohyrax dorsalis nigricans) are present
in forest habitats, the former associated with gallery forests and nearby miombo in
eastern Angola (Machado 1969; Crawford-Cabral and Veríssimo 2005). The
394 P. Beja et al.
Western Tree Hyrax is only known from one confirmed record obtained in gallery
forest in Cabinda (Peters 1879; Crawford-Cabral and Veríssimo 2005), but recent
observations have suggested that the species’ may be present on the central escarp-
ment region (Vaz Pinto, unpublished data). The Angolan Bush Hyrax is likely an
endemic taxon and the Kaokoveld Rock Hyrax has a global distribution restricted to
the Namib Desert between coastal Namibia and Angola, but to date no study has
specifically addressed Angolan hyraxes.
Lagomorpha (Hares)
A recent review recognises only two species of lagomorphs in Angola (Smith et al.
2018), but taxonomic uncertainties, controversial identifications and lack of recent
studies in the country, have blurred the matter and it is likely that at least three spe-
cies occur. The African Savanna Hare (Lepus victoriae) is the most widespread spe-
cies, occurring throughout the country except the arid southwest and the forested
northwest (Smith et al. 2018). There are also records of additional Lepus species,
but at least some of these may refer to misidentifications and thus need further con-
sideration. Genest-Villard (1969) recorded the presence of two species of hares
based on specimens collected by Crawford-Cabral, with Cape Hare (Lepus capen-
sis) occuring in semi-desertic areas and L. crawshayi (= L. victoriae) in less arid
areas. Actually, hares collected in Moçâmedes were initially classified as a distinct
species, L. salai Jentink 1880, and then ascribed to a subspecies of Cape Hare (L. c.
salai) occurring in subdesertic areas of southwestern Angola and western Namibia
(Petter and Genest 1965). In the mammal collection of IICA, currently housed in
ISCED-Huíla, various skins from coastal Namibe were identified as L. capensis
(Crawford-Cabral and Veríssimo, unpublished data), and recent field observations
suggest the Cape Hare to be common in Iona NP and along the southwestern arid
coastal plain into Benguela province (Vaz Pinto, unpublished data). A recent record
of Cape Scrub Hare (Lepus saxatilis) in the Malanje province by Moraes and Putzke
(2014) is probably a misidentification, as the species is a narrow endemic to the
Cape province of South Africa (Smith et al. 2018). Jameson’s Red Rock Hare
(Pronolagus randensis) occurs in southern Angola, corresponding to the northern
tip of a larger range extending into central Namibia, which is disjunct from another
population with its core distribution in Zimbabwe, eastern Botswana and northeast
South Africa (Smith et al. 2018). Interestingly, the Rock Hare in Angola appears to
be present in two subpopulations segregated by altitude. One subpopulation is pres-
ent in altitude on top of the escarpment, often above 2000 m in Huíla Province, and
has been tentatively ascribed by Crawford-Cabral to (P. r. waterbergensis)
(Crawford-Cabral and Veríssimo, unpublished data). The other subpopulation is
found on rocky outcrops and inselbergs on the coastal plain, and distinct coloration
patterns led the same author to suggest a new taxon (P. r. moçamedensis) (Crawford-
Cabral and Veríssimo, unpublished data). No subspecies of Red Rock Hare are cur-
rently recognised (Happold 2013), but the Angolan populations and their distribution
pattern remain unresolved. There have also been claims suggesting the occurrence
of Bunyoro Rabbit (Poelagus marjorita) in Angola (Petter 1972), but these were
subsequently challenged and dismissed in recent publications, which consider the
species to be restricted to relatively small disjunct populations in the Central African
Republic, South Sudan, and Uganda (Happold and Wendelen 2006; Happold 2013;
Smith et al. 2018). Nevertheless, a couple of museum specimens collected in 1941
on the Angolan Escarpment near Gabela and attributed to this species (RBINS
2017) are intriguing and require further verification, particularly as the region is
known as an endemism hotspot (Hall 1960; Happold and Wendelen 2006; Clark
et al. 2011; Svensson et al. 2017).
Macroscelidea (Sengis)
Three species of Macroscelideae have been collected in Angola (Crawford-Cabral

and Veríssimo, unpublished data). Short-snouted Sengi (Elephantulus brachyrhyn-
chus) is reportedly distributed across the plateau, associated with grasslands and
thickets in miombo woodlands (Monard 1935; Hill and Carter 1941; Crawford-
Cabral and Veríssimo, unpublished data). The species is possibly represented by
three subspecies in Angola, E. b. brachyrhynchus, E. b. brachyurus and E. b. schinzi
(Crawford-Cabral and Veríssimo, unpublished data), but a clinal variation has also
been suggested and Perrin (2013) recognises no subspecies. Bushveld Sengi
(Elephantulus intufi) is present in the semi-arid savannas of the southwest and often
associated with mopane woodlands, with the subspecies E. i. mossamedensis occur-
ring on the coastal plain, while E. i. alexandri is found inland (Hill and Carter 1941;
Crawford-Cabral and Veríssimo, unpublished data). Four-toed Sengi (Petrodromus
tetradactylus tordayi) has been collected from gallery forests in Lunda-Norte, and it
is likely that the nominate race P. t. tetradactylus also occurs in eastern Moxico
(Hayman 1963; Crawford-Cabral and Veríssimo, unpublished data; Rathbun 2013).
As no studies have focused on Angolan sengis, and considering the known range of
sengis in neighbouring countries, it is possible that future surveys may add new spe-
cies to the country list, such as the Western Rock Sengi (Elephantulus rupestris) in
the arid southwest region.
Perissodactyla (Rhinoceros, Zebras)
The Order Perissodactyla is represented in Angola by Black Rhinoceros (Diceros

bicornis), and two species of zebra. Although White Rhinoceros (Ceratotherium
simum) has occasionally been included on the Angolan mammal lists, this is not
supported by any hard data. Claims referring White Rhinoceros to having been pres-
ent in the southeastern regions were not based in any collected specimens, but rather
on poor and indirect evidence or old unsubstantiated witness reports (Shortridge
396 P. Beja et al.
1934; Newton da Silva 1970; Crawford-Cabral and Veríssimo 2005). Most authors
agree that even if they had once occurred, which is possible but remains unproven,
by the mid-twentieth century they had long been extirtpated (Hill and Carter 1941;
Newton da Silva 1970; Crawford-Cabral and Veríssimo 2005).
Family Equidae
The zebras native to Angola are Hartmann’s Mountain Zebra (Equus zebra hart-
mannae) and Plains Zebra (Equus quagga). A zebra collected in the nineteenth cen-
tury by Penrice 70 km north of Moçamedes led Oldfield Thomas to describe a new
species of mountain zebra (Thomas 1900) as E. penricei, but these were later reval-
uated and synonymised with Hartmann’s Zebra (Hill and Carter 1941; Crawford-
Cabral and Veríssimo 2005). In Angola, Hartmann’s Zebras were found in the arid
southwest, mostly in Namibe Province but its distribution may have once extended
north to southern Benguela (Shortridge 1934; Crawford-Cabral and Veríssimo
2005). The stronghold of the species in Angola was in Iona NP where a healthy
population was present in the 1970s (Huntley 1973c), although 20 years later they
were on the verge of extinction and their status remained unaltered until the end of
the war (Huntley and Matos 1992; Novellie et al. 2002; Crawford-Cabral and
Veríssimo 2005; Penzhorn 2013). Nevertheless a few herds survived the war in Iona
NP, and aerial surveys in the park have reported an increase in estimated numbers
from 263 to 434 individuals between 2003 and 2016 (Kolberg and Kilian 2003; van
der Westhuizen et al. 2017). On the other hand, recent observations and circumstan-
tial evidence strongly suggest that some herds in Iona NP may have been hybridiz-
ing with feral donkeys, as various individuals with intermediate phenotypes have
been photographed in recent years (Vaz Pinto, unpublished data). Genetic confirma-
tion of the hybridisation is lacking and the extent of the phenomenon remains
unknown.
Angolan Plains Zebras have been tentatively ascribed to various subspecies,
however recent molecular studies failed to distinguish among traditional subspecies
(Lorenzen et al. 2008). Plains Zebra used to be relatively common and widespread
in Angola across the southern half of the country and possibly also present in east-
ern Moxico (Newton da Silva 1970; Crawford-Cabral and Veríssimo 2005). A six-
teenth century report has even suggested that zebras in those days may have extended
along the coast to Ambriz, but the claim remains controversial (Crawford-Cabral
and Veríssimo 2005). In the 1970s they were still numerous at least in protected
areas such as Iona NP and Bicuar NP, and in the Cuando Cubango province (Huntley
1973c, 1974; Crawford-Cabral and Veríssimo 2005). It is likely that Plains Zebra
were more affected by war-time poaching than most other large ungulates, and
already by 1992 they were feared extinct in Angola (Huntley and Matos 1992).
Some surveys after the end of the war have failed to record the species in southern
Angola (Kolberg and Kilian 2003; Veríssimo 2008; van der Westhuizen et al. 2017),
but recent reports suggest that a few animals may still linger in Bicuar NP (Overton
et al. 2017). Small numbers were confirmed in general surveys conducted in
Luengue-Luiana NP (Chase and Schlossberg 2016; Funston et al. 2017).
Family Rhinocerotidae
Black Rhinoceros were known in Angola from two disjunct populations probably
corresponding to different subspecies, although the distinctions among black rhino
subspecies remain controversial (Crawford-Cabral and Veríssimo 2005; Rookmaaker
2005; Emslie and Adcock 2013). A population of arid-adapted rhinos occurring in
Angola to the west of the Cubango (Okavango) river, has been ascribed to D. b.
minor (Ansell 1972; Crawford-Cabral and Veríssimo 2005), but it is now generally
recognised to represent instead the former northern limit for the typical race D. b.
bicornis (Emslie and Brooks 1999; Emslie and Adcock 2013). The other population
used to extend to the east of the Cuito River in southeastern Angola, and was once
considered as D. b. chobiensis (Ansell 1972; Crawford-Cabral and Veríssimo 2005).
However, this putative race has been more often synonymised with D. b. minor
(Emslie and Brooks 1999; Rookmaaker 2005; Emslie and Adcock 2013).
No research has specifically focused on Angolan rhinos, and the existing knowl-
edge is based in the few specimens collected during early expeditions, scattered
reports from trophy hunters and the work by ecologists in the 1970s (Hill and Carter
1941; Newton da Silva 1970; Huntley 1973c, 1974; Crawford-Cabral and Veríssimo
2005). Black Rhinoceros were likely always scarce in numbers throughout histori-
cal times (Huntley 1973c, 1974; Crawford-Cabral and Veríssimo 2005), and in the
1970s they were estimated at around 30 in Iona NP with small populations in south-
ern Cuando Cubango (Huntley 1973c). The situation deteriorated fast during the
armed conflict that followed independence, and by 1992 they were already gone or
on the verge of extinction (Huntley and Matos 1992; Crawford-Cabral and Veríssimo
2005). By the turn of the millennium rhinos were considered extinct in Angola
(Emslie and Brooks 1999), and have remained since, in spite of occasional
unconfirmed sightings that suggest the possibility of a few scattered individuals
surviving in remote locations. Recent general surveys in regions where they used to
occur have consistently failed to record the species (e.g. Veríssimo 2008; Chase and
Schlossberg 2016; Funston et al. 2017; Overton et al. 2017; NGOWP 2018).
Pholidota (Pangolins)
Three species of pangolins occur in Angola (Crawford-Cabral and Veríssimo,

unpublished data), but no specific studies have been conducted regarding the status
or ecology of these species within the country. White-bellied Pangolin (Phataginus
tricuspis) has been recorded, and collected (Hill and Carter 1941; Trense 1959) in
the provinces of Cabinda, Lunda-Norte, Zaire and Cuanza-Norte, and although
assumed overall scarce, this species is expected to be widely distributed across
northern Angola and the upper plateau, and likely extending south along the escarp-
ment forests (Crawford-Cabral and Veríssimo, unpublished data). The species has
also been recorded recently in Cangandala NP, Malanje Province (Vaz Pinto, unpub-
lished data), and it has been found in Angolan bushmeat markets (Svensson et al.
2014b). Temminck’s Ground Pangolin (Smutsia temminckii) is the most common
398 P. Beja et al.
species of pangolin in Angola, with a wide distribution in the central and southern
areas of the country. It has been recorded in the provinces of Cuanza-Sul, Benguela,
Bié, Huíla and adjacent areas of Namibe, and Cuando Cubango (Crawford-Cabral
and Veríssimo, unpublished data), although the only known material available in
natural history collections is housed in the AMNH (Hill and Carter 1941) and
ISCED (Crawford-Cabral and Veríssimo, unpublished data). Giant Ground Pangolin
(Smutsia gigantea) is restricted to the forests of Cabinda, and its occurrence there
has been known since the mid-1970s (Huntley 1973e). More recently, the species
has been re-confirmed in the Maiombe region of the enclave (Ron 2005).
Primates (Monkeys, Apes, Pottos, Galagos)
Angola has a great diversity of primates, including up to 15 diurnal and possibly 8

nocturnal species. The vast majority of Angola’s primate species are found in the
rainforests and riverine forest-woodland mosaics in the north, including Cabinda,
within the biodiversity-rich Guinea-Congolian biome (Huntley 1973e; Kuedikuenda
and Xavier 2009; IUCN 2018). The montane forests/dry woodland mosaics of the
Angolan Escarpment along the coast, which connect to the rainforests in the north,
are also rich in primates, including at least four diurnal and four nocturnal species
(Bersacola et al. 2015).
Family Cercopithecidae
The Cercopithecidae is the most speciose group of primates occurring in Angola,

including two baboons, one mangabey, six guenons, two talapoin monkeys, and one
colobus. The two baboons are Chacma Baboon (Papio ursinus), ranging in the arid
southwestern regions (Benguela, Namibe, Huíla and Cunene Provinces), and Kinda
Baboon (P. kindae), occupying the central and northeast of the country (Machado
1969). Chacma Baboon is adapted to woodland, sub-desert, savanna and montane
habitats, whereas the Kinda Baboon tends to be present in miombo woodland,
deciduous and semi-deciduous forests, savanna, gallery and riverine forests habitats
(Kingdon 2016; Rowe and Myers 2016). Up-to-date information on baboon distri-
bution in Angola is not available. Black Crested Mangabey (Lophocebus aterrimus)
inhabit the tropical forests south of the River Congo. The subspecies occurring in
Angola, the Southern Black Crested Mangabey (L. a. opdenboschi) was recorded in
the gallery forests of Lunda-Norte (Machado 1969) and just on the eastern side of
the Cuango River in the Lunda-Norte province (Hart et al. 2008). Little is known
about the current status of this subspecies (Hart et al. 2008; Rowe and Myers 2016).
Red-tailed Monkey (Cercopithecus ascanius) is represented by three subspecies
in Angola, namely Black-nosed Red-tailed Monkey (Cercopithecus a. atrinasus),
Black-cheeked Red-tailed Monkey (C. a. ascanius), and Katanga Red-tailed
Monkey (C. a. katangae), all ranging in the north of the country (Machado 1969;
Oates et al. 2008a, b; Rowe and Myers 2016). The existence of Black-nosed Red-
tailed Monkey in Angola is based on only nine individuals collected in the 1960s
(Machado 1969; Sarmiento et al. 2001; Oates et al. 2008b). Red-tailed Monkeys are
typically found in a wide range of habitats, including rainforests, swamp, riverine
and montane forests, and deciduous and semi-deciduous forests (Sarmiento et al.
2001; Rowe and Myers 2016). All three subspecies of Red-tailed Monkey have been
known to hybridize among each other in Angola (Machado 1969; Detwiler et al.
2005). Blue Monkey (Cercopithecus mitis) occurs in two disjunct populations in
Angola, corresponding to two subspecies. The Pluto Monkey (Cercopithecus mitis
mitis) is endemic to Angola and ranges along the Angolan Escarpment areas in the
west part of the country (Machado and Crawford-Cabral 1999; Kingdon 2008a;
Lawes et al. 2013). In 2013, this was the second most commonly occurring species
in a bushmeat survey (Bersacola et al. 2014). The Rump-spotted Blue Monkey (C.
mitis opisthostictus), ranges in a small part of eastern Angola, corresponding to the
edge of a much wider distribution in East Africa (Kingdon 2008b). Black-footed
Crowned Monkey (Cercopithecus pogonias nigripes) supposedly occurs in the
Cabinda Province, likely occurring in the Maiombe NP. The species uses high veg-
etation strata, mainly occurring in primary and lowland tropical forests, as well as
savanna, gallery forests, mature secondary forests and montane forests (Zinner et al.
2013). Moustached Monkey (Cercopithecus cephus) was common in Cabinda
(Machado 1969; Huntley 1973e). The species is mainly found in lowland tropical
rainforest, but also in secondary habitats (Gautier-Hion et al. 1999). Machado
(1969) also recorded Putty-nosed Monkey (C. nictitans) in Cabinda, a species found
in lowland and montane tropical moist forests, as well as gallery and secondary
forests (Oates and Groves 2008). De Brazza’s Monkey (C. neglectus) was common
in Lunda-Norte (Machado 1969). The species is mainly found in riverine forest
habitats, in lowland and submontane semi-deciduous or tropical moist forest, as
well as in swamp forest (Struhsaker et al. 2008). The Malbrouck Monkey
(Chlorocebus cynosuros) occurs throughout Angola, in various habitats and eleva-
tion gradients (Huntley 1973c; Sarmiento 2013). This species is mainly present in
open woodland, savanna and forest-grassland mosaic, and tends to occur close to
water sources (Butynski 2008; Sarmiento 2013). However, it is also able to occupy
both rural and urban environments (Butynski 2008).
The two Talapoin Monkeys of Angola are Southern Talapoin Monkey
(Miopithecus talapoin) and Northern Talapoin Monkey (M. ogouensis). The first
species occurs along the Angolan Escarpment, including Quiçama, Cumbira Forest,
up to the Congo River (Machado 1969; Gautier-Hion 2013a; Groom et al. 2018).
The second species is present in Cabinda (Gautier-Hion 2013b). The Southern
Talapoin Monkey is one of Africa’s least studied primates, but it is assumed to be
ecologically similar to the Northern Talapoin Monkey, therefore preferring dense
forest environments, such as riverine forest (Machado 1969). Sclater’s Angolan
Colobus (Colobus angolensis angolensis) was known to occur in the northeast of
Angola (Malanje, Lunda-Norte; Machado 1969), where it was recorded in 2009
near Lóvua (Pedro Vaz Pinto, unpublished data) and in riverine forests in the Lagoa
Carumbo area in 2011 (Huntley and Francisco 2015).
400 P. Beja et al.
Family Galagidae
The Galagidae are represented by six species in Angola. Four of these species have
been confirmed in recent surveys, namely Thick-tailed Greater Galago (Otolemur
crassicaudatus), Southern Lesser Galago (Galago moholi), Demidoff’s Dwarf
Galago (Galagoides demidoff), and importantly, a new, recently-described Dwarf
Galago believed to be endemic to Angola, the Angolan Dwarf Galago (Galagoides
kumbirensis) (Bersacola et al. 2015; Svensson et al. 2017). The Angolan Dwarf
Galago was named after Cumbira Forest where it was first observed (Svensson et al.
2017), but it appears to be adapted to a wide range of habitats (Bersacola et al.
2015). The geographical distribution of this new species is not yet established,
though it might range as far as the Congo River in DRC (Svensson et al. 2017).
Thick-Tailed Greater Galago occurs throughout Angola, except the extreme
south (Bearder 2008). It is typically associated with open woodland and savanna
habitats as well as in forest edges and thickets, using mid to high strata (Bearder
et al. 2003; Bearder and Svoboda 2013). Its occurrence was confirmed in semi-arid
savanna environments and Cumbira Forest (Cuanza-Sul), and in miombo wood-
lands in Malanje Province (Bersacola et al. 2015). Southern Lesser Galago is wide-
spread in the miombo woodlands of Angola (Huntley 1973c; Bersacola et al. 2015).
It is known to use all strata in open woodland, savanna, forest edges and other semi-
arid habitats (Bearder et al. 2003; Bearder et al. 2008; Pullen and Bearder 2013).
Demidoff’s Dwarf Galago is known to occur across central to northeastern parts of
Angola, including Cuanza-Sul, Cuanza-Norte, Malanje, Lunda-Sul and Lunda-
Norte Provinces (Machado 1969; Svensson 2017). The occurrence of this species in
semi-arid savanna zones of the Angolan Escarpment extended the species’ range c.
190 km further southwest (Svensson 2017). This is the smallest of all the galagos
and is typically associated with forest habitats, including deciduous and semi-
deciduous forests, evergreen and gallery forests, mainly in the edge and understory
habitats (Bearder et al. 2003; Ambrose and Butynski 2013). Their relative abun-
dance in Angola was correlated with undergrowth density, canopy cover and tree
density (Bersacola et al. 2015).
Two additional species have been reported from Angola, but their current status
is unknown. Thomas’s Dwarf Galago (Galagoides thomasi) is believed to range in
the northern parts of Angola, but this still needs confirmation (Bersacola et al.
2015). Bersacola et al. (2015) proposed that competitive exclusion between the
Thomas’s and the Angolan Dwarf Galagos could explain why the former species
was not observed in the Angolan Escarpment forests. The Southern Needle-clawed
Galago (Euoticus elegantulus) was listed as possibly occurring in Cabinda (Huntley
1973e). The species is known to occur in both primary and secondary forests at low-
to medium-altitude, including in deciduous and semi-deciduous, evergreen and lit-
toral forests (Ambrose and Butynski 2013).
Family Hominidae
The two species of the family Hominidae in Angola are the Western Lowland
Gorilla (Gorilla gorilla gorilla) and the Central Chimpanzee (Pan troglodytes trog-
lodytes), which are confined to the Cabinda Province (Maisels et al. 2016a, b). Both
species were known to be present in the area currently included in the Maiombe NP,
a c. 2000 km2 area consisting of mainly tropical forest ecosystems which are part of
the Guinea-Congolian biome. The landscape in Cabinda is characterised by semi-
deciduous tropical forests in the northeast (including Maiombe), agroforest mosaics
largely covering the south, as well as mangrove and flooded swamp forests along
the coast. Western Lowland Gorilla occur in different types of forest environments
(Robbins et al. 2004; Tutin and Fernandez 1984). Across their range chimpanzees
occupy a great variety of habitats, from tropical rainforests to semi-arid savanna
environments (Boesch and Boesch-Achermann 2000; Pruetz 2006). Chimpanzees
show high socioecological flexibilities in human-dominated environment (Hockings
et al. 2012; McLennan 2013; Bessa et al. 2015). Considering the high socioecologi-
cal flexibility of chimpanzees across their range, this species’ range in Cabinda is
likely to include human-dominated areas in the south of the province. Despite the
urgency for surveys highlighted previously (Tutin et al. 2005), population estimates
for the two great apes in Cabinda remain unavailable. In the province, chimpanzees
are likely targeted for the commercial bushmeat trade (Ron and Golan 2010), but the
scale of this trade is poorly known. Huntley (2017) records that gorilla and chim-
panzee were by tradition not included among bushmeat species in Cabinda in 1973,
while Bersacola et al. (2014) reports on chimpanzees occuring in the pet trade in
Angola that were believed to originate from Cabinda. Future studies to assess the
distribution and population status of the two great apes in Cabinda, including in-
depth investigation on the human-great ape interactions in this region should be
considered a priority.
Family Lorisidae
Milne-Edwards’s Potto (Perodicticus edwardsi) and Golden Angwantibos

(Arctocebus aureus) are the two species of the family Lorisidae reported for Angola
(Huntley 1973e; Bersacola et al. 2015; Svensson 2017). Milne-Edwards’s Potto is
the largest of the potto species, generally occurring in both primary and secondary
forests, from low altitude to montane forests (Butynski and de Jong 2007; Pimley
2009; Oates 2011; Pimley and Bearder 2013). The species was reported from
Cabinda by Huntley (1973e), while recent surveys confirmed its occurrence in the
tall, sub-humid forests of the Cuanza-Norte Province, and for the first time it was
observed in the agro-forest mosaics in Cumbira Forest (Cuanza-Sul), extending the
previously known geographical range c. 320 km further south (Bersacola et al.
2015). The Golden Angwantibos was reported from Cabinda by Huntley (1973e),
but no information has been obtained thereafter.
402 P. Beja et al.
Proboscidae (Elephants)
African elephants (genus Loxodonta) have traditionally been considered monotypic

and to comprise two subspecies, but recent genetic studies, albeit confirming
hybridisation along a contact zone, have also provided compelling evidence to sus-
tain the validity of those two forms as full species (Grubb et al. 2000; Roca et al.
2001, 2015; Palkopoulou et al. 2018), namely Savanna Elephant (L. africana) and
Forest Elephant (L. cyclotis). Interestingly, the earliest morphological studies sug-
gesting that both species should be recognised result from the efforts of Frade
(1933, 1936, 1955) and were based on the analyses of specimens collected in differ-
ent parts of Angola. Although the studies by Frade reflect the occurrence of both
species, since then no material from Angola has been critically evaluated and
records were often assumed by default to refer to Savanna Elephant. On the other
hand, some authors, have distinguished among various ‘types’ including potential
subspecies of Savanna Elephants (Hill and Carter 1941), and some hunters even
suggested the occurrence in the north of a dwarf elephant (Crawford-Cabral and
Veríssimo 2005), but these likely correspond to geographical variants of either
species.
Forest Elephant was once likely common to abundant in moist forested habitats
across northwestern Angola, and including Cabinda, and a large number of records
from hunters suggest their former presence throughout the provinces of Zaire, Uíge
and Cabinda (Crawford-Cabral and Veríssimo 2005; Crawford-Cabral and
Veríssimo, unpublished data). Soon after the end of the civil war, Forest Elephants
were still not uncommon in the Maiombe rainforest of Cabinda (Heffernan 2005)
but more recent accounts are lacking. There remains little doubt that their numbers
have plummeted in recent decades as a result of hunting and habitat destruction, and
Forest Elephant may have been extirpated from large parts of its former range in
Angola. Surprisingly, a few herds of what appear to be Forest Elephant still seem to
linger in forest blocks in Cuanza-Norte and Bengo Provinces (Vaz Pinto, unpub-
lished data). If confirmed, this pocket of Forest Elephant in Cuanza-Norte and
Bengo may consist of an isolated, southernmost population of the species, which
would much increase its conservation importance. The possibility of a hybrid zone
in northern Angola where Savanna Elephant also used to occur (Crawford-Cabral
and Veríssimo 2005) cannot be excluded, but this hypothesis remains untested.
Unfortunately, and paradoxically considering the large interest and funding chan-
nelled to research and conservation of elephants in general, very little is known
about the Angolan Forest Elephant, including its taxonomic status and relationships
with other populations, ecology, and numbers.
Savanna Elephant used to have a wide distribution in Angola, including along the
coastal plain and in the east and northeast, but the core was in the southern half of
the country. An extrapolation of figures based on aerial counts suggested the popu-
lation in the southeast province of Cuando Cubango to reach up to 23,000 animals
(Hall-Martin and Pienaar 1992), but the ensuing civil war prevented reliable counts.
Quantification of elephant populations for the whole country was also attempted in
the 1990s but based on little ground data, and yet suggesting a steep reduction in
numbers from 50,000 to less than 10,000 (Anstey 1991, 1993). As a result of the
armed conflict, the Savanna Elephant populations were most affected and may have
disappeared completely from extensive regions, particularly along the coastal plain,
while in other regions they may have survived in much reduced pockets. Following
the end of the war, they may have recolonised extensive regions of the southeast, as
migratory routes have been reopened, allowing the dispersal of Savanna Elephants
coming from neighbouring countries, particularly Botswana (Chase and Griffin
2011), although more recent evidence suggest negative trends as a result of increased
poaching pressure and human encroachment (Chase and Schlossberg 2016; Funston
et al. 2017; Schlossberg et al. 2018). A small contingent of Savanna Elephant were
introduced into Quiçama NP in 2000 and 2001, and these have since increased from
32 to about 90 (Carmignani 2015), but a small number of the original population
may also have survived in the park (Groom et al. 2018). Much of what is currently
known on the distribution and status of elephants in Angola is summarised in the
IUCN African Elephant Status Report (Thouless et al. 2016).
Rodentia (Mole Rats, Mice, Dormice, Rats, Voles, Squirrels)
The rodents are a vast group with at least 85 species currently recognised to occur
in Angola (Monadjem et al. 2015; Taylor, unpublished data Taylor et al. (2018c).
Rodents show a high degree of endemism in Angola, with at least 13 endemic or
near-endemic species. However, there are considerable uncertainties regarding the
taxonomy of African rodents, and in the future it is likely that many species will be
split after the development of detailed taxonomic and genetic studies, thus increas-
ing the number of endemics or near endemics (Monadjem et al. 2015). For instance,
while Taylor (2016) treats the African Marsh Rat (Dasymys incomtus) as a single
widespread species, this may indeed be a complex of several similar species, some
of which may have restricted distributions (Monadjem et al. 2015). Continued field
surveys are also likely to increase the rodent list, as several species have been
recorded on or close to the borders of Angola and will probably be shown to occur
there in the future.
Family Anomaluridae
There are only two species of Anomaluridae in Angola, both of which are known
from just a few records (Monadjem et al. 2015). Lord Derby’s Scaly-tailed Squirrel
(Anomalurus derbianus) occurs in the northern half of Angola. It is a nocturnal and
arboreal species, which is mostly associated with the forest zone of tropical Africa,
though it also occurs in miombo woodlands. Beecroft’s Scaly-tailed Squirrel (A.
beecrofti) is also a species from the forests of tropical Africa, which in Angola
occurs mainly in Cabinda, though there are also records south of the Congo River
(Happold 2013).
404 P. Beja et al.
Family Bathyergidae
There are two species of Bathyergidae recorded in Angola (Monadjem et al. 2015).
Bocage’s Mole Rat (Fukomys bocagei) is near endemic, occurring widely in the
west of Angola and extending narrowly into northern Namibia (Faulkes et al. 2016).
Mechow’s Mole Rat (F. mechowi) is a highly adaptable species often found in vil-
lages and croplands, and which occurs in central and northeast Angola. The popula-
tion in western Angola seems to be separated by a large gap from those in eastern
Angola, Zambia and DRC, but this may reflect the paucity of surveys. Damara Mole
Rat (F damarensis) is known in Angola from only two specimens collected in
1964 in the southeast (Orrell and Hollowell 2018), but it has not been recorded
thereafter. The presence of this species thus needs confirmation, though this is likely
because it is known from neighbouring areas in Namibia and Zambia. Old records
refer to the presence of African Mole Rat (C. hottentotus or C. h. bocagei) in Angola
(e.g., Conroy 2018, MNHN 2018), but these were more likely Bocage’s Mole Rats.
Family Gliridae
There are five Gliridae in Angola (Monadjem et al. 2015). Angolan African
Dormouse (Graphiurus angolensis) is a near endemic savanna species, which is
restricted to Angola and to a small disjunct area in western Zambia. Kellen’s
Dormouse (G. kelleni) has a range in central Angola that extends into western
Zambia, but that is disjunct from other populations across the savanna zone of sub-
Saharan Africa. Monard’s Dormouse (G. monardi) is a species associated with
miombo woodland, with a small distribution in northeastern Angola and northwest-
ern Zambia. Stone Dormouse (G. rupicola) is a rupicolous species with its core
distribution in a narrow belt along the escarpment of Namibia, and marginally into
southwest Angola and South Africa. Lorrain Dormouse (G. lorraineus) is a forest
species with a known distribution in Angola restricted to the northeast, though the
species occurs in Zambia close to the border of Angola.
Family Hystricidae
Cape Porcupine (Hystrix africaeaustralis) is the sole Hystricidae occurring in

Angola (Monadjem et al. 2015). It is an eclectic species that occurs in most habitats
except dense forests, and that it is widespread throughout the country.
Family Muridae
The Muridae include 48 native species recorded in Angola, of which eight are
endemic (Monadjem et al. 2015). Endemics are mainly associated with the central
plateau, such as Thomas’s Rock Rat (Aethomys thomasi), Angolan Marsh Rat
(Dasymys nudipes), and Angolan Vlei Rat (Otomys anchietae), or the western high-
lands, such as Angolan Multimammate Mouse (Myomyscus angolensis) and Cuanza
Vlei Rat (O. cuanzensis). Coetzee’s Praomys (Praomys coetzeei) is a species
recently described that is known from just a few specimens collected in northern
Angola (van der Straeten 2008). Angolan Hylomyscus (Hylomyscus carillus) is
only known from Angola, but may also occur in neighbouring DRC. Heinrich’s
Hylomyscus (H. heinrichorum) was very recently described from specimens col-
lected in 1954 at Mount Moco and Mount Soque (Carleton et al. 2015). Another
seven species are near endemics, occurring in Angola and neighbouring countries.
Bocage’s Rock Rat (A. bocagei), Griselda’s Single-striped Mouse (Lemniscomys
griselda), Angolan Brush-furred Rat (Lophuromys angolensis), Callewaert’s Mouse
(Mus callewaerti), and Bell Groove-toothed Swamp Rat (Pelomys campanae) are
restricted to Angola and DRC. Shortridge’s Mastomys (Mastomys shortridgei) is
known from just a few scattered localities in Angola, the Caprivi Strip (Namibia)
and the extreme northwestern region of Botswana. Cabral’s Marsh Rat (D. cabrali)
and Setzer’s Mouse (M. setzeri) are endemic to a narrow area in southeastern
Angola, northwestern Botswana and northeastern Namibia. The latter species was
only recently recorded in the Okavango source lakes region of Angola (Taylor et al.
2018c).
The Muridae species with the widest distribution in Angola are those able to
thrive in association with agricultural fields and homesteads, including for instance
the Natal multimammate mouse (Mastomys natalensis). There are also widespread
species associated with the woodlands and grasslands of the central plateau, though
some of these are associated with moister tropical conditions and have their distri-
butions biased towards the north and/or the west, while others are more associated
with drier savanna habitats and have their distribution biased towards the south and
east. Overall, this is a large group of species including Marsh Rat (D. cf incomtus,
sensu Monadjem et al. 2015), Savanna Gerbil (Gerbilliscus validus), Bushveld
Gerbil (G. leucogaster), Woodland Thicket Rat (Grammomys dolichurus), Pygmy
Mouse (M. minutoides), Gray-bellied Pygmy Mouse (M. triton), Thomas’s Pygmy
Mouse (M. sorella), Creek Groove-toothed Swamp Rat (P. fallax), Angoni Vlei Rat
(O. angoniensis), Mesic Four-striped Grass Rat (Rhabdomys dilectus), and
Hildegarde’s Broad-headed Mouse (Zelotomys hildegardeae). Some of these spe-
cies, however, are known from just a few scattered records, including for instance
Thomas’s Pygmy Mouse and Hildegarde’s Broad-headed Mouse.
Some rodent species have relatively restricted distributions in Angola, because
they are associated with habitats represented only marginally in the country. This is
the case of species associated with rainforests and other moist tropical habitats,
which occur mainly in Cabinda and/or relatively small areas in the north of the
country, including African Wading Rat (Colomys goslingi), Shining Thicket Rat (G.
poensis), Typical Striped Grass Mouse (L. striatus), Dollman’s Brush-furred Rat (L.
rita), Jackson’s Soft-furred Mouse (P. jacksoni) and Big-eared Swamp Rat
(Malacomys longipes). Some species largely restricted to the north may penetrate
southwards along the Angolan Escarpment, as it is the case of Rufous-nosed Rat
(Oenomys hypoxanthus). In contrast, species associated with deserts, arid and semi-
406 P. Beja et al.
arid habitats occur mainly in the southwest of Angola, including Cape Short-eared
Gerbil (Desmodillus auricularis), Hairy-footed Gerbil (G. paeba), Setzer’s Gerbil
(G. setzeri), Black-tailed Tree Rat (Thallomys nigricauda), and Striped Mouse
(Rhabdomys bechuanae). Other species with marginal distributions in Angola due
possibly to environmental or biogeographic constraints include Kaiser’s Rock Rat
(A. kaiseri), Nyika Rock Rat (A. nyikae) and Least Groove-toothed Swamp Rat (P.
minor), in the northeast, and Red Rock Rat (A. chrysophilus), Highveld Gerbil (G.
brantsii), and Woosnam’s Broad-headed Mouse (Zelotomys woosnami), in the
south. The latter species was only confirmed in recent surveys (Taylor et al. 2018c).
Records of Desert Pygmy Mouse (Mus indutus) in southeast Angola need to be
confirmed through molecular data. Namaqua Rock Rat (Micaelamys namaquensis)
is a species widespread in southern Africa, which penetrates northwards through
western Angola.
Besides native species, the rodents of Angola also include three non-native inva-
sives, namely House Mouse (Mus musculus), a widespread commensal species,
Brown Rat (Rattus norvegicus), mainly occurring in coastal cities, and Black Rat
(Rattus rattus), widespread throughout the country. It is likely that future surveys
will increase the list of native murids in the country, including tropical species that
are known to occur close to the border of Cabinda and the north of Angola, such as
Congo Forest Mouse (D. ferrugineus), Ansorge’s Brush-furred Rat (L. ansorgei),
Peter’s Striped Mouse (Hybomys univittatus), Beaded Wood Mouse (H. aeta),
Ansell’s Wood Mouse (H. anselli), African Groove-toothed Rat (Mylomys
dybowskii), Petter’s Praomys (Praomys petteri), and Target Rat (Stochomys
longicaudatus).
Family Nesomyidae
The Nesomyidae include 15 species recorded in Angola, of which four are endemic
or near endemic to the country (Monadjem et al. 2015). Angolan Gray African
Climbing Mouse (Dendromus leucostomus) is known only from its type locality
(Caluquembe) in the highlands of Angola, but some authors treat it as conspecific
with the Gray African Climbing Mouse (D. melanotis). Vernay’s Climbing Mouse
(D. vernayi) is only known at present from a series of specimens collected near
Chitau in the central Angolan highlands. Bocage’s Fat Mouse (Steatomys bocagei)
is restricted to northern Angola and neighbouring regions of the DRC. Shortridge’s
Rock Mouse (Petromyscus shortridgei) is known only from a few scattered locali-
ties in northwestern Namibia and southwestern Angola. Another two species have
largely isolated populations in Angola. Nyika Climbing Mouse (D. nyikae) occurs
along the Angolan Escarpment, well separated from other patchy populations in
Central and East Africa. Tiny Fat Mouse (S. parvus) occurs widely in East Africa
and the northern savannas of southern Africa, but the population in southwestern
Angola seems to be largely isolated.
Six Nesomyidae are associated with savanna woodlands and/or grasslands
throughout their range, and are widespread in Angola. This includes Northern Giant
Pouched Rat (Cricetomys ansorgei), Chestnut Climbing Mouse (D. mystacalis),

Gray African Climbing Mouse (D. melanotis), Southern African Pouched Mouse
(Saccostomus campestris), Fat Mouse (S. pratensis), and Kreb’s Fat Mouse (S.
krebsii). In contrast to these species, the Forest Giant Pouched Rat (Cricetomys
emini) is associated with tropical rainforests, and in Angola its presence is vouched
in Cabinda by Musser and Carleton (2005) and Monadjem et al. (2015), but the spe-
cies does not appear to occur as widely in northern Angola as indicated in Happold
(2013). Gerbil Mouse (Malacothrix typica) and Pygmy Rock Mouse (P. collinus)
are restricted to arid environments in the southwest of the country.
Banana Climbing Mouse (D. messorius) was recorded by three specimens from
the Field Museum from Dundo in the extreme northeast Angola collected by Barros
Machado in 1948 (Grant and Ferguson 2018), but there are no known records close
to this (Monadjem et al. 2015). Previous comments on these specimens by Hayman
(1963) suggest they may be African Climbing Mouse (D. mystacalis) (Taylor
unpublished data). The monotypic Dendroprionomys, Velvet Climbing Mouse (D.
rousselotti), is known only from the type locality Brazzaville which is close to
Cabinda and may be shown to occur there.
Family Pedetidae
Spring Hare (Pedetes capensis) is the sole Pedetidae in Angola (Monadjem et al.
2015). It occurs throughout the country, except in the arid southwest, and in the
moist and forested areas of the north.
Family Petromuridae
Dassie Rat (Petromus typicus) is the single species of Petromuridae. It has been
recorded in a small area in southwest Angola, with its range expanding southwards
through Namibia and into the Northern Cape Province of South Africa (Monadjem
et al. 2015; Cassola 2016a). The species is confined to the western escarpment and
adjoining mountainous areas, as well as inselbergs.
Family Sciuridae
There are nine species of Sciuridae in Angola (Monadjem et al. 2015). Rope
Squirrels of the Funisciurus genus typically have distributions towards the north of
the country, probably due to their association with moist tropical forests. Congo
Rope Squirrel (Funisciurus congicus) is the most widespread species, ranging
widely in the Congo basin, south through the western provinces of Angola and into
northwestern Namibia. Ribboned Rope Squirrel (F. lemniscatus) occurs widely in
the Lower Guinea Forest zone of Central Africa but in Angola has only been
recorded in Cabinda, while Fire-footed Rope Squirrel (F. pyrropus) is widely
408 P. Beja et al.
distributed in the forest zone of tropical Africa but in Angola is restricted to Cabinda
and the northwest. Lunda Rope Squirrel (F. bayonii) is globally restricted to north-
ern and northeastern Angola and neighbouring areas of DRC. This species is associ-
ated with moist savanna mosaics, sandy woodlands, and low to medium elevation
moist forests (Thorington et al. 2012). Thomas’s Rope Squirrel (F. anerythrus) has
been recorded near the border of Cabinda and could possibly occur there, but this is
yet to be confirmed.
Gambian Sun Squirrel (Heliosciurus gambianus) is widely distributed in moister
savannas of tropical Africa, occurring in central and northeastern Angola. African
Giant Squirrel (Protoxerus stangeri) is the largest squirrel in Africa, having a wide
distribution across the rainforest belt of tropical Africa. In Angola it occurs in the
northwest, penetrating to the south along the escarpment, corresponding to the
endemic subspecies P. s. loandae (Happold 2013). Red-legged Sun Squirrel
(Heliosciurus rufobrachium) is not known to occur south of the Congo River but a
doubtful record from ‘Raca Camele, north of Quionlungo’ was attributed to a speci-
men from Yale Peabody Museum identified by A Heinrich. Boehm’s Bush Squirrel
(Paraxerus boehmi) has been reported for Angola based on old records from
Cabinda (Wendelen and Noé 2017) and Benguela (MNHN 2018), quite far from the
core of species distribution in tropical forests of Central Africa. Two other species
occur in the south of the country, with Damara Ground Squirrel (Xerus princeps)
occurring in a small area in southwest Angola, corresponding to the northern tip of
a larger distribution in western Namibia, and Smith’s Bush Squirrel (Paraxerus
cepapi) occurring in southern savannas.
Family Thryonomyidae
The family Thryonomyidae is represented in Angola by the Greater Cane Rat

(Thryonomys swinderianus), which is a habitat generalist and occurs throughout the
country (Monadjem et al. 2015). There are also three old records of Lesser Cane Rat
(T. gregorianus), but they are dubious because the species is difficult to distinguish
from Greater Cane Rat and the closest known records of this species are from cen-
tral DRC and western Zambia (Happold 2013; Monadjem et al. 2015). These
records refer to three specimens from Mount Moco collected by GH Heinrich in
1954 (Grant and Ferguson 2018)
Sirenia (African Manatee)
The African Manatee (Trichechus senegalensis) occurs in Angola at the southern

limit of its global distribution (Powell 1996), where it is associated with mangroves
along the lower sections of large rivers in northern Angola. The species has been
confirmed from Cabinda and various rivers between the Congo and the Cuanza
(Hatt 1934; Crawford-Cabral and Veríssimo 2005; Morais et al. 2006a, b; Dodman
et al. 2008; Collins et al. 2011). There seems to be some uncertainty regarding its
current distribution. The southernmost records have often been suggested to be the
Longa or the Queve rivers (Crawford-Cabral and Veríssimo 2005), but recent sur-
veys found no evidence of their existence south of the Cuanza (Morais et al. 2006a).
It has also been suggested that they might occur throughout the Angolan coast,
including in coastal lagoon systems such as Mussulo and as far south as the Cunene
River (Powell 1996). However, these claims lack supporting data and the habitat
present is not adequate, and so they these reports should therefore be treated with
caution (Dodman et al. 2008).
Soricomorpha (Shrews)
There are 15 species of Soricomorpha thus far recognised to occur in Angola, all
from the genus Crocidura and Suncus (Hill and Carter 1941; Hayman 1963,
Crawford-Cabral and Veríssimo unpublished data). For many of these species, how-
ever, there are only a few old records and their occurrence in Angola needs to be
confirmed. Greater Gray-brown Musk Shrew (C. luna) is known to occur in the
northeast of Angola, although until now only validated by a single specimen from
Lunda-Norte (Hayman, 1963). Reddish-gray Musk Shrew (C. cyanea) has a very
restricted range in southwestern Angola, representing the northwestern tip of a
much wider distribution in Namibia, South Africa, and elsewhere in the eastern part
of southern Africa (Baxter et al. 2016). Roosvelt’s Shrew (C. roosevelti) is a species
occurring in moist savanna around the Congo Basin forest block, which in Angola
seems to be restricted to the northeast (Hutterer and Peterhans 2016). Records of
two additional species, Lesser Gray-brown Musk Shrew (C. silacea) and Dent’s
Shrew (C. denti), are either doubtful or in need of further re-identification because
their known range is nowhere near Angola (Happold and Happold 2013). The only
potentially valid Angolan record for the Lesser Gray-brown Musk Shrew is a speci-
men in alcohol in the Museum of Dundo (Lunda-Norte), identified by Heim de
Balsac and quoted by Hayman (1963). However, this species was not considered in
the checklist, because the location of this single record is many hundreds of kilome-
ters from other known records, and difficulties in identification at the time may
signify misidentification. Dent’s Shrew record was an undated record from the
Natural History Museum with no recorded locality. The endangered Ansell’s Shrew
(C. ansellorum) is known only from two locations in gallery forests of northwestern
Zambia, close to the Angolan border where the species may also occur (Kennerley
2016).
Two of the Crocidura species recorded in Angola are endemics with restricted
distributions. The Heather Shrew (C. erica) is a poorly known species found in
Western Angola (Gerrie and Kennerley 2016), with records collected in the prov-
inces of Cuanza-Norte, Malanje, Huambo, Benguela, Huíla (Crawford-Cabral and
Veríssimo, unpublished data), while Blackish White-toothed Shrew (C. nigricans)
occurs in the southwest (Crawford-Cabral 1987; Hutterer 2016), particularly
410 P. Beja et al.
in localities along the mountainous western belt of the Angolan plateau (Huambo,
Benguela, Huíla, Cunene) (Crawford-Cabral and Veríssimo, unpublished data). The
only Crocidura shrew that is thought to be widespread throughout Angola is Oliver’s
Shrew (C. olivieri) (Crawford-Cabral and Veríssimo, unpublished data; Cassola
2016c) which represents a species complex and is in urgent need of revision.
However, Tumultuous Shrew (C. turba), African Black Shrew (C. nigrofusca) and
Swamp Musk Shrew (C. mariquensis) are also widespread in some regions of the
north and along the Angolan Escarpment. Small-footed Shrew (C. parvipes) is
known to occur in the provinces of Bié and Huíla (Hill and Carter 1941; Crawford-
Cabral and Veríssimo, unpublished data). Other species in Angola have their distri-
butions associated with their much wider ranges in Africa, including Lesser Red
Musk Shrew (C. hirta) recorded from Lunda-Norte, Lunda-Sul, Cuanza-Sul,
Huambo and Huíla, and may occur everywhere on the Angolan highlands, being
locally rather common, and likely representing the subspecies C. hirta luimbalensis
(Crawford-Cabral and Veríssimo, unpublished data). In the south (Huíla and Cuando
Cubango) there were two specimens identified as Desert Lesser Red Shrew (C.
deserti), which may be a subspecies of Lesser Red Musk Shrew (Cassola 2016d).
Bicolored Musk Shrew (C. fruscomurina) has been recorded from Bengo/Luanda,
Cuanza-Norte, Cuanza-Sul, Malanje, Lunda-Norte, Huambo, Huíla, Namibe and
Cuando Cubango, and is thus thought to occur throughout the country (Crawford-
Cabral and Veríssimo, unpublished data).
Both Greater Dwarf Shrew (S. lixus) and Climbing Shrew (S. megalura) are
known to occur in Angola (Happold and Happold 2013; Crawford-Cabral and
Veríssimo, unpublished data). Greater Dwarf Shrew is known to occur in the east,
with records in Lunda-Norte (Heim de Balzac and Meester 1977). Climbing Shrew
has only been reported in Angola from Cuanza-Sul and Lunda-Norte provinces, yet
very scarcely, but it may occur throughout most of northern Angola. Recent field
collections in the Okavango source lakes area in 2016 added an additional species
for Angola, the Lesser Dwarf Shrew (S. varilla) (Taylor et al. 2018c). The species
has a sparse distribution and was previously known from southeast DRC so its
occurrence in central Angola is not surprising.
Tubulidentata (Aardvark)
The Aardvark (Orycteropus afer) is the only Tubulidentata occurring in Angola,

where it seems to be widespread, although it is known from relatively few and scat-
tered records due to its cryptic nature (Hill and Carter 1941; Crawford-Cabral and
Veríssimo 2005). No studies have focused on this species, and although it is often
hunted for bushmeat, it is likely not threatened due to its widespread distribution
and elusive habits. The species has also been frequently reported in general mam-
mal surveys conducted in protected areas in southern Angola (Veríssimo 2008;
Funston et al. 2017; Overton et al. 2017; NGOWP 2018).
Zoogeographic Outline
A quantitative regionalisation of Africa based on plants and vertebrates was carried

out by Linder et al. (2012), providing information on the biogeographic position of
Angola in the context of the African continent. In the analysis based on mammals,
this study located most of the country in a wide Zambezian band across south-
central Africa, crossing the continent from the Atlantic coast of Angola to the Indian
Ocean coasts of Tanzania and Mozambique. This band was bordered to the north by
the Guinean-Congolian region, corresponding to the tropical moist forests of the
Congo Basin and West Africa, which encompasses the enclave of Cabinda and a
narrow fringe in north and northeast Angola. To the south, the Zambezian region
was bordered by a Southern African Region, which extends into a narrow strip in
southern Angola. This rather coarse regionalisation was refined in analyses using a
dataset combining all plants and vertebrates. This analysis recognised a broad
southern transition zone (the Shaba subregion) between the Congolian and the
Zambezian regions, which forms an arc from the Angolan Atlantic coast to the
southern Ugandan uplands, and that expands southwards in Angola along the coastal
escarpment. Also, it split the Southern African Region in several subregions, includ-
ing the small biogeographic unit of Southwest Angola. In this new analysis the
Zambezian zone was bordered in the south by the Kalahari subregion.
Early efforts to undertake a zoogeographic analysis of Angola based on the
mammalian fauna were made by Crawford-Cabral (1982, 1997) and Feiler (1990).
More recently, the theme was revisited by Rodrigues et al. (2015), aiming to refine
the broad scale analysis of Linder et al. (2012) and to understand the environmental
determinants of biogeographic patterns (Fig. 15.2). The quantitative regionalisation
developed by Rodrigues et al. (2015) focused solely on Angola (excluding Cabinda)
and used data on ungulates (Cetartiodactyla, Perissodactyla, Hyracoidea), carni-
vores and small mammals available in the literature (Crawford-Cabral and Simões
1987, 1988; Crawford-Cabral 1998; Crawford-Cabral and Veríssimo 2005). The
study retrieved four main biogeographical units, which were particularly clear in the
analysis focusing on ungulates (Fig. 15.2): one northern region (Zaire-Lunda-
Cuanza), one central region (Central Plateau) and two regions in the south (Namibe
and Cunene-Cuando Cubango). This biogeographical pattern was strongly affected
by environmental factors, reflecting the dominant climate gradients in this region of
Africa (Le Houérou 2009), and the associated variation in soil and vegetation types.
The regions identified also closely matched the strong north–south gradient in
closed canopy forest cover (Hansen et al. 2013) with a progressive southwards tran-
sition to savannas (Murphy and Bowman 2012).
In the north, the Zaire-Lunda-Cuanza Region largely matched the Angolan por-
tion of the Shaba Region (Linder et al. 2012), and was mainly characterised by
indicator species that have their core range within the Congo Basin forests and
reach their southern limit in Angola, such as Forest Buffalo, Yellow-backed Duiker,
Blue Duiker, Black-fronted Duiker, and Bay Duiker. The Central Plateau corre-
sponds roughly to the Zambezian region of Linder et al. (2012) and encompasses to
412 P. Beja et al.
Fig. 15.2 Zoogeographic regions of Angola identified with a cluster analysis of quarter degree
grid cells (approximately 25 × 25 km) characterised using the occurrences of ungulate species
(Cetartiodactyla, Perissodactyla, Hyracoidea). Analysis used information documented by
Crawford-Cabral and Veríssimo (2005), corresponding to data collected mainly in 1930–1980. As
there was no information in many areas, grid cells were converted into a network of Thiessen
polygons, each containing the centre of a single grid cell with occurrence records for five or more
species (black dots). Colours are used to differentiate regions; A – Zaire-Lunda-Cuanza; B –
Central Plateau; C – Cunene-Cuando Cubango; D – Namibe. (Redrawn from Rodrigues et al.
(2015) and published under license by John Wiley and Sons)
a large extent the WWF Ecoregion of the Angolan miombo woodlands (Olson et al.
2001). It intergrades to some extent with the Zaire-Lunda-Cuanza region, suggest-
ing that there is a north–south gradient in mammalian assemblages, rather than two
well-defined regions. Indicator species of this region were Oribi, Roan Antelope,
Eland, Common Warthog, Bush Duiker, and the local endemic Giant Sable Antelope.
In the south, the Cunene-Cuando Cubango region was clearly identified in analy-
sis, corresponding to a savanna band running along the southern border of Angola
with Namibia, eastward of the Namibe region and continuing farther north along the
border with Zambia. The region corresponds to the northern limit of the Kalahari
subregion of Linder et al. (2012), though extending further north into Angola than
previously recognised. It also corresponds to the Zambezian Baikiaea woodlands
and the Angolan mopane woodlands defined by WWF (Olson et al. 2001). The indi-
cators of this region are species such as Wildebeest, Giraffe, Sable Antelope, Black
Rhino, Common Tsessebe, Buffalo, Hartebeest and Impala, which are widespread
in savannas of southern and eastern Africa, but that have restricted distributions
elsewhere in Angola. Finally, the Namibe region matches closely the south-western
Angola region of Linder et al. (2012), and the WWF Ecoregion of the Kaokoveld
desert (Olson et al. 2001), representing the northern part of the vast Namib Desert,
and the Namibian savanna woodlands. The main indicators of this region were spe-
cies such as Rock Hyrax, Yellow-spotted Rock Hyrax, Black-faced Impala, Kirk’s
Dik-dik, Klipspringer, Springbok, Gemsbok and Mountain Zebra.
Although this biogeographic regionalisation emerged from the analysis of occur-
rence data for ungulates, it is likely that similar patterns would be obtained for other
groups, though a quantitative analysis was not possible due to the scarcity of infor-
mation. However, the recent reviews on African bats and rodents by (Monadjem
et al. 2010a, 2015), Schoeman et al. (2013) and Cooper-Bohannon et al. (2016),
clearly suggest marked north-south distribution gradients, with species from
Guinean and Congolian tropical forests penetrating southwards in northern Angola,
species from arid areas occurring in the southwest, and savanna species occurring in
a southern band. The same seems to happen with other groups such as primates.
As described in ‘The Mammal Fauna’ section above, Angola has a number of
endemic and near endemic species, which are particularly numerous in the case of
rodents. The number of endemics recognised has been growing in recent years,
mainly due to the use of molecular techniques to understand phylogenetic relation-
ships among taxa, new field surveys, and the re-examination of specimens collected
several decades ago (Carleton et al. 2015; Svensson et al. 2017). In addition, there
are several endemic and near endemic subspecies, though their taxonomic status is
often uncertain. Finally, there are in Angola some isolated or otherwise disjunct
populations, often far from the main distribution range of the corresponding species,
which in the future may prove to warrant taxonomic recognition (Monadjem et al.
2010a, 2015). Despite this richness, however, there has been no systematic attempt
to identify the regions or habitats where such endemisms occur in Angola, or on the
phylogeographic processes that have driven their divergence from sister taxa.
Nevertheless, analogies with other taxonomic groups suggest that in Angola there
are well-defined regions that are centres of endemism, and thus merit further sur-
veys and conservation attention. These are mostly concentrated along the Angolan
Escarpment and Afromontane forests of western Angola, which are known to have
high prevalence of plant, invertebrate and vertebrate endemisms (Hall 1960;
Figueiredo et al. 2009; Clark et al. 2011; Mills et al. 2011, 2013), including mam-
mals (Carleton et al. 2015).
414 P. Beja et al.
Conservation
The global conservation status of 95.2% of the 290 native mammal species known
to occurr in Angola has been evaluated by IUCN (2018), of which there are 2
Critically Endangered, 2 Endangered, 11 Vulnerable, 14 Near-Threatened, and 12
Data Deficient species (Appendix). The order Carnivora has the largest number of
threatened species (5), but there are also threatened species in the orders Pholidota
(3), Cetartiodactyla (2), Perissodactyla (2), Primates (2) and Sirenia (1). The
Critically Endangered species are Black Rhinoceros, which is likely to be currently
extinct in the country, and Western Gorilla, which seems to be restricted to a very
small area in the Maiombe forests of Cabinda. There is also one Critically
Endangered subspecies, the Giant Sable Antelope, though its parental species is
considered Not Threatened. The Endangered species are the African Wild Dog,
which in Angola maintains some populations that may be relevant for species con-
servation at the global scale (Veríssimo 2008; Overton et al. 2017; Fabiano et al.
2017; Funston et al. 2017; Monterroso et al., unpublished data), and Chimpanzee,
which like the Western Gorilla is restricted to small areas in the forests of Cabinda.
The Vulnerable category includes a diverse array of species, including Cheetah,
African Golden Cat, Lion, Leopard, Giraffe, Hippopotamus, Mountain Zebra,
White-bellied Pangolin, Giant Ground Pangolin, Temminck’s Ground Pangolin and
African Manatee. Although classified as Vulnerable by IUCN, the Elephant was not
considered in this group because we assumed the scientifically well-established
division in two separate species, Forest and Savanna Elephants, which have hitherto
not been evaluated by IUCN. Given the precipituous decline of Forest Elephant due
to poaching and its very low intrinsic growth rate, the species may qualify to the
Endangered or even Critically Endangered status (e.g., Cerling et al. 2016; Poulsen
et al. 2017; Turkalo et al. 2017). Black-faced Impala is a subspecies considered
Vulnerable, though its parental species is classified as Not Threatened. Regarding
Near Threatened species, these include four carnivores (Brown Hyena, African
Clawless Otter, Congo Clawless Otter, Spotted-necked Otter), six Cetartiodactyla
(Bay Duiker, White-bellied Duiker, Yellow-backed Duiker, Waterbuck, Southern
Lechwe, Puku), three bats (Striped leaf-nosed Bat, Large-eared Free-tailed Bat,
Angolan Epauletted Fruit Bat), and one primate (Black Crested Mangabey).
There is very little information on the current status in Angola of most threatened
and near-threatened species, but many of them are feared extinct or at the brink of
extinction. This is mainly the case for large carnivores and herbivores, which were
heavily hunted during and in the years following the civil war, and for which virtu-
ally no information on distribution and numbers has been collected for over three
decades (Huntley 2017). A few recent surveys confirm this situation, showing that
many species once common in Angola only persist at present in remote areas, usu-
ally having small and fragmented populations (Veríssimo 2008; Overton et al. 2017;
Fabiano et al. 2017; Funston et al. 2017; Monterroso et al., unpublished data). This
is illustrated for instance by the critical condition of the iconic Giant Sable Antelope,
which has declined to very small numbers over the past decades (e.g., Vaz Pinto
et al. 2016 and Vaz Pinto 2019). Another iconic species, Savanna Elephant, also
appeared to have declined precipitously, even after the end of the civil war (Milliken
et al. 2006; Chase and Griffin 2011; Schlossberg et al. 2018), and the Forest Elephant
may be on the verge of extinction in the country. Poaching and habitat destruction
are likely the main threats, and a major cause of concern is the illegal trade of ivory,
channelled through the capital Luanda, and which is fuelling most of the poaching
activities (Milliken et al. 2006; Svensson et al. 2014b). African Manatee is another
globally vulnerable species that is likely on the verge of extinction in Angola, result-
ing from unsustainable hunting associated with the bushmeat trade (Morais et al.
2006a, b; Collins et al. 2011). Information is even scarcer for smaller and less char-
ismatic species, and it is noteworthy that many of the data deficient mammal taxa
listed by IUCN for Angola are endemics or near-endemics to the country. These
include for instance Lesser Angolan Epauletted Fruit Bat, Hayman’s Dwarf
Epauletted Fruit Bat, Angolan Long-eared Bat, Black-nosed Red-tailed Monkey,
Pluto Monkey, Southern Black Crested Mangabey, Angolan African Dormouse,
Monard’s Dormouse, Angolan Shaggy Rat, Vernay’s Climbing Mouse, Lunda Rope
Squirrel, and Heather Shrew, all of which have rather restricted distribution and may
qualify to a threatened or near-threatened category upon further investigation.
The problems affecting the conservation of biodiversity in Angola and some of
their potential solutions are discussed at length in Huntley et al. (2019). The same
considerations broadly apply to the mammal species. It should be noted, however,
that mammals will inevitably be at the forefront of biodiversity conservation in
Angola, as this group includes some of the most threatened species at the global
scale, as well as some of the species that have suffered the most from decades of
persecution and management neglect. Furthermore, the key threats to many of the
most endangered species are likely to continue or even intensify in the country
(Huntley 2017), due to a detrimental combination of factors such as commercial and
illegal wildlife trade, bushmeat harvesting, and habitat destruction through defores-
tation, agricultural expansion, and infrastructure development. At the same time,
however, there are encouraging signs for biodiversity conservation in Angola, many
of which involving efforts to preserve endangered mammals. Despite multiple prob-
lems, it has been possible to secure the critically endangered populations of the
Giant Sable (e.g., Vaz Pinto et al. 2016 and Vaz Pinto 2019), while new surveys have
revealed the potential of recovery of other iconic species such as Lion and Wild
Dog, among others (Veríssimo 2008; Overton et al. 2017; Fabiano et al. 2017;
Funston et al. 2017; Monterroso et al., unpublished data). For at least some of these
species, Angola may play an important role for conservation efforts, by securing
relevant populations and thus reducing risks at the global scale (e.g., Riggio et al.
2013). Many of these species have persisted in areas that are now protected by
national legislation, and it is expected that current conservation efforts, albeit yet
modest, will help them to recover in range and numbers. It is now necessary to move
forward, increasing the conservation efforts for these threatened species, at the same
time that new surveys are carried out to obtain a more complete appreciation of the
diversity and conservation needs of the overall mammal fauna of Angola (Fig. 15.3).
416 P. Beja et al.
Fig. 15.3 Angolan mammals. 1 Pack of African Wild Dogs (Lycaon pictus) in Luando Strict
Nature Reserve; 2 Herd of Forest Buffalo (Syncerus caffer nanus) in Quiçama National Park; 3
Yellow-backed Duiker (Cephalophus silvicultor ruficrista) in Luando Strict Nature Reserve; 4
Black Rhinoceros (Diceros bicornis bicornis) in Iona National Park; 5 Angolan Bush Hyrax
(Heterohyrax brucei bocagei) at Serra da Neve; 6 White-bellied Pangolin (Phataginus tricuspis) in
Cangandala National Park; 7 Angolan Dwarf Galago (Galagoides kumbirensis) in Cumbira Forest;
8 Pluto Monkey (Cercopithecus mitis mitis) in Quiçama National park. (Photo Credits: 1, 4 – Brian
J. Huntley, 2 – Merle Huntley: 1970s personal archive; 3, 5, 6, 8 – Pedro Vaz Pinto; 7 – Elena
Bersacola)
Appendix
Checklist of the native mammals of Angola. The table provides all species with
occurrence confirmed in the country (e.g., collected specimens, photos, reliable
sightings), as well as all subspecies (in grey font) confirmed in the country and that
are evaluated separately in the IUCN Red List (IUCN 2018). The Order, Family,
Latin name (author, date), English name, IUCN conservation status (CS), and key
references (Ref) confirming species presence are provided for each taxa. Taxa are
arranged by alphabetic order of Order, Family and Latin name
Species English name CSa Ref.b

Afrosoricida Chrysochloridae
Huetia leucorhina (Huet, 1885) Congo Golden Mole DD 21
Afrosoricida Tenrecidae
Potamogale velox (Du Chaillu, 1860) Giant Otter Shrew LC 3-5
Carnivora Canidae
Canis adustus (Sundevall, 1847) Side-striped Jackal LC 12
Canis mesomelas (Schreber, 1775) Black-backed Jackal LC 12
Lycaon pictus (Temminck, 1820) African Wild Dog EN 13
Otocyon megalotis (Desmarest, 1822) Bat-eared Fox LC 12
Vulpes chama (A. Smith, 1833) Cape Fox LC 12
Carnivora Felidae
Acinonyx jubatus (Schreber, 1775) Cheetah VU 13
Caracal aurata (Temminck, 1827) African Golden Cat VU 16
Caracal caracal (Schreber, 1776) Caracal LC 12
Felis silvestris (Schreber, 1777) Wild Cat LC 12
Leptailurus serval (Schreber, 1776) Serval LC 12
Panthera leo (Linnaeus, 1758) Lion VU 13
Panthera pardus (Linnaeus, 1758) Leopard VU 13
Carnivora Herpestidae
Atilax paludinosus (G.[Baron] Cuvier, 1829) Marsh Mongoose LC 12
Bdeogale nigripes (Pucheran, 1855) Black-legged Mongoose LC 12
Crossarchus ansorgei (Thomas, 1910) Ansorge’s Cusimanse LC 12
Cynictis penicillata (G.[Baron] Cuvier, 1829) Yellow Mongoose LC 12
Helogale parvula (Sundevall, 1847) Common Dwarf Mongoose LC 12
Herpestes flavescens (Bocage, 1889) Kaokoveld Slender Mongoose LC 10
Herpestes ichneumon (Linnaeus, 1758) Egyptian Mongoose LC 12
Herpestes sanguineus (Rüppell, 1835) Common Slender Mongoose LC 10
Ichneumia albicauda (G.[Baron] Cuvier, 1829) White-tailed Mongoose LC 12
Mungos mungo (Gmelin, 1788) Banded Mongoose LC 12
Paracynictis selousi (de Winton, 1896) Selous’s Mongoose LC 12
Suricata suricatta (Schreber, 1776) Meerkat LC 12
Carnivora Hyaenidae
Crocuta crocuta (Erxleben, 1777) Spotted Hyaena LC 13
(continued)
418 P. Beja et al.

Parahyaena brunnea (Thunberg, 1820) Brown Hyaena NT 13
Proteles cristata (Sparrman, 1783) Aardwolf LC 13
Carnivora Mustelidae
Aonyx capensis (Schinz, 1821) African Clawless Otter NT 12
Aonyx congicus (Lönnberg, 1910) Congo Clawless Otter NT 12
Hydrictis maculicollis (Lichtenstein, 1835) Spotted-necked Otter NT 12
Ictonyx striatus (Perry, 1810) Striped Polecat LC 12
Mellivora capensis (Schreber, 1776) Honey Badger LC 12
Poecilogale albinucha (Gray, 1864) African Striped Weasel LC 12
Carnivora Nandiniidae
Nandinia binotata (Gray, 1830) African Palm Civet LC 12
Carnivora Otariidae
Arctocephalus pusillus Brown Fur Seal LC 36
Carnivora Viverridae
Civettictis civetta (Schreber, 1776) African Civet LC 12
Genetta angolensis Bocage, 1882 Miombo Genet LC 12
Genetta genetta (Linnaeus, 1758) Common Genet LC 12
Genetta maculata (Gray, 1830) Large-spotted Genet LC 12
Cetartiodactyla Bovidae
Aepyceros melampus (Lichtenstein, 1812) Common Impala LC 14
Aepyceros melampus ssp. melampus (Lichtenstein, Common Impala LC 14
1812)
Aepyceros melampus ssp. petersi (Bocage, 1879) Black-faced Impala VU 14
Alcelaphus buselaphus (Pallas, 1766) Hartebeest LC 14
Alcelaphus buselaphus ssp. lichtensteinii (Peers, Lichtenstein’s Hartebeest LC 14
1849)
Alcelaphus buselaphus ssp. caama (É. Geoffroy Red Hartebeest LC 14
Saint-Hilaire, 1803)
Antidorcas marsupialis (Zimmermann, 1780) Springbok LC 14
Cephalophus dorsalis (Gray, 1846) Bay Duiker NT 14
Cephalophus leucogaster (Gray, 1873) White-bellied Duiker NT 14
Cephalophus nigrifrons (Gray, 1871) Black-fronted Duiker LC 14
Cephalophus silvicultor (Afzelius, 1815) Yellow-backed Duiker NT 14
Connochaetes taurinus (Burchell, 1823) Common Wildebeest LC 14
Damaliscus lunatus (Burchell, 1823) Topi LC 14
Hippotragus equinus (É. Geoffroy Saint-Hilaire, Roan Antelope LC 14
1803)
Hippotragus niger (Harris, 1838) Sable Antelope LC 14
Hippotragus niger ssp. variani (Thomas, 1916) Giant Sable Antelope CR 14
Kobus ellipsiprymnus (Ogilbyi, 1833) Waterbuck NT 14
Kobus ellipsiprymnus ssp. defassa (Ruppell, 1835) Defassa Waterbuck NT 14
Kobus ellipsiprymnus ssp. ellipsiprymnus (Ogilbyi, Common Waterbuck LC 14
1833)
Kobus leche (Gray, 1850) Southern Lechwe NT 14
Kobus leche leche (Gray, 1850) Red Lechwe NT 14
(continued)

Kobus vardonii (Livingstone, 1857) Puku NT 14
Madoqua kirkii (Günther, 1880) Kirk’s Dik-dik LC 14
Oreotragus oreotragus (Zimmermann, 1783) Klipspringer LC 14
Oryx gazella (Linnaeus, 1758) Gemsbok LC 14
Ourebia ourebi (Zimmermann, 1783) Oribi LC 14
Philantomba monticola (Thunberg, 1789) Blue Duiker LC 14
Raphicerus campestris (Thunberg, 1811) Steenbok LC 14
clinginRedunca arundinum (Boddaert, 1785) Southern Reedbuck LC 14
Sylvicapra grimmia (Linnaeus, 1758) Common Duiker LC 14
Syncerus caffer (Sparrman, 1779) Forest Buffalo LC 14
Tragelaphus oryx (Pallas, 1766) Common Eland LC 14
Tragelaphus scriptus (Pallas, 1766) Bushbuck LC 14
Tragelaphus spekii (Speke, 1863) Sitatunga LC 14
Tragelaphus strepsiceros (Pallas, 1766) Greater Kudu LC 14
Cetartiodactyla Giraffidae
Giraffa camelopardalis (Linnaeus, 1758) Giraffe VU 14
Cetartiodactyla Hippopotamidae
Hippopotamus amphibius (Linnaeus, 1758) Hippopotamus VU 14
Cetartiodactyla Suidae
Phacochoerus africanus (Gmelin, 1788) Common Warthog LC 14
Potamochoerus larvatus (F. Cuvier, 1822) Bushpig LC 14
Potamochoerus porcus (Linnaeus, 1758) Red River Hog LC 14
Cetartiodactyla Tragulidae
Hyemoschus aquaticus (Ogilby, 1841) Water Chevrotain LC 14
Chiroptera Emballonuridae
Coleura afra (Peters, 1852) African Sheath-tailed Bat LC 8, 29
Saccolaimus peli (Temminck 1853) Pel’s Pouched Bat LC 8, 29
Taphozous mauritianus (E. Geoffroy, 1818) Mauritian Tomb Bat LC 8, 29
Chiroptera Hipposideridae
Hipposideros caffer (Sundevall, 1846) Sundevall’s Leaf-nosed Bat LC 8, 29
Hipposideros vittatus (Peters, 1852) Striped Leaf-nosed Bat NT 8, 29
Hipposideros ruber (Noack, 1893) Noack’s leaf-nosed Bat LC 8, 29
Macronycteris gigas (Wagner, 1845) Giant Leaf-nosed Bat LC 8, 29
Triaenops afer (Peters, 1877) African Trident Bat LC 20
Chiroptera Minipteridae
Miniopterus natalensis (A. Smith 1833) Natal Long-fingered Bat LC 8,29
Chiroptera Molossidae
Chaerephon ansorgei (Thomas, 1913) Ansorge’s Free-tailed Bat LC 8, 29
Chaerephon chapini (J.A. Allen, 1917) Pale Free-tailed Bat LC 8, 29
Chaerephon nigeriae (Thomas, 1913) Nigerian Free-tailed Bat LC 8, 29
Chaerephon pumilus (Cretzschmar, 1826) Little Free-tailed Bat LC 8, 29
Mops condylurus (A. Smith, 1833) Angolan Free-tailed Bat LC 8, 29
Mops midas (Sundevall, 1843) Midas Free-tailed Bat LC 8, 29
Mops niveiventer (Cabrera and Ruxton, 1926) White-bellied Free-tailed Bat LC 8, 29
(continued)
420 P. Beja et al.

Otomops martiensseni (Matschie, 1897) Large-eared Free-tailed Bat NT 8, 29
Sauromys petrophilus (Roberts, 1917) Roberts’s Flat-headed Bat LC 8
Tadarida aegyptiaca (E. Geoffroy St.-Hilaire, Egyptian Free-tailed Bat LC 8, 29
1818)
Chiroptera Nycteridae
Nycteris arge (Thomas, 1903) Bate’s Slit-faced Bat LC 8, 29
Nycteris hispida (Schreber, 1775) Hairy Slit-faced Bat LC 8, 29
Nycteris intermedia (Aellen, 1959) Intermediate Slit-faced Bat LC 8, 29
Nycteris macrotis (Dobson, 1876) Large-eared Slit-faced Bat LC 8, 29
Nycteris nana (K. Andersen, 1912) Dwarf Slit-faced Bat LC 8, 29
Nycteris thebaica (E. Geoffroy St.-Hilaire, 1818) Egyptian Slit-faced Bat LC 8, 29
Chiroptera Pteropodidae
Eidolon helvum (Kerr, 1792) Straw-coloured Fruit Bat LC 8, 29
Epomophorus angolensis (Gray, 1870) Angolan Epauletted Fruit Bat NT 8, 29
Epomophorus crypturus (Peters, 1852) Peters’s Epauletted Fruit Bat LC 8, 29
Epomophorus grandis (Sanborn, 1950) Lesser Angolan Epauletted DD 8, 29
Fruit Bat
Epomophorus wahlbergi (Sundevall, 1846) Wahlberg’s Epauletted Fruit LC 8, 29
Bat
Epomops dobsoni (Bocage, 1889) Dobson’s Epauletted Fruit Bat LC 8, 29
Epomops franqueti (Tomes, 1860) Franquet’s Epauletted Fruit LC 8, 29
Bat
Hypsignathus monstrosus (H. Allen, 1862) Hammer-headed Fruit Bat LC 8, 29
Megaloglossus woermanni (Pagenstecher, 1885) Woermann’s Long-tongued LC 8, 29
Fruit Bat
Micropteropus intermedius (Hayman, 1963) Hayman’s Dwarf Epauletted DD 8, 29
Fruit Bat
Micropteropus pusillus (Peters, 1868) Peters’s Dwarf Epauletted LC 8, 29
Fruit Bat
Myonycteris angolensis (Bocage, 1898) Angolan Soft-furred Fruit Bat LC 8, 29
Myonycteris torquata (Dobson, 1878) Little Collared Fruit Bat LC 8, 29
Plerotes anchietae (Seabra, 1900) Anchieta’s Fruit Bat DD 8, 29
Rousettus aegyptiacus (E. Geoffroy St.-Hilaire, Egyptian Rousette LC 8, 29
1810)
Chiroptera Rhinolophidae
Rhinolophus damarensis (Roberts, 1946) Damara Horseshoe Bat LC 1
Rhinolophus denti (Thomas, 1904) Dent’s Horseshoe Bat LC 8
Rhinolophus eloquens (K. Andersen, 1905) Eloquent Horseshoe Bat LC 19
Rhinolophus fumigatus (Rüppell, 1842) Rüppell’s Horseshoe Bat LC 8, 29
Rhinolophus lobatus (Peters, 1852) Peters’s Horseshoe Bat NE 33
Chiroptera Vespertilionidae
Cistugo seabrai (Thomas, 1912) Angolan Hairy Bat LC 8, 29
Eptesicus hottentotus (A. Smith, 1833) Long-tailed Serotine LC 8, 29
Glauconycteris argentata (Dobson, 1875) Common Butterfly Bat LC 8, 29
(continued)

Glauconycteris beatrix (Thomas, 1901) Beatrix’s Butterfly Bat LC 8, 29
Glauconycteris variegata (Tomes, 1861) Variegated Butterfly Bat LC 8, 29
Hypsugo anchietae (Seabra, 1900) Anchieta’s Pipistrelle LC 8, 29
Hypsugo crassulus (Thomas, 1904) Broad-headed Pipistrelle LC 20
Kerivoula argentata (Tomes, 1861) Damara Woolly Bat LC 34
Kerivoula lanosa (A. Smith, 1847) Lesser Woolly Bat LC 29
Laephotis angolensis (Monard, 1935) Angolan Long-eared Bat DD 8, 29
Laephotis botswanae (Setzer, 1971) Botswana Long-eared Bat LC 26
Mimetillus moloneyi (Thomas, 1891) Moloney’s Flat-headed Bat LC 20
Mimetillus thomasi (Hinton, 1920) Thomas’s Flat-headed Bat NE 29
Myotis bocagii (Peters, 1870) Bocage’s Mouse-eared Bat LC 8, 29
Myotis welwitschii (Gray, 1866) Welwitsch’s Mouse-eared Bat LC 8, 29
Neoromicia capensis (A. Smith, 1829) Cape Serotine LC 8, 29
Neoromicia grandidieri (Dobson, 1876) Dobson’s Pipistrelle DD 29
Neoromicia nana (Peters, 1852) Banana Bat LC 8, 29
Neoromicia tenuipinnis (Peters, 1872) White-winged Serotine LC 8, 29
Neoromicia zuluensis (Roberts, 1924) Zulu Serotine LC 29
Nycticeinops schlieffeni (Peters, 1859) Schlieffen’s Bat LC 8, 29
Pipistrellus hesperidus (Temminck, 1840) Dusky Pipistrelle LC 20
Pipistrellus rueppellii (J. Fischer, 1829) Rüppell’s Pipistrelle LC 8, 29
Pipistrellus rusticus (Tomes, 1861) Rusty Pipistrelle LC 34
Scotoecus hindei (Thomas, 1901) Thomas’s House Bat NE 29
Scotophilus dinganii (A. Smith, 1833) Yellow-bellied House Bat LC 8, 29
Scotophilus leucogaster (Cretzschmar, 1826) White-bellied House Bat LC 29
Scotophilus viridis (Peters, 1852) Green House Bat LC 8, 29
Erinaceomorpha Erinaceidae
Atelerix frontalis (A. Smith, 1831) Southern African Hedgehog LC 7
Hyracoidea Procaviidae
Dendrohyrax arboreus (A. Smith, 1827) Southern Tree Hyrax LC 14
Dendrohyrax dorsalis (Fraser, 1855) Western Tree Hyrax LC 14
Heterohyrax brucei (Gray, 1868) Bush Hyrax LC 14
Procavia capensis (Pallas, 1766) Kaokoveld Rock Dassie LC 14
Lagomorpha Leporidae
Lepus capensis (Linnaeus, 1758) Cape Hare LC 18
Lepus victoriae (Thomas, 1893) African Savanna Hare LC 31
Pronolagus randensis (Jameson, 1907) Jameson’s Red Rock Hare LC 31
Macroscelidea | Macroscelididae
Elephantulus brachyrhynchus (A. Smith, 1836) Short-snouted Elephant-shrew LC 15
Elephantulus intufi (A. Smith, 1836) Bushveld Elephant-shrew LC 15
Petrodromus tetradactylus (Peters, 1846) Four-toed Elephant-shrew LC 15
Perissodactyla Equidae
Equus quagga (Boddaert, 1785) Plains Zebra LC 14
Equus zebra (Linnaeus, 1758) Mountain Zebra VU 14
(continued)
422 P. Beja et al.

Equus zebra ssp. hartmannae (Matschie, 1898) Hartmann’s Mountain Zebra VU 14
Perissodactyla Rhinocerotidae
Diceros bicornis (Linnaeus, 1758) Black Rhino CR 14
Diceros bicornis ssp. bicornis (Linnaeus, 1758) South-western Black Rhino VU 14
Diceros bicornis ssp. minor (Drummond, 1876) Southern-central Black Rhino CR 14
Pholidota Manidae
Phataginus tricuspis (Rafinesque, 1821) White-bellied Pangolin VU 22
Smutsia gigantea (Illiger, 1815) Giant Ground Pangolin VU 22
Smutsia temminckii (Smuts, 1832) Temminck’s Ground Pangolin VU 25
Primates Cercopithecidae
Cercopithecus ascanius (Audebert, 1799) Red-tailed Monkey LC 27
Cercopithecus ascanius ssp. atrinasus (Machado, Black-nosed Red-tailed DD 27
1965) Monkey
Cercopithecus cephus (Linnaeus, 1758) Moustached Monkey LC 27
Cercopithecus mitis (Wolf, 1822) Blue Monkey LC 28
Cercopithecus mitis ssp. mitis (Wolf, 1822) Pluto Monkey DD 28
Cercopithecus mitis ssp. opisthostictus (Sclater, Rump-spotted Blue Monkey LC 28
1894)
Cercopithecus pogonias (Bennett, 1833) Crowned Monkey NE ?
Cercopithecus pogonias ssp. nigripes (Du Chaillu, Black-footed Crowned LC ?
1860) Monkey
Cercopithecus neglectus (Schlegel, 1876) De Brazza’s Monkey LC 27
Cercopithecus nictitans (Linnaeus, 1766) Putty-nosed Monkey LC 27
Chlorocebus cynosuros (Scopoli, 1786) Malbrouck Monkey LC 23
Colobus angolensis (P. Sclater, 1860) Angola Colobus LC 27
Colobus angolensis ssp. angolensis (P. Sclater, Sclater’s Angolan Colobus LC 27
1860)
Lophocebus aterrimus (Oudemans, 1890) Black Crested Mangabey NT 27
Lophocebus terrimus ssp. opdenboschi Southern Black Crested DD 27
(Schouteden, 1944) Mangabey
Miopithecus ogouensis (Kingdon, 1997) Northern Talapoin Monkey LC 17
Miopithecus talapoin (Schreber, 1774) Southern Talapoin Monkey LC 27
Papio kindae (Lönnberg, 1919) Kinda Baboon LC 27
Papio ursinus (Kerr, 1792) Chacma Baboon LC 27
Papio ursinus ssp. ursinus (Kerr, 1792) Southern Chacma Babbon LC 27
Primates Galagidae
Euoticus elegantulus (Le Conte, 1857) Southern Needle-clawed LC 24
Galago
Galago moholi (A. Smith, 1836) Southern Lesser Galago LC 2
Galagoides demidoff (G. Fischer, 1806) Demidoff’s Dwarf Galago LC 27
Galagoides kumbirensis (Svensson et al. 2017) Angolan Dwarf Galago NE 32
Galagoides thomasi (Elliot, 1907) Thomas’s Dwarf Galago LC 2
Otolemur crassicaudatus (É. Geoffroy Saint- Garnett’s Greater Galago LC 2
Hilaire, 1812)
(continued)

Primates Hominidae
Gorilla gorilla (Savage, 1847) Western Gorilla CR 24
Gorilla gorilla ssp. gorilla (Savage, 1847) Western Lowland Gorilla CR 24
Pan troglodytes (Blumenbach, 1799) Chimpanzee EN 24
Pan troglodytes ssp. troglodytes (Blumenbach, Central Chimpanzee EN 24
1799)
Primates Lorisidae
Arctocebus aureus de (Winton, 1902) Golden Potto LC 2
Perodicticus edwardsi (Bouvier, 1879) Milne-Edwards’s Potto LC 2
Proboscidea Elephantidae
Loxodonta africana (Blumenbach, 1797) Savanna Elephant NE 14
Loxodonta cyclotis (Matschie, 1900) Forest elephant NE 14
Rodentia Anomaluridae
Anomalurus beecrofti (Fraser, 1853) Beecroft’s Scaly-tailed LC 30
Squirrel
Anomalurus derbianus (Gray, 1842) Lord Derby’s Scaly-tailed LC 30
Squirrel
Rodentia Bathyergidae
Fukomys bocagei (de Winton, 1897) Bocage’s Mole Rat LC 30
Fukomys damarensis (Ogilby, 1838) Damara Mole Rat LC 30
Rodentia Gliridae
Graphiurus angolensis (de Winton, 1897) Angolan African Dormouse DD 30
Graphiurus kelleni (Reuvens, 1890) Kellen’s Dormouse LC 30
Graphiurus lorraineus (Dollman, 1910) Lorrain Dormouse LC 30
Graphiurus monardi (St. Leger, 1936) Monard’s Dormouse DD 30
Graphiurus rupicola (Thomas & Hinton, 1925) Stone Dormouse LC 30
Rodentia Hystricidae
Hystrix africaeaustralis (Peters, 1852) Cape Porcupine LC 30
Rodentia Muridae
Aethomys bocagei (Thomas, 1904) Bocage’s Rock Rat LC 11,
30
Aethomys chrysophilus (de Winton, 1897) Red Rock Rat LC 11,
30
Aethomys kaiseri (Noack, 1887) Kaiser’s Rock Rat LC 11,
30
Aethomys nyikae (Thomas, 1897) Nyika Rock Rat LC 30
Aethomys thomasi (de Winton, 1897) Thomas’s Rock Rat LC 11,
30
Colomys goslingi (Thomas & Wroughton, 1907) African Wading Rat LC 11,
30
Dasymys cabrali (Verheyen et al. 2003) Cabral’s Marsh Rat NE 30
Dasymys cf incomtus African Marsh Rat LC 30
Dasymys nudipes (Peters, 1870) Angolan Shaggy Rat DD 11,
30
(continued)
424 P. Beja et al.

Desmodillus auricularis (A. Smith, 1834) Cape Short-eared Gerbil LC 11,
30
Gerbilliscus brantsii (A. Smith, 1836) Highveld Gerbil LC 11,
30
Gerbilliscus leucogaster (Peters, 1852) Bushveld Gerbil LC 11,
30
Gerbilliscus paeba (A. Smith, 1836) Hairy-footed Gerbil LC 11,
30
Gerbilliscus setzeri (Schlitter, 1973) Setzer’s Hairy-footed Gerbil LC 11,
30
Gerbilliscus validus (Bocage, 1890) Savanna Gerbil LC 11,
30
Grammomys dolichurus (Smuts, 1832) Woodland Thicket Rat LC 11,
30
Grammomys poensis (Eisentraut, 1965) Shining Thicket Rat NE 11,
30
Hylomyscus carillus (Thomas, 1904) Angolan Wood Mouse LC 11,
30
Hylomyscus heinrichorum (Carleton et al. 2015) Heirich’s Hylomyscus NE 6
Lemniscomys griselda (Thomas, 1904) Griselda’s Striped Grass LC 11
Mouse
Lemniscomys striatus (Linnaeus, 1758) Typical Striped Grass Mouse LC 11,
30
Lophuromys angolensis (Verheyen et al. 2000) Angolan’s Brush-furred Rat NE 30
Lophuromys rita (Dollman, 1910) Dollman’s Brush-furred Rat NE 30
Malacomys longipes (Milne-Edwards, 1877) Big-eared Swamp Rat LC 11,
30
Mastomys natalensis (Smith, 1834) Natal Multimammate Mouse LC 11,
30
Mastomys shortridgei (St. Leger, 1933) Shortridge’s Multimammate LC 11,
Mouse 30
Micaelamys namaquensis (A. Smith, 1834) Namaqua Rock Rat LC 11,30
Mus callewaerti (Thomas, 1925) Callewaert’s Mouse LC 11,30
Mus indutus (Thomas, 1910) Desert Pygmy Mouse LC 30
Mus minutoides (Smith, 1834) Pygmy Mouse LC 30
Mus setzeri (Petter, 1978) Setzer’s Mouse LC 30
Mus sorella (Thomas, 1909) Thoma’s Mouse LC 30
Mus triton (Thomas, 1909) Gray-bellied Pygmy Mouse LC 11,
30
Myomyscus angolensis (Bocage, 1890) Angolan Multimammate LC 11,
Mouse 30
Oenomys hypoxanthus (Pucheran, 1855) Rufous-nosed Rat LC 11,
30
Otomys anchietae (Bocage, 1882) Angolan Vlei Rat LC 11,
30
Otomys angoniensis (Wroughton, 1906) Angoni Vlei Rat LC 11,
30
(continued)

Otomys cuanzensis (Hill & Carter, 1937) Kuanza Vlei Rat LC 30
Pelomys campanae (Huet, 1888) Bell Groove-toothed Swamp LC 11,
Rat 30
Pelomys fallax (Peters, 1852) Creek Groove-toothed Swamp LC 11,
Rat 30
Pelomys minor (Cabrera & Ruxton, 1926) Least Groove-toothed Swamp LC 11,
Rat 30
Praomys coetzeei (Van der Straeten, 2008) Coetzee Praomys NE 30,
35
Praomys jacksoni (de Winton, 1897) Jackson’s Soft-furred Mouse LC 11,
30
Rhabdomys bechuanae (Thomas, 1893) Thoma’s Four-striped Grass NE 30
Mouse
Rhabdomys dilectus (de Winton, 1897) Mesic Four-striped Grass Rat NE 11,
30
Thallomys nigricauda (Thomas, 1882) Black-tailed Tree Rat LC 11,
30
Zelotomys hildegardeae (Thomas, 1902) Hildegarde’s Broad-headed LC 11,
Mouse 30
Zelotomys woosnami (Schwann, 1906) Woosnam’s Broad-headed LC 34
Mouse
Rodentia Nesomydae
Cricetomys ansorgei (Thomas, 1904) Southern Giant Pouched Rat LC 30
Cricetomys emini (Wroughton, 1910) Forest Giant Pouched Rat LC 30
Dendromus leucostomus (Monard, 1933) Gray African Climbing Mouse LC 30
Dendromus melanotis (A. Smith, 1834) Gray African Climbing Mouse LC 30
Dendromus mystacalis (Heuglin, 1863) Chestnut Climbing Mouse LC 30
Dendromus nyikae (Wroughton, 1909) Nyika Climbing Mouse LC 30
Dendromus vernayi (Hill & Carter, 1937) Vernay’s Climbing Mouse DD 30
Malacothrix typica (A. Smith, 1834) Gerbil Mouse LC 30
Petromyscus collinus (Thomas & Hinton, 1925) Pygmy Rock Mouse LC 30
Petromyscus shortridgei (Thomas, 1926) Shortridge’s Rock Mouse LC 30
Saccostomus campestris (Peters, 1846) Southern African Pouched LC 30
Mouse
Steatomys bocagei (Thomas, 1892) Bocage’s Fat Mouse LC 30
Steatomys krebsii (Peters, 1852) Kreb’s Fat Mouse LC 30
Steatomys parvus (Rhoads, 1896) Tiny Fat Mouse LC 30
Steatomys pratensis (Peters, 1846) Fat Mouse LC 30
Rodentia Pedetidae
Pedetes capensis (Forster, 1778) Spring Hare LC 30
Rodentia Petromuridae
Petromus typicus (A. Smith, 1831) Dassie Rat LC 30
Rodentia Sciuridae
Funisciurus bayonii (Bocage, 1890) Lunda Rope Squirrel DD 30
(continued)
426 P. Beja et al.

Funisciurus congicus (Kuhl, 1820) Congo Rope Squirrel LC 30
Funisciurus lemniscatus (Le Conte, 1857) Ribboned Rope Squirrel LC 30
Funisciurus pyrropus (F. Cuvier, 1833) Fire-footed Rope Squirrel LC 30
Heliosciurus gambianus (Ogilby, 1835) Gambian Sun Squirrel LC 30
Paraxerus boehmi (Reichenow, 1886) Boehm’s Bush Squirrel LC 30
Paraxerus cepapi (A. Smith, 1836) Smith’s Bush Squirrel LC 30
Protoxerus stangeri (Waterhouse, 1842) African Giant Squirrel LC 30
Xerus princeps (Thomas, 1929) Damara Ground Squirrel LC 30
Rodentia Thryonomydae
Thryonomys swinderianus (Temminck, 1827) Greater Cane Rat LC 30
Sirenia Trichechidae
Trichechus senegalensis (Link, 1795) African Manatee VU 14
Soricomorpha Soricidae
Crocidura cyanea (Duvernoy, 1838) Reddish-gray Musk Shrew LC 25
Crocidura erica (Dollman, 1915) Heather Shrew DD 15
Crocidura fuscomurina (Heuglin, 1865) Bicolored Musk Shrew LC 15
Crocidura hirta (Peters, 1852) Lesser Red Musk Shrew LC 15
Crocidura luna (Dollman, 1910) Greater Gray-brown Musk LC 21
Shrew
Crocidura mariquensis (A. Smith, 1844) Swamp Musk Shrew LC 15
Crocidura nigricans (Bocage, 1889) Blackish White-toothed LC 9
Shrew
Crocidura nigrofusca (Matschie, 1895) African Black Shrew LC 15
Crocidura olivieri (Lesson, 1827) African giant shrew LC 15
Crocidura parvipes (Osgood, 1910) Small-footed Shrew LC 15
Crocidura roosevelti (Heller, 1910) Roosvelt’s Shrew LC 25
Crocidura turba Dollman, 1910 Turbo Shrew LC 15
Suncus lixus (Thomas, 1898) Greater Dwarf Shrew LC 15
Suncus megalura (Jentink, 1888) Climbing Shrew LC 15
Suncus varilla (Thomas, 1895) Lesser Dwarf Shrew LC 34
Tubulidentata Orycteropodidae
Orycteropus afer (Pallas, 1766) Aardvark LC 14
a
IUCN Conservation Status categories. CR critically endangered, EN endangered, VU vulnerable,
NT near threatened, LC least concern, DD data deficient, NE not evaluated
b
References – 1. Monadjem et al. (2017); 2. Bersacola et al. (2015); 3. du Bocage (1865); 4. du
Bocage (1882); 5. du Bocage (1890); 6. Carleton et al. (2015); 7. Cassola (2016b); 8. Crawford-
Cabral (1986); 9. Crawford-Cabral (1987); 10. Crawford-Cabral (1996); 11. Crawford-Cabral
(1998); 12. Crawford-Cabral and Simões (1987); 13: Crawford-Cabral and Simões (1988); 14.
Crawford-Cabral and Veríssimo (2005); 15. Crawford-Cabral and Veríssimo, unpublished data; 16:
Errol de Beer, unpublished data; 17. Gautier-Hion (2013b); 18. Genest-Villard (1969); 19. Grant
and Ferguson (2018); 20. Happold and Happold (2013); 21. Hayman (1963); 22. Hill and Carter
(1941); 23. Huntley (1973c); 24. Huntley (1973e); 25. IUCN RedList Map; 26. Monadjem et al.
(2010a); 27. Machado (1969); 28. Machado and Crawford-Cabral (1999); 29. Monadjem et al.
(2010a); 30. Monadjem et al. (2015); 31. Smith et al. (2018); 32. Svensson et al. (2017); 33. Taylor
et al. (2018c); 34. Taylor (in press); 35. Van der Straeten (2008); 36. Meÿer (2007)
References
Adams WB (2013) Against extinction: the story of conservation. Routledge, London

Ambrose L, Butynski TM (2013) Galagoides demidovii Demidoff’s Dwarf Galago. In: Butynski
TM, Kingdon J, Kalina J (eds) The mammals of Africa. Vol II (Primates). Bloomsbury
Publishing, London, pp 459, 556 pp–461
Angelici FM, Do Linh San E (2015) Crossarchus ansorgei. The IUCN Red List of threatened spe-
cies 2015:e.T41594A45205422. Downloaded on 10 April 2018
Angelici FM, Gaubert P, Do Linh San E (2016) Genetta maculata. The IUCN Red List of threat-
ened species 2016:e.T41699A45218948. Downloaded on 12 April 2018
Ansell WFH (1972) Part 15. Order Artiodactyla. In: Meester JA, Setzer HW (eds) The mammals
of Africa: an identification manual. Smithsonian Institution Press, Washington, DC, pp 1, 432
pp–93
Anstey S (1991) Plano de Conservação do Elefante para Angola. Unpublished report. Ministério
da Agricultura. Instituto de Desenvolvimento Florestal, Luanda
Anstey S (1993) Angola: elephants, people and conservation – a preliminary assessment of the
status and conservation of elephants in Angola. Unpublished report. IUCN Regional Office for
Southern Africa, Harare
Bahaa-el-din L, Mills D, Hunter L, Henschel P (2015) Caracal aurata. The IUCN Red List of
threatened species 2015:e.T18306A50663128. Downloaded on 12 April 2018
Barnett R, Yamaguchi N, Shapiro B et al (2014) Revealing the maternal demographic history of
Panthera leo using ancient DNA and a spatially explicit genealogical analysis. BMC Evolut
Biol 14:70
Bauer H, Chapron G, Nowell K et al (2015) Lion (Panthera leo) populations are declining rapidly
across Africa, except in intensively managed areas. Proc Natl Acad Sci U S A 112:14894–14899
Baxter R, Hutterer R, Griffin M, Howell K (2016) Crocidura cyanea (errata version published in
2017). The IUCN Red List of threatened species 2016:e.T40625A115176043. Downloaded on
10 May 2018
Bearder S (2008) Otolemur crassicaudatus. The IUCN Red List of threatened species 2008:e.
T15643A4943752. Downloaded on 20 March 2018
Bearder SK, Svoboda NS (2013) Otolemur crassicaudatus Large-eared Greater Galago. In:
Butynski TM, Kingdon J, Kalina J (eds) The mammals of Africa. Vol II (Primates). Bloomsbury
Bearder SK, Ambrose L, Harcourt C et al (2003) Species-typical patterns of infant contact, sleep-
ing site use and social cohesion among nocturnal primates in Africa. Folia Primatologica
74:337–254
Bearder S, Butynski TM, Hoffmann M (2008) Galago moholi. The IUCN Red List of threatened
species 2008:e.T8788A12932349. Downloaded on 20 March 2018
Benda P, Vallo P (2009) Taxonomic revision of the genus Triaenops (Chiroptera: Hipposideridae)
with description of a new species from southern Arabia and definitions of a new genus and
tribe. Folia Zoologica 58(Monograph 1):1–45
Bersacola E, Svensson MS, Bearder SK et al (2014) Hunted in Angola: surveying the bushmeat
trade. SWARA 2014(January–March):31–36
Bersacola E, Svensson MS, Bearder SK (2015) Niche partitioning and environmental factors
affecting abundance of strepsirrhines in Angola. Am J Primatol 77:1179–1192
Bessa J, Sousa C, Hockings KJ (2015) Feeding ecology of chimpanzees (Pan troglodytes verus)
inhabiting a forest-mangrove-savanna-agricultural matrix at Caiquene-Cadique, Cantanhez
National Park, Guinea-Bissau. Am J Primatol 77:651–665
Biggs D, Courchamp F, Martin R et al (2013) Legal trade of Africa’s rhino horns. Science
339:1038–1039
Blaine G (1922) Notes on the Zebras and some Antelopes of Angola. J Zool 92:317–339
Blaine G (1925) New subspecies of Connochaetes taurinus. Ann Mag Nat Hist 15(Series
9):129–130
428 P. Beja et al.
Boesch C, Boesch-Achermann H (2000) The Chimpanzees of the Taï Forest: behavioural ecology
and evolution. Oxford University Press, New York, 328 pp
Bohm T, Höner OR (2015) Crocuta crocuta. The IUCN Red List of threatened species 2015:e.
T5674A45194782. Downloaded on 11 April 2018
Bohm C, Jonsson C (2017) Vertebrates of the Gothenburg Natural History Museum (GNM).
Version 4.2. Gothenburg Natural History Museum, Gothenburg Accessed via GBIF.org on 24
Feb 2018
Brashares JS, Arcese P (2013) Ourebia ourebi Oribi. In: Kingdon J, Hoffmann M (eds) The
mammals of Africa. Vol VI (Hippopotamuses, pigs, deer, giraffe and bovids). Bloomsbury
Broom J, Milton S, Davis C et al (1974) Expedition to South-Western Angola June/July 1974.
Unpublished report. University of Cape Town/Wild Life Society, Cape Town
Brotherton PNM (2013) Madoqua (kirkii) Kirk’s Dik-Dik Species Group. In: Kingdon J, Hoffmann
M (eds) The mammals of Africa. Vol VI (Hippopotamuses, pigs, deer, giraffe and bovids).
Bloomsbury Publishing, London, pp 327, 680 pp–333
Butynski TM (2008) Chlorocebus cynosuros. The IUCN Red List of threatened species 2008:e.
Butynski TM, De Jong YA (2007) Distribution of the potto Perodicticus potto (Primates: Lorisidae)
in Eastern Africa, with a description of a new subspecies from Mount Kenya. J East Afr Nat
Hist 96:113–147
Carleton MD, Banasiak RA, Stanley WT (2015) A new species of the rodent genus Hylomyscus
from Angola, with a distributional summary of the H. anselli species group (Muridae: Murinae:
Praomyini). Zootaxa 4040:101–128
Carmignani E (2015) Elephant assessment and report on the status on Quissama National Park’s
Special Conservation Area October – November 2015. Unpublished report. Instituto Nacional
de Biodiversidade e Áreas de Conservação, Luanda
Carr T, Carr N, David JHM (1985) A record of the sub-Antarctic fur seal Arctocephalus tropicalis
in Angola. Afr Zool 20:77
Cassola F (2016a) Petromus typicus. The IUCN Red List of threatened species 2016:e.
T16776A22240649. Downloaded on 19 April 2018
Cassola F (2016b) Atelerix frontalis (errata version published in 2017). The IUCN Red List of
Cassola F (2016c) Crocidura olivieri (errata version published in 2017). The IUCN Red List of
threatened species 2016:e.T41348A115180235. Downloaded on 01 May 2018
Cassola F (2016d) Crocidura hirta (errata version published in 2017). The IUCN Red List of
threatened species 2016:e.T41323A115178068. Downloaded on 10 May 2018
Cerling TE, Barnette JE, Chesson LA et al (2016) Radiocarbon dating of seized ivory confirms
rapid decline in African elephant populations and provides insight into illegal trade. Proc Natl
Acad Sci U S A 113:13330–13335
Chardonnet P, Crosmary W (2013) Hippotragus equinus Roan Antelope. In: Kingdon J, Hoffmann
Chase MJ, Griffin CR (2011) Elephants of south-east Angola in war and peace: their decline, re-
colonization and recent status. Afr J Ecol 49:353–361
Chase MJ, Schlossberg S (2016) Dry-season fixed-wing aerial survey of elephants and other large
mammals in Southeast Angola. Unpublished report. Elephants Without Borders, Kasane,
Botswana
Ciofolo I, Le Pendu Y (2013) Giraffa camelopardalis Giraffe. In: Kingdon J, Hoffmann M (eds)
The mammals of Africa. Vol VI (Hippopotamuses, pigs, deer, giraffe and bovids). Bloomsbury
for biodiversity exploration. Biodiver Conserv 20:2543
Collins T, Keith L, Rosembaum H (2011) Inshore and Congo River Marine Mammals. Final
Report. Unpublished report, Ocean Giants Program. Wildlife Conservation Society, New York
Conroy C (2018) MVZ mammal collection (Arctos). Version 35.13. Museum of Vertebrate
Zoology, Berkeley. Accessed via GBIF.org on 2018-04-19
Cooper-Bohannon R, Rebelo H, Jones G et al (2016) Predicting bat distributions and diversity
hotspots in southern Africa. Hystrix 27:38–48
Crawford-Cabral J (1961) Considerações em torno de Equus quagga intermedia Taborda Morais.
Boletim do Instituto de Angola 15:77–79
Crawford-Cabral J (1966a) Some new data on Angolan Muridae. Zoologica Africana 2:193–203
Crawford-Cabral J (1966b) Quatro formas de mamíferos novas para Angola. Boletim do Instituto
de Investigação Científica de Angola 3:137–148
Crawford-Cabral J (1967) Mamíferos da Reserva do Luando. Boletim do Instituto de Investigação
Científica de Angola 4:33–44
Crawford-Cabral J (1968) Notas sobre a variação geográfica da pelagem de alguns carnívoros.
Boletim do Instituto de Investigação Científica de Angola 5:15–122
Crawford-Cabral J (1969a) As Genetas de Angola. Boletim do Instituto de Investigação Científica
de Angola 6:25–26
Crawford-Cabral J (1969b) A study of the Giant Sable (Hippotragus niger variani). News Bulletin
of the Zoological Society of Southern Africa 10:1–7 [Erratum: ibid 10: 32]
Crawford-Cabral J (1970a) Alguns aspectos da ecologia da Palanca real. Boletim do Instituto de
Investigação Científica de Angola 7:7–42
Crawford-Cabral J (1970b) As Genetas da África Central (República do Zaire, Ruanda e Burundi).
Boletim do Instituto de Investigação Científica de Angola 7:3–23
Crawford-Cabral J (1971) A Suricata do Iona, subspécie nova. Boletim do Instituto de Investigação
Científica de Angola 8:65–83
Crawford-Cabral J (1982) Esboço zoogeográfico de Angola em ordem à fauna de mamíferos ter-
restres. Unpublished report
Crawford-Cabral J (1986) A list of Angolan Chiroptera with notes on their distribution. Boletim do
Instituto de Investigação Científica de Angola 13:7–48
Crawford-Cabral J (1987) The taxonomic status of Crocidura nigricans Bocage, 1889 (Mammalia,
Insectivora). Garcia de Orta, Série de Zoologia 14:3–12
Crawford-Cabral J (1992) Parapatry as a secondary event. Garcia de Orta, Série de Zoologia 19:1–6
Crawford-Cabral J (1996) The species of Galerella (Mammalia: Carnivora: Herpestinae) occur-
ring in the southwestern corner of Angola. Garcia de Orta, Série de Zoologia, Lisboa 21:7–17
Crawford-Cabral J (1997) A zoogeographical division of Western Angola (Africa), based on the
distribution of Muroidea (Rodentia). In: Ulrich H (ed) Tropical biodiversity and systematics:
Proceedings of the international symposim on biodiversity and systematics in tropical ecosys-
tems, Bonn, 1994. Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn,
pp 221–227
Crawford-Cabral J (1998) The Angola rodents of the superfamily Muroidea. An account on their
distribution. Estudos, Ensaios e Documentos do Instituto de Investigação Científica Tropical
161:1–222
Crawford-Cabral J, Fernandes CA (2001) The Rusty-spotted Genets as a group with three species
in Southern Africa (Carnivora: Viverridae). In: Denys C, Granjon L, Poulet A (eds) African
small mammals/Petits Mammifères Africains. IRD Éditions, Paris, pp 65, 570 pp–80
Crawford-Cabral J, Simões AP (1987) Distributional data and notes on Angolan carnivores
(Mammalia: Carnivora) I – small and medium-sized species. Garcia de Orta, Série de Zoologia
14:3–27
Crawford-Cabral J, Simões AP (1988) Distributional data and notes on Angolan carnivores
(Mammalia: Carnivora) II – larger species. Garcia de Orta, Série de Zoologia 15:9–20
Crawford-Cabral J, Veríssimo LN (2005) The ungulate fauna of Angola: systematic list, distri-
bution maps, database report. Estudos, Ensaios e Documentos do Instituto de Investigação
Científica Tropical 163:1–277
430 P. Beja et al.
Cumming DHM (2013) Phacochoerus africanus Common Warthog. In: Kingdon J, Hoffmann
Detwiler KM, Burrell AS, Jolly CJ (2005) Conservation implications of hybridization in African
cercopithecine monkeys. Int J Primatol 26:661–684
Dodman T, Diop NMD, Sarr K (2008) Conservation strategy for the West African Manatee. UNEP
and Wetlands International Africa, Nairobi/Dakar, 128 pp
Drake-Brockman RE (1909) VI. – On a new species and a new subspecies of the genus Madoqua
and a new subspecies of the genus Rhynchotragus. J Nat Hist 4:48–51
Drake-Brockman RE (1930) 4. A review of the Antelopes of the Genera Madoqua and
Rhynchotragus. J Zool 100:51–57
du Bocage JVB (1865) 3. Sur quelques Mammifères rares ou peu connus d’Afrique occidentale
qui se trouvent au Muséum de Lisbonne. Proc Zool Soc Lond 1865:401–402
du Bocage JVB (1869) Sur une espèce de ‘Cephalophus’ à taille plus forte, d’Afrique occiden-
tale, qui parait identique au ‘C. longiceps’ Gray. Jornal de Sciências Mathemáticas, Physicas
e Naturaes 2:220–222
du Bocage JVB (1878) Liste des Antilopes d’Angola. Proc Zool Soc Lond 1878:741–745
du Bocage JVB (1879) Subsídios para a Fauna das possessões portuguesas d’Africa occidental.
Jornal de Sciências Mathemáticas, Physicas e Naturaes 7:85–96
du Bocage JVB (1882) Liste des mammifères envoyés de Caconda «Angola» par M. D’Anchieta.
Jornal de Sciências Mathemáticas, Physicas e Naturaes 9:25–29
du Bocage JVB (1889a) Chiroptères africains nouveaux, rares ou peu connus. Jornal de Sciências
Mathemáticas, Physicas e Naturaes, Segunda Série 1:1–7
du Bocage JVB (1889b) Les Damans d’Angola. Jornal de Sciências Mathemáticas, Physicas e
Naturaes, Segunda Série 1:186–196
du Bocage JVB (1890) Mammifères d’Angola et du Congo (Suite). Jornal de Sciências
du Bocage JVB (1897) Mammiferos, Reptis e Batrachios d’Africa de que existem exemplares
typicos no Museu de Lisboa. Jornal de Sciências Mathemáticas, Physicas e Naturaes, Segunda
Série 4:187–206
du Bocage JVB (1898) Sur une nouvelle espèce de Cynonycteris d’Angola. Jornal de Sciências
du Bocage JVB (1902) Les Antilopes d’Angola. Jornal de Sciências Mathemáticas, Physicas e
Naturaes, Segunda Série 4:234–242
Du Toit JT (2013) Raphicerus campestris Steenbok. In: Kingdon J, Hoffmann M (eds) The
East R (1999) African Antelope Database 1998. Occasional Paper of the IUCN Species Survival
Commission No. 21. IUCN, Gland, 434 pp
Emslie RH, Adcock K (2013) Diceros bicornis Black Rhinoceros. In: Kingdon J, Hoffmann
M (eds) The mammals of Africa. Vol V (Carnivores, Pangolins, Equids and Rhinoceros).
Emslie R, Brooks M (1999) African Rhino: status survey and conservation action plan. IUCN,
Gland, 92 pp
Estes RD (2013) Hippotragus niger Sable Antelope. In: Kingdon J, Hoffmann M (eds) The
Estes RD, Estes RK (1974) The biology and conservation of the giant sable antelope, Hippotragus
niger variani Thomas, 1916. Proc Acad Nat Sci U S A 126:73–104
Fabiano EC, Álvares F, Kosmas S et al (2017) The conservation status of the Endangered African
Wild Dogs in Angola: an historical and contemporary perspective. Final progress report
(Project No.162513063). Unpublished report. University of Namibia, Katima Mulilo
Faulkes C, Maree S, Griffin M (2016) Fukomys bocagei. The IUCN Red List of threatened species
2016:e.T5752A22184407. Downloaded on 19 April 2018
Feiler A (1986) Zur faunistik and biometrie angolanisher Fledermause. Zoologische
Abhandlungen, Staatliches Museum für Tierkunde Dresden 42:65–77
Feiler A (1989) Individuelle variation bei Buschböcken (Tragelaphus scriptus) aus Angola
(Mammalia, Artiodactyla: Bovidae). Zoologische Abhandlungen, Staatliches Museum für
Tierkunde Dresden 44:149–154
Feiler A (1990) Distribution of mammals in Angola and notes on biogeography. In: Peters G,
Hutterer R (eds) Vertebrates in the tropics: Proceedings of the international symposium on ver-
tebrate biogeography and systematics in the tropics, Bonn, June 5–8, 1989. Alexander Koenig
Zoological Research Institute and Zoological Museum, Bonn, pp 221–236
Fennessy J, Bidon T, Reuss F et al (2016) Multi-locus analyses reveal four giraffe species instead
of one. Curr Biol 26:2543–2549
Fenykovi J (1953) Angola, en el visor del rifle y de la cámara. Cayrel Ediciones, Madrid, 255 pp
Figueira R (2017) IICT Colecção Zoológica. Version 4.2. Instituto de Investigação Científica
Tropical, Lisbon. Accessed via GBIF.org on 23 Feb 2018
Figueiredo E, Smith GF, César J (2009) The flora of Angola: first record of diversity and ende-
Flagstad Ø, Syversten PO, Stenseth NC et al (2001) Environmental change and rates of evolution:
the phylogeographic pattern within the hartebeest complex as related to climatic variation. Proc
R Soc Lond B 268:667–677
Foley NM, Thong VD, Soisook P et al (2015) How and why overcome the impediments to resolu-
tion: lessons from rhinolophid and hipposiderid bats. Mol Biol Evolut 32:313–333
Frade F (1933) Eléphants d’Angola. Bulletin de la Société Portugaise des Sciences Naturelles
11:319–333
Frade F (1936) Distribution géographique des eléphants d’Afrique. Compte Rendu du XII Congres
International de Zoologie, Lisbonne 1935:1191–1202
Frade F (1955) Ordre des Proboscidiens (Proboscidea Illiger, 1811). In: Grassé PP (ed) Traité de
Zoologie. Anatomie, Systématique, Biologie, Tome XVII, 1° fascicule. Masson et Cie, Paris,
pp 715–783
Frade F (1956) Reservas naturais de Angola – I (alguns mamíferos da Reserva da Quiçama). Anais
da Junta de Investigações do Ultramar 11:228–245
Frade F (1958) Mesures adoptées pour la protection de l’hippotrague géant en Angola. Mammalia
22:476–477
Frade F (1959a) Medidas para a protecção da Palanca gigante de Angola (Hippotragus niger vari-
ani Thomas). Memórias da Junta de Investigações do Ultramar (Série 2) 8:11–18
Frade F (1959b) Breve notícia a propósito da Reserva da Quiçama. Garcia de Orta 4:215–223
Frade F (1960) Os animais na etnologia ultramarina. Estudos, Ensaios e Documentos da Junta de
Frade F (1963) Linhas gerais da distribuição dos Vertebrados em Angola. Memórias da Junta de
Frade F, Sieiro DM (1960) Palanca preta gigante de Angola. Garcia de Orta 8:21–38
França P (1967) Sur la présence d’Arctocephalus pusillus (Schreber) (Otariidae) et de Mirounga
leonina (Linne) (Phocidae) au sud de l’Angola. Mammalia 31:50–54
Fritz H, Bourgarel M (2013) Aepyceros melampus Impala. In: Kingdon J, Hoffmann M (eds) The
Funston P, Henschel P, Petracca L et al (2017) The distribution and status of lions and other large
carnivores in Luengue-Luiana and Mavinga National Parks, Angola. KAZA TFCA Secretariat,
Kasane
Galvão HCF, Montês A (1943–1945) A Caça no Império Português, 2 vols. Editorial Primeiro de
Janeiro, Porto, 639 pp
432 P. Beja et al.
Gaubert P, Fischer C, Hausser Y et al (2016) Genetta angolensis. The IUCN Red List of threatened
species 2016:e.T41696A45218468. Downloaded on 10 March 2018
Gautier-Hion A (2013a) Miopithecus talapoin Southern talapoin monkey. In: Butynski TM,
Kingdon J, Kalina J (eds) The mammals of Africa. Vol II (Primates). Bloomsbury Publishing,
London, pp 252, 556 pp–253
Gautier-Hion A (2013b) Miopithecus ogouensis – Northern talapoin monkey. In: Butynski TM,
London, pp 253, 556 pp–256
Gautier-Hion A, Colyn M, Gautier J-P (1999) Histoire Naturelle des Primates d’Afrique Centrale.
ECOFAC, Libreville, 162 pp
Genest-Villard H (1969) Particularités des lièvres du Sud-Ouest de l’Angola. Mammalia
33:124–132
Gerrie R, Kennerley R (2016) Crocidura erica (errata version published in 2017). The IUCN Red
List of threatened species 2016:e.T5626A115078377. Downloaded on 01 May 2018
Gilchrist JS, Jennings AP, Veron G, Cavallini P (2009) Family Herpestidae. In: Wilson DE,
Mittermeier RA (eds) Handbook of the mammals of the world, Vol I (Carnivores). Lynx
Edicions, Barcelona, pp 262, 728 pp–328
Gosling LM, Capellini I (2013) Alcephalus buselaphus Hartebeest. In: Kingdon J, Hoffmann
Grant S, Ferguson A (2018) Field Museum of Natural History (Zoology) Mammal Collection.
Version 9.3. Field Museum, Chicago. Accessed via GBIF.org on 19 Apr 2018
Gray JE (1868) VI. – Revision of the species of hyrax, founded on the specimens in the British
Museum. Ann Mag Nat Hist Ser 4 1:35–51
Gray JE (1869) New species of hyrax. Ann Mag Nat Hist Ser 4 3:242–243
Groom R, Elizalde S, Elizalde D, de Sá S, Alexandre G (2018) Large and medium sized terrestrial
mammals survey in Quiçama National Park. Preliminary results report based on species pres-
ence and bushmeat. Instituto Nacional de Áreas de Conservação, Luanda, 17 pp
Groves C, Grubb P (2011) Ungulate taxonomy. Johns Hopkins University Press, Baltimore, 336 pp
Grubb P (1972) Variation and incipient speciation in the African buffalo. Zeitschrift für
Säugetierkunde 37:121–144
Grubb P (1993) The afrotropical suids (Phacochoerus, Hylochoerus and Potamochoerus). In:
Oliver WLR (ed) Status survey and conservation plan – pigs, peccaries and hippos. IUCN,
Gland, pp 66, 202 pp–75
Grubb P, Groves CP, Dudley JP, Shoshani J (2000) Living African elephants belong to two species:
Loxodonta africana (Blumenbach 1797) and Loxodonta cyclotis (Matschie 1900). Elephant
2:1–4
Hall-Martin A, Pienaar D (1992) A note on the elephants of Southeast Angola. Unpublished report.
African Elephant and Rhino Specialist Group, Nairobi
cover change. Science 342:850–853
Happold DCD (2013) The mammals of Africa. Vol III (Rodents, hares and rabbits). Bloomsbury
Publishing, London, 789 pp
Happold M, Happold DCD (2013) The mammals of Africa. Vol IV (Hedgehogs, shrews and bats).
Bloomsbury Publishing, London, 800 pp
Happold DCD, Wendelen W (2006) The distribution of Poelagus marjorita (Lagomorpha:
Leporidae) in central Africa. Mammalian Biology-Zeitschrift für Säugetierkunde 71:377–383
Hart JA, Kingdon J (2013) Philatomba monticola Blue Duiker. In: Kingdon J, Hoffmann M (eds)
Hart J, Groves CP, Ehardt C (2008) Lophocebus aterrimus ssp. opdenboschi. The IUCN Red List
of threatened species 2008:e.T12311A3334719. Downloaded on 21 March 2018
Hassanin A, Delsuc F, Ropiquet A et al (2012) Pattern and timing of diversification of

Cetartiodactyla (Mammalia, Laurasiatheria), as revealed by a comprehensive analysis of mito-
chondrial genomes. Comptes Rendus Biologies 335:32–50
Hatt RT (1934) A manatee collected by the American Museum Congo Expedition: with observa-
tions on the recent manatees. Bull Am Mus Nat Hist 66:553–566
Hayman RW (1951) Notes on some Angolan Mammals. Publicações Culturais da Companhia de
Hayman RW (1963) Mammals from Angola, mainly from the Lunda District. Publicações
Culturais da Companhia de Diamantes de Angola 66:81–139
Heffernan J (2005) Elephants of Cabinda; Mission report, Angola, April 2005. Unpublished report.
Fauna & Flora International & United Nations Development Programme in co-operation with
the Deptartment of Urban Affairs & Environment, Cabinda
Heim de Balzac H, Meester J (1977) Part 1. Order Insectívora. In: Meester JA, Setzer HW (eds)
The mammals of Africa: an identification manual. Smithsonian Institution Press, Washington,
DC
Hill JE (1941) A collection of mammals from Dondi, Angola. J Mammal 22:81–85
Hill JE, Carter TD (1941) The mammals of Angola, Africa. Bull Am Mus Nat Hist 78:1–211
Hinton MAC (1921) Klipspringers of Rhodesia, Angola and Northern Nigeria. Ann Mag Nat Hist
Ser 9 8:129–133
Hoare R (2015) Lessons from 20 years of human–elephant conflict mitigation in Africa. Hum
Dimens Wildl 20:289–295
Hockings KJ, Anderson JR, Matsuzawa T (2012) Socioecological adaptations by chimpanzees, Pan
troglodytes verus, inhabiting an anthropogenically impacted habitat. Anim Behav 83:801–810
Hoffmann M, Wilson V (2013) Raphicerus sharpei Sharpe’s Grysbok. In: Kingdon J, Hoffmann
Hofmeyr GJG (2015) Arctocephalus pusillus. The IUCN Red List of threatened species 2015:
e.T2060A45224212. Downloaded on 10 May 2018
Huntley BJ (1972a) Relatório do Ecólogo Sobre a Ocupação do Parque Nacional da Quiçama
pela Pecuária da Barra do Cuanza. Unpublished report. Direcção Provincial dos Serviços de
Veterinária, Luanda, 11pp
Huntley BJ (1972b) Plano para o Futuro da Palanca Real em Angola. Unpublished report. Direcção
Provincial dos Serviços de Veterinária, Luanda
Huntley BJ (1972c) Report on visit to Iona National Park. 24 February to 25 March 1972.
Unpublished report. Direcção Provincial dos Serviços de Veterinária, Luanda, 13 pp
Huntley BJ (1973a) Aspectos Gerais da Conservação do Bravio em Angola. Relatório N° 15.
Unpublished report. Direcção Provincial dos Serviços de Veterinária, Luanda, 30 pp
Huntley BJ (1973b) Reordenamento da População Humana no Parque Nacional da Quiçama.
Relatório N° 19. Unpublished report. Direcção Provincial dos Serviços de Veterinária, Luanda,
10 pp
Huntley BJ (1973c) Distribuição e Situação da Grande Fauna Selvagem de Angola com Referência
Especial às Espécies Raras e em Perigo de Extinção – Primeiro Relatório Sobre o Estado
Actual. Relatório N°. 21. Unpublished report. Direcção Provincial dos Serviços de Veterinária,
Luanda, 37 pp
Huntley BJ (1973d) Parque Nacional do Iona: Administração, Maneio, Investigação e Turismo.
Relatório N°. 23. Unpublished report. Direcção Provincial dos Serviços de Veterinária, Luanda,
37 pp
Huntley BJ (1973e) Proposta para a Criação de uma Reserva Natural Integral na Floresta do
Maiombe, Cabinda. Unpublished report. Direcção Provincial dos Serviços de Veterinária,
Luanda, Angola, 9 pp
Huntley BJ (1973f) Distribution of larger mammals of Angola according to vegetation types of Dr.
Grandvaux Barbosa’s Map. Unpublished report
Huntley BJ (1974) Outlines of wildlife conservation in Angola. S Afr J Wildl Res 4:157–166
434 P. Beja et al.
Huntley BJ (2017) Wildlife at war in Angola. The rise and fall of an African Eden. Protea Book
Huntley BJ, Francisco P (eds) (2015) Avaliação Rápida da Biodiversidade de Região da Lagoa
Carumbo, Lunda-Norte – Angola/Rapid Biodiversity Assessment of the Carumbo Lagoon
Area, Lunda-Norte – Angola. Ministério do Ambiente, Luanda, 219 pp
Huntley BJ, Matos E (1992) Angola environment status Quo Assessment report. Unpublished
report. IUCN Regional Office for Southern Africa, Harare, 255 pp
Huntley BJ, Beja P, Vaz Pinto P et al (2019) Biodiversity conservation: history, protected areas and
hotspots. In: Huntley BJ, Russo V, Lages F, Ferrand N (eds) Biodiversity of Angola. Science &
conservation: a modern synthesis. Springer, Berlin
Hutterer R (2016) Crocidura nigricans. The IUCN Red List of threatened species 2016:e.
T41345A22310112. Downloaded on 01 May 2018
Hutterer R, Peterhans JK (2016) Crocidura roosevelti (errata version published in 2017). The
IUCN Red List of threatened species 2016: e.T41355A115181119. Downloaded on 10 May
2018
iNaturalist.org (2018a) iNaturalist research-grade observations. Occurrence dataset https://
doi.org/10.15468/ab3s5x. Accessed via GBIF.org on 27 June 2018. https://www.gbif.org/
occurrence/1802602131
iNaturalist.org (2018b) iNaturalist research-grade observations. Occurrence dataset https://
doi.org/10.15468/ab3s5x. Accessed via GBIF.org on 27 June 2018. https://www.gbif.org/
occurrence/1135207782
INBAC – Instituto Nacional da Biodiversidade e Áreas de Conservação (2016) Plano de Acção
Nacional de Conservação da Chita e Mabeco em Angola. Unpublished report. Ministério do
Ambiente, República de Angola, Luanda, 30pp
IUCN (2018) The IUCN Red List of threatened species v. 2017.3. IUCN, Gland Accessed 20 Mar
2018. www.redlist.org
Jacques H, Reed-Smith J, Davenport C, Somers MJ (2015a) Aonyx congicus. The IUCN Red List
of threatened species 2015:e.T1794A14164772. Downloaded on 13 April 2018
Jacques H, Reed-Smith J, Somers MJ (2015b) Aonyx capensis. The IUCN Red List of threatened
species 2015:e.T1793A21938767. Downloaded on 13 April 2018
Jenkins R (2013) Kobus vardonii Puku. In: Kingdon J, Hoffmann M (eds) The mammals of Africa.
Vol VI (Hippopotamuses, pigs, deer, giraffe and bovids). Bloomsbury Publishing, London,
pp 445, 680 pp–449
Jentink FA (1887) On mammals from Mossamedes. Notes Leyden Mus 9:171–180
Jentink FA (1893) On some mammals from Cahama. Notes Leyden Mus 15:262–265
Jentink FA (1900) The species of the antelope – genus Pediotragus. Notes Leyden Mus 22:33–43
Jentink FA (1901) The antelopes in the Leyden Museum. Notes Leyden Mus 23:17–31
Jordan NR, Do Linh San E (2015) Suricata suricatta. The IUCN Red List of threatened species
2015:e.T41624A45209377. Downloaded on 30 May 2018
Juste J, Carballo C (1992) Proyecto Para la Definición y Justificación de la Recuperación y
Conservación de los Parques Nacionales de Quissama, Bikuar, Mupa, Iona y la Reserva Parcial
de Namibe (Angola). Asociacíon Amigos del Coto de Doñana, Sevilla, 79 pp
Keesing F, Young TP (2014) Cascading consequences of the loss of large mammals in an African
Savanna. BioScience 64:487–495
Kennerley R (2016) Crocidura ansellorum (errata version published in 2017). The IUCN Red List
of threatened species 2016:e.T5558A115073943. Downloaded on 10 May 2018
Kingdon J (2008a) Cercopithecus mitis ssp. mitis. The IUCN Red List of threatened species 2008:
e.T136943A4351535. Downloaded on 20 March 2018
Kingdon J (2008b) Cercopithecus mitis ssp. opisthostictus. The IUCN Red List of threatened spe-
cies 2008:e.T136850A4346858. Downloaded on 20 March 2018
Kingdon J (2016) Papio kindae. The IUCN Red List of threatened species 2016:e.
Kingdon J, Hoffmann M (2013) Redunca arudinum Southern Reedbuck. In: Kingdon J, Hoffmann
Kingdon J, Lahm SA (2013) Cephalophus silvicvultor Yellow-backed Duiker. In: Kingdon J,
Hoffmann M (eds) The mammals of Africa. Vol VI (Hippopotamuses, pigs, deer, giraffe and
bovids). Bloomsbury Publishing, London, pp 288, 680 pp–293
Kingdon J, Happold D, Butynski T et al (2013) The mammals of Africa. Vols 1–6. Bloomsbury
Publishing, London
Kingswood SC, Kumamoto AT (1997) Madoqua kirkii. Mamm Species 569:1–10
Kitchener AC, Breitenmoser-Würsten C, Eizirik E et al (2017) A revised taxonomy of the Felidae.
The final report of the Cat Classification Task Force of the IUCN/SSC Cat Specialist Group.
Cat News Spec Issue 11:1–80
Klingel H (2013) Hippopotamus amphibius Common Hippopotamus. In: Kingdon J, Hoffmann
Knight M (2013) Oryx gazella Gemsbok (Southern Oryx). In: Kingdon J, Hoffmann M (eds)
Kock D, Amr Z, Mickleburgh S et al (2008) Hipposideros caffer. The IUCN Red List of threatened
species 2008:e.T10115A3166805. Downloaded on 24 April 2018
Kolberg H, Kilian W (2003) Report on an aerial survey of Iona National Park, Angola, 6 to 14 June
2003. Directorate of Scientific Services, Ministry of Environment and Tourism, Windhoek,
22 pp
Kuedikuenda S, Xavier NG (2009) Framework report on Angola’s Biodiversity. Unpublished
report. Republic of Angola, Ministry of Environment, Luanda, 60 pp
Kumamoto AT, Kingswood SC, Hugo W (1994) Chromosomal divergence in allopatric popula-
tions of Kirk’s dik-dik, Madoqua kirki (artiodactyla, Bovidae). J Mammal 75:357–364
Kunz TH, Braun de Torrez E, Bauer D et al (2011) Ecosystem services provided by bats. Ann N
Y Acad Sci 1223:1–38
Laurance WF, Sayer J, Cassman KG (2014) Agricultural expansion and its impacts on tropical
nature. Trends Ecol Evolut 29:107–116
Lawes MJ, Cords M, Lehn C (2013) Cercopithecus mitis Gentle monkeys. In: Butynski TM,
London, pp 354, 556 pp–362
Le Houérou HN (2009) Bioclimatology and biogeography of Africa. Springer, Heidelberg, 240 pp
Leus K, Vercammen P (2013) Potamochoerus porcus Red River Hog. In: Kingdon J, Hoffmann
Linder HP, de Klerk HM, Born J et al (2012) The partitioning of Africa: statistically defined bio-
geographical regions in sub-Saharan Africa. J Biogeogr 39:1189–1205
Lindsey PA, Petracca LS, Funston PJ et al (2017) The performance of African protected areas for
lions and their prey. Biol Conserv 209:137–149
Lorenzen ED, Arctander P, Siegismund HR (2006) Regional genetic structuring and evolutionary
history of the impala Aepyceros melampus. J Hered 97:119–132
Lorenzen ED, Arctander P, Siegismund HR (2008) High variation and very low differentiation
in wide ranging plains zebra (Equus quagga): insights from mtDNA and microsatellites. Mol
Ecol 17:2812–2824
Loveridge AJ, Kuiper T, Parry RH et al (2017) Bells, bomas and beefsteak: complex patterns of
human-predator conflict at the wildlife-agropastoral interface in Zimbabwe. PeerJ 5:e2898
Lydekker R (1899) The great and small game of Africa. Rowland Ward, London, 642 pp
Lydekker R (1903) Hutchinson’s animal life, vol 2. Hutchinson, London
Lydekker R (1904) On the subspecies of Giraffa camelopardalis. Proc Zool Soc Lond 1904:202–227
436 P. Beja et al.
Lydekker R, Blaine G (1913–1916) Catalogue of Ungulate mammals in the British Museum

(Natural History), 5 vols. British Museum (Natural History), London
Machado AB (1952) Generalidades acerca da Lunda e da sua exploração biológica. Publicações
Machado AB (1968) A exploração biológica da Lunda. Memórias da Academia de Ciências de
Lisboa 12:35–71
Machado AB (1969) Mamíferos de Angola ainda não citados ou pouco conhecidos. Publicações
Machado AB, Crawford-Cabral J (1999) As subespécies de Cercopithecus mitis Wolf, 1922
(Primates, Cercopithecidae) existentes em Angola. Garcia de Orta, Série de Zoologia 23:99–117
Maisels F, Strindberg S, Breuer T et al (2016a) Gorilla gorilla ssp. gorilla (errata version published
in 2016). The IUCN Red List of threatened species 2016:e.T9406A102328866. Downloaded
on 21 March 2018
Maisels F, Strindberg S, Greer D et al (2016b) Pan troglodytes ssp. troglodytes (errata version
published in 2016). The IUCN Red List of threatened species 2016:e.T15936A102332276.
Downloaded on 21 March 2018
Malbrant R, Maclatchy A (1949) Faune de l’Équateur Africain Français. Tome 2: Mammifères.
Lechavalier, Paris, 324 pp
Malhi Y, Doughty CE, Galetti M et al (2016) Megafauna and ecosystem function from the
Pleistocene to the Anthropocene. Proc Natl Acad Sci U S A 113:838–846
Matschie P (1900) Über Equus penricei. Sitzungsberichte der Gesellschaft Naturforschender
Freunde zu Berlin 1900:231
Matschie P (1906) Einige noch nicht beachrieben des Arten africanischen Büffels. Sitzungsberichte
der Gesellschaft Naturforschender Freunde zu Berlin 7:161–179
McLennan MR (2013) Diet and feeding ecology of chimpanzees (Pan troglodytes) in Bulindi,
Uganda: foraging strategies at the forest–farm interface. Int J Primatol 34:585–614
McNutt JW, Stein AB, McNutt LB, Jordan NR (2017) Living on the edge: characteristics of human–
wildlife conflict in a traditional livestock community in Botswana. Wildl Res 44:546–557
Meÿer MA (2007) The first aerial survey of Cape Fur Seal numbers at Baia dos Tigres, south-
ern Angola. In: Kirkman SP (ed) Final report of the BCLME (Benguela current large marine
ecosystem). Project on top predators as biological indicators of ecosystem change in the
BCLME. Avian Demography Unit, Cape Town, pp 307–308, 382 pp
MHNG (2018). Mammals housed at MHNG, Geneva. Muséum d’Histoire Naturelle de la Ville de
Genève. Accessed via GBIF.org on 2018-04-23
Milliken T, Pole A, Huongo A (2006) No peace for elephants: unregulated domestic ivory mar-
kets in Angola and Mozambique. TRAFFIC online report series no. 11. Traffic East/Southern
Africa, Harare, 46 pp
Mills MS, Olmos F, Melo M, Dean WRJ (2011) Mount Moco: its importance to the conservation
of Swierstra’s Francolin Pternistis swierstrai and the Afromontane avifauna of Angola. Bird
Mills MS, Melo M, Vaz A (2013) The Namba mountains: new hope for Afromontane forest birds
MNHN (2018) The mammals collection (ZM) of the Muséum National d’Histoire Naturelle.
Version 43.58. Museum National d’Histoire Naturelle, Paris Accessed via GBIF.org on 19 Apr
2018
Monadjem A, Taylor PJ, Cotterill W, Schoeman MC (2010a) Bats of southern and Central Africa:
a biogeographic and taxonomic synthesis. Wits University Press, Johannesburg, p 596
Monadjem A, Schoeman MC, Reside A et al (2010b) A recent inventory of the bats of Mozambique
with documentation of seven new species to the country. Acta Chiropterologica 12:371–391
Monadjem A, Richards L, Taylor PJ, Denys C, Dower A, Stoffberg S (2013a) Diversity of
Hipposideridae in the Mount Nimba massif, West Africa, and the taxonomic status of
Hipposideros lamottei. Acta Chiropterologica 15:341–352
Monadjem A, Goodman SM, Stanley WT et al (2013b) A cryptic new species of Miniopterus from
south-eastern Africa based on molecular and morphological characters. Zootaxa 3746:123–142
Monadjem A, Taylor PJ, Denys C, Cotterill FP (2015) Rodents of Sub-Saharan Africa: a biogeo-
graphic and taxonomic synthesis. de Gruyter, Berlin, 1102 pp
Monadjem A, Jacobs D, Taylor P, Cohen L, MacEwan K, Richards LR, Sethusa T
(2017) Rhinolophus damarensis. The IUCN Red List of threatened species 2017:
e.T67369846A67369914. Downloaded on 25 April 2018
Monard A (1930) Mission Scientifique Suisse dans l’Angola. Résultats scientifiques. Mammifères.
Part I: Ongulés. Bulletin de la Société Neuchâteloise des Sciences Naturelles 54:73–102
Part I: Carnivores. Bulletin de la Société Neuchâteloise des Sciences Naturelles 55:51–71
Part IV: Ongulés (Suite). Bulletin de la Société Neuchâteloise des Sciences Naturelles 57:45–66
Arquivos do Museu Bocage 6:1–103
Monsarrat S, Kerley GI (2018) Charismatic species of the past: biases in reporting of large mam-
mals in historical written sources. Biol Conserv 223:68–75
Moodley Y, Bruford MW (2007) Molecular biogeography: towards an integrated framework for
conserving pan-African biodiversity. PLoS One 2:e454
Moodley Y, Bruford MW, Bleidorn C et al (2009) Analysis of mitochondrial DNA data reveals
non-monophyly in the bushbuck (Tragelaphus scriptus) complex. Mammalian Biology-
Zeitschrift für Säugetierkunde 74:418–422
Moraes J, Putzke J (2013) Ocorrência de Lepus saxatilis F. Cuvier, 1823 na província de Malanje,
norte de Angola. Cad Pesqui 25:40–43
Morais M, Velasco L, Carvalho E (2006a) Avaliação da Condição e Distribuição do Manatim
Africano (Trichechus senegalensis) ao longo do Rio Cuanza. Unpublished report. Universidade
Agostinho Neto & Ministério do Urbanismo e Ambiente, Luanda
Morais M, Torres MOF, Martins MJ (2006b) Biodiversidade Marinha e Costeira em Angola.
Projecto de Estratégia e Plano de Acção Nacionais para a Biodiversidade (NBSAP). Ministério
do Urbanismo e Ambiente, Luanda, Angola
Murphy BP, Bowman DMJS (2012) What controls the distribution of tropical forest and savanna?
Ecol Lett 15:748–758
Musser GG, Carleton MD (2005) Superfamily Muroidea. In: Wilson DE, Reeder DM (eds)
Mammal species of the world: a taxonomic and geographic reference. The Johns Hopkins
University Press, Baltimore, pp 894–1531
Newton da Silva S (1970) A Grande Fauna Selvagem de Angola. Direcção Provincial dos Serviços
de Veterinária, Luanda, 151 pp
NGOWP (2018) National Geographic Okavango Wilderness Project. Initial Findings from
Exploration of the Upper Catchments of the Cuito, Cuanavale and Cuando Rivers in Central
Novellie P, Lindeque M, Lindeque P et al (2002) Status and Action Plan for the Mountain Zebra
(Equus zebra). In: Moehlman PDR (ed) Equids: zebras, asses, and horses. Status Survey and
Conservation Action Plan. IUCN, Gland, pp 28, 190 pp–42
Oates JF (2011) Primates of West Africa: a field guide and natural history. Conservation
International, Arlington, 556 pp
Oates JF, Groves CP (2008) Cercopithecus nictitans. The IUCN Red List of threatened species
Oates JF, Hart J, Groves CP et al (2008a) Cercopithecus ascanius. The IUCN Red List of threat-
ened species 2008. Downloaded on 30 May 2018
Oates JF, Hart J, Groves CP et al (2008b) Cercopithecus ascanius ssp. atrinasus. The IUCN Red
List of threatened species 2008:e.T136869A4347775. Downloaded on 20 March 2018
Oboussier H (1962) Zur Kenntnis des Kaffernbüffels (Syncerus caffer Sparrman, 1779). Hirn
und Hypophyse. Ergebnisse einer Forshungsreisen nach Süd-Angola. Boletim do Instituto de
Investigação Científica de Angola 1:39–47
438 P. Beja et al.
Oboussier H (1963) Die Pferdeantilope (Hippotragus equinus cottoni Dollman and Burlace,
1928). Ergebnisse der Forschungsreisen nach Sud-Angola. Zeitschrift fuer Morphologie und
Oekologie der Tiere 52:688–713
Oboussier H (1964) Ein ungewöhlnliches Warzenschwein (Phacochoerus aethiopicus shortridgei
St. Leger, 1932). Säugetierkundliche Mitteilungen 12:94–97
Oboussier H (1965) Zur Kenntnis der Schwarzfersenantilope (Impala) Aepyceros melampus unter
besonderer Berücksichtigung des Grosshirnfurchenbildes und der Hypophyse. Ergebnisse der
Forschungsreisen nach Süd-Angola und Ostafrika. Zeitschrift fuer Morphologie und Oekologie
der Tiere 54:531–550
Oboussier H (1966) Das Grosshirnfurchenbild als Merkmal der Evolution. Untersuchungen an
Boviden II (Subfamilien Cephalophinae und Antilopinae nach Simpson 1945). Mitteilungen
Oboussier H (1972) Morphologische und quantitative Neocortexuntersuchungen bei Boviden, ein
Beitrag zur Phylogenie dieser Familie. II. Formen geringen Körpergewichts (3 kg – 25 kg)
aus den Subfamilien Cephalophinae und Antilopinae. Mitteilungen aus dem Hamburgischen
Zoologischen Museum und Institut 68:231–269
Oboussier H (1976) Zur Kenntnis der Moorantilopen (Mammalia, Bovidae, Reduncini).
Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 73:281–294
Oboussier H, Von Tyszka H (1964) Beiträge zür Kenntnis der Reduncini (Hippotraginae – Bovidae)
Süd-Angolas Hirnfurchenbild und Hypophyse. Ergebnisse der Forschungsreisen von Prof. Dr.
H. Oboussier nach Angola 1959 und 1961. Zeitschrift fuer Morphologie und Oekologie der
Tiere 53:362–386
new map of life on Earth. BioScience 51:93–938
Orrell T, Hollowell T (2018) NMNH extant specimen records. Version 1.16. National Museum of
Natural History, Smithsonian Institution, Washington, DC Accessed via GBIF.org on 19 Apr
2018
Overton J, Fernandes S, Elizalde D et al (2017) A large mammal Survey of Bicuar and Mupa
National Parks, Angola. Unpublished report. Instituto Nacional da Biodiversidade e Áreas de
Conservação, Luanda
Owen-Smith N (2013) Tragelaphus strepsiceros Greater Kudu. In: Kingdon J, Hoffmann M (eds)
Palkopoulou E, Lipson M, Mallick S et al (2018) A comprehensive genomic history of extinct and
living elephants. Proc Natl Acad Sci U S A 115:E2566–E2574
Penzhorn B (2013) Equus zebra Mountain Zebra. In: Kingdon J, Hoffmann M (eds) The mam-
mals of Africa. Vol V (Carnivores, pangolins, equids and rhinoceros). Bloomsbury Publishing,
London, pp 438, 544 pp–443
Perrin M (2013) Elephantulus brachyrhynchus Short-snouted Sengi. In: Kingdon J, Happold D,
Hoffmann M et al (eds) The mammals of Africa. Vol I (Introductory chapters and afrotheria).
Peters WC (1865) Note on the Mammalia observed by Dr. Welwitsch in Angola. Proc Zool Soc
Lond 1865:400–401
Peters WC (1870) Lista dos Mammiferos das possessões portuguesas d’Africa occidental e
diagnose d’algumas especies novas. Jornal de Sciências Mathemáticas, Physicas e Naturaes,
Primeira Série 1:123–127
Peters WC (1879) Eine Neue Art der Säugethiergattung Hyrax (H. nigricans) aus Chinchoxo
und über eine neue Eidechse, Platysaurus torquatus, aus Mossambique. Sitzungsberichte der
Gesellschaft Naturforschender Freunde zu Berlin 1879:10–11
Petter F (1972) Part 5. Lagomorpha. In: Meester J, Setzer HW (eds) The mammals of Africa: an
identification manual. Smithsonian Institution Press, Washington, DC, pp 1–7
Petter F, Genest H (1965) Variation morphologique et répartition géographique de Lepus capensis
dans le Sud-Ouest Africain. Mammalia 27:238–255
Pimley ER (2009) A survey of nocturnal primates (Strepsirrhini: Galaginae, Perodictinae) in

southern Nigeria. Afr J Ecol 47:784–787
Pimley ER, Bearder SK (2013) Perodicticus potto – Potto. In: Butynski TM, Kingdon J, Kalina
J (eds) The mammals of Africa. Vol II (Primates). Bloomsbury Publishing, London, pp 393,
556 pp–398
Poulsen JR, Koerner SE, Moore S et al (2017) Poaching empties critical Central African wilder-
ness of forest elephants. Curr Biol 27:R134–R135
Powell JA (1996) The distribution and biology of the West African manatee (Trichechus senega-
lensis Link, 1795). Unpublished report. United Nations Environment Programme, Regional
Seas Programme, Oceans and Coastal Areas, Nairobi, 68 pp
Prins HHJ, Sinclair ARE (2013) Syncerus caffer African Buffalo. In: Kingdon J, Hoffmann M (eds)
Pruetz JD (2006) Feeding ecology of savanna chimpanzees (Pan troglodytes verus) at Fongoli,
Senegal. In: Hohmann G, Robbins MM, Boesch C (eds) Feeding ecology in apes and other
primates. Cambridge University Press, Cambridge, pp 161, 540 pp–182
Pullen S, Bearder SK (2013) Galago moholi Southern Lesser Bushbaby. In: Butynski TM, Kingdon
J, Kalina J (eds) The mammals of Africa. Vol II (Primates). Bloomsbury Publishing, London,
pp 430, 556 pp–433
Purchase GK, Marker L, Marnewick K et al (2007a) Regional assessment of the status, distribution
and conservation needs of cheetahs in southern Africa. Cat News 3:44–46
Purchase GK, Mateke C, Purchase D (2007b) A review of the status and distribution of carnivores,
and levels of human carnivore conflict, in the protected areas and surrounds of the Zambezi
Basin. Unpublished report. The Zambezi Society, Bulawayo, 79 pp
Rapson S, Rathbun GB (2015) Herpestes flavescens. The IUCN Red List of threatened species
Rapson SA, Goldizen AW, Seddon J (2012) Species boundaries and hybridization between the
black mongoose (Galerella nigrata) and the slender mongoose (Galerella sanguinea). Mol
Phylogenet Evolut 65(3):831–839
Rathbun GB (2013) Petrodromus tetradactylus Four-toed Sengi. In: Kingdon J, Happold D,
Hoffmann M et al (eds) The mammals of Africa. Vol I (Introductory chapters and afrotheria).
Rathbun GB, Cowley TE (2008) Behavioural ecology of the black mongoose (Galerella nigrata)
in Namibia. Mammalian Biology-Zeitschrift für Säugetierkunde 73:444–450
RBINS (2017) RBINS DaRWIN. Royal Belgian Institute of Natural Sciences, Brussels Accessed
via GBIF.org on 19 Apr 2018
Reed-Smith J, Jacques H, Somers MJ (2015) Hydrictis maculicollis. The IUCN Red List of threat-
Riggio J, Jacobson A, Dollar L et al (2013) The size of savannah Africa: a lion’s (Panthera leo)
view. Biodiver Conserv 22:17–35
Robbins MM, Bermejo M, Cipolletta C et al (2004) Social structure and life-history patterns in
western gorillas (Gorilla gorilla gorilla). Am J Primatol 64:145–159
Roberts SC (2013) Oreotragus oreotragus Klipspringer. In: Kingdon J, Hoffmann M (eds) The
Roca AL, Georgiadis N, Pecon-Slattery J et al (2001) Genetic evidence for two species of elephant
in Africa. Science 293:1473–1477
Roca AL, Ishida Y, Brandt AL et al (2015) Elephant natural history: a genomic perspective. Annu
Rev Anim Biosci 3:139–167
Rodrigues P, Figueira R, Vaz Pinto P et al (2015) A biogeographical regionalization of Angolan
mammals. Mamm Rev 45:103–116
Rodrigues P, Figueira R, Beja P (2018) Bibliographic records of Angola mammals. Version 2.2.
Instituto de Investigação Científica Tropical, Lisboa Accessed via GBIF.org on 24 Apr 2018
440 P. Beja et al.
Ron T (2005) The Maiombe forest in Cabinda: conservation efforts, 2000–2004. Gorilla J 30:18–21
Ron T, Golan T (2010) Angolan Rendezvous: man and nature in the shadow of war. 30 Degrees
South Publishers, Kwa-Zulu Natal, 272 pp
Rookmaaker LC (2005) The black rhino needs a taxonomic revision for sound conservation. Int
Zoo News 52:280–282
Rowe N, Myers M (2016) All the world’s primates. Pogonias Press, Charlestown, 777 pp
Sanborn CC (1950) Chiroptera from Dundo, Lunda, northeastern Angola. Publicações Culturais da
Companhia de Diamantes de Angola 10:51–62
Sarmiento EE (2013) Chlorocebus cynosures Malbrouck monkey. In: Butynski TM, Kingdon J,
Kalina J (eds) The mammals of Africa. Vol II (Primates). Bloomsbury Publishing, London,
pp 284, 556 pp–286
Sarmiento EE, Stiner EO, Brooks EGE (2001) Red-tail monkey Cercopithecus ascanius distin-
guishing characters and distribution. Afr Primate 5:18–24
Schlossberg S, Chase MJ, Griffin CR (2018) Poaching and human encroachment reverse recov-
ery of African savannah elephants in south-east Angola despite 14 years of peace. PLoS One
13:e0193469
Schoeman MC, Cotterill FPDW, Taylor PJ et al (2013) Using potential distributions to explore
environmental correlates of bat species richness in southern Africa: Effects of model selection
and taxonomy. Curr Zool 59:279–293
Seabra A (1898a) Sobre a determinação dos géneros da família Pteropodidae fundada nos carac-
teres extrahidos da fórma, disposição e numero das pregas do paladar e lista das especies d’esta
familia, existentes nas collecções do Museu de Lisboa. Jornal de Sciências Mathemáticas,
Physicas e Naturaes, Segunda Série 5:163–171
Seabra A (1898b) Sobre um caracter importante para a determinação dos generos e especies
dos ‘Microchiropteros’ e lista das espécies d’este grupo existentes nas collecções do Museu
Nacional. Jornal de Sciências Mathemáticas, Physicas e Naturaes, Segunda Série 5:247–258
Seabra AF (1905) Mammiferos e aves da exploração de F. Newton em Angola. Jornal de Sciências
Searchinger TD, Estes L, Thornton K et al (2015) High carbon and biodiversity costs from convert-
ing Africa’s wet savannahs to cropland. Nat Clim Chang 5:481–486
Seiler N, Robbins MM (2016) Factors influencing ranging on community land and crop raiding by
mountain gorillas. Anim Conserv 19:176–188
Seydack AHW (2013) Potamochoerus larvatus Bushpig. In: Kingdon J, Hoffmann M (eds) The
Shortridge GC (1934) The mammals of South West Africa, 2 vols. Heinemann, London
Simões AP, Crawford-Cabral J (1988) Notice on a large hoofed steenbok, Raphicerus campestris
(Mammalia: Artiodactyla), from Angola. Garcia de Orta, Série de Zoologia 15:1–8
Sirami C, Jacobs DS, Cumming GS (2013) Artificial wetlands and surrounding habitats provide
important foraging habitat for bats in agricultural landscapes in the Western Cape, South
Africa. Biol Conserv 164:30–38
Skinner JD (2013) Antidorcas marsupialis Springbok. In: Kingdon J, Hoffmann M (eds) The
Sliwa A (2013) Felis nigripes Black-footed Cat. In: Kingdon J, Hoffmann M (eds) The mam-
mals of Africa. Vol V (Carnivores, pangolins, equids and rhinoceros). Bloomsbury Publishing,
London, pp 203, 544 pp–206
Smith AT, Johnston CH, Alves PC et al (2018) Lagomorphs: pikas, rabbits, and hares of the world.
John Hopkins University Press, Baltimore, p 280
Sokolowski A (1903) Die Antilopenarten der von der Kunene- Sambesi Expedition durchzogenen
Gebiete auf Grund der von der Expedition mitgebrachten Gehörne. In: Baum H (ed) Kunene-
Zambesi expedition. Verlag des Kolonial-Wirtschaftlichen Komitees, Berlin, pp 517–593
St. Leger J (1936) Dr. Karl Jordan’s Expedition to South-West Africa and Angola: Mammals.
Novitates Zoologicae 40:75–81
Statham JCB (1922) Through Angola: a coming colony. W. Blackwood & Sons, London, 388 pp
Statham JCB (1926) With my wife across Africa by Canoe and Caravan. Simpkin Marshall
Hamilton Kent & Co Ltd, London, 323 pp
Struhsaker T, Oates JF, Hart J et al (2008) Cercopithecus neglectus. The IUCN Red List of threat-
Sunquist M, Sunquist F (2009) Family Felidae (Cats). In: Wilson DE, Mittermeier RA (eds)
Handbook of the mammals of the world, Vol I (Carnivores). Lynx Edicíons, Barcelona, pp 54,
728 pp–168
Svensson MS (2017) Conservation and ecology of nocturnal primates: night monkeys, galagos,
pottos and Angwantibos as case studies. PhD thesis, Oxford Brookes University, Oxford
Svensson MS, Bersacola E, Bearder SK et al (2014a) Open sale of elephant ivory in Luanda,
Angola. Oryx 48:13
Svensson MS, Bersacola E, Bearder SK (2014b) Pangolins in Angolan bushmeat markets. IUCN/
SSC Pangolins Specialist Group Newsletter. www.pangolinsg.org/news
Svensson MS, Bersacola E, Mills MS et al (2017) A giant among dwarfs: a new species of galago
(Primates: Galagidae) from Angola. Am J Phys Anthropol 163:30–43
Swart PS (1967) New data on the black-faced impala (Aepyceros melampus petersi Bocage).
Cimbebasia 20:3–18
Taylor ME (2013) Herpestes flavescens Kaokoveld Slender Mongoose (Angolan Slender
Mongoose). In: Kingdon J, Hoffmann M (eds) The mammals of Africa. Vol V (Carnivores,
pangolins, equids and rhinoceroses). Bloomsbury Publishing, London, pp 304, 680 pp–306
Taylor PJ (2016) Dasymys incomtus (errata version published in 2017). The IUCN Red List of
Taylor ME, Goldman CA (1993) The taxonomic status of the African mongooses, Herpestes san-
guineus, H. nigratus, H. pulverulentus and H. ochraceus (Carnivora: Viverridae). Mammalia
57:375–391
Taylor PJ, Grass I, Alberts AJ et al (2018a) Economic value of bat predation services – a review
and new estimates from macadamia orchards. Ecosyst Serv 30:372–381
Taylor PJ, Macdonald A, Goodman SM et al (2018b) Integrative taxonomy resolves three new
cryptic species of small southern African horseshoe bats (Rhinolophus). Zool J Linn Soc
https://doi.org/10.1093/zoolinnean/zly024
Taylor PJ, Neef G, Keith M et al (2018c) Tapping into technology and the biodiversity informatics
revolution: updated terrestrial mammal list of Angola, with new records of mammals from the
Okavango Basin. ZooKeys 2018:51–88
Themido AA (1931) Catalogue des ongulés et siréniens existants dans les collections du Muséum
Zoologique de Coimbra. Memórias e Estudos do Museu Zoológico da Universidade de
Coimbra, Série 1(49):5–22
Themido AA (1946) Mamíferos das colónias portuguesas (catálogo das colecções do Museu
Zoológico de Coimbra). Memórias e Estudos do Museu Zoológico da Universidade de
Coimbra, Série 1(174):1–52
Thomas O (1892) On the species of the Hyracoidea. Proc Zool Soc Lond 1892:50–76
Thomas O (1900) On Equus penricei, a representative of the Mountain Zebra (Equus zebra, L.)
discovered by Mr. W. Penrice in Angola. Ann Mag Nat Hist Ser 7 6:456–466
Thomas O (1904) On the mammals from northern Angola collected by Dr. W. J. Ansorge. Ann
Mag Nat Hist Ser 7 13:405–421
Thomas O (1916) A new Sable Antelope from Angola. Proc Zool Soc Lond 1916:298–301
Thomas O (1926) On Mammals from Ovamboland and the Cunene River, obtained during Capt.
Shortridge’s third Percy Sladen and Kaffrarian Museum Expedition into South-West Africa.
Proc Zool Soc Lond 1926:285–312
Thomas O, Wroughton RC (1905) On a second collection of Mammals obtained by Dr. W. J.
Ansorge in Angola. Ann Mag Nat Hist Ser 7 16:169–178
442 P. Beja et al.
Thorington RW Jr, Koprowski JL, Steele MA et al (2012) Squirrels of the world. The John Hopkins
University Press, Baltimore, p 472
Thouless C (2013) Tragelaphus oryx Common Eland. In: Kingdon J, Hoffmann M (eds) The
Publishing, London, pp 191-198, 680 pp
Thouless C, Dublin HT, Blanc J et al (2016) African elephant status report 2016. Occasional paper
of the IUCN species survival commission no. 60. IUCN, Gland, 37 pp
Timberlake J, Chidumayo E (2011) Miombo ecoregion vision report. Occasional publication in
biodiversity no. 20. Biodiversity Foundation for Africa, Famona, Bulawayo, 76 pp
Trense W (1959) Die Saugetiere Angolas, ihre Beziehungen zucinander, zu den benachbarten
Gebieten und ihre Geschichte (Nach den Ergebnissen der Hamburgischen Angola-Expedition,
1952–1954). Unpublished report. Centro de Zoologia, Instituto de Investigação Científica
Tropical, Lisboa, 211 pp
Trombone T (2016) AMNH mammal collections. American Museum of Natural History, New York
Accessed via GBIF.org on 23 Feb 2018
Tromp S (2011) The effects of past major climatic fluctuations on the genetic structures of fauna
endemic to Namibia’s Granite Inselbergs. PhD thesis, The University of Queensland, Brisbane
Turkalo AK, Wrege PH, Wittemyer G (2017) Slow intrinsic growth rate in forest elephants indi-
cates recovery from poaching will require decades. J Appl Ecol 54:153–159
Tutin CE, Fernandez M (1984) Nationwide census of gorilla (Gorilla g. gorilla) and chimpanzee
(Pan t. troglodytes) populations in Gabon. Am J Primatol 6:313–336
Tutin C, Stokes E, Boesch C et al (2005) Regional action plan for the conservation of Chimpanzees
and Ggorillas in Western Equatorial Africa. IUCN/SSC Primate Specialist Group, Conservation
International, Washington, DC
Vallo P, Guillén-Servent A, Benda P et al (2008) Variation of mitochondrial DNA in the
Hipposideros caffer complex (Chiroptera: Hipposideridae) and its taxonomic implications.
Acta Chiropterologica 10:193–206
van der Straeten E (2008) Notes on the Praomys of Angola with the description of a new species
(Mammalia: Rodentia: Muridae). Stuttgarter Beiträge zur Naturkunde A, Neue Serie 1:123–131
van der Westhuizen J, Thomas J, Haraes L et al (2017) An aerial photographic wildlife survey
of the Iona National Park, Angola, November 2016 to February 2017. Unpublished report.
Ministry of Environment of Angola, Luanda, 31 pp
van Velden J, Wilson K, Biggs D (2018) The evidence for the bushmeat crisis in African savannas:
a systematic quantitative literature review. Biol Conserv 221:345–356
Varian HF (1953) Some African milestones. George Ronald, Oxford, 272 pp
Vaz Pinto P (2018) Evolutionary history of the critically endangered giant sable antelope
(Hippotragus niger variani). Insights into its phylogeography, population genetics, demogra-
phy and conservation. PhD thesis, University of Porto, Porto
Vaz Pinto P (2019) The giant sable antelope: Angola’s national icon. In: Huntley BJ, Russo V,
Springer, Cham
Vaz Pinto P, Veríssimo L (2016) Yellow-backed duiker in miombo woodland in Angola. Gnusletter
35:13
Vaz Pinto P, Beja P, Ferrand N et al (2016) Hybridization following population collapse in a criti-
cally endangered antelope. Sci Rep 6:18788
Veríssimo LN (2008) Mucusso reserve. Larger mammals assessment. Preliminary report.
Unpublished report. United States Agency International Development, Washington, DC, 56 pp
Veron G, Patou M-L, Jennings AP (2018) Systematics and evolution of the mongooses (Herpestidae,
Carnivora). In: Do Linh San E, Sato JJ, Belant JL et al (eds) Small carnivores: evolution, ecol-
ogy, behaviour and conservation. Wiley-Blackwell, Oxford
Wasser SK, Brown L, Mailand C et al (2015) Genetic assignment of large seizures of elephant
ivory reveals Africa’s major poaching hotspots. Science 349:84–87
Weir CR (2019) The Cetaceans (Whales and Dolphins) of Angola. In: Huntley BJ, Russo V,
Wendelen W, Noé N (2017) African Rodentia. Belgian Biodiversity Platform, Brussels Accessed
via GBIF.org on 19 Apr 2018
Wilhelm JH (1933) Das Wild des Okawangogebietes und des Caprivizipfels. J South-West Afr Sci
Soc 6:51–74
Wilkie DS, Wieland M, Boulet H et al (2016) Eating and conserving bushmeat in Africa. Afr J Ecol
54:402–414
Wilson VJ (2013) Sylvicapra grimmia Common Duiker. In: Kingdon J, Hoffmann M (eds) The
Wilson DE, Reeder DM (2005) Mammal species of the world: a taxonomic and geographic refer-
ence, 3rd edn. Johns Hopkins University Press, Baltimore, 2000 pp
Wolf C, Ripple WJ (2016) Prey depletion as a threat to the world’s large carnivores. R Soc Open
Sci 3:160252
Wozencraft WC (2005) Order Carnivora. In: Wilson DE, Reeder DM (eds) Mammal species of
the world: a taxonomic and geographic reference, 3rd edn. Johns Hopkins University Press,
Baltimore, pp 532, 2000 pp–628
Zhang YP, Ryder OA (1995) Different rates of mitochondrial DNA sequence evolution in Kirk’s
Dik-dik (Madoqua-kirkii) populations. Mol Phylogenet Evol 4:291–297
Zinner D, Fickenscher GH, Roos C (2013) Cercopithecidae (old world monkeys). In: Mittermeier
RA, Rylands AB, Wilson DE (eds) Handbook of the mammals of the World. Vol III (Primates).
Lynx Edicíons, Barcelona, pp 550, 952 pp–753
Zukowsky L (1924) Beitrag zur Kenntnis der Säugetiere der nördlichen Teile Deutsch-
Südwestafrikas unter besonderer Berücksichtigung des Großwildes. Archiv für Naturgeschichte,
Abteilung 1:29–164
Zukowsky L (1964) Die Systematic der Gattung Diceros Gray, 1821. Zoologische Garten 30:1–178
Zukowsky L, Haltenorth T (1957) Das Erdferkel (Orycteropus afer) aus Angola, eine eigene
Unterart? Säugetierkundliche Mitteilungen 5:24–126
Chapter 16
The Cetaceans (Whales and Dolphins)
of Angola
Caroline R. Weir
Abstract The history of whale and dolphin (cetacean) research in Angolan waters
is scant. Prior to the 2000s it primarily consisted of information from historical
(1700s to the 1920s) and modern (1920s–1970s) whaling catches, from which
baleen whales and the sperm whale were confirmed. Very few species were added
to Angola’s cetacean checklist between the whaling era and the 2000s. However,
observations since 2003 have confirmed Angola as a range state for at least 28 spe-
cies, comprising seven baleen whales, two sperm whale species, at least two beaked
whales, and at least 17 delphinids. There is potential for approximately seven more
species to be identified in the region based on their known worldwide distributions.
Angola has one of the most diverse cetacean faunas in Africa, and indeed world-
wide, due to its varied seabed topography and transitional ocean climate which
supports both (sub)tropical species and those associated with the Benguela Current.
While no cetacean species are truly endemic to Angola, the country is one of few
confirmed range states for the Critically Endangered Atlantic humpback dolphin
and the Benguela-endemic Heaviside’s dolphin. Those species, together with endan-
gered baleen whales and breeding populations of sperm and humpback whales, are
highlighted as conservation priorities.
Keywords Benguela current · Checklist · Conservation · Endangered species ·

Endemism · Whaling
Introduction
The occurrence of cetaceans along the west coast of Africa in the eastern tropical
Atlantic (ETA) is poorly-studied, due to factors including remoteness, the history of
political unrest in many countries, deficiencies in funding and logistical support
C. R. Weir (*)
Ketos Ecology, Kingsbridge, Devon, UK
e-mail: caroline.weir@ketosecology.co.uk

https://doi.org/10.1007/978-3-030-03083-4_16
446 C. R. Weir
(especially for marine work requiring boats), and a lack of training programmes to
support local marine scientists (Jefferson et al. 1997; Weir 2010a, 2011a,b). Located
at the southern limit of the ETA, Angola is expected to support a diverse cetacean
community due to its varied marine environment. This chapter provides the history
of Angolan cetacean research, reviews cetacean biodiversity and identifies priorities
for future research and conservation options.
Methods
Study Area
Angolan waters are defined as marine habitat from the coast to the 200 nautical mile
seaward limit of the Exclusive Economic Zone (EEZ), which is located in oceanic
habitat over 4000 m deep (Fig. 16.1). They extend from the southern border with
Namibia (17°15′S) northwards to the border with the Republic of Congo in Cabinda
(5°02′S), but excluding the Democratic Republic of the Congo (DRC) EEZ which
divides Angola from the exclave of Cabinda. Some maritime areas in the northernmost
EEZ are the subject of disputed ownership with neighbouring countries (Fig. 16.1),
but are included here in the non-political context of assessing cetacean occurrence.
Weir (2011a) described the oceanography of the Angolan EEZ as habitat for
cetaceans. The Angolan continental shelf is widest in the north, extending to 80 km
from the coast off Soyo where it is intersected by the deep Congo Canyon at the
mouth of the Congo River. In the southern part of the country, the shelf is narrow
and depth increases strongly, bringing deep waters (>1000 m) to within 15 km of the
coast in places. The region is predominantly tropical, with warm (>24 °C) nutrient-
poor water flowing southward from the Gulf of Guinea as the Angola Current.
However, the Benguela Current influences the southern area, bringing nutrient-rich
cold water northwards from Namibia. The two currents converge at latitudes of
between 14° and 16°S (depending on season) to form the Angola–Benguela Front
(Fig. 16.1).
Data
Published (and some available unpublished) papers and reports were reviewed for
information on Angolan cetaceans (see Weir 2011a). Whaling catch statistics were
acquired from the International Whaling Commission (IWC). Since 2003, Marine
Mammal Observers (MMOs), sometimes supported by Passive Acoustic Monitoring
(PAM), have been used during seismic surveys by the oil and gas industry to miti-
gate the potential impacts of airgun sound on cetaceans (Weir 2008). With the
exception of published subsets, MMO data are not publicly available and are there-
fore not included here.
16 The Cetaceans (Whales and Dolphins) of Angola 447
Fig. 16.1 Angolan waters showing the places and major current systems mentioned in this chap-
ter. Hatched areas show some areas of disputed ownership of the Exclusive Economic Zone (EEZ)
with neighbouring countries
Species Identification
Cetaceans are often seen briefly and only partially by an observer, and there are
morphological similarities between many species in the ETA region (e.g. within
Stenella dolphins, beaked whales and Balaenoptera whales) that causes confusion.
High potential for species misidentification exists, even for established cetacean
observers and trained MMOs (many of whom lack previous field experience with
448 C. R. Weir
the particular species occurring off Angola). Published records therefore require
careful evaluation (e.g. Best 2001; Fertl et al. 2003; Weir et al. 2014), particularly
records originating prior to the 2000s, after which knowledge of key identification
features increased markedly with the advent of digital photography, modern field
guides and genetic work. Consequently, some Angolan records were not considered
sufficiently well-supported for inclusion (e.g., Brown 1959, Mörzer Bruyns 1971,
Tormosov et al. 1980).
History of Cetacean Research in Angola
Angola’s Whaling Era
Whaling has been practiced since prehistoric times, and whaling data provides the
earliest information available on the species identification, distribution, migrations
and population status of whale stocks around the world. Whaling also generated
much of the best-available information on the life histories, morphology and diet of
large whales. Consequently, the whaling era is still considered a prime source of
scientific data on the larger baleen whales and the sperm whale (Physeter
macrocephalus).
It was not until the 1700s that American pelagic whalers first visited the west
coast of Africa in search of the relatively slow-moving and oil-rich sperm whales
and southern right whales (Eubalaena australis). They reached the coast of Angola
by 1770 (Best 1981), and catches from this period onward provide the earliest docu-
mentation of whale species in Angola. The distribution of certain whales as shown
by logbook records of American whale ships, published by Charles Haskins
Townsend in 1935, included the capture locations of over 50,000 whales taken dur-
ing American pelagic whaling between 1761 and 1920, including three species from
Angolan waters (sperm whales, southern right whales and humpback whales,
Megaptera novaeangliae: Fig. 16.2). Similar and expanded analyses of whaling
logbook catch datasets including Angolan waters have also been published by other
authors (e.g., Richards 2009; Smith et al. 2012).
Whaling changed drastically from the mid-1800s with the development of
exploding harpoon guns, modern steam-driven whaling boats (‘catcher boats’),
cannon-fired bow-mounted harpoons and the technique of inflating dead whales
with air to keep them afloat (Harmer 1928; Mackintosh 1965; Tønnessen and
Johnsen 1982). Species that had previously been inaccessible to whalers, especially
the Balaenoptera whales that were fast-swimming and sank after death, could now
be harvested, and were either towed to shore stations or processed at factory vessels
moored in coastal bays. Shore-based whaling stations were established in several
African countries during the early 1900s (Tønnessen and Johnsen 1982; Best 1994).
Summarised statistics on whale catches worldwide since 1900 (together with some
incomplete information on catches taken in the late 1800s) are maintained by the
IWC (Allison 2016a). There is also a catch database for individual captures that
Fig. 16.2 Distribution of whale catch positions in the Angola EEZ. MV Sierra catches from
the IWC database (Allison 2016b). Digitised Townsend (1935) charts are available from
https://canada.wcs.org/wild-places/global-conservation/townsend-whaling-charts.aspx
contains date, length, sex, foetus details, stomach contents and location (when those
are available; Allison 2016b). These databases are continually updated (Allison and
Smith 2004), and consequently the total species catches reported by various sources
has altered over time (e.g. Best 1994; Figueiredo and Weir 2014; this chapter).
Catches of whales in Angola since 1900 are presented in Table 16.1.
The first modern coastal whaling operation in Angola was established at Tômbwa
(formerly Porto Alexandre), with the moored Norwegian factory ship Ambra taking
around 237 whales in 1909 (number revised by the IWC from 270 whales in earlier
sources; Figueiredo 1960; Tønnessen and Johnsen 1982; Best 1994). The Ambra
returned to Tômbwa in 1910 and took 650 whales, with a second operation (shore
station and a Portuguese catcher vessel) commencing at Moçâmedes and taking
450 C. R. Weir
Table 16.1 Estimated whale catches in Angola from the International Whaling Commission
databases (Allison 2016a, b)
Sei/
Year Locations Blue Fin Sperm Humpback Bryde’s Right Minke Total
Shore-based operations
1909 Tômbwa 1 0 0 236 0 0 0 237
1910 Tômbwa, Moçâmedes 2 1 0 718 0 0 0 721
1911 Tômbwa, Lobito, Baía 2 2 0 2281 4 0 0 2289
dos Elefantes, Baía
dos Tigres,
Moçâmedes
1912 Tômbwa, Baía dos 0 0 18 3417 0 0 0 3435
Elefantes, Baía dos
Tigres, Moçâmedes
1913 Tômbwa, Baía dos 121 38 39 2419 700 1 0 3318
Tigres, Moçâmedes
1914 Tômbwa, Baía dos 542 200 138 596 102 0 0 1578
Tigres, Moçâmedes
1915 Tômbwa, Baía dos 360 260 79 201 79 0 0 979
Elefantes, Moçâmedes
1916 Baía dos Elefantes, 118 85 26 65 26 0 0 320
Moçâmedes
1923 Pelagic floating 168 26 17 2 0 0 0 213
factory just outside of
territorial waters
1924 Baía dos Elefantes 75 17 17 47 274 0 0 430
1925 Baía dos Elefantes 134 42 27 17 68 0 0 288
1926 Moçâmedes 303 40 14 6 33 0 0 396
1927 Moçâmedes 186 73 3 3 305 0 0 570
1928 Moçâmedes 58 32 141 37 246 0 0 514
Total shore landings 2070 816 1837 10,045 519 1 0 15,288
Pelagic operations
1934 Angola EEZ 1 21 44 7 10 0 0 83
1936 Angola EEZ 0 1 2 17 6 0 0 26
1956 Angola EEZ 0 0 20 0 0 0 0 20
1971 Angola EEZ 0 0 44 0 234 0 0 278
1972 Angola EEZ 0 0 0 0 10 0 1 11
1973 Angola EEZ 0 0 0 0 228 0 2 230
1974 Angola EEZ 0 0 0 0 221 0 2 223
1975 Angola EEZ 0 0 42 0 100 0 0 142
1976 Angola EEZ 0 0 48 0 0 0 0 48
Total pelagic catches 1 22 200 24 809 0 5 1061
around 70 whales (Mackintosh 1942; Best 1994; Allison 2016a, b). Operations
increased during 1911, with the issuing of five licenses to Norwegian floating fac-
tory ships (based at Tômbwa, Lobito, Baía dos Elefantes and Baía dos Tigres) at the
end of 1910, and the continuation of the Portuguese operation at Moçâmedes
(Figueiredo 1960, Allison 2016a, b). Whaling in Angola boomed between 1911 and
1914, capturing over 10,000 animals (mostly humpbacks: Table 16.1). However, the
catch in 1914 was half that of 1912 and 1913, and a collapse in whale stocks was
suggested (Figueiredo 1960). The combination of declining whale stocks and the
occurrence of the First World War meant that no whales were caught off Angola
between 1917 and 1922 (Best 1994).
Whaling was re-established off Angola in 1923, with a Norwegian floating fac-
tory ship operating just outside of territorial waters, and coastal operations resuming
at Baía dos Elefantes and Moçâmedes between 1924 and 1928. This second period
did not yield sufficient captures to be profitable (Table 16.1), and marked the end of
coastal whaling from Angolan shore stations (Figueiredo 1960; Tønnessen and
Johnsen 1982).
The 1920s saw the development of new ocean-going factory ships (fitted with a
stern slipway and a flensing station to process whales) that could operate for long
periods with a fleet of smaller catcher vessels and allowed whaling to move into
offshore waters. Between 1934 and 1937 the Norwegian factory ships Pioner,
Haugar and Norskhavet operated in the ETA including Angola. Catches in Angolan
waters over this period included one blue (Balaenoptera musculus), 22 fin (B. phy-
salus), 24 humpback, 16 sei/Bryde’s (B. borealis/B. edeni), and 46 sperm whales
(Allison 2016b). In later decades factory ships opportunistically took sperm whales
encountered while transiting through Angolan waters. For example, the Olympic
Challenger caught 20 in March 1956, the Peder Huse took 41 in early 1971, and the
Sovetskaya Ukraina took 90 in 1975 and 1976 (Mikhalev et al. 1981a; Allison
2016b).
Most recently, the combined catcher/factory vessel MV ‘Run/Sierra’ operated
year-round between South Africa and the Gulf of Guinea during the 1970s. The
IWC database includes 801 whales taken by the vessel in the Angolan EEZ between
1971 and 1975, comprising five minke whales, three sperm whales and 793 ‘sei
whales’ (Fig. 16.2; Allison 2016a,b). However, the Run/Sierra ‘sei whale’ catches
are now considered to predominantly comprise Bryde’s whales (Tønnessen and
Johnsen 1982; Best 1996, 2001).
The composition of whaling catch data altered over time as each species declined
to levels where protection in the Southern Hemisphere was introduced by the IWC,
beginning with the southern right whale in the 1930s, continuing with the blue and
humpback whales in the 1960s, fin and sei whales in the 1970s, and finally with the
worldwide ban on the exploitation of all whale species under the 1986 moratorium.
Consequently, the whaling era in Angola was ended in the 1970s by the protection
of most Southern Hemisphere whale stocks.
452 C. R. Weir
Opportunistic Sightings and Specimen Records
Weir (2011a) recognised a ‘stranding and specimen era’ of cetacean research in the
ETA (1950s–1970s), during which new information emerged on the taxonomy,
morphometry and distribution of many small cetaceans (see Cadenat 1959, Jefferson
et al. 1997). However, the majority of this work was carried out by French scientists
in Mauritania, Senegal and Côte d’Ivoire, and the only information emerging from
Angola during this period appears to be the 1972 paper of Bree and Purves, which
included a single skull from Angola in an evaluation of the Delphinus genus. Some
opportunistic sightings in Angolan waters by the Dutch sea captain Mörzer Bruyns
were also published (Mörzer Bruyns 1968, 1971), although the species identifica-
tion for many of his records cannot be confirmed. Effort has been made to locate
cetacean specimens that may have been captured off Angola during this period and
preserved by naturalists in Lisbon museum collections. However, it appears that no
cetaceans from Angola are present in Portuguese collections (Cornelis Hazevoet,
pers. comm.). The dearth of papers from Angola in this period was also noted in the
compilation of African cetacean research by Elwen et al. (2011).
During the 1980s and 1990s a few publications from the wider Atlantic region
included opportunistic at-sea sightings (species identifications unsupported) from
Angolan waters, for example Tormosov et al. (1980), Mikhalev et al. (1981b) and
Wilson et al. (1987). In 1997 Jefferson et al. published a review of dolphin and por-
poise records off West Africa, but their study area (to 6°S) included only the exclave
of Cabinda and not the rest of Angola. The only ‘Angolan’ cetacean records located
by Jefferson et al. (1997) were common dolphins (Delphinus sp.) reported
by Simmons (1968). However, careful reading of Simmons (1968) indicates that the
observations were actually recorded off Cape Palmas in Liberia rather than Angola.
Targeted At-Sea Cetacean Surveys
Although instability related to the Angolan civil war from 1975 to 2002 is known to
have interrupted field studies of terrestrial fauna (other chapters, this volume), dedi-
cated cetacean research had still not yet developed prior to the outbreak of war. In
fact, the first dedicated field study of cetaceans in Angolan waters began during the
final period of the war in September 1998, when the Whale Unit of the Mammal
Research Institute in South Africa was invited to northern Angola (6°52′S) by an oil
company to conduct a preliminary investigation into large numbers of humpback
whales reported in the area. This initial field study was successful in acquiring acous-
tic and behavioural data, photographing whale tail flukes for photo-identification and
acquiring 13 genetic samples via biopsy sampling (Best et al. 1999). Although the
authors recommended that a full survey programme should be initiated to assess the
distribution, abundance and status of humpback whales in Angolan waters using
aerial surveys and small boat work, such work never developed.
During the early 2000s, some cetacean data were collected concurrently with
pelagic fish abundance assessments in Angolan waters as part of an agreement
between the Norwegian Institute of Marine Research (IMR) and the Angolan
Instituto de Investigação Pesqueira e Marinha (INIP). The IMR research vessel Dr
Fridtjof Nansen surveyed a series of transects across the continental shelf in the
Angolan EEZ. These investigations were carried out in cooperation with the
Benguela Current Large Marine Ecosystem (BCLME) research programme. Cruise
reports outlining the fish stock results are available from the IMR website, and ceta-
cean observations are included for surveys (all between July and September) in
2003 (Krakstad et al. 2003), in 2004 (Axelsen et al. 2004), in 2005 (Axelsen et al.
2005, Roux et al. 2007), and in 2015 (Michalsen et al. 2015).
The Best et al. (1999) study was the first of several suites of cetacean survey
work in Angola to be associated with, and funded by, the burgeoning oil and gas
industry. From 2003 many oil companies began to use MMOs during their seismic
surveys in Angolan waters, leading to a sudden increase in the potential for biolo-
gists to use geophysical survey vessels as ‘platforms of opportunity’ to collect data
on cetacean occurrence. This was a landmark development in the documentation of
Angola’s cetacean biodiversity, since many seismic surveys covered deep, oceanic
waters that had previously been inaccessible to cetacean scientists. A resulting surge
of information on Angolan cetacean occurrence was published from 2006 to 2014
including: (1) the documentation of species records for Angola (Weir 2006a, b, c,
Weir et al. 2008, 2010, 2014); (2) evaluations of seasonal relative abundance and
spatial distribution (Weir 2007, 2011a, b); (3) examinations of morphology and tax-
onomy (Weir and Coles 2007, Weir et al. 2014); (4) assessment of habitat prefer-
ences (Weir et al. 2012); and (5) studies of behaviour (Weir 2008). Weir (2010a)
also published a comprehensive review of cetacean records in the Angola to Gulf of
Guinea region, which together with her fieldwork on oceanic cetaceans and Atlantic
humpback dolphins was published as the first doctoral thesis focused on Angolan
cetaceans (Weir 2011a).
Between 2008 and 2009, some marine mammal survey work was also carried out
in association with the construction of a Liquefied Natural Gas (LNG) terminal at
the mouth of the Congo River at Soyo, including the use of Marine Autonomous
Recording Units (MARUs) between March and December 2008 at two locations
along the edge of the Congo Canyon (6°S). The MARUs recorded singing hump-
back whales between June and early December (Cerchio et al. 2014), and blue
whale calls on one date in October (Cerchio et al. 2010).
The year 2008 saw the onset of independent (non-industry) cetacean field
research, when Weir (2009, 2011a) visited Namibe Province in southern Angola
during two seasons to conduct an ecological study of the Atlantic humpback dol-
phin. That work provided the first comprehensive assessment of an Atlantic hump-
back dolphin population, collecting information on abundance (via
photo-identification), distribution, movements, seasonality and behaviour ( including
vocal behaviour: Weir 2010b). The study also produced information on several
other cetacean species in coastal waters (Weir 2010c).
454 C. R. Weir
Cetacean Species Recorded in Angola
A checklist of Angolan cetacean species is provided in Table 16.2 and some images

of the most frequently-recorded species are shown in Fig. 16.3. The SMM (2018)
currently recognises 89 species of cetacean worldwide, of which 28 (including
unidentified beaked whales of the Mesoplodon genus, and only accounting for a
single species of common dolphin) have been confirmed to occur in Angola to date.
At least seven further species might potentially be added to Angola’s fauna in the
future.
Baleen Whales
Southern right whale—The majority of Angolan records are from Baía dos Tigres
(17°S), which was the northernmost ground for southern right whale catches in the
1700s and 1800s; over 30 were taken there in 1801 (Best 1981; Richards 2009).
Catches occurred predominantly in June and July (and thus likely represent a winter
breeding presence: Best 1981). The northernmost record in Angola is at approxi-
mately 6°S to the southwest of the Congo River mouth (Townsend 1935), but may
be atypical. An animal taken off Tômbwa in 1913 is the only record in the 1900s
(Table 16.1; Allison 2016a). Best (1990) reported that a catch of 17 right whales at
Baía dos Elefantes during 1925 was probably erroneous and actually related to
Bryde’s whales.
Blue whale―A comprehensive review of blue whale records in Angolan waters
was provided by Figueiredo and Weir (2014). Over 2000 blue whales were captured
off Angola between 1909 and 1928 (Table 16.1; Allison 2016a), and all were landed
at stations in the southern half of Angola (south of 13°S). A single animal was also
taken close to Baía dos Tigres in 1934 (Figueiredo and Weir 2014). Several blue
whale calls were recorded on an acoustic device off the Congo River mouth (6°S) in
October 2008 (Cerchio et al. 2010). Four photographically-verified sightings of
blue whales were recently reported from deep waters (>1000 m) off central Angola,
at latitudes between 11 and 12°30′S (Figueiredo and Weir 2014). The presence of
calves in whaling catches and one sighting indicates the potential use of Angolan
waters as a calving or nursery ground (Figueiredo and Weir 2014).
Fin whale―Primarily documented from whaling catches, with over 800 animals
captured off Angola between 1910 and 1928, and an additional 22 taken by pelagic
whalers between 1934 and 1936 (Table 16.1; Allison 2016a, b). Four sightings were
reported off Angola between 2003 and 2006 (Weir 2007); however, two of those
were downgraded after subsequent evaluation (Weir 2011a, b). The two remaining
sightings occurred in deep-water (>1500 m) during winter (August).
Sei and Bryde’s whales―It is considered that the majority of reported ‘sei
whale’ catches in the ETA were misidentifications and more likely comprised
Bryde’s whales (Harmer 1928, Ruud 1952, Best 1994, 1996, 2001). An estimated
total of 1837 sei/Bryde’s whales were landed at Angolan shore stations between
Table 16.2 Cetacean species confirmed in Angola. IUCN (November 2018) conservation status:
DD (Data Deficient); LC (Least Concern); NT (Near Threatened); VU (Vulnerable); EN
(Endangered); CR (Critically Endangered)
English name Portuguese name Scientific name IUCN
Southern right Baleia-franca-austral Eubalaena australis LC
whale
Blue whale Baleia-azul Balaenoptera musculus EN
Fin whale Baleia-comum Balaenoptera physalus VU
Sei whale Baleia-sardinheira Balaenoptera borealis EN
Bryde’s whale Baleia-de-Bryde Balaenoptera brydei/ LC
B. edeni
Antarctic minke Baleia-anã-antártica Balaenoptera bonaerensis NT
whale
Humpback whale Baleia-de-bossa Megaptera novaeangliae LC
Sperm whale Cachalote Physeter macrocephalus VU
Dwarf sperm Cachalote-anão Kogia sima DD
whale
Cuvier’s beaked Zífio; Baleia-de-bico-de-Cuvier Ziphius cavirostris LC
whale
Mesoplodon sp. Baleia-de-bico Mesoplodon sp. DD
Killer whale Orca Orcinus orca DD
Short-finned pilot Baleia-piloto-tropical Globicephala LC
whale macrorhynchus
False killer whale Falsa-orca Pseudorca crassidens NT
Melon-headed Cabeça-de-melão Peponocephala electra LC
whale
Atlantic Golfinho-de-bossa do Atlântico; Sousa teuszii CR
humpback dolphin Golfinho-corcunda do Atlântico
Rough-toothed Caldeirão; Golfinho-de-bico-comprido Steno bredanensis LC
dolphin
Dusky dolphin Golfinho-cinzento Lagenorhynchus obscurus DD
Risso’s dolphin Grampo; Golfinho-de-Risso Grampus griseus LC
Common Roaz-corvineiro; Golfinho-roaz Tursiops truncatus LC
bottlenose dolphin
Pantropical Golfinho-malhado-pantropical Stenella attenuata LC
spotted dolphin
Atlantic spotted Golfinho-malhado do Atlântico; Stenella frontalis LC
dolphin Golfinho-pintado
Spinner dolphin Golfinho-fiandeiro-de-bico-comprido; Stenella longirostris LC
Golfinho-fiandeiro
Clymene dolphin Golfinho-fiandeiro-de-bico-curto Stenella clymene LC
Striped dolphin Golfinho-riscado Stenella coeruleoalba LC
Common dolphin Golfinho-comum Delphinus sp. LC/
sp. DD
Fraser’s dolphin Golfinho-de-Fraser; Lagenodelphis hosei LC
Golfinho-do-Bornéu
Heaviside’s Golfinho-de-Heaviside Cephalorhynchus NT
dolphin heavisidii
Fig. 16.3 Photographs of the 10 most frequently-recorded cetacean species in Angolan waters
(>55 records; Weir 2011a, b): (a) Bryde’s whale; (b) humpback whale; (c) sperm whale; (d) short-
finned pilot whale; (e) Atlantic humpback dolphin; (f) Risso’s dolphin; (g) bottlenose dolphin; (h)
Atlantic spotted dolphin; (i) striped dolphin; and (j) common dolphin. All photographs taken in
Angolan waters by the author
1911 and 1928, with a further 809 animals taken by pelagic whalers between 1934
and 1975 (Table 16.1; Harmer 1928; Best 1994; Allison 2016a, b). The majority of
pelagic catches were included in the comprehensive assessment of the distribution,
migration and diet of ETA Bryde’s whales by Best (1996, 2001). Only one sighting
of sei whales has been reported for Angola; two animals observed in deep water
southwest of Soyo during August 2004 (Weir 2007). In contrast, 63 sightings of
Bryde’s whales were recorded off northern Angola, mostly in oceanic waters of
1000 to 3000 m depth (Weir 2007, 2011a, b). Bryde’s whales have also been con-
firmed in central and southern Angola, from the Nansen surveys (Axelsen et al.
2004, 2005), during coastal dolphin surveys off Namibe Province (Weir 2010c),
north of Baía dos Tigres (Dyer 2007), and off Tômbwa and Lobito (Olsen 1913).
Best (1996, 2001) described a seasonal migration of the offshore Bryde’s whale
population in and out of Angolan waters. However, sightings have been reported
year-round (Weir 2007, 2010c, 2011a, b), although seasonal fluctuations occur. For
example, Weir (2010c) only recorded Bryde’s whales during the summer in coastal
Namibe Province, while most sightings from northern Angola are in winter and
spring (August and September; Weir 2011a, b).
Minke whale―While there are unspecific mentions of minke whales off Angola
in several sources (e.g. Mörzer Bruyns 1971; Stewart and Leatherwood 1985), the
number of verified records is very low. The vessel Run/Sierra caught five Antarctic
minke whales at latitudes of 5°S to 16°S (Allison 2016b). An Antarctic minke whale
stranded at the Coroca River mouth near Tômbwa (15°45′S) during March 1970
(photograph held in the Museu do Mar, Cascais, Portugal; Peter Best pers. comm.),
also confirming this species in Angolan waters (Best 2007).
Humpback whale―Townsend (1935) noted that the region between the equator
and 12°S produced the highest nineteenth century humpback catches on the west
coast of Africa, particularly between June and October. Angolan whaling catches
from 1909 to 1928 included over 10,000 humpbacks, with a strong peak between
1911 and 1913 (Table 16.1; Best 1994; Allison 2016a). No new information on
humpback whales emerged until the 1998 field study off northern Angola by Best
et al. (1999), which recorded many surface-active groups, cow-calf pairs and singing
males, and led those authors to conclude that the area was (or was very close to) a
breeding ground. Acoustic monitoring off northern Angola (6°S) during 2008
recorded humpback whale singing activity, which was also considered indicative of
breeding behaviour (Cerchio et al. 2014). Numerous sightings of humpback whales
have been recorded during sighting surveys, including in southern Angola (Axelsen
et al. 2004; Dyer 2007; Weir 2010c), central regions (Krakstad et al. 2003; Axelsen
et al. 2005; Roux et al. 2007; Michalsen et al. 2015), and the northern areas off Soyo
and Cabinda (Weir 2007, 2011a, b). The highest densities occur over the shelf, but
sightings also occur far offshore (to at least 4000 m depth: Weir 2011b). Strong sea-
sonality is evident in Angolan waters, with all captures, sightings and acoustic
records occurring between May and January, and with a strong peak between July
and October (Weir 2011a, b; Cerchio et al. 2014). The humpback whales using
Angolan waters originate from Southern Hemisphere IWC stock B (Rosenbaum
et al. 2009), and migrate between breeding areas in the ETA and summer Antarctic
feeding grounds.
458 C. R. Weir
Synopsis―Both whaling data and sightings surveys indicate that humpback and
Bryde’s whales are the most numerous baleen whale species in the region, with the
remaining species either naturally less common or still to recover from whaling
exploitation. The timing of catches (Allison 2016a, b), and observations from year-
round sighting surveys (Weir 2011a, b), indicate that most baleen whales exhibit
strong seasonality in Angolan waters, occurring during the austral winter and spring
(June to October) which corresponds with the breeding period of Southern
Hemisphere whale stocks. There is evidence for breeding in Angolan waters of at
least humpback whales and blue whales. Many humpback whales may also use
Angolan waters as a migratory corridor to reach well-established calving grounds
off Gabon and in the Gulf of Guinea (Rosenbaum et al. 2009). The Bryde’s whale is
one of few baleen whale species that inhabit warm waters year-round (Best 2001),
and its seasonal movements in Angolan waters more likely relate to prey availabil-
ity. Although there are no confirmed records to date in Angola, three additional
baleen whale species may be recorded in the future including two documented else-
where from warm Atlantic Ocean waters (Common minke whale Balaenoptera acu-
torostrata and Omura’s whale B. omuraii) and one cool water species that has been
recorded further south off northern Namibia (19°28′S; pygmy right whale Caperea
marginata; Leeney et al. 2013) and could extend into the Benguela-influenced
waters of southern Angola.
Sperm Whales
Sperm whale―The whaling charts of Townsend (1935) reveal numerous sperm

whale captures on the ‘Coast of Africa’ whaling ground (3–23°S), including the
entire coast of Angola. Over 500 sperm whales were landed at Angolan shore sta-
tions between 1912 and 1928, with an additional 200 taken by pelagic fleets from
the 1930s to the 1970s (Table 16.1; Harmer 1928; Mikhalev et al. 1981a; Best 1994;
Allison 2016a, b). Sighting surveys in Angolan waters found that the sperm whale
was one of the most frequently-recorded cetacean species (Weir 2011a, b). Sightings
were distributed exclusively in deep waters from 800 to 3800 m and usually com-
prised singletons or nursery schools of ≤20 animals, although loose aggregations of
up to 65 animals have been observed (Weir 2011a, b). Sperm whales are present in
Angolan waters year-round, but there may be fine-scale spatio-temporal fluctuations
in their occurrence and an overall preference for warmer waters where sea surface
temperatures (SSTs) exceed 23 °C (Weir et al. 2012).
Dwarf sperm whale―Twenty-six sightings of this species were reported by Weir
(2011a, b) from Angolan waters, comprising small groups of one to three animals
seen in deep waters in the 1000–2000 m range. The closely-related pygmy sperm
whale (Kogia breviceps) has not been confirmed off Angola to date, but may be
expected to occur based on its worldwide distribution (Caldwell and Caldwell 1989).
Beaked Whales
Of the 22 currently-recognised beaked whale species (SMM 2018), only the

Cuvier’s beaked whale (Ziphius cavirostris) has been positively-confirmed in
Angolan waters to date, with four sightings in slope waters of 847–2040 m depth
(Weir 2006a, 2011a, b). Eleven additional sightings of unidentified beaked whales
(including Mesoplodon species) are documented off Angola in deep waters exceed-
ing 730 m (Weir 2006a, 2011a, b). Mörzer Bruyns (1968) also observed three
unidentified Mesoplodon whales off Angola in July 1966. There is one record of a
stranded adult male Gervais’ beaked whale (Mesoplodon europaeus) from the
mouth of the Cunene River (on the Angola–Namibia border) in 1997. Although
considered a Namibian record (Griffin and Coetzee 2005), this stranding is highly-
supportive of an occurrence in Angolan waters. The warm Atlantic distribution of
Blainville’s beaked whale (M. densirostris; MacLeod et al. 2006) is also indicative
of a likely occurrence off Angola.
Delphinids
Killer whale―Records in Angola include observations south of Moçâmedes during

July 1966 (Mörzer Bruyns 1971), from a pelagic whaler (Mikhalev et al. 1981b),
and from the Nansen surveys (Axelsen et al. 2005). Weir et al. (2010) provided
information on 18 sightings from Angolan waters between 1991 and 2008. An addi-
tional two sightings were reported in 2009 (Weir 2011a, b). Sightings have com-
prised 1 to 12 animals observed at latitudes of 5°S to 12°S and in water depths
ranging from very shallow coastal waters to well over 2000 m. In January 2005, a
group of five killer whales was seen attacking sperm whales off northern Angola
(Weir et al. 2010).
Short-finned pilot whale―All pilot whales observed in northern Angola to date
have been conclusively identified as the short-finned pilot whale (Globicephala
macrorhynchus). However, it is likely that long-finned pilot whales (G. melas) also
occur in the Benguela Current-influenced areas and will be confirmed in the future.
Pilot whales were the third most frequently-observed species in Angolan waters
(perhaps partly because they are easy to identify at distance), with 125 sightings
reported by Weir (2011a, b). Over 94% of sightings consisted of ≤50 animals, and
all records were located over the slope or in oceanic waters (400–4000 m depth).
This species was also reported by Krakstad et al. (2003), Axelsen et al. (2004, 2005)
and Dyer (2007).
False killer whale―Thirteen sightings of false killer whales were reported in
oceanic habitat (1400–2600 m depth) off northern Angola, comprising groups of
2–50 animals (Weir 2011a, b).
460 C. R. Weir
Melon-headed whale―Four sightings of melon-headed whales have been

reported in oceanic waters (>1300 m depth) off the northern half of Angola (Weir
2011a, b). Three of the schools were large, comprising 100–300 animals.
Atlantic humpback dolphin―First documented in Angola from a photograph
taken near Tômbwa in 2004 (Van Waerebeek et al. 2004). The ‘numerous reports’
cited in Van Waerebeek et al. (2004) from opportunistic observers in northern
Angola and Cabinda have not been upheld by subsequent scientific fieldwork in
those areas (Weir 2009, 2011a; Weir and Collins 2015), and are considered likely
misidentifications. Dedicated photo-identification surveys in Namibe Province in
January and June/July 2008 revealed a very small population of 10 humpback dol-
phins that inhabit nearshore (<1.4 km) waters along a small 40 km stretch of coast
year-round, and use the area for both feeding and calving (Weir 2009, 2010c).
Published information on the whistles of this species represents one of few cetacean
acoustic studies in Angola to date (Weir 2010b).
Rough-toothed dolphin―Weir (2006b) reported three sightings of rough-
toothed dolphins from Angolan waters in 2004 and 2005, while Weir (2011a, b)
added an additional 15 sightings up until 2009. All records were seaward of the
shelf (700–2200 m), and usually comprised ≤60 animals although several larger
groups were observed. An interesting account of an interaction between rough-
toothed dolphins and a sport fishing tournament off Luanda was described by Weir
and Nicolson (2014), with dolphins taking bait from the fishing lines of several
vessels.
Dusky dolphin―Two were photographed off Lobito (12°22′S: Kramer 1961;
Findlay et al. 1992; Best and Meÿer 2009). A group of 40 was reported by Axelsen
et al. (2004) at 16°48′S off Baía dos Tigres, while four schools of 6–40 animals
were recorded during August 2005 south of 16°06′S (Axelsen et al. 2005; Roux
et al. 2007). Dyer (2007) observed a group of six at 15°40′S just north of Tômbwa.
Dusky dolphins inhabit cool Benguela Current-influenced waters along the west
coast of Africa, and are likely limited to southern Angola.
Risso’s dolphin―A total of 75 Angolan sightings was described in Weir (2011a,
b), and included in the global review of Jefferson et al. (2013). Sightings occurred
in slope and oceanic habitat from 900 to 2500 m depth. Group size was generally
≤10 animals, but some larger groups of 35–75 animals were recorded.
Bottlenose dolphin―Fifty-six sightings were reported in Angolan waters by
Weir (2011a, b), occurring in water depths varying from 10 m by the coast to 3700 m
in oceanic areas. Group size in Angola is typically small at 15 or fewer animals, and
in oceanic regions they frequently form mixed-species associations with pilot
whales (Weir 2011a, b). They have also been regularly reported during the Nansen
surveys, including mixed groups with pilot whales (Krakstad et al. 2003, Axelsen
et al. 2004, 2005). Weir (2010c) reported 24 sightings (1–50 animals) in the coastal
waters between Tômbwa and Moçâmedes in 2008, with more frequent sightings
during the winter.
Pantropical spotted dolphin―Weir (2011a, b) reported four sightings from

Angola in slope and oceanic habitat (≥820 m depth) north of 8°40’S. The groups
ranged from 50 to 200 animals.
Atlantic spotted dolphin―A total of 101 sightings was recorded by Weir
(2011b), making it the most commonly recorded species of the Stenella genus in
Angola. Water depth ranged from 800 to 3000 m, and group sizes were 1–500
animals.
Spinner dolphin―A single sighting exists for Angola, comprising three animals
in 1000 m depth off northern Angola in 2004 (Weir 2007, 2011a, b). There have
been 11 additional sightings of animals identified as either spinner or Clymene dol-
phins, but too distant to confirm (Weir 2011a).
Clymene dolphin―The first record for Angola was reported by Weir (2006c). A
comprehensive review of Clymene dolphins in the ETA was conducted by Weir
et al. (2014) and included 16 records for Angola from 6°S off the Congo River to
14°S. Clymene dolphins in Angola were sighted in water depths ranging from 466
to 2362 m, and in groups of 12 to 1000 animals (Weir et al. 2014).
Striped dolphin―Two sightings were reported by Wilson et al. (1987; No’s
40082 and 40083) at 13°59′S and 09°15′S off central Angola in October 1974. A
total of 66 sightings were reported from the northern half of Angola by Weir (2011a,
b), occurring in slope and oceanic waters from 800 to 2700 m depth.
Common dolphin―The taxonomic status of Delphinus dolphins worldwide
remains unresolved (Cunha et al. 2015). A few Angolan common dolphin skulls
have been included in morphological analyses of the Delphinus genus (Bree and
Purves 1972), identifying ‘short-beaked’ and ‘long-beaked’ forms (Van Waerebeek
1997). However, these may be morphotypes of a single species (Cunha et al. 2015).
The external appearance of Angolan animals appears intermediate between short-
beaked (D. delphis) and long-beaked (D. capensis) common dolphins (Weir and
Coles 2007; Weir 2011a), and until their taxonomy is better clarified then they are
referred to simply as ‘common dolphin’. The surveys by Weir (2011a, b) reported
62 sightings of common dolphins off Angola, including in shelf, slope and oceanic
habitat (to 2600 m depth), and in group sizes of up to 500 animals. Sightings have
been reported as far south as Moçâmedes (15°20′S: Axelsen et al. 2004). Weir et al.
(2012) identified a preference for cooler SSTs (≥22.1 °C) in Angola, suggesting the
species is associated with areas of upwelling.
Fraser’s dolphin―The occurrence of Fraser’s dolphins off Angola was first
described by Weir et al. (2008) from two sightings recorded in 2007 and 2008. An
additional record was added by Weir (2011a, b). All sightings have occurred at lati-
tudes of around 07°30’S off northern Angola, and in deep waters exceeding 1300 m.
Heaviside’s dolphin―Two animals were caught by a trawler approximately
12 km north of the Cunene River mouth near the Angola-Namibia border (17°09’S:
Findlay et al. 1992; Peter Best pers. comm.). Another was caught in a fishing net off
the Cunene River mouth just south of Angola during January 1982 (Windhoek
Museum specimen WM 11708; Peter Best pers. comm.), supporting an occurrence
in southern Angolan waters. Two Heaviside’s dolphin sightings were recorded
462 C. R. Weir
during the Nansen 2004 surveys, in water depths of 20–120 m and at latitudes south
of 16°48′S between Baía dos Tigres and the Namibian border (Axelsen et al. 2004,
Best 2007). This species appears to inhabit water temperatures ≤15 °C (Best and
Abernethy 1994), and is likely restricted to Benguela-influenced regions in the far
south of Angola (Best 2007).
Synopsis―At least 17 delphinid species have been confirmed in Angolan waters
(assuming only one species of Delphinus). Most are likely to occur year-round,
although there may be seasonal fluctuations in the distributions of some species
depending on the extent of the Benguela Current influence. This applies particularly
to dusky dolphins and Heaviside’s dolphins, which reach the northern limits of their
African distribution range in the southern part of Angola. Sighting surveys indicated
that some delphinid species were relatively more common than others off Angola,
with Atlantic spotted and common dolphins being frequently-sighted, while pan-
tropical spotted and spinner dolphins were far less common. The relative frequency
of dolphin species likely relates to (at least) water temperature, water depth and
productivity, with some niche partitioning evident (Weir et al. 2012). Despite large
amounts of survey effort in suitable habitat, there are no published (verified) sight-
ings of the pygmy killer whale (Feresa attenuata) to date in Angola. This species is
likely to be added to the Angola cetacean list in the future, along with the long-
finned pilot whale.
Endemism
As highly mobile oceanic predators, none of the reported cetacean species are
endemic to Angolan waters. However, four species are endemic to the Atlantic
Ocean, including the Atlantic spotted dolphin, Clymene dolphin, Atlantic hump-
back dolphin and the Heaviside’s dolphin. The latter two species have restricted
geographic ranges, with the Atlantic humpback dolphin occurring only in nearshore
waters of the ETA (Weir and Collins 2015), and the Heaviside’s dolphin occupying
cool shelf waters of the Benguela Current system (Best and Abernethy 1994).
Consequently, Angolan waters are of particular relevance for those species in terms
of their very limited global range.
Cetacean Biodiversity and the Marine Environment
The occurrence of cetacean species is strongly related to seabed topography (i.e.

depth, slope) and oceanographic variables such as SST, turbidity, salinity and chlo-
rophyll (e.g. Davis et al. 2002; Hamazaki 2002). Consequently, cetacean biodiver-
sity in Angola varies according to habitat (Weir et al. 2012).
Large marine ecosystems (LMEs) have been recognised worldwide based on eco-
logical criteria including bathymetry, hydrography, productivity, and
trophically-dependent populations, with the majority of the Angolan EEZ situated

within the Benguela Current LME (Fig. 16.1; Sherman 2014). The Angola Front at 5°S
forms the northern limit of the Benguela Current LME, and the waters off Cabinda
therefore fall into the tropical Guinea Current LME. A biogeographic system to classify
marine regions was also developed by Spalding et al. (2007) for coastal waters. In this
system, the majority of the Angolan EEZ is situated in the Angolan ecoregion of the
Gulf of Guinea province in the Tropical Atlantic realm (Fig. 16.1). However, the north-
ernmost area (north of 6°30′S) falls into the more tropical Gulf of Guinea South ecore-
gion, while the area south of 15°45′S is recognised as an entirely different biogeographic
region located in the Namib ecoregion of the Benguela province in the Temperate
Southern Africa realm (Spalding et al. 2007). Consequently, both the LME (Sherman
2014) and marine ecoregion (Spalding et al. 2007) approaches support transition zones
within the Angolan EEZ between tropical and temperate (Benguela-influenced) biomes.
Cetacean species in Angola can be broadly classified into communities, based on
their occurrence in shelf (less than 200 m depth) versus oceanic (greater than 200 m
depth) waters and on their distribution according to marine ecoregion (which
broadly corresponds with water temperatures). Using this method, three distinct
communities are apparent, with the most diverse comprising the warm water species
found in oceanic waters (Fig. 16.4). A second community inhabits cool shelf waters
in the south of the study area, while the Atlantic humpback dolphin occupies a
unique niche being found only in warm waters on the shelf. There are also six spe-
SHELF (<200 m depth) SHELF AND OCEANIC OCEANIC (>200 m depth)
Cuvier's beaked whale

Mesoplodon sp.
Dwarf sperm whale
Short-finned pilot whale
(SUB)TROPICAL False killer whale
(Gulf of Guinea Melon-headed whale
Atlantic humpback dolphin Rough-toothed dolphin
South and Angolan
ecoregions) Pantropical spotted dolphin
Atlantic spotted dolphin
Spinner dolphin
Clymene dolphin
Striped dolphin
Fraser's dolphin
Humpback whale
Bryde's whale Blue whale
(SUB)TROPICAL TO Antarctic minke whale Fin whale
TEMPERATE Killer whale Sei whale
Bottlenose dolphin Sperm whale
Common dolphin Risso's dolphin
TEMPERATE Southern right whale

(Namibe ecoregion) Dusky dolphin
Heaviside's dolphin
Fig. 16.4 Classification of Angolan cetacean communities. Some species have wider ecological
niches than shown here; for example, blue, fin and sei whales are found in shelf waters in some
geographic regions, while right whales and dusky dolphins may also be oceanic. However, the
information is based solely on documented occurrence in Angola to date. The species in the grey
box are those with the most cosmopolitan distributions. The Risso’s dolphin is included as a tem-
perate species due to additional sightings of this species during survey work offshore of Lobito
(Weir unpublished data)
464 C. R. Weir
cies that might be expected to occur throughout the temperature range, primarily
comprising the migrating baleen whales and several very cosmopolitan species (e.g.
killer whales, bottlenose dolphin and common dolphin) that occupy wide habitat
ranges (Fig. 16.4). This system is a useful starting point for considering the underly-
ing drivers of cetacean biodiversity off Angola, and further research into species
distribution and environmental parameters should narrow down the habitat prefer-
ences for some species in the future. The seasonally variable and transitional ocean-
ographic environment off Angola explains the high cetacean biodiversity recorded
relative to most other (solely tropical) ETA countries (Weir 2010a, 2011a).
The association of particular cetacean communities with oceanographic biomes
means that species diversity in central and southern Angola will fluctuate on a sea-
sonal basis. The Angola-Benguela Front exhibits spatio-temporal variation over the
year as the Benguela Current strengthens and weakens, and Weir (2011a) showed
corresponding seasonal SST variations of over 7 °C along the Angolan coast.
Consequently, species with preferences for cold or tropical waters may shift in dis-
tribution northwards or southwards in response to seasonal changes in
oceanography.
Environmental parameters also influence the relative abundance of different spe-
cies in Angolan waters. For example, in the genus Stenella the prevalence of Atlantic
spotted dolphins, striped dolphins and Clymene dolphins off Angola in comparison
with very few sightings of pantropical and spinner dolphins, may be the result of the
productive Benguela-influence. Pantropical spotted and spinner dolphins are more
characteristic of tropical oligotrophic waters (Au and Perryman 1985), and are
replaced in more productive, slightly cooler areas by the other members of the genus.
The specific use of Angolan waters by some cetacean species also relates to envi-
ronmental conditions. For example, Cabinda is located in the tropical Gulf of
Guinea LME in the far north of Angola, and has consistently warmer SSTs during
the winter than further south. This may explain why humpback whale calving and
singing behaviour (i.e. breeding activity) has only been confirmed to date in that
region of Angola (Best et al. 1999; Cerchio et al. 2014).
Conservation
There are few published accounts of the conservation issues facing cetaceans in
Angolan waters, but identified threats in other ETA regions include directed takes
(i.e. for human food as ‘marine bushmeat’), bycatch in fishing gear, entanglement,
prey reduction due to over-fishing, habitat loss and degradation (including noise
disturbance and pollution), vessel strikes, marine ecotourism and live captures for
display in aquaria (review by Weir and Pierce 2013).
In 1986, the International Whaling Commission’s moratorium effectively ended
commercial whaling in Angolan waters, but there is also evidence for the capture of
small cetaceans. Brito and Vieira (2009) found reports of catches of ‘toninhas’
(unidentified dolphins) in Angola between 1940 and 1954 in the national fishing
books kept in the National Institute of Statistics in Lisbon, with an average of 20

dolphins landed annually. Those authors considered it likely that bow-riding dol-
phins were purposefully harpooned by hand for their meat (Brito and Vieira 2009).
There are no specific published records of cetacean fisheries bycatch in Angolan
waters, but bycatch affects small cetaceans worldwide and its absence in the litera-
ture can be considered a lack of reporting rather than a lack of occurrence in Angola.
Weir et al. (2011) reported high numbers of artisanal gillnets deployed in nearshore
waters in Namibe Province, and identified them as a major threat to coastal dolphins
in the area. Weir and Nicolson (2014) described the potential for bycatch of dol-
phins during depredation of recreational and commercial fisheries.
Several studies have reported the potential for seismic survey operations to dis-
turb cetaceans in Angola, including spatial avoidance (Weir 2008) and reductions in
singing by humpback whales (Cerchio et al. 2014).
The lack of population size information and the absence of quantitative data on
impacts on Angolan cetaceans make it impossible to currently assess status and
conservation threats. However, the small population of humpback dolphins identi-
fied in Namibe Province is clearly of high conservation concern (Weir 2009; Weir
et al. 2011), especially given the recent upgrading of the species to Critically
Endangered by the IUCN (2018).
Research in Angola: What Next?
Cetacean research in Angola is still in its infancy. Although the species checklist is
more complete for Angola than many other ETA countries (Weir 2010a, 2011a), this
relates predominantly to MMO data collected during offshore seismic surveys.
MMO data can provide information on ‘presence’, species composition and group
sizes, but cannot provide robust ‘absence’ data due to the unknown potentially-
adverse affects of airgun sound on species occurrence and the fact that sightings
often remain unidentified to species level due to the lack of ability to approach
animals.
Most survey effort and records of cetaceans to date have originated from the
(sub)tropical waters between Luanda and Cabinda, where the oil and gas industry is
most active (Weir 2011a, b). With the exception of several short periods of effort
(e.g. Axelsen et al. 2004, 2005; Roux et al. 2007; Weir 2009, 2010c), the waters in
the southern half of Angola have not been surveyed for cetaceans. Consequently,
establishing the year-round species composition, distribution and abundance of
cetaceans in the Benguela-influenced region south of Lobito should be a priority for
future research, especially since whaling captures and recent sightings indicate that
region may be most important for large endangered whales (e.g. blue whale;
Figueiredo and Weir 2014).
Information on population sizes, population structure (via genetic sampling),
spatio-temporal distribution, movements and diet are required for all cetacean spe-
cies in Angolan waters. Critical to this is the development of comprehensive ongo-
466 C. R. Weir
ing training programmes for local biologists in species identification and techniques
such as photo-identification, genetic sampling, necropsying of dead animals and
passive acoustic monitoring. In particular the field identification of cetacean species
takes significant training and field experience, and building this capacity within
Angola will be fundamental to the success of long-term population monitoring.
The collection of quantitative data to assess threats is also highlighted as a
research priority, and could be achieved through a trained bycatch observer pro-
gramme for fishing communities, and monitoring at artisanal and commercial land-
ing sites.
Species priorities for Angolan research include Endangered large whale species
(Table 16.2) and the Critically Endangered Atlantic humpback dolphin. A decade
has passed since Weir’s (2009) study of humpback dolphins in Namibe Province,
and the current status of the species in nearshore waters along the entire coast
requires urgent assessment if it is to be conserved in future decades (Weir et al.
2011). Additionally, Angolan waters are potentially of global importance for breed-
ing populations of sperm whale (Weir 2011a, b), and the waters off Cabinda appear
to comprise a calving area for humpback whales (Best et al. 1999, Weir 2011a, b,
Cerchio et al. 2014). A systematic research programme would be valuable for
informing the management of both of those species.
References
Allison C (2016a) IWC summary catch database Version 6.1; Date: 18 July 2016. Available from
the International Whaling Commission, Cambridge, UK
Allison C (2016b) IWC individual catch database Version 6.1; Date: 18 July 2016. Available from
the International Whaling Commission, Cambridge, UK
Allison C, Smith TD (2004) Progress on the construction of a comprehensive database of twentieth
century whaling catches. Paper SC/56/O27 presented to the International Whaling Commission
Au DWK, Perryman WL (1985) Dolphin habitats in the eastern tropical Pacific. Fish Bull
83:623–643
Axelsen BE, Lutuba-Nsilulu H, Zaera D, et al. (2004) Surveys of the fish resources of Angola.
Survey of the pelagic resources 28 July–27 August 2004. Cruise Reports Dr Fridtjof Nansen.
Available at: http://hdl.handle.net/11250/107204
Axelsen BE, Luyeye N, Zaera D et al. (2005) Surveys of the fish resources of Angola. Survey of
the pelagic resources 16 July–24 August 2005. Cruise Reports Dr Fridtjof Nansen. Available
at: http://hdl.handle.net/11250/107244
Best PB (1981) The status of right whales (Eubalaena glacialis) off South Africa, 1969–1979.
Investigational Report of the Sea Fisheries Institute, South Africa 123:1–44
Best PB (1990) The 1925 catch of right whales off Angola. Rep Int Whaling Commission
40:381–382
Best PB (1994) A review of the catch statistics for modern whaling in southern Africa, 1908-1930.
Reports of the International Whaling Commission 44:467–485
Best PB (1996) Evidence of migration by Bryde’s whales from the offshore population in the
Southeast Atlantic. Rep Int Whaling Commission 46:315–322
Best PB (2001) Distribution and population separation of Bryde’s whale Balaenoptera edeni off
southern Africa. Mar Ecol Progr Ser 220:277–289
Best PB (2007) Whales and dolphins of the southern African Subregion. Cambridge University
Press, Cape Town
Best PB, Abernethy RB (1994) Heaviside’s dolphin Cephalorhynchus heavisidii (Gray, 1828).
In: Ridgway SH, Harrison R (eds) Handbook of marine mammals, volume 5, the first book of
dolphins. Academic, San Diego, pp 289–310
Best PB, Meÿer MA (2009) Neglected but not forgotten: southern Africa’s dusky dolphins. In:
Würsig B, Würsig M (eds) The dusky dolphin: master acrobats off different shores. Elsevier
Science & Technology, Oxford, pp 291–312
Best PB, Reeb D, Morais M, et al. (1999) A preliminary investigation of humpback whales off
northern Angola. Paper SC/51/CAWS33 presented to the IWC Scientific Committee, 12 pp
Bree PJH v, Purves PE (1972) Remarks on the validity of Delphinus bairdii (Cetacea, Delphinidae).
J Mammal 53:372–374
Brito C, Vieira N (2009) Captures of “toninhas” in Angola during the 20th century. Paper SC/61/
SM18 presented to the Scientific Committee of the International Whaling Commission,
Madeira, Portugal, 2009
Brown SG (1959) Whales observed in the Atlantic Ocean: notes on their distribution. Norsk
Hvalfangst-Tidende 48:289–308
Cadenat J (1959) Rapport sur les petits cétacés ouest-africains: résultats des recherches entreprises
sur ces animaux juqu’au mois de mars 1959. Bulletin de l’Institut Français D’Afrique Noire
Série A, Sciences Naturelles 21:1367–1409
Caldwell DK, Caldwell MC (1989) Pygmy sperm whale Kogia breviceps (de Blainville, 1838):
Dwarf sperm whale Kogia sima Owen, 1866. In: Ridgway SH, Harrison R (eds) Handbook
of marine mammals, volume 4, river dolphins and the larger toothed whales. Academic, San
Diego, pp 235–260
Cerchio S, Collins T, Mashburn S et al. (2010) Acoustic evidence of blue whales and other
baleen whale vocalizations off northern Angola. Paper SC/62/SH13 presented to the Scientific
Committee of the International Whaling Commission, Agadir, June 2010. 8 pp. (Available from
the IWC Secretariat, Cambridge, UK)
Cerchio S, Strindberg S, Collins T et al (2014) Seismic surveys negatively affect humpback whale
singing activity off northern Angola. PLoS One 9(3):e86464
Cunha HA, de Castro RL, Secchi ER et al (2015) Molecular and morphological differentiation of
common dolphins (Delphinus sp.) in the Southwestern Atlantic: testing the two species hypoth-
esis in sympatry. PLoS One 10(11):e0140251
Davis RW, Ortega-Ortiz JG, Ribic CA et al (2002) Cetacean habitat in the northern oceanic Gulf
of Mexico. Deep-Sea Res I Oceanogr Res Pap 49:121–142
Dyer BM (2007) Report on top-predator survey of southern Angola including Ilha dos Tigres,
20–29 November 2005. In Kirkman SP (ed) Final Report of the BCLME (Benguela Current
Large Marine Ecosystem) Project on Top Predators as Biological Indicators of Ecosystem in
the BCLME. BCLME Project LMR/EAF/03/02. 381 pp. Available at: http://www.adu.uct.
ac.za/adu/projects/sea-shore-birds/communication/report-bclmetpp
Elwen SH, Findlay KP, Kiszka J et al (2011) Cetacean research in the southern African subregion:
a review of previous studies and current knowledge. Afr J Mar Sci 33:469–493
Fertl D, Jefferson TA, Moreno IB et al (2003) Distribution of the Clymene dolphin Stenella cly-
mene. Mamm Rev 33:253–271
Figueiredo JM (1960) Pescarias de baleia nas províncias africanas Portuguesas. Boletim da Pesca
66:29–37
Figueiredo I, Weir CR (2014) Blue whales (Balaenoptera musculus) off Angola: recent sightings
and evaluation of whaling data. Afr J Mar Sci 36(2):269–278
Findlay KP, Best PB, Ross GJB et al (1992) The distribution of small odontocete cetaceans off the
coasts of South Africa and Namibia. S Afr J Mar Sci 12:237–270
Griffin M, Coetzee CG (2005) Annotated checklist and provisional national conservation status of
Namibian mammals. Technical Reports of Scientific Services No. 4. Directorate of Scientific
Services, Ministry of Environment and Tourism, Windhoek, 207 pp
468 C. R. Weir
Hamazaki T (2002) Spatiotemporal prediction models of cetacean habitats in the mid-western

North Atlantic Ocean (from Cape Hatteras, North Carolina, U.S.A. to Nova Scotia, Canada).
Mar Mamm Sci 18:920–939
Harmer SF (1928) History of whaling. Proc Linnaean Soc Lond 140:51–95
IUCN (2018) The IUCN Red List of Threatened Species. Version 2017–3. www.iucnredlist.org.
Downloaded on 20 January 2018
Jefferson TA, Curry BE, Leatherwood S et al (1997) Dolphins and porpoises of West Africa: a
review of records (Cetacea: Delphinidae, Phocoenidae). Mammalia 61:87–108
Jefferson TA, Weir CR, Anderson RC et al (2013) Global distribution of Risso’s dolphin Grampus
griseus: a review and critical evaluation. Mamm Rev 44:56–68
Krakstad J-O, Vaz-Velho F, Axelsen BE et al. (2003) Surveys of the fish resources of Angola.
Survey of the pelagic resources 20 July–19 August 2003 (Including observations of marine
seabirds and mammals). Cruise Reports Dr Fridtjof Nansen. Available at: http://hdl.handle.
net/11250/107254
Kramer MO (1961) Dolphins have the laugh on us … as far as speed goes. South African Yachting
News 1961:28–30
Leeney RH, Post K, Best PB et al (2013) Pygmy right whale Caperea marginata records from
Namibia. Afr J Mar Sci 35(1):133–139
Mackintosh NA (1942) The southern stocks of whalebone whales. Discovery Rep 22:197–300
Mackintosh NA (1965) The stocks of whales. Fishing News (Books) Ltd, London
Macleod CD, Perrin WF, Pitman RL et al (2006) Known and inferred distributions of beaked whale
species (Ziphiidae: Cetacea). J Cetacean Res Manag 7(3):271–286
Michalsen K, Alvheim OB, Zaera-Perez D et al. (2015) Surveys of the fish resources of Angola.
Cruise Report N0. 8/2015. Surveys of the pelagic fish resources of Angola, 15 August–13
September 2015. Cruise reports Dr. Fridtjof Nansen. EAF—N2015/8. Available at: http://hdl.
handle.net/11250/2374573
Mikhalev JA, Savusin VP, Kishiyan NA et al (1981a) To the problem of the feeding of sperm
whales from the Southern Hemisphere. Rep Int Whaling Commission 31:737–745
Mikhalev YA, Ivashin MV, Savusin VP et al (1981b) The distribution and biology of killer whales
in the Southern Hemisphere. Rep Int Whaling Commission 31:551–565
Mörzer Bruyns WFJ (1968) Sight records of cetacea belonging to the genus Mesoplodon Gervais,
1850. Zeitschrift für Säugetierkunde 33:106–107
Mörzer Bruyns WFJ (1971) Field guide of whales and dolphins. CA Mees, Amsterdam
Olsen Ø (1913) On the external characters and biology of Bryde’s whale (Balaenoptera brydei), a
new rorqual from the coast of South Africa. Proc Zool Soc Lond 1913:1073–1090
Richards R (2009) Past and present distributions of southern right whales (Eubalaena australis).
N Z J Zool 36(4):447–459
Rosenbaum HC, Pomilla C, Mendez M et al (2009) Population structure of humpback whales from
their breeding grounds in the South Atlantic and Indian oceans. PLoS One 4(10):e7318
Roux J-P, Dundee BL, da Silva J (2007) Seabirds and marine mammals distributions and patterns
of abundance. Chapter 41 in Kirkman, S.P. (ed.) Final report of the BCLME (Benguela Current
Large Marine Ecosystem) Project on Top Predators as Biological Indicators of Ecosystem in
the BCLME. BCLME Project LMR/EAF/03/02. 381 pp. Available at: http://www.adu.uct.
ac.za/adu/projects/sea-shore-birds/communication/report-bclmetpp
Ruud JT (1952) Catches of Bryde-whale off French Equatorial Africa. Norsk Hvalfangst-Tidende
12:662–663
Sherman K (2014) Toward ecosystem-based management (EBM) of the world’s large marine eco-
systems during climate change. Environ Dev 11:43–66
Simmons DC (1968) Purse seining off Africa’s west coast. Commer Fish Rev 30:21–22
Smith TD, Reeves RR, Josephson EA et al (2012) Spatial and seasonal distribution of American
whaling and whales in the age of sail. PLoS One 7(4):e34905
SMM (2018) Committee on Taxonomy. List of marine mammal species and subspecies, updated
2017. Society for Marine Mammalogy: www.marinemammalscience.org, consulted on 20
January 2018
Stewart BS, Leatherwood S (1985) Minke whale Balaenoptera acutorostrata Lacépède, 1804. In:
Ridgway SH, Harrison R (eds) Handbook of marine mammals, volume 3, the sirenians and
baleen whales. Academic, San Diego, pp 91–136
Tønnessen JN, Johnsen AO (1982) The history of modern whaling. University of California Press,
Berkeley/Los Angeles
Tormosov DD, Budylenko GA, Sazhinov EG (1980) Biocenoological aspects in the investigations
of sea mammals. Paper SC/32/02 presented to the IWC Scientific Committee, July 1980, 9 pp
Townsend CH (1935) The distribution of certain whales as shown by logbook records of American
whaleships. Zoologica 19:3–50
Van Waerebeek K (1997) Long-beaked and short-beaked common dolphins sympatric off central-
West Africa. Paper SC/49/SM46 presented to the IWC Scientific Committee, October 1997,
4 pp
Van Waerebeek K, Barnett L, Camara A et al (2004) Distribution, status, and biology of the Atlantic
humpback dolphin, Sousa teuszii (Kükenthal, 1892). Aquat Mamm 30:56–83
Weir CR (2006a) Sightings of beaked whales (Cetacea: Ziphiidae) including first confirmed
Cuvier’s beaked whales Ziphius cavirostris from Angola. Afr J Mar Sci 28:173–175
Weir CR (2006b) Sightings of rough-toothed dolphins (Steno bredanensis) off Angola and Gabon,
South-east Atlantic Ocean. Abstracts of the 20th Annual Conference of the European Cetacean
Society, Gdynia, Poland, 2–7 April 2006
Weir CR (2006c) First confirmed records of Clymene dolphin Stenella clymene (Gray, 1850) from
Angola and Congo, South-East Atlantic Ocean. Afr Zool 41:297–300
Weir CR (2007) Occurrence and distribution of cetaceans off northern Angola, 2004/05. J Cetacean
Res Manag 9:225–239
Weir CR (2008) Overt responses of humpback whales (Megaptera novaeangliae), sperm whales
(Physeter macrocephalus), and Atlantic spotted dolphins (Stenella frontalis) to seismic explo-
ration off Angola. Aquat Mamm 34:71–83
Weir CR (2009) Distribution, behaviour and photo-identification of Atlantic humpback dolphins
(Sousa teuszii) off flamingos, Angola. Afr J Mar Sci 31:319–331
Weir CR (2010a) A review of cetacean occurrence in West African waters from the Gulf of Guinea
to Angola. Mamm Rev 40:2–39
Weir CR (2010b) First description of Atlantic humpback dolphin (Sousa teuszii) whistles, recorded
off Angola. Bioacoustics 19:211–224
Weir CR (2010c) Cetaceans observed in the coastal waters of Namibe Province, Angola, during
summer and winter 2008. Mar Biodivers Rec 3:e27
Weir CR (2011a) Ecology and conservation of Cetaceans in the waters between Angola and the
Gulf of Guinea, with focus on the Atlantic Humpback Dolphin (Sousa teuszii). PhD thesis,
University of Aberdeen, Aberdeen
Weir CR (2011b) Distribution and seasonality of cetaceans in tropical waters between Angola and
the Gulf of Guinea. Afr J Mar Sci 33:1–15
Weir CR, Coles P (2007) Morphology of common dolphins (Delphinus spp.) photographed off
Angola. Abstracts of the 17th Biennial Conference of the Society for Marine Mammalogy,
Cape Town, South Africa, 29 November–3 December 2007. Society for Marine Mammalogy,
San Diego
Weir CR, Collins T (2015) A review of the geographical distribution and habitat of the Atlantic
humpback dolphin (Sousa teuszii). Adv Mar Biol 72:79–117
470 C. R. Weir
Weir CR, Nicolson I (2014) Depredation of a sport fishing tournament by rough-toothed dolphins
(Steno bredanensis) off Angola. Aquat Mamm 40(3):297–304
Weir CR, Pierce GJ (2013) A review of the human activities impacting cetaceans in the eastern
tropical Atlantic. Mammal Rev 43(4):258–274
Weir CR, Debrah J, Ofori-Danson PK et al (2008) Records of Fraser’s dolphin Lagenodelphis
hosei Fraser, 1956 from the Gulf of Guinea and Angola. Afr J Mar Sci 30:241–246
Weir CR, Collins T, Carvalho I et al (2010) Killer whales (Orcinus orca) in Angolan and Gulf of
Guinea waters, tropical West Africa. J Mar Biol Assoc U K 90:1601–1611
Weir CR, Van Waerebeek K, Jefferson TA et al (2011) West Africa’s Atlantic humpback dolphin
(Sousa teuszii): endemic, enigmatic and soon endangered? Afr Zool 46:1–17
Weir CR, MacLeod CD, Pierce GJ (2012) Habitat preferences and evidence for niche partitioning
amongst cetaceans in the waters between Gabon and Angola, eastern tropical Atlantic. J Mar
Biol Assoc U K 92:1735–1749
Weir CR, Coles P, Ferguson A et al (2014) Clymene dolphins (Stenella clymene) in the eastern trop-
ical Atlantic: distribution, group size, and pigmentation pattern. J Mammal 95(6):1289–1298
Wilson CE, Perrin WF, Gilpatrick JW et al (1987) Summary of worldwide locality records of the
striped dolphin, ‘Stenella coeruleoalba’. NOAA-TM-NMFS-SWFC-90. 72 pp
Chapter 17
The Giant Sable Antelope: Angola’s
National Icon
Pedro Vaz Pinto
Abstract The giant sable antelope Hippotragus niger variani is the most widely
recognised representative of Angolan biodiversity, owing to its endemic status, rar-
ity and physical attributes. One of the last large mammals to be described in Africa,
it is confined to the upper Cuanza basin, in central Angola. Studies on the biology
of giant sable were mostly conducted in the 1970s, but ongoing efforts using mod-
ern tools such as DNA analyses, GPS tracking, camera trapping and satellite imag-
ery are improving our knowledge. Past explanations for the extent of the isolation
and relationships with other sable populations have been controversial. Molecular
studies have only recently made significant contributions to interpret the evolution-
ary history of giant sable. Although much pursued by hunters during the first half of
the twentieth century, the conservation needs of giant sable were recognised early
on, with the proclamation of two protected areas and the setting in place of strict
regulations. Park management and efficient protection was enforced in the 1960s,
but these protected areas were abandoned soon after the country’s independence,
leading to population crashes and interspecific hybridization, which left the subspe-
cies on the verge of extinction. The giant sable is currently the main focus of a
conservation programme supervised by the Angolan Government that is success-
fully promoting its recovery.
Keywords Cangandala · Conservation · Cuanza · Evolutionary history ·

Extinction · Hybridisation · Luando · Population collapse · Trap cameras
P. Vaz Pinto (*)
do Porto, Campus de Vairão, Vairão, Portugal

https://doi.org/10.1007/978-3-030-03083-4_17
472 P. Vaz Pinto
Introduction and Background
Having captured the imagination of naturalists and the general public for over one
hundred years, the giant sable antelope Hippotragus niger variani is the undisputed
icon of Angola’s natural heritage (Fig. 17.1). Its cultural significance extends from
local totemic status among resident communities (where it is known as ‘kolo’ or
‘sumbakaloko’) to global recognition as an antelope symbol and flagship for con-
servation. Soon after its discovery, the giant sable was elevated to a high pedestal
among the hunting community as one of the most sought after trophy prizes, and
fuelled the lust of big game hunters from all over the world. In Angola, the giant
sable was the first animal to receive full legal protection and was soon embraced as
an icon during colonial rule. Since Angola’s independence in 1975, its status has
been reinforced. The importance of the current unanimous recognition of the giant
sable as a national symbol should not be underestimated, constituting a key factor
uniting Angola’s people regardless of their different ethnic groups, religious beliefs
or political ideologies, and thus contributing to social cohesion and national pride.
Fig. 17.1 A splendid

Giant Sable Antelope bull.
(Photo: P. Vaz Pinto)
17 The Giant Sable Antelope: Angola’s National Icon 473
Observations on the biology of giant sable were reported soon after its scientific
description by Thomas (1916), resulting from the many expeditions undertaken to
collect material for museums in Europe and the United States of America. Of par-
ticular importance were very detailed morphological observations, with taxonomic
implications and accurate ecological descriptions, made by Gilbert Blaine (1922).
In addition, behavioural observations were made by trophy hunters (e.g. Statham
1922; Gray 1930, 1933; Powell-Cotton 1932; Curtis 1933), while concerns
expressed by the subspecies’ discoverer himself led to timely conservation interven-
tions (Varian 1953). For a long time these reports would remain as the most reliable
sources of knowledge on the taxonomy and biology of giant sable, although several
other publications addressed its conservation status (Harper 1945; Heim 1954;
Newton da Silva 1958). In the late 1950s and during the following decade, the first
systematic efforts focusing on giant sable addressed its biology and related conser-
vation issues, and were undertaken by Portuguese researchers working for the
Centro de Zoologia da Junta de Investigação do Ultramar or Instituto de Investigação
Científica de Angola (e.g. Frade 1958, 1967; Frade and Sieiro 1960; Sieiro 1962;
Crawford-Cabral 1965, 1966, 1969, 1970). Later, Huntley (1972) made important
contributions on conservation planning for giant sable while adding ecological
insights. Published in 1972, the book A Palanca Real (Silva 1972) provided a com-
prehensive and appealing photographic compilation, including a few ecological and
behavioural observations. The researcher who became inextricably linked with the
giant sable, the famous biologist Richard Estes, spent one full year studying the
giant sable in Luando Nature Strict Reserve between 1969 and 1970. Estes is still
the most relevant contributor to the current knowledge on the biology of the species
(e.g. Estes and Estes 1970, 1972, 1974). Another invaluable source is the book A
Certain Curve of Horn (Walker 2004), in which the author gives a detailed and vivid
account of the various explorations, studies and conservation initiatives around the
giant sable, a history spanning over 100 years. In recent years, molecular studies
have addressed giant sable looking into phylogenetic relationships, genetic diversity
or hybridisation (e.g. Pitra et al. 2006; Jansen van Vuuren et al. 2010; Vaz Pinto
et al. 2015, 2016). Currently, comprehensive ongoing research is being developed
on the giant sable and addressing a wide range of topics, including evolutionary his-
tory, biology and conservation (Vaz Pinto 2018).
Scientific Discovery
The giant sable was not discovered and described until the twentieth century, but
two intriguing and obscure previous records are worth mentioning. The oldest
proven giant sable material is a single horn, named ‘The Florence Horn’ after the
museum in Italy where it was deposited in 1873 (Walker 2004; Vaz Pinto 2018).
Nothing is known about the provenance details of the Florence Horn. Although
recognised early on as extraordinary, and suspected later to have been obtained from
the Angolan relic population (Thomas 1916; Walker 2004), only recently has a
474 P. Vaz Pinto
molecular study provided convincing evidence of it being a giant sable horn (Vaz
Pinto 2018). The Florence Horn may, however, have been preceded by a skull col-
lected by the famous Austrian botanist Friedrich Welwitsch in Angola between 1853
and 1861, and classified by Bocage as Hippotragus niger (Bocage 1878, 1890;
Thomas 1916). As with so many other priceless biological material from Angola,
this specimen was tragically lost in the fire that destroyed the Museu Bocage in
1978, and it was never possible to attribute it to subspecies, even though the reported
length of the horns suggest that it was a giant sable. The specimen was stated as
having been collected by Welwitsch inland from Moçâmedes (Bocage 1878, 1890;
Thomas 1916; Hill and Carter 1941). The exact collecting locality remains in doubt,
but it is known that Welwitsch collected in the Malanje region (Crawford-Cabral
and Mesquitela 1989), which could have given him access to giant sable.
The discovery and scientific description of this taxon had to wait another half-
century, and followed the efforts of the chief engineer overseeing the construction
of the Benguela Railway, the British citizen Frank Varian (Varian 1953; Walker
2004). The first mention of a sable with extraordinary horns from the Cuanza dis-
trict was made by Varian in 1909 and simply based on one photograph and witness
accounts, but it was met with disbelief in Europe (Varian 1953; Walker 2004). The
first material was secured in 1911, but it was only in 1916 that skulls and skins were
shipped by Varian to the British Museum and led to the formal description of
Hippotragus niger variani (Thomas 1916), thus honouring the discoverer.
Description
The giant sable is a large, compact and muscular antelope carrying massive scimitar-
shaped horns. The original description was based on a giant sable skull with horns
that measured 57 inches in length along the curvature and 11 inches in circumfer-
ence at the base (Thomas 1916). Horn length is a distinctive trademark for giant
sable as they are usually above 50 and often above 60 inches, while bulls from all
other sable antelope populations only very rarely surpass 50 inches (Halse 1998;
Vaz Pinto 2018).
The size of horns alone made the type specimen stand out, but equally striking
was the darker face, in which unlike other sable, the ante-orbital white spots are not
connected by a white streak to the sides of the muzzle (Thomas 1916). These fea-
tures proved consistent as more specimens were analysed, but other peculiarities
were revealed to be distinctive, as described in detail by Gilbert Blaine (1922). Very
clear differences have been found in the skulls, both structurally and from measure-
ments, and these helped to sustain a claim for elevating giant sable to specific status
(Blaine 1922). Giant sable have longer and narrower forefaces and less prominent
forehead compared to typical sables, and relatively small ears in adult males, while
the neck in mature bulls is short, massive, oval in section and wedge shaped (Blaine
1922). Other published measurements on both skulls (Groves and Grubb 2011) and
teeth (Klein 1974) have relied on very small sample sizes, and therefore added little
to Blaine’s observations.
The combination of skull and body build, crowned by the massive arched horns,
must have greatly contributed to the ‘giant’ epithet attributed to this taxon, but as no
specimen was ever weighted and few were measured (Blaine 1922; Harper 1945;
Estes 1982, 2013), doubts remain regarding the size of giant sable in comparison
with other sable antelope populations. Giant sable, particularly bulls, have often been
considered as much larger in size and weight than other sable (Blaine 1922; Statham
1922; Harper 1945), but this view was not shared by Estes (1982, 2013). Nevertheless,
recent observations made on immobilised animals suggest that mature bulls may be
at least heavier than other sable subspecies (Vaz Pinto Unpublished Data).
Sexual dimorphism is pronounced in sable, but appears to be more extreme in
giant sable (Fig. 17.2), possibly as a by-product of their sedentary nature and the
mesic environmental in which they occur (Estes 2013; Vaz Pinto 2018). The distinct
characteristics observed in males and female giant sable is obvious in body size,
horn length, and pelage colouration. Various physical and colouration features were
described in great detail by Blaine (1922). In short, mature giant sable bulls are
glossy black with white bellies, white facial markings, reddish hocks and yellowish
back to the ears (Blaine 1922; Frade and Sieiro 1960; Estes 2013). In females, the
black colour of bulls is replaced by brown tones which have been described as dif-
ferently as bright golden chestnut (Blaine 1922), light to dark brown-chocolate
(Silva 1972) or deep chestnut to blackish brown (Groves and Grubb 2011). Female
colouration has been used to assist differentiation among sable subspecies (Groves
and Grubb 2011), but pelage colouration of giant sable females is variable irrespec-
tive of age (Vaz Pinto Unpublished Data).
Taxonomy
While examining the Angolan material, Oldfield Thomas, the mammal curator of
the British Museum, was so impressed by some peculiarities and obvious differ-
ences when comparing Varian’s material with ordinary sable that he considered the
possibility of awarding specific status to the giant sable (Thomas 1916; Blaine
1922; Walker 2004). Subsequently, and having assembled additional material,
Gilbert Blaine (1922) found it justifiable to recognise giant sable as a full species on
its own Hippotragus variani, a view that was followed by some authors (e.g. Harper
1945). Nevertheless, Blaine’s claim eventually fell into disuse and the giant sable
became widely accepted as one of four to five subspecies of sable antelope (e.g. Hill
and Carter 1941; Ansell 1972; Groves and Grubb 2011; Estes 2013).
With the advent of molecular tools, the taxonomic placement of giant sable could
be revisited. The first molecular studies to include giant sable samples were based
on small mitochondrial fragments and reported paraphyly for giant sable with
respect to sable antelope across the southern parts of their range. These findings cast
doubt on the recognition of H. n. variani as a valid subspecies (Mathee and Robinson
476 P. Vaz Pinto
Fig. 17.2 Sexual dimorphism in antelopes is unique to the Sable Antelope but particularly pro-
nounced in the more sedentary Giant Sable of Angola, where mature bulls are jet black (bottom)
and cows have a rich cinnamon pelage. (Photos: P Vaz Pinto)
1999; Pitra et al. 2006). The results, in combination with the fact that sable collected
in western Zambia often present similar facial markings, were further interpreted as
blurring the differences between giant and western Zambian sable antelope (Wessels
2007). However, clear genetic differences exist between these western Zambian
sable and giant sable, confirming that the giant sable represents a distinct mitochon-
drial evolutionary lineage (Jansen van Vuuren et al. 2010).
Much progress has recently been achieved allowing the clarification of the taxo-
nomic status of giant sable, following more ambitious molecular efforts that
included the use of mitogenomics and nuclear markers (Vaz Pinto 2018). The full
sequencing of mitochondria on a large dataset pooled from across the full range of
sable distribution, concluded that giant sable constitutes a reciprocally monophy-
letic group, one of six geographically discrete sable clusters (Vaz Pinto 2018).
Confirming previous suggestions (Pitra et al. 2006; Jansen van Vuuren et al. 2010),
the giant sable maternal lineage proved to be more closely related to some west
Tanzanian lineages, than to those found elsewhere in southern Africa (Vaz Pinto
2018). A population approach resorting to 57 autosomal microsatellites applied to
an even more comprehensive dataset showed giant sable as being the most divergent
of five clearly identified and geographically coherent populations (Fig. 17.3, Vaz
Pinto 2018). The molecular results combining mitogenomics and nuclear markers
Fig. 17.3 Sable Antelope populations (and sub-species) as determined through nuclear DNA
analysis (Vaz Pinto 2018). Eastern Sable Hippotragus niger roosevelti; Tanzanian Sable (possibly
H. n. subsp. nov); Zambian Sable H. n. kirki; Angolan Sable H. n. variani; Southern Sable H. n.
niger
478 P. Vaz Pinto
provide compelling evidence to support the uniqueness of giant sable, and are there-
fore concordant with its unequivocal recognition as a separate endemic taxon (Vaz
Pinto 2018).
Evolutionary History
The origin of giant sable has remained intriguing and subject to different interpreta-
tions, and only recently by resorting to modern molecular tools, a more robustly
sustained and coherent picture is emerging. Attempting to explain the rarity of giant
sable and its oddly restricted distribution, Huntley (1972) argued it not to be a recent
artefact driven by anthropogenic causes, but rather a result of very specific habitat
requirements that forced this population to a long-standing confinement in central
Angola. Other authors have suggested alternative views such as the possibility of H.
n. kirkii from western Zambian having had at some point a continuous distribution
into the Angolan plateau before receding and a subpopulation becoming isolated in
the Cuanza Basin (Crawford-Cabral and Veríssimo 2005; Wessels 2007).
The first molecular studies to include giant sable, based on small mitochondrial
fragments and limited sampling (Mathee and Robinson 1999; Pitra et al. 2006)
found H. n. variani samples to cluster within other sables, and results suggested
recent connectivity with other southern African populations. The suggestion that
giant sable may share a closer ancestry with some (but not all of) west Tanzanian
lineages rather than with the geographically closer west Zambian populations (Pitra
et al. 2006; Jansen van Vuuren et al. 2010) was difficult to interpret and explain. It
was speculated that central Angola and southern Tanzania could have been colo-
nised from a common source population (Pitra et al. 2006). Furthermore, a tentative
date for this split was set at approximately 200,000 years ago (Pitra et al. 2006). In
an effort to disentangle this phylogenetic riddle, Groves and Grubb (2011) sug-
gested that H. n. variani could have migrated from Angola to southern Tanzania and
hybridised with H. n. roosevelti, but this scenario is difficult to justify and, if any-
thing, would complicate things even further.
A more recent molecular study based on mitogenomics revealed the phylogeo-
graphic patterns that may have shaped the evolutionary history of sable antelope,
influenced by a complex interplay between Pleistocene climatic oscillations and
geomorphological features (Vaz Pinto 2018). The study also provided credible time
estimations for time splitting events. It was hypothesised that during the dry climatic
period of the Elster Glaciation, corresponding to the Marine Isotopic Stage (MIS) 6
and estimated at approximately between 130,000 and 186,000 years ago, an ances-
tral sable lineage evolved in Central Congo and separated from two other lineages in
eastern and southern Africa (Vaz Pinto 2018). When the climate became warmer and
receding savannas were replaced by forests, at around 120,000 years ago, the Central
Congo lineage would have split into an Angolan lineage and another that ended up
in the rift region (Vaz Pinto 2018). Subsequently, the Angolan lineage may have
since evolved confined to the Cuanza river basin and adapted to particular habitat
conditions (Vaz Pinto 2018). The population approach using autosomal nuclear
markers confirmed the uniqueness of the giant sable population, with low levels of
gene flow for a long time, and as a result being genetically very distinctive (Vaz
Pinto 2018). In addition it was possible to genetically differentiate the populations
from Luando and Cangandala, even though the genetic signal may have been
enhanced by an extreme recent bottleneck affecting the latter (Vaz Pinto 2018). It
seems possible that the giant sable evolved in isolation in central Angola since the
beginning of the late Pleistocene until the present day, and exposed to relatively rare
male-mediated gene flow events with neighbouring sable populations (Vaz Pinto
2018).
Habitat and Ecology
As with other sable races, the giant sable is a specialist of miombo, a type of wood-
land and mesic savanna that occurs on poor dystrophic soils dominated by trees
from the genus Brachystegia, Julbernardia and Isoberlinia (Estes 2013, Fig. 17.4).
Giant sable are ecotone species, showing a preferential use of the edge between
woodland and grassland (Estes and Estes 1974; Estes 2013). A peculiar feature in
the gently undulating giant sable region are vast termite-infested open savannas
covered by fire-prone geoxyle vegetation, and known by the local name of anharas
(Barbosa 1970; Estes and Estes 1974; Zigelski et al. 2019). A mosaic of woodland
and anharas seems to constitute the prime habitat for giant sable herds both in
Luando Strict Nature Reserve (LSNR) and Cangandala National Park (CNP) (Estes
and Estes 1974). It has also been suggested that the presence of these anharas, char-
acterised by unique vegetation types and influenced by poor soils and unique local
climatic conditions, may explain the current pattern of giant sable distribution (Vaz
Pinto 2018). Giant sable are also water-dependant, and the availability of water, in
streams or water holes, during the dry season, is a key component determining habi-
tat value and can become a limiting factor affecting their distribution patterns (Estes
and Estes 1974).
Giant sable are herbivores and mostly selective grazers, with preference for
perennial grasses such as Brachiaria, Digitaria, Panicum or Setaria spp., and typi-
cally biting off only the tender outer portion of the plants (Estes and Estes 1974).
Although predominantly grazers, they also browse frequently, in particular the
locally abundant shrub species Diplorhynchus condylocarpon which seems to be
favoured by giant sable all year round (Estes and Estes 1974; Vaz Pinto Unpublished
Data), to the point of being referred by locals as ‘palanca bush’ (Statham 1922).
Also browsed often are the tree Julbernardia paniculata, and the dwarf shrubs
Mucana stans, Cryptosepalum maraviense and Dolichus sp (Statham 1922;
Crawford-Cabral 1970; Silva 1972; Estes and Estes 1974; Vaz Pinto 2018). In addi-
tion it has been found that giant sable resort often to geophagy, eating soil excavated
on some ancient termite mounds, a habit likely evolved as a consequence of very
low nutrient soils (Estes and Estes 1974; Baptista et al. 2013).
480 P. Vaz Pinto
Fig. 17.4 Aerial view of the preferred habitat of the Giant Sable Antelope: a mosaic of grassland
and miombo woodland – the latter pictured in the bottom photograph. (Photos: P Vaz Pinto)
In common with most other social antelopes, giant sable are gregarious and are
structured according to three social classes: the breeding or nursery herds, bachelor
groups, and territorial bulls (Estes and Estes 1974; Estes 2013). The matriarchal
herd, composed of breeding cows, calves and young, constitute the main unit (Estes
and Estes 1974; Estes 2013). Numbers and composition of giant sable breeding
herds change seasonally and sometimes even daily, and different average figures
have been obtained by various authors, usually ranging from eight to 24 animals
(Blaine 1922; Crawford-Cabral 1966, 1970; Estes and Estes 1974; Vaz Pinto
Unpublished Data). The nursery herds are sedentary and tend to perpetuate their
home ranges across generations (Estes and Estes 1974; Estes 2013). Young males
will be tolerated within the herd until around 3 years of age, before dispersing where
after they either initiate solitary life or temporarily join other males to form bachelor
groups (Estes and Estes 1974). Around their sixth year of age, bulls become territo-
rial, and will demarcate their own territory by scrapping, defecating and bush-
thrashing with their horns (Estes and Estes 1974). Dominant bulls typically display
aggressive behaviour towards intruders, exerting domination by physical intimida-
tion and chasing, and only exceptionally will the confrontation lead to a serious
fight (Estes and Estes 1974).
Giant sable are seasonal breeders. The rutting coincides with miombo springtime
and has been stated to start around late August (Estes and Estes 1974), although
recent observations suggest the mating season to peak in September and October
(Vaz Pinto 2018). The gestation likely follows what has been found for other sable
populations, estimated at 8.5–9 months (Wilson and Hirst 1977). The calving sea-
son for giant sable matches the onset of the dry season, peaking during a 2-month
period between May and July (Estes and Estes 1972), but off-season calving is not
unusual (Estes and Estes 1974; Vaz Pinto Unpublished Data). As the calving season
approaches, the breeding herds tend to break and the most heavily pregnant cows
become isolated (Estes and Estes 1974). Giant sable are ‘hiders’, meaning that
females will calve alone and hide their calves, attending them at irregular intervals
for several days or weeks, before re-joining the herd with their offspring, and form-
ing crèches with other calves of similar age (Estes and Estes 1974; Estes 2013).
Giant sable herds have been found to make use of home ranges of varying size,
with one of two giant sable herds monitored for 1 year in Luando Strict Nature
Reserve staying within 12 km2 while the other moved a distance of 15 km to use
different areas in the dry and rainy seasons, raising the size of the annual home
range of one of those groups to 40 km2 (Pedrosa 1971; Estes and Estes 1974). The
same authors found home range size to be affected by food availability and season-
ality (Estes and Estes 1974). Herds will tend to break up after the onset of rains, and
move from the anharas into the woodlands (Estes and Estes 1974). During the wet-
test periods, sables will avoid waterlogged areas such as floodplains, and spend
most of the time on high ground within the woodland (Crawford-Cabral 1970; Estes
and Estes 1974). Nevertheless, recent data obtained with GPS transmitters, have
found relatively little seasonal variation, and yet a huge contrast in home range size,
ranging from 14 to 110 km2 measured by Minimum Convex Polygon (MCP) (Vaz
Pinto 2018). The daily movements of herds is reported to be modest, typically vary-
ing from one to two km (Estes and Estes 1974), and this finding is consistent with
GPS data (Vaz Pinto Unpublished Data). In general, herd movement patterns can be
summarised as concentrations in open areas during the dry season, followed by
group partition as the rain starts and confinement of smaller stable groups in wooded
482 P. Vaz Pinto
parts of their range, and then increased movements towards the end of the rains and
further group fragmentation prior to the calving season (Estes and Estes 1974).
Different herds will not overlap in home ranges and are frequently separated by
several km of seemingly suitable habitat (Estes and Estes 1974). Sable bulls have
been reported to hold relatively small territories separated from neighbours by 1 to
3 km apart, while spending most of their time within 3–4 km2, and are able to
expand their area to at least 10–12 km2 when accompanying breeding herds (Estes
and Estes 1974). However, preliminary data obtained from GPS monitoring of sev-
eral giant sable bulls over a period of 5 years suggest a very different spatial use, as
bulls tend to move a lot more than previously thought, and with overlapping territo-
ries ranging above 200 km2 measured as MCP’s (Vaz Pinto 2018).
Historical Distribution and Abundance
Soon after the description of H. n. variani, it was assumed that they only occurred
within the Cuanza watershed, and especially confined to the lowlands between the
upper Cuanza and its tributary the Luando (Blaine 1922; Statham 1922; Hill and
Carter 1941; Varian 1953). This region, also known as the ‘land between two rivers’
(Walker 2004), is a narrow strip of land stretching across 200 km oriented NW – SE,
and spanning up to 60 km at its widest. Most of the hunters and naturalists exploring
the region in search of giant sable entered the sable lands from the Benguela
Railroad, heading northwards and thus focusing on the southern region within the
species' distribution (Walker 2004). Nevertheless, one of the earliest giant sable
explorations, by Col. Statham, entered from the north, and traversed across the land
between the two rivers on foot, and was able to confirm the presence of giant sable
up to the confluence of the Luando and Cuanza rivers (Statham 1922). Effectively
limited by the two massive rivers to the north, to the south giant sable seemed to be
contained by three features: swamps, an inland escarpment, and a desolate territory
to the east crossed by the railway and known as the ‘Hungry Country’ (Varian 1953;
Walker 2004). Within the ‘land between two rivers’, sable appear to prefer the
Luando and Lingoio sub-basins while avoiding the remaining Cuanza drainage
(Blaine 1922; Statham 1922; Vaz Pinto Unpublished Data). The land between two
rivers was set aside as a conservation area in 1938, first as the Giant Sable Reserve,
and since 1955 as Luando Strict Nature Reserve (Huntley 1971), and covering
approximately 828,000 hectares.
The possible existence of a giant sable population north of the Luando River,
near the village of Cangandala, was first suggested by Statham (1922). This was
based on trophies acquired by Statham from a Portuguese settler. However, Statham
seems to have been deliberately misled by local tribal chiefs who denied that giant
sable occurred elsewhere than to the south of the Luando River (Statham 1922;
Walker 2004). More than 30 years would pass before the existence of a second,
albeit much smaller, sable population north of the Luando was eventually confirmed
near Cangandala between the Cuque and Cuije rivers (Frade 1958), and a mere
50 km south of Malanje town. As result of this find, Cangandala Strict Nature
Reserve was proclaimed in 1963, and elevated to a National Park status in 1970. The
occurrence of giant sable outside the boundaries of Luando Reserve and Cangandala
National Park was often claimed but never proven conclusively. Witness accounts
mentioned sable to be present between the Cuanza River and its western tributary
the Cutato (Blaine 1922), but by the 1970’s their persistence in the region was at
best doubtful (Estes and Estes 1974; Huntley Unpublished Data). Other uncon-
firmed records were reported from the ‘Hungry Country’ and from the areas that lie
in between Cangandala National Park and Luando Strict Nature Reserve (Estes and
Estes 1974; Crawford-Cabral and Veríssimo 2005; Huntley Unpublished Data).
Two of the most geographically extreme giant sable records, falling outside the
Cuanza basin, were from Quibala in Cuanza-Sul, and from Baixa de Cassanje in
Lundas, but only males were reported (Estes and Estes 1974; Huntley Unpublished
Data). A third case was a trophy obtained near Lupire in Cuando Cubango in 1966
and described by the collector as a very old lonely bull in poor condition (Francisco
Sousa Machado pers. comm.). The trophy would have become the world record if
accepted as a typical sable (Halse 1998; Wessels 2007), but a recent molecular study
has established it as giant sable (Vaz Pinto 2018). Other reports of giant sables from
as far from Luando as Katanga in the Democratic Republic of Congo or western
Zambia (Schouteden 1947; Wessels 2007) are likely erroneous and lack molecular
support (Ansell 1972; Jansen van Vuuren et al. 2010; Vaz Pinto 2018).
As bulls may disperse far and unpredictably, it is fair to conclude that the histori-
cal distribution of giant sable has remained well contained within the Cuanza Basin,
and likely centred in Luando Reserve (Estes and Estes 1974), with relatively small
population pockets reaching Cangandala National Park and surrounding areas.
The first estimations of giant sable populations were based on limited knowledge
of the species distribution and lacked reliable quantifiable data, suggesting the total
numbers to reach a few hundred individuals. Based on guestimates provided by
Portuguese naturalists, numbers were set at less than 750–800 (Harper 1945) or at
around 700 (Heim 1954). By partially surveying approximately one third of Luando
Reserve, 159 giant sable were counted (Frade and Sieiro 1960; Newton da Silva
1970), leading Fernando Frade to suggest that the total number could be even less
than 500 (Frade 1958, 1967). As conservation measures increased and biologists
renewed their interest in the giant sable, subsequent efforts raised the estimations to
range between 1500 and 2500 animals (Crawford-Cabral 1970; Huntley 1972,
1973). Nevertheless, and as pointed out by Richard Estes, these estimations should
be taken with care as they were not based on actual counts (Estes and Estes 1972,
1974). In the early 1970s a few aerial counts were attempted by plane and helicop-
ter, but the results did not contribute much for determining the population size
(Estes and Estes 1970, 1974, Pedrosa 1971). With the bulk of giant sable present in
Luando Reserve, the population in Cangandala NP was estimated to range between
100 and 150 animals (Crawford-Cabral 1970; Huntley 1973; João Serôdio
484 P. Vaz Pinto
Unpublished Data). The existing estimations suggest therefore an average giant

sable density of 0.0025 sable/ha.
Collapse, Rediscovery, Hybridisation
The civil war that ravaged Angola following the country’s independence in 1975
had a dramatic impact on the populations of giant sable. As result of the war, man-
agement was abandoned in protected areas and infrastructure destroyed, while
accounts reported on the widespread killing of giant sable (Walker 2004). A brief
visit made in 1982 by Richard Estes, then chairman of the IUCN Antelope Specialist
Group, was able to confirm various herds of giant sable that were still being moni-
tored by a park warden in Cangandala (Estes 1982). But by then the Luando Reserve
had been taken over by UNITA (União Nacional para a Independência Total de
Angola) military and could not be surveyed by Estes, and the reports were worri-
some (Walker 2004). Soon after Estes’ visit the warden was forced to flee Cangandala
as UNITA took control of the park and in Luando the rangers that stayed behind
were killed. When a short peace allowed a national biodiversity assessment to be
carried out in 1992, the situation was stated as mostly unknown in spite of recent
sightings (Huntley and Matos 1992), and as the armed conflict resumed and grew in
intensity many questioned if the survival of giant sable was even a realistic possibil-
ity (Walker 2004). Efforts to find the species were put on hold, and could only be
implemented when peace and security were restored in the bush (Walker 2004).
In November 2001, a few months before the end of the war, the Kissama
Foundation, a local biodiversity Non-Governmental Organisation, made a flight over
Cangandala National Park on a large military helicopter without results (Walker
2004). A subsequent and more ambitious effort was carried out in August 2002, soon
after the war ended, by the Kissama Foundation and again with strong support from
the Angolan military. On this occasion, Cangandala was subject to a one-day survey
by a large party on foot, and a couple of flights on a military helicopter were
done over Luando Reserve (Walker 2004). No animals were seen from the air, but
brief observations were reported in Cangandala of what could have been giant sable,
although they could not be substantiated (Walker 2004). Ground surveys were then
implemented in Cangandala by the Centro de Estudos e Investigação Científica
(CEIC) of the Catholic University of Angola starting in 2003, and in the following
year surveys extended into Luando Reserve where an aerial survey was also per-
formed. No giant sable were seen, but Hippotragus dung was collected in both pro-
tected areas. In 2004, a monitoring programme with trap cameras was also initiated.
Finally, in early 2005, it was possible to prove the survival of giant sable when DNA
extracted from dung samples revealed that the mitochondria was typical of H. n.
variani, and trap camera photographs showed sable herds in Cangandala (Pitra et al.
2006).
Although giant sable had proved resilient enough to persist in Cangandala National
Park, the situation proved to be much worse than anticipated. As the trap camera
records increased and additional ground observations were made in Cangandala, an
alarming scenario emerged: only one herd of giant sable was present, no sable bulls
could be found, a roan bull was seen escorting the sable herd, and based on morpho-
logical features alone, calves and young animals appeared to be hybrids (Vaz Pinto
2007). The possibility of interspecific hybridisation was subsequently confirmed
with modern molecular tools (Vaz Pinto et al. 2016). The occurrence and extent of
this phenomenon was unexpected, and allowed for a quite unprecedented study
among mammals, as it was possible to document in detail the underlying mecha-
nisms that led to such an extreme outcome (Vaz Pinto et al. 2016). By 2009, the last
sable herd in Cangandala included nine old pure females and nine hybrids (Vaz Pinto
et al. 2016). In spite of being naturally sympatric and with deep independent evolu-
tionary histories, roan and sable were not only able to hybridise but also produced at
least two confirmed second generation hybrids (Vaz Pinto et al. 2016).
In Luando, a combination of ground surveys, trap camera monitoring and four
aerial surveys with helicopter between 2009 and 2016, suggested that as result of
the war, giant sable had been extirpated from approximately 75% of their historical
range, but a few herds did manage to survive in the remaining area, and may have
totalled around 100 animals at the end of the war in 2002 (Vaz Pinto 2018).
The extreme population crashes that affected the giant sable populations, were
further unravelled by an extensive genetic study using mitogenomes and nuclear
DNA, applied on a very extensive dataset that included dozens of modern samples
and pre-war material obtained from natural history collections in museums around
the world (Vaz Pinto 2018). The population collapse in Cangandala, caused a severe
loss of heterozygosity that ranks among the lowest ever recorded in mammals (Vaz
Pinto 2018). In Luando, the heterozygosity was only moderately reduced as result
of the bottleneck, but the loss of mitochondrial diversity was extreme, when 11
haplotypes were found in samples dating from the early twentieth century and a
single haplotype became fixed in the extant population (Vaz Pinto 2018). Such a
pattern is consistent with a scenario in which at least two subpopulations coexisted
in Luando with gene flow maintained by male dispersal, before the more ancient
nucleus that used to function as repository of maternal diversity became extinct dur-
ing the war (Vaz Pinto 2018).
Conservation
Being an endemic population, restricted to the north-central Angolan plateau, and

extremely rare, the giant sable has always been a species of conservation concern,
firstly listed in the 1933 London Convention for the Preservation of Fauna and
Flora, as a Class A species worthy of absolute formal protection (Walker 2004). It
also featured as Endangered in the IUCN Red List of Threatened Species since its
creation in 1964, with the status revised to Critically Endangered in 1996 and
486 P. Vaz Pinto
maintained since then (IUCN 2017). The giant sable is also included in Appendix I
of the Convention on International Trade of Endangered Species (CITES) since its
creation in 1975.
The earliest known giant sable conservation policies, in the form of regional
hunting bans, were adopted by the District Governor of Moxico in 1913 just prior to
the formal species description, and by the high commissioner for the colony of
Angola, General Norton de Matos in 1922 (Varian 1953; Walker 2004). On both
occasions, these bans intended to counteract excessive hunting of giant sable, and
must be credited to the vision and perseverance of Frank Varian, without whom the
species may not have lasted long (Walker 2004). This however, did not prevent
many trophy hunters from travelling to Angola to shoot giant sable in the 1920s and
1930s (Walker 2004). Following the listing as a Class A protected species by the
London Convention of 1933, giant sable hunting and the possession of trophies was
prohibited in 1934 (Heim 1954).
Nevertheless, these regulations were poorly enforced, and many specimens were
still being collected by foreign hunters, Portuguese traders and local communities
(Harper 1945). In 1938 the ‘Land between Two Rivers’ was demarcated as a hunt-
ing reserve, and named Reserva da Caça do Luando, thus laying the ground for the
implementation of future integrated conservation policies. Recommendations were
made by the International Union for the Conservation of Nature and Natural
Resources (IUCN) for the creation of a National Park (Heim 1954), and in response
the colonial government upgraded the category of Luando from a hunting reserve to
a strict nature reserve in 1955 (Huntley 1971). In the same year, a new law regulat-
ing hunting and nature conservation in Angola underlined the giant sable as priority
species worthy of full protection, and regulations published in 1957 criminalised the
killing of giant sable, setting the fine at 100,000 escudos (Frade and Sieiro 1960),
which would be equivalent to more than US $10,000 at current value. Although
giant sable conservation finally started to be taken seriously and hunting had been
almost terminated, one last trophy hunting expedition was authorised in 1961 fol-
lowing negotiations at the highest level (Agundis 1965; Vaz Pinto 2018).
Conservation measures for the giant sable were discussed in the late 1950s (Newton
da Silva 1958; Frade 1958; Frade and Sieiro 1960), while for the first time the need
to protect the small pocket population found near the town of Cangandala was
addressed (Frade 1958). The recognition of this second population would lead to the
creation of Cangandala Strict Nature Reserve in 1963, and then elevated to a
National park in 1970 (Huntley 1971).
By the early 1970s the giant sable was relatively well protected and benefited
from conservation management practices in both Cangandala and Luando (Pedrosa
1971; Huntley 1973; Estes and Estes 1974). This was achieved in spite of relatively
moderate budgets and small management teams, comprising two senior rangers and
seven assistant-rangers in Luando, and one senior ranger with four assistant rangers
in Cangandala (Huntley 1971). As the direct harvest of giant sable was much
reduced and virtually eliminated with increased enforcement of protection and man-
agement, other threats gained in perceived relevance, particularly habitat deteriora-

tion due to slash and burn agriculture (Crawford-Cabral 1970; Huntley 1972; Estes
and Estes 1974). Luando Reserve had a human resident population estimated at
18,000 in 1971 in addition to 800 people living inside Cangandala National Park,
mostly resorting to tree-cutting to plant manioc, and thus negatively affecting the
natural miombo woodland (Huntley 1971; Pedrosa 1971; Estes and Estes 1974).
The possible relocation of human populations, conversion of agricultural practices,
and even translocation of giant sable to safe havens was then suggested (Huntley
1972, 1973; Estes and Estes 1974). Nevertheless, such conservation concerns would
soon become irrelevant.
During the Angolan civil war the situation deteriorated rapidly, and if conserva-
tion measures were still in place in Cangandala until 1982 (Estes 1982), soon after
this all management and protection ceased across the giant sable areas. Conservation
initiatives could only be reinstated in recent years, following the arrival of peace and
the location of the last surviving population pockets (Vaz Pinto 2018). In the absence
of formal management in Luando and Cangandala, the Giant Sable Conservation
Project was launched in 2003 by the Catholic University of Angola, and since 2010
has been led by the Kissama Foundation. The Giant Sable Project has been assisting
the Angolan Government in the implementation of conservation practices and man-
agement in both protected areas. Members of resident communities were trained and
appointed as conservation agents, and some have already been transferred to the
park management as government rangers. In Cangandala National Park, the Giant
Sable Project has rehabilitated park infrastructure, deployed equipment and built
fences (Vaz Pinto 2018). Extraordinary conservation measures were adopted in
2009 to tackle the hybridisation crisis in Cangandala National Park, leading to trans-
locations, sterilisation of hybrids and the constitution of a breeding nucleus (Vaz
Pinto et al. 2016).
The most critical issue that has affected the giant sable in recent years, causing
the populations to crash and risking compromising their recovery, is widespread
uncontrolled poaching driven by the bushmeat trade (Vaz Pinto 2018). Although
giant sables do not appear to be specifically targeted by poachers, they are still shot
at, but the most pervasive and negative impact is the large scale use of snares and
foot traps that are having a huge toll, affecting mostly young females and immature
sables (Vaz Pinto 2018).
The Giant Sable Project has installed a network of trap cameras covering a good
part of the reserves, which has allowed regular monitoring and the identification of
individuals, and was instrumental in detecting and documenting the hybridisation
phenomenon in Cangandala (Vaz Pinto et al. 2016; Vaz Pinto 2018). Between 2009
and 2016, a total of 74 giant sable and nine hybrids were darted and marked, 65 of
them released with tracking collars, which included 32 GPS collars, allowing much
increased surveillance power and the gathering of knowledge on the biology of the
species (Vaz Pinto 2018).
488 P. Vaz Pinto
The Angolan Government has been boosting law enforcement measures, and in
2016 the fine for killing one giant sable was increased to a value equivalent to
roughly US$100,000, although no one has been prosecuted for such an offense in
recent decades. Currently, the conservation of giant sable is understandably focused
on anti-poaching, relying on an increase of surveillance, close monitoring of ani-
mals and strengthening park management. By 2018, the giant sable populations had
recovered to around 70 animals in Cangandala National Park and an estimated
150 in Luando.
The Way Forward
The unquestionable importance of securing the future of a critically endangered

taxon, which also happens to be a flagship species and a national icon, must frame
current and future activities. In addition to the obvious need to implement more
effective law enforcement measures and enhance park management through infra-
structure rehabilitation and staff recruitment and training, some specific issues
deserving consideration include erecting new fences and making relocations to
recover parts of the historical giant sable distribution.
As tools ultimately benefitting giant sable conservation and management, several
lines of research can be developed, either as completely new approaches, or by
building on previous work. A study of the food preferences of giant sable is a crucial
topic which can now be addressed in detail with modern molecular tools, comple-
mented with remote tracking of movements. Studies on the use of other local
resources may also prove critical, such as water and natural salt licks. A better
understanding of factors that affect the breeding success and calf mortality, is also
needed. An epidemiological study is required, including a study on parasites poten-
tially affecting the giant sable, and a monitoring programme should be implemented
and extended to other species and domestic animals in the region. The impact of
frequent dry-season fires, and how they reflect on vegetation and on the movements
of giant sable should be assessed, and the use of fire as a management tool should
be explored. The fact that many giant sable have been collared with GPS satellite
transmitters and more will probably be in the future, opens up unique opportunities
to develop research on the spatial use of herds and bulls, address territoriality, use of
local resources, breeding and response to extrinsic factors. Existing molecular tools
should continue to be employed to address individual identification, parameters of
genetic diversity, with clear application on the management of existing populations.
Comparisons with other subspecies and historical giant sable material, and develop-
ment of more advance molecular tools such as with genomics, will also greatly
improve our knowledge, and may assist future breeding efforts, and help preserve
some critical and unique genetic features of this magnificent antelope.
References
Agundis TM (1965) El Llamado de la Montaña, Viajes de Cacería, Angola-Tanzania-Alaska.

Rustica Editorial, Mexico
Ansell WFH (1972) Part 15: order Artiodactyla. In: Meester JAJ, Setzer HW (eds) The mammals
of Africa: an identification manual. Smithsonian Institution Press, Washington, DC, pp 1–93
Baptista SL, Pinto PV, Freitas MDC et al (2013) Geophagy by African ungulates: the case of the
critically endangered giant sable antelope of Angola (Hippotragus niger variani). Afr J Ecol
51(1):139–146
Angola, Luanda
Blaine G (1922) Notes on the zebras and some antelopes of Angola. Proc Zool Soc London
92(2):317–339
Bocage JVB du (1890) Mammifères d’Angola et du Congo (Suite). Jornal de Sciencias,
Mathemáticas, Physicas e Naturaes, Lisboa, Segunda Série 1(1):8–32
Crawford-Cabral J (1965) A palanca preta gigante, sua situação e medidas a adoptar. Luanda,
Unpublished mimeograph
Crawford-Cabral J (1966) A palanca preta gigante, aditamentos e correcções ao relatório do ano
anterior. Luanda, Unpublished mimeograph
Crawford-Cabral J (1969) A study of the giant sable. The zoological society of Southern Africa.
News Bull 10(2):1–7
Crawford-Cabral J (1970) Alguns aspectos da ecologia da palanca real (Hippotragus niger variani
Thomas). Boletim do Instituto de Investigação Científica de Angola 7:5–38
Crawford-Cabral J, Mesquitela LM (1989) Índice toponímico de colheitas zoológicas em Angola
(Mammalia, Aves, Reptilia e Amphibia). Estudos, Ensaios e Documentos 151:1–206
Crawford-Cabral J, Veríssimo LN (2005) The ungulate fauna of Angola: systematic list, distribu-
tion maps, database report. Lisboa: Instituto de Investigação Científica Tropical, Lisboa
Curtis CP (1933) Giant sable antelope. In: Grinnell GB, Roosevelt K (eds) Hunting trails on three
continents. Boone and Crocket Club, New York, pp 237–252
du Bocage JVB (1878) Liste des Antilopes d’Angola. Proc Zool Soc London 1878:741–745
Estes RD (1982) The giant sable and wildlife conservation in Angola. Report to IUCN/Species
Survival Commission, Gland, Switzerland
Estes RD (2013) Hippotragus niger sable antelope. In: Kingdon J et al (eds) Mammals of Africa,
vol 6. Bloomsbury, London, pp 556–565
Estes RD, Estes RK (1970) Preliminary report on the giant sable. Unpublished manuscript, p 22
Estes RD, Estes RK (1972) The giant sable antelope Hippotragus niger variani. Summary report
and recommendations. Unpublished manuscript, p 55
Estes RD, Estes RK (1974) The biology and conservation of the giant sable antelope, Hippotragus
niger variani Thomas, 1916. Proc Acad Natl Sci Phila 26:73–104
Frade F (1958) Mesure adoptées pour la protection de l’Hippotrague géant en Angola. Mammalia
22(3):476–477
Frade F (1967) Palanca Preta Gigante, Relíquia da Fauna de Angola. Unpublished mimeograph,
p 3
Frade F, Sieiro D (1960) Palanca preta gigante de Angola. Garcia de Orta 8:21–38
Gray PN (1930) African game lands. The Sportsman 8(4):1–34
Gray PN (1933) Along the livingstone trail. In: Grinnell GB, Roosevelt K (eds) Hunting trails on
three continents. Boone and Crocket Club, New York, pp 103–143
Groves C, Grubb P (2011) Ungulate taxonomy. Johns Hopkins University Press, Baltimore, p 317
Halse ARD (ed) (1998) Rowland Ward’s records of big game, XXV edn. Rowland Ward
Publications, Johannesburg
490 P. Vaz Pinto
Harper F (1945) Extinct and vanishing mammals of the old world. American Committee for
International Wildlife Protection, New York
Heim F (1954) Les Fossiles des Demain: treize mammifères menaces d’extinction. Étudiés par le
“Service de Sauvegarde”. Union Internationale pour la Protection de la Nature, p 112
Hill JE, Carter TD (1941) The mammals of Angola Africa. Bull Am Mus Nat Hist 78(1):1–211
Huntley BJ (1971) Guia Preliminar dos Parques e Reservas de Angola. Relatório 3. Repartição
Técnica da Fauna, Direcção Provincial dos Serviços de Veterinária, Luanda, Mimeograph
report, p 17
Huntley BJ (1972) Plano para o futuro da palanca real de Angola. Relatório 11. Repartição Técnica
da Fauna, Direcção Provincial dos Serviços de Veterinária, Luanda, Mimeograph report, p 9
Huntley BJ (1973) Distribuição e situação da grande fauna selvagem da Angola com referência
especial às espécies raras e em perigo de extinção – primeiro relatório sobre o estado actual.
Relatório 21. Repartição Técnica da Fauna, Direcção Provincial dos Serviços de Veterinária,
Luanda, Mimeograph report, p 14
Huntley BJ, Matos EM (1992) Biodiversity: Angolan environmental status quo assessment report.
IUCN Regional Office for Southern Africa, Harare
IUCN SSC Antelope Specialist Group (2017) Hippotragus niger ssp. variani. The IUCN red list of
threatened species 2017: e.T10169A50188611
Jansen van Vuuren BJ, Robinson TJ, Vaz Pinto P et al (2010) Western Zambian sable: are they a
geographic extension of the giant sable antelope? S Afr J Wildl Res 40(1):35–42
Klein RG (1974) On the taxonomic status, distribution and ecology of the blue antelope,
Hippotragus leucophaeus Pallas 1766. Ann S Afr Mus 65(4):99–143
Mathee CA, Robinson TJ (1999) Mitochondrial DNA population structure of roan and sable ante-
lope: implications for the translocation and conservation of the species. Mol Ecol 8(2):227–238
Newton da Silva S (1958) A Caça e a Protecção da Fauna em Angola. Author’s Edition, Lisboa,
p 174
Newton da Silva S (1970) A Grande Fauna Selvagem de Angola. Direcção Provincial dos Serviços
de Veterinária, Luanda
Pedrosa V (1971) Deslocação à Reserva Natural e Integral do Luando. Direcção dos Serviços de
Veterinária, Luanda, Unpublished manuscipt, p 24
Pitra C, Vaz Pinto P, O’Keeffe BW et al (2006) DNA-led rediscovery of the giant sable antelope in
Angola. Eur J Wildl Res 52(3):145–152
Powell-Cotton PHG (1932) Angola. In: Maydon HC (ed) Big game shooting in Africa. Seeley,
London, p 445
Schouteden H (1947) De zoogdieren van Belgisch-Congo en van Ruanda-Urundi (Les mam-
mifères du Congo Belge et du Ruanda-Urundi), III. Ungulata (2) Rodentia. Annales du Musée
du Congo Belge Série 2 3(3):333–576
Sieiro D (1962) Novas observações acerca da palanca preta gigante Ozanna grandicornis variani
(Thomas). Boletim do Instituto de Investigação Científica de Angola 1:49–57
Silva JA (1972) A palanca real – contribuição para o estudo bioecológico da palanca real
(Hippotragus niger variani). Junta de Investigações do Ultramar, Lisboa
Statham JCB (1922) Through Angola: a coming colony. Blackwood & Sons, London
Thomas O (1916) A new sable antelope from Angola. Proc Zool Soc Lond 1916:298–301
Varian HF (1953) Some African milestones. Books of Rhodesia, Bulawayo, p 272
Vaz Pinto P (2007) Hybridization in giant sable. A conservation crisis in a critically endangered
Angolan icon. IUCN/SSC Antelope Spec Group Gnuslett 26:47–58
Vaz Pinto P (2018) Evolution history of the critically endangered giant sable antelope (Hippotragus
niger variani) – insights into its phylogeography, population genetics, demography and conser-
vation. PhD thesis. University of Porto, Porto
Vaz Pinto P, Lopes S, Mourão S et al (2015) First estimates of genetic diversity for the highly endan-
gered giant sable antelope using a set of 57 microsatellites. Eur J Wildl Res 61(2):313–317
Vaz Pinto P, Beja P, Ferrand N et al (2016) Hybridization following population collapse in a criti-
cally endangered antelope. Sci Rep 6:18788
Walker JF (2004) A certain curve of horn: the hundred-year quest for the giant sable antelope of
Angola. Grove/Atlantic Inc, New York
Wessels J (2007) Western Zambian sable: a giant sable look-alike or the real thing. Game Hunt
13:32–36
Wilson DE, Hirst SM (1977) Ecology and factors limiting roan and sable antelope populations in
South Africa. Wildl Monogr 54:3–111
Part V
Research and Conservation Opportunities
Chapter 18
Biodiversity Conservation: History,
Protected Areas and Hotspots
Brian J. Huntley, Pedro Beja, Pedro Vaz Pinto, Vladimir Russo,

Luís Veríssimo, and Miguel Morais
Abstract Angola is a large country of great physiographic, climatic and habitat

diversity, with a corresponding richness in animal and plant species. Legally pro-
tected areas (National Parks and Game Reserves) were established from the 1930s
and occupied 6% of the country’s terrestrial area at the time of independence in
1975. As a consequence of an extended war, the Protected Areas were exposed to
serious neglect, poaching and land invasions. Many habitats of biogeographic
importance, and many rare and endemic species came under threat. The recently
strengthened administration gives cause for optimism that a new era for biodiversity
conservation is at hand. The Protected Areas system was greatly expanded in 2011,
and increasing resources are being made available towards achieving management
effectiveness.
B. J. Huntley (*)
P. Beja
e-mail: pbeja@cibio.up.pt
P. Vaz Pinto
CIBIO-InBIO Centro de Investigação em Biodiversidade e Recursos Genéticos,
V. Russo · L. Veríssimo
e-mail: vladyrusso@gmail.com; vladimir.russo@holisticos.co.ao; lmnverissimo@gmail.com
M. Morais
Faculdade de Ciências, Universidade Agostinho Neto, Luanda, Angola
e-mail: dikunji@yahoo.com.br

https://doi.org/10.1007/978-3-030-03083-4_18
496 B. J. Huntley et al.
Keywords Angola · Bushmeat · Ecoregions · Marine ecosystems · Protected areas

· Threatened species · Wildlife trade
Introduction: Wildlife Conservation During the Colonial Era
In common with most colonial territories in Africa, conserving wildlife was not a
general consideration in Angola until the twentieth century. However, the first
expression of concern regarding wildlife numbers came much earlier, and from
none other than the most famous zoological collector to work in Angola – José
Anchieta. In correspondence with the great Barbosa du Bocage, Anchieta (1869, in
Andrade 1985:87) noted that inland of Luanda “the big game, abundant until fifty
years ago, has moved into the interior because of the increased population and gen-
eral use of firearms.” But worse was to come. In 1880, the ‘Angola Boers’ arrived in
Humpata, having crossed the Kalahari on their fateful Thirstland Trek (Stassen
2016). The Boers’ hunting depredations soon spread across the country. Professional
hunters such as William Chapman (Stassen 2010) described the wealth of game in
the southwest, and personally contributed to its depletion.
Globally, by the end of the nineteenth century many repentant hunters were
becoming alarmed at the fate of the once abundant herds and mobilised action to
address the problem. The first international agreement on nature conservation was
the Convention for the Preservation of Wild Animals, Birds and Fish in Africa, oth-
erwise known as the London Convention, held in London in 1900. Attended by 11
European powers, the Convention was not ratified by several countries, including
Portugal, and was abandoned with the onset of World War I (Carruthers 2017).
Interestingly, the convention was the brain-child of the German hunter/explorer/
military officer Hermann von Wissman, who with Paul Pogge collected in Malange
and the Lundas in the early1880s, before he crossed Africa on the first of two trans-
continental expeditions.
By the early twentieth century the impact of Boer biltong hunters had become
notorious, and Thomas Varian, who introduced the Giant Sable Antelope to science,
convinced the Governor of Moxico in 1913, and the then Portuguese High
Commissioner, Norton de Matos, to close the sable lands to hunting (Varian 1953).
The fame of the Giant Sable drew numerous trophy-hunting and scientific expedi-
tions to Angola through the 1920s and 1930s (Walker 2004) and the zoological col-
lections they made contributed much to our knowledge of Angola’s biodiversity.
The London Convention of 1900 was followed in 1933 by the Convention
Relative to the Preservation of Fauna and Flora in their Natural State – also known
as the London Convention. Whereas the 1900 convention focused on hunting regu-
lations, the 1933 convention promoted the idea that each colonial power should
establish national parks and reserves in their colonial territories, following the
model of the Kruger National Park established by South Africa in 1926. The 1933
convention also required states to give special protection to an internationally
selected list of species – a list that included Giant Sable Antelope and the enigmatic
desert plant Welwitschia mirabilis. Interest in protecting Angola’s fauna was rising,
18 Biodiversity Conservation: History, Protected Areas and Hotspots 497
and hunters such as Henrique Galvão and Teodósio Cabral, administrators such as
Norton da Matos and Abel Pratas, and scientists such as Fernando Frade and Luis
Carrisso, immediately championed the National Park model. Portugal set about cre-
ating Angola’s National Parks and Game Reserves even though it did not ratify the
1933 convention until 1950. The first of these protected areas was Iona, established
as a Game Reserve by decree on 2 October 1937, followed by Cameia, Quiçama,
Bicuar and Luando proclaimed on 16 April 1938. The first four of these game
reserves were raised to National Park status in the 1950 and 1960s.
The good work of the 1930s and 1940s was reversed by the caça livre (free hunt-
ing) period of the early 1950s, when the wildlife populations of the cattle ranching
region of the southwest were decimated because of concerns regarding stock dis-
eases carried by wild species. The voices of reason were raised by a younger gen-
eration – Luis Carmo, Armando Malacriz and Newton da Silva (Newton da Silva
1952, 1970) and by 1955 Angola had a new and detailed legislative instrument, the
Decreto 40,040 (Regulamento sobre a Protecção do Solo, Flora e Fauna) which
was only revoked in 2017. Wildlife conservation was given formal status as a public
concern by the establishment of the Conselho de Protecção da Natureza (CPN) in
1965, chaired by the Governor General. The CPN played a pivotal role in the expul-
sion of cattle ranches from Quiçama in the 1970s (Huntley 2017). The increasing
support of the Portuguese government for conservation came to a head when a
major conference of its African territories was convened in Lubango in 1972. Titled
Reunião para o Estudo dos Problemas da Fauna Selvagem e Protecção da Natureza
no Ultramar Português, the meeting ran for 2 weeks and was attended by 73 dele-
gates. It prepared 53 recommendations for action to improve the protection of nature
throughout Angola, leading the government to double the budget of the department
responsible for National Parks – the Repartição Técnica da Fauna.
Post-independence History of Conservation in Angola
Following the ‘Carnation Revolution’ of 25 April 1974 in Portugal, and soon after
gaining independence, Angola entered a period of increasing difficulty and ulti-
mately war, which only ended in February 2002. The impact of this period of vio-
lence and displacement on the wildlife and protected areas of Angola is described
elsewhere (Walker 2004; Huntley 2017). During the war years, efforts to bring pub-
lic support to the National Parks were made through convening annual Semanas do
Ambiente (Environment Weeks) led by a small network of Angolans, most notably
Carlos Pinto Nogueira, Serôdio d’Almeida and Vladimir Russo. Most of the pro-
tected areas were abandoned and the wildlife populations decimated during the early
years of the war. In 1992 the International Union for the Conservation of Nature
(IUCN) led an international study of the situation (IUCN 1992) that concluded:
Since 1975, most, if not all populations of large mammals have been severely reduced, if
not eliminated. Wholesale slaughter of elephant, rhino, eland, roan, oryx, springbok, zebra,
bushbuck, reedbuck, lechwe and many other species occurred in all parks and reserves. It is
possible that some nucleus herds still survive, sufficient to recover if given effective
protection.
In an ironic twist of wildlife conservation practice, in 1995 the Kissama Foundation

(KF) was established, led by a group of conservation-minded military generals.
Wishing to support Quiçama National Park on its road to recovery, the KF raised
funds to re-introduce species that had been severely reduced during the first decades
of the war. Unfortunately the initiative, promoted as ‘Operation Noah’s Ark’ in
2000 introduced many species never previously known to occur in Quiçama. Despite
international concern regarding the introductions, the programme was continued
and expanded in 2014 by the then Minister of Environment, as an essentially private
effort to create a mixed collection of animals in the tiny encampment – ca. 1% of
Quiçama –that formed the ‘Special Protected Area’. Sadly, the remaining 99% of
Quiçama has since been left to the ravages of the bushmeat trade and illegal land
occupation. Species never previously recorded in Quiçama but introduced with
Ministerial approval in 2000 and 2014 included Plains Zebra, Giraffe, Kudu, Nyala,
Common Waterbuck, Blue Wildebeest, Red Hartebeest, Blesbok, Oryx, and
Common Impala. Nyala and Blesbok have never been recorded in Angola, or within
2300 km of Quiçama. Only two of the species introduced, Savanna Elephant and
Eland, were native to the park, but the poorly documented animals introduced by
wildlife dealers were from different genepools to the original Quiçama
populations.
During the early 2000s, international interest in Angola led to several initiatives,
most notably those of the Global Environment Facility (GEF), to support conserva-
tion in the country. A fundamental step supported by the GEF was the development
of a National Biodiversity Strategy and Action Plan (NBSAP), (GoA 2006) that
gave direction to policy established in the Base Law for the Environment (GoA
1998). In support of the objectives of the NBSAP, a survey of the Cuando Cubango
paved the way to the proclamation of the Luengue-Luiana National Park (Bergman
and Verissimo 2008). The proclamation of the Maiombe National Park in Cabinda
resulted from the Mayombe Transfrontier Conservation initiative. GEF funding was
raised to help rehabilitate and expand the protected areas system of Angola, and this
and other initiatives continue to support the government in its programme.
The Protected Areas System
Angola’s protected areas system, proposed in 1936, with the first reserve estab-
lished in 1937, expanded rapidly through to the 1970s, by which time 13 Protected
Areas (PAs) had been established, totaling 75,267 km2 or 6,0% of national territory.
During the early 1970s, extensive surveys were undertaken to identify key biodiver-
sity hotspots or other areas deserving inclusion in an expanded conservation net-
work (Huntley 1974a, b, c, d, 2010). The objective was to increase the representation
of Angola’s vegetation types and faunal species diversity within the PA system.
Unfortunately the interruption of war and the weakness of governance systems
delayed the consideration and approval of the recommendations until 2011, when
the Conselho de Ministros not only approved the proposals of 1974 but added
Table 18.1 Terrestrial protected areas of Angola

Name Category Date established Area 1, km2 Area 2, km2
Iona National Park 1937 15,150 15,196
Cameia National Park 1938 14,450 14,688
Quiçama National Park 1938 9960 9227
Mupa National Park 1938 6600 6039
Bicuar National Park 1938 7900 6748
Cangandala National Park 1963 650 637
Mavinga National Park 2011 Unknown Unknown
Luengue-Luiana National Park 2011 45,818 22,720
Maiombe National Park 2011 1930 2074
Chimalavera Regional Nature Park 1971 150 102
Luando Integral Nature Reserve 1938 8280 9930
Ilhéu dos Pássaros Integral Nature Reserve 1973 2 1.5
Búfalo Partial Reserve 1974 400 405
Namibe Partial Reserve 1957 4450 4642
Total Area, km2 115,740 92,409.5
Two game reserves established in the 1930s – Ambriz, of 1125 km2, and Milando, of 6150 km2 –
and since deproclaimed – are not included in this listing. Furthermore, the boundaries of Mavinga
await clarification. Sources for Area – 1: GoA 2018; 2: Veríssimo Unpublished Data 2018
several new areas. In terms of Law 38/11 of 29 December 2011, the PA system
increased to over 115,000 km2 of national territory in one step, Table 18.1, Fig. 18.1.
However, some debate continues regarding the definition of the boundaries of the
individual PAs, with recent estimates by Verissimo (2018 Unpublished Data) pro-
viding new insights. While the area proclaimed as PAs was nearly doubled, the
budget has remained on a very low plateau. Most National Parks still lack the most
basic management capacity and effectiveness, despite the wealth of legislation pro-
mulgated since the Base Law for the Environment was approved in 1998.
The contradictions of global conservation policy, on one hand pressuring govern-
ments to reach a target of 17% of national territory under protected areas by 2020
(CBD 2010) and on the other, expecting the governments of developing countries to
provide funds to effectively manage such PAs, is well illustrated by the situation in
Angola. The drive to double the area under legislation has been accompanied by the
neglect of iconic protected areas such as Quiçama, Iona and Luando. Since peace
was achieved in 2002, the illegal occupation of the vulnerable Quiçama coastline by
tourism lodges, fishing villages, oil exploration infrastructure, commercial enter-
prises and quarries, and by cattle ranches and commercial agricultural schemes, in
addition to the rampant activities of bushmeat poachers and charcoal producers, has
continued unabated. Iona, once a pristine desert environment, is now occupied by
nomadic pastoralists who have invaded the heart of the park, supported by govern-
ment sponsored water points which give permanence to the occupation. While some
of the land invasions were a consequence of the war, most have occurred since the
peace of 2002.
Fig. 18.1 Protected areas of Angola: • 1 Maiombe • 2 Quiçama • 3 Cangandala • 4 Cameia • 5 Iona
• 6 Bicuar • 7 Mupa • 8 Luengue-Luiana • 9 Luando • 11 Chimalavera • 12 Búfalo • 13 Namibe.
(Mavinga is not indicated on this map due to incomplete details regarding its boundaries in its
gazettement)
The difficulties attending limited budgets, weak technical capacity and poorly
trained human resources suggests that a triage approach should be considered to
bring a focus to where the government’s limited conservation resources should be
targeted (Huntley 2017). Recent government policy has been to expand the PA
estate, regardless of the management capacity of such ‘paper parks’. Fortunately,
despite the reverses of the past decades, each protected area still includes areas of
sufficient dimension that can, with effective management, achieve significant biodi-
versity conservation goals. Since 2017, the new government leadership gives prom-
ise for a revitalised and energetic approach to conservation in Angola, as

demonstrated in the recent Strategic Plan for the Conservation Areas System of
Angola (GoA 2018).
Wildlife Populations
In contrast to most southern African countries, where reliable statistics of wildlife

population dynamics have been recorded over many decades, the data sets for
Angola are extremely sparse. Estimates made during the 1970s tended to be conser-
vative, but indicated robust populations of Elephant (600), Forest Buffalo (6000),
Eland (3000) and Roan Antelope (3000) in Quiçama (Huntley 1971). These species
were extinct or nearly so in the park by 1992 (IUCN 1992). The populations of
Giant Sable Antelope and Red Lechwe in Luando, each estimated at 2000 in 1972
(Huntley 1972), had dropped to less than 100 Giant Sable and with Lechwe on the
verge of extinction by 2017 (Vaz Pinto 2018, 2019). Savanna Elephant, Blue
Wildebeest and Eland, abundant in Bicuar in the 1970s, had fallen to low numbers
by 2017 (Beja et al. 2019). Across Angola, wildlife populations declined precipi-
tously after 1974, but remarkably, very small but tenacious surviving populations of
most species, including top predators such as Lion, Leopard, Cheetah and Wild
Dog, have held out (Beja et al. 2019). Of considerable conservation concern is the
number of large mammal species for which no recently confirmed records are avail-
able, including Gorilla, Black Rhinoceros, Puku, Red Hartebeest and Lichtenstein’s
Hartebeest (Beja et al. 2019).
The wildlife population densities and biomasses of Angolan protected areas have
never been comparable to those in eastern and southern Africa. While this might be
a factor of hunting pressure, more fundamental ecological factors are at play. As
demonstrated by Bell (1982), herbivore population density and biomass in Africa is
related to rainfall and soil nutrients, and more directly to the ratio of soluble to
structural carbohydrates in plant material available to herbivores. The vast area of
Angola covered by miombo woodlands with low-nutrient grasses, shrubs and trees
accounts for the notoriously low game populations of central Angola. Only in the
more arid savannas of the southwest and southeast were relatively large populations
of herbivores found in colonial times. The popular perception of vast populations of
game across Angola in the nineteenth century is an illusion, certainly if compared
with eastern Africa, as manifest by the much lower volumes of ivory exported from
Angola relative to Kenya throughout the period (Walker 2009). The highest bio-
masses for ungulates during the 1970s, based on estimates from field surveys, were
the western littoral grasslands of Quiçama, occupied by Eland, Roan and cattle, and
the northern ‘Baixa dos Elefantes’ forests and floodplains of the Cuanza, occupied
by Savanna Elephant, Forest Buffalo and Hippopotamus. Ungulate biomass in
Cangandala, Luando and Bicuar, in dense miombo woodlands, was very low, as was
that of Iona. The richest hunting areas (Coutadas) of the southeastern Cuando
Cubango (Mucusso and Luiana) possibly had biomasses approximating those of

similar nutrient poor mixed miombo woodlands of eastern Africa, such as Selous
Game Reserve (Huntley Unpublished Data).
pecies Richness, Endemism, Threatened Species

S
and Biodiversity Hotspots
The seminal paper on Systematic Conservation Planning by Margules and Pressey

(2000) triggered the wide adoption of objective measures for the identification of
biodiversity conservation priorities. The process has been effectively applied in
southern Africa, where fine-scale spatial data on species distribution and status are
available, such as that required by IUCN categories of threat (Raimondo et al.
2009), vegetation and habitat maps (Mucina and Rutherford 2006), and surveys of
marine ecosystems and their dynamics (Kirkman et al. 2016; Holness et al. 2014).
These rich data sets have been used to produce detailed national and regional biodi-
versity conservation management plans (Driver et al. 2012; Kirkman et al. 2016)
which provide models for future work in Angola.
A preliminary survey of the conservation status of Angolan mammals (Huntley
1973) gave subjective estimates for 70 species, none of which were considered
threatened with extinction but several, in particular Gorilla, Chimpanzee and Black
Rhinoceros, were deemed vulnerable. A summary of recent assessments of rarity
and threat in various taxonomic groups in Angola are presented in Table 18.2. More
specific details of conservation status or threats are provided in the sources for each
major taxonomic group referenced in Table 18.2.
In an early objective assessment of habitats, the areas of 32 vegetation units
mapped by Barbosa (1970) were measured to evaluate the proportional representa-
tion of each unit in the protected areas system (Huntley 1974a). The results were
then used to focus attention on under-represented types, taking into consideration
faunal as well as floral distribution and status (Huntley 1974c). Of the 32 vegetation
types described by Barbosa (1970), only 11 fell within protected areas in 1974. The
disparity of protection afforded to representatives of the major biogeographic divi-
sions was considerable. The Karoo-Namib, represented by Barbosa vegetation types
27, 28 and 29, which occupy 2.6% of the country’s land surface, had 50.6% of its
area conserved, while the Guineo-Congolian forest/savanna mosaic, comprising
25.7% of Angola’s total area, and holding probably over 70% of its biodiversity, was
not represented in any protected area. The small relict fragments of Afromontane
forest, without doubt the most threatened of all ecosystems in Angola, and currently
reduced to less than 1000 ha in area, were also without protection. Both the
Afromontane forests (Humbert 1940; Hall and Moreau 1962) and the Angolan
Escarpment Zone (Hall 1960) have long been regarded as key centres of avifaunal
speciation and floristic importance. But both remain unmapped and unprotected.
Table 18.2 Species richness, endemism and threatened status for selected taxa
Endemic
species
Group Total species n° % IUCN status Source
Plants 6850 indigenous 997 14.6 399 species have been formally 1–3
species assessed, of which: 36 threatened:
230 naturalised 32 vulnerable,
species 4 endangered,
49 threatened or near-threatened
Butterflies & 792 57 7.2 Not evaluated 4
Skippers
Dragonflies & 260 16 6.1 1 vulnerable 5
Damselflies 4 near threatened
16 data deficient
6 not evaluated
Fishes 358 78 22 0 6
Amphibians 111 21 19.3 Not evaluated 7
Reptiles 278 Not evaluated 8
Birds 940 29 3.1 Not evaluated 9
Mammals 291 12 4.1 2 critically endangered, 10
2 endangered
11 vulnerable
14 near threatened
12 data deficient
235 least concern
1: Figuerido and Smith (2008), 2: Goyder and Gonçalves (2019), 3: IUCN (2018), 4: Mendes et al.
(2019), 5: Kipping et al. (2019), 6: Skelton Unpublished Data, 7: Baptista et al. (2019), 8: Branch
et al. (2019), Dean et al. (2019), 10: Beja et al. (2019)
The identification of sites of high biodiversity importance (in terms of ende-

mism, species richness, and threat) – popularly termed biodiversity ‘hotspots’
(Myers 1988; Myers et al. 2000) – was the focus of the Angolan Protected Areas
Expansion Strategy (Huntley 2010) submitted to the Minister of Environment and
adopted, with additional recommendations in 2011 (GoA 2011, 2018). The sites
recommended for future gazettement included examples of:
• Guineo-Congolian Forest and Savanna: (Maiombe – Cabinda; Serra Pingano –
Uíge; Lagoa Carumbo – Lunda-Norte; Serra Mbango – Malange; Gabela forest –
Cuanza-Sul; Cumbira forest – Cuanza-Sul;
• Afromontane Forest and Grassland: (Mount Namba – Cuanza-Sul; Mount
Moco – Huambo; Serra da Neve – Namibe; Serra da Chela – Huíla Province);
• Zambezian Flooded Grassland: (Luiana – Cuando Cubango).
The inclusion of these proposals in the Angolan protected areas system would
effectively address the asymmetry of ecosystem representation, with the number of
Barbosa vegetation units increasing from 11 to 23. To date, Maiombe forest and
Luiana (with adjacent Luengue, and Mavinga) have been gazetted as additional
National Parks.
Recent studies by the National Geographic Okavango Wilderness Project have
identified further biodiversity hotspots in the upper reaches of the Cuando and
Cubango drainages (NGOWP 2018). Field surveys in Huíla (Mendelsohn
Unpublished Data) and Zaire (Vaz Pinto Unpublished Data) and Cuanza-Norte
(Hines Unpublished Data) have identified sites of high biodiversity interest that are
also deserving of further study and evaluation as future protected areas. As biodiver-
sity surveys become more inclusive of Angola’s less accessible areas, more sites of
conservation merit will undoubtedly be added to the list of priorities.
Coastal and Marine Ecosystems
At the vast scale of marine environments, the recently concluded multinational pro-
gramme of research in the Benguela Current Large Marine Ecosystem (BCLME)
has provided very detailed assessments of the demersal fish biodiversity hotspots
and the dynamics of the oceanic and climatic systems that influence this biodiver-
sity (Kirkman et al. 2013, 2016; Kirkman and Nsingi 2019). These researchers
found that hotspots of species richness were associated with greater water depths
and cooler bottom temperatures. From consideration of the relevance of measured
climate changes, they concluded that range shifts in species associated with warm-
ing temperatures could conceivably affect the spatio-temporal persistence of
hotspots in the long term (Kirkman et al. 2013).
In a detailed analysis of the spatial characterisation of the BCLME, based on the
physical driving forces, primary and secondary production, trophic structures and
species richness, Kirkman et al. (2016) found four different sub-systems, of which
two fall within Angolan waters. The first lies to the north of the Angola-Benguela
Front and the second between the Angola-Benguela Front and Luderitz. Using the
products of the BCLME projects, Holness et al. (2014) used Systematic Conservation
Planning concepts and approaches to identify potential marine protected areas for
the benthic and coastal ecosystems of Angola, Namibia and South Africa. A total of
248 distinct ecosystem types within the BCLME of these countries were mapped
and classified according to Ecosystem Threat Assessments and Ecosystem Protection
Level Assessments. In Angola, five ecosystem types were found to be both Critically
Endangered and Not Protected, mainly situated in areas subject to intensive coastal
development, in the oil and gas fields in the north, or in particular inshore areas
subject to more intense fishing pressure. If the Endangered and Poorly Protected
categories are also included, there are an additional 23 priority ecosystem types for
protection in Angola. The BCLME studies (Kirkman et al. 2013, 2016; Holness
et al. 2014) provide excellent models for the application of Systematic Conservation
Planning meriting replication across the terrestrial ecosystems of Angola.
The coastal ecosystems of Angola are particularly vulnerable to human distur-

bance, both directly through over-exploitation of living resources and indirectly
through urbanisation and industrialisation within coastal environments (Weir et al.
2007; Morais et al. 2005, 2008, 2016). The marine turtle species that depend on
Angola’s sandy beaches for nesting are particularly vulnerable. Despite these chal-
lenges, Angola remains a very important sea turtle conservation nation, with Olive
Ridley (Lepidochelys olivacea), Leatherback (Dermochelys coriacea) and Green
(Chelonia mydas) sea turtles nesting regularly during the summer (Morais 2016,
2017). Loggerhead (Caretta caretta) nest very sporadically, while Hawksbill
(Eretmochelys imbricata) sea turtles are not known to nest on Angolan shores
although juveniles have been recorded on the Soyo and Cabinda coast (Morais
2016). Recent studies estimate that between 33,000 and 102,000 Olive Ridley Sea
Turtles made use of the Angolan coast to nest during the 2015/2016 summer, show-
ing a decrease from between 38,000 and 110,000 estimated during the 2014/2015
season. These figures demonstrate that the coast of Angola is one of the most impor-
tant nesting regions for this species in the Eastern Atlantic (Morais 2016; Kitabanga
Project 2017). Leatherback Sea Turtle are much less abundant, with estimates of
between 495 and 1320 animals nesting along the entire coast of Angola during the
2015/2016 breeding season (Morais 2016). Angola provides the southern extension
of the Gabon nesting grounds, where 6000 to 7000 females nest annually (Billes
et al. 2006). As such, Angola may be second in importance on the Eastern Atlantic
coastline for the nesting of this species. Inadequate data are available to determine
trends in Green Sea Turtle populations on the Angolan coast (Morais 2015, 2016).
Drivers of Species Loss
One of the immediate causes of population declines and species loss in vertebrates
since 1975 has been hunting for bushmeat during the prolonged war, undertaken by
rural communities faced with starvation, or by soldiers seeking to supplement very
limited rations. Moreover, the illegal trade in wildlife products (ivory, rhino horn
and teak) became significant during the war as the leaders of UNITA (União
Nacional para a Independencia Total de Angola) sought funds to purchase arms
(Breytenbach 2015). Luanda has long provided an open market for the illegal trade
in wildlife products (Milliken et al. 2006; Svensson et al. 2014) and was described
by Martin and Vigne (2014) as the biggest ivory market in Africa. The Mercado do
Artesanato in Luanda has openly traded in ivory (mainly sourced from the DRC),
leopard skins and other wildlife products, in the full knowledge of the Angolan
authorities. Following international condemnation of the practice, trading in ivory
has been banned in Angola since 2017. Despite the proclamation of two mega-parks
in the Cuando Cubango in 2011, the poaching of elephants for ivory has increased
in the parks and the elephant population is estimated to have decreased by 21% from
2005 to 2015 (Schlossberg et al. 2018). The inclusion of the area within the
much-publicised Kavango-Zambezi Transfrontier Conservation Area, promoted as

the largest TFCA on Earth (Peace Parks Foundation 2016) has yet to demonstrate
conservation benefits.
A more pervasive impact than ivory poaching, recorded across the country
despite the prohibition of hunting since the late 1970s, is the informal trade in bush-
meat (Bersacola et al. 2014). During a survey in September 2013, travelling 1700 km
along the Angolan Escarpment, Bersacola and colleagues stopped at 13 market
places and counted 71 specimens of 15 prey species. The surveys were mostly in
forested areas, where mammals have been more resilient to poaching pressures than
in open savannas and woodlands. The most numerous species found were Blue
Duikers (45%), Blue (Pluto) Monkeys (11%), Bush Hyraxes (10%) and Yellow-
backed Duikers (8%). “For 25 fresh carcasses, the hunting technique was evident. A
total of 84% of these fresh carcasses were hunted with shotguns, 16% were trapped
using metal or string snares.” The National Geographic Okavango Wilderness
Project has described ‘industrial-scale’ bushmeat harvesting operations in many
areas of the Cuando Cubango (NGOWP 2018).
While the illegal trade in wildlife products has been documented for animal spe-
cies, a much larger trade in timber products has exploded over the past 5 years but
without any measurement or monitoring. In an effort to stimulate alternative foreign
income streams following the global collapse of oil prices, the then Angolan presi-
dent signed decrees in 2016 that facilitated the rapid issue of concessions for timber
extraction over much of the country. Chinese agents have mobilised the massive
extraction of hardwoods from across Angola, accelerating the deforestation of vast
areas, even in the previously near-pristine woodlands of Moxico and Cuando
Cubango provinces (Mendelsohn 2019 Unpublished Data).
Land transformation, as described by Mendelsohn (2019), is perhaps the most
potent of all drivers of biodiversity loss, but like the timber trade, its impacts on
biodiversity have not been quantified at a species level. Mendelsohn and Mendelsohn
(2018) draw attention to the transformation of rural to urban economies, and from
subsistence to cash-based economies. The result has been the demand of the newly
urbanised populations for cash to purchase goods and services previously provided
by rural ecosystems. For rural dwellers, cash is now derived from the sale of bush-
meat and charcoal, not of fruit and vegetables.
Another insidious driver of species loss is that of invasive alien species. The pres-
ence of potentially invasive alien fish species introduced for aquaculture has been
reported for Oreochromis mossambicus in the Cuanza and Oreochromis niloticus in
Cabinda, and recently in the upper Cubango (Skelton 2019). Invasive alien plants
have already become established over extensive areas of western Angola. Rejmánek
et al. (2017) conducted a rapid assessment of invasive plant species across 13 pri-
mary vegetation types (Barbosa 1970) in western Angola and recorded populations
of 44 naturalised plant species, 19 of which are conclusively invasive (spreading far
from introduction sites). They found that dense invasive populations of Chromolaena
odorata, Inga vera and Opuntia stricta pose the greatest threats. Opuntia stricta has
invaded large areas of the arid coastal plain northwards from Dombe Grande, and
along the Chela escarpment. Inga vera is widespread in the moist ‘coffee forests’ of
the central escarpment, while Chromolaena odorata is prevalent in the northern
escarpment. These species have become major environmental and economic

problems elsewhere in Africa and the lack of any control actions in Angola is cause
for concern.
Science and Protected Area Management
The development during the nineteenth century of pragmatic wildlife management

practices into a sophisticated conservation science is reflected in the histories of the
protected areas systems of South Africa, Namibia and Tanzania (Carruthers 2017).
Angola has had very limited investment in research in its national parks and reserves
until the present decade. Notwithstanding limited resources, biologists from the
Instituto de Investigação Científica de Angola (IICA) and the Instituto de
Investigação Agronómica de Angola conducted important surveys of birds (Pinto
1983), mammals (Frade 1956, 1960; Crawford-Cabral 1970, 1971) and vegetation
(Teixeira et al. 1967; Teixeira 1968; Barbosa 1970) in various parks during the
1960s and 1970s. Estes and Estes (1974) undertook detailed behavioural studies of
Giant Sable in Luando in 1970/71. Huntley undertook general ecological surveys in
the protected areas and across most of Angola (Huntley 1973, 1974d, 2017), while
Dean (2000) studied the avifauna of Angola in the field and in the key collections of
museums of Angola, Europe and the USA. But it was not until the present century
that more detailed studies were initiated in the protected areas of Angola, such as
the long-term studies of Giant Sable in Cangandala and Luando by Vaz Pinto (2018)
and of sea turtles on the Angolan coast (Kitabanga Project 2017). Nevertheless,
there have recently been important surveys of the remnant populations of large
mammals in the protected areas of Angola (Beja et al. 2019), and on the threatened
and endemic avifauna of the escarpment (Dean et al. 2019).
Despite these recent advances, the need for full-time biologists posted to and liv-
ing in Angola’s protected areas is urgent. This is a vacuum that should be filled by
young Angolan researchers, with the guidance and support of mentors from across
the globe, as the successful models of many other African countries have demon-
strated. The modern tools of remote sensing, geographic information systems,
immobilisation drugs, radio-tracking collars, trap-cameras, drones, genetic finger-
printing, and much more are readily available. The opportunities are endless and the
difficult challenges of the past are being resolved each year as access to Angola
improves and both international and national government support increases.
Key Priorities for Biodiversity Conservation
Priorities for the conservation of species within different taxonomic groups (plants,
invertebrates, vertebrates) are summarised by Russo et al. 2019). Here we focus on
generic issues of concern.
The effective management of protected areas is one of the key mechanisms that
governments have available to achieve biodiversity conservation goals (CBD 2010).
With over 10 million ha of Protected Areas (PAs) gazetted (GoA 2018), Angola has
a considerable proportion of its terrestrial landscape under formal legislation. This
provides the potential for a broad base to the PA estate, with many species and
ecoregions represented in the system. However, many of the biodiversity hotspots
identified in successive PA expansion strategies (Huntley 1973, 2010; GoA 2011,
2018) are yet to be accurately surveyed, described and gazetted. A first priority
would be to ensure that legislative protection is given to Angola’s most critically
endangered biodiversity hotspots, such as the forests of the escarpment, the central
highlands and the northern borders with the Democratic Republic of Congo.
As urgently important as legal protection is effective management of PAs. The
existing network of extensive PAs such as Iona, Quiçama, Cangandala, Luando,
Bicuar, and Luengue-Luiana lack adequate resources, and these need reinforcement
through provision of personnel, training, equipment and operational budgets. The
options of joint ventures with international conservation organisations and public/
private partnerships such as those that have succeeded in Botswana, Mozambique,
Namibia, Zambia and other southern African countries needs consideration. Field
training of rangers and researchers with ongoing mentoring is a fundamental pro-
cess for professional development. At a national scale, Angola has excellent conser-
vation strategies (GOA 2006, 2018), and several parks already have pragmatic
‘emergency’ management plans (Huntley 1974b, 2003; Anderson and Morkel
2009). These need adaptation and implementation rather than repetition. For many
PAs, a triage approach to zonation and investment is appropriate where land inva-
sions, illegal infrastructure developments and other irreversible developments have
taken place (Huntley 2017).
Concluding Remarks
The engagement of the public at large in conservation is a first priority for Angola’s
biodiversity agenda. The use of social media has already born unexpected results.
The Facebook forum Angola Ambiente has over 1000 members and the posting of
dragonfly photos on its page has led to 12 species being identified as new to science
(Chris Hines, pers. comm.). The conservation of flagship species that attract public
attention at national and international scales is also of the utmost importance. A
well-publicised example is the conservation project that has successfully saved the
Giant Sable Antelope in Cangandala and Luando protected areas, as described by
Vaz Pinto (2019). Another noteworthy example is the Kitabanga Project of
Agostinho Neto University, which has monitored sea turtle populations and under-
taken conservation actions since 2013 (Projecto Kitabanga 2017). The project
involves research and environmental education on sea turtles, with specific empha-
sis on protecting the nesting beaches. The Kitabanga Project provides an excellent
model of a locally driven conservation research and education initiative and deserves
replication in Angola. The National Geographic Okavango Wilderness Project

(NGOWP 2018) has also brought wide attention to Angola’s biodiversity, and stim-
ulated young Angolans to join biodiversity exploration and research initiatives.
These and other projects that will be developed in the future contribute effectively
to leveraging conservation action in Angola, attracting funders and the public
administration to initiatives with high visibility and meaningful impact. The conser-
vation of Angola’s remarkably rich biodiversity is first and last an Angolan respon-
sibility, to be led to success by Angolans.
References
Anderson JL, Morkel PV (2009) Parque Nacional da Quiçama. Status quo, as ameaças e a neces-
sidade de accões eficazes. Report to Ministry of Environment, Luanda, 24 pp
Andrade AA (1985) O Naturalista José de Anchieta. Instituto de Investigação Científica Tropical,
Lisboa, 187 pp
Angola. Science & conservation: A modern synthesis. Springer, Cham
Lages F, Ferrand N (eds) Biodiversity of Angola. Science & conservation: A modern synthesis.
Springer, Cham
Bell RHV (1982) The effect of soil nutrient availability on community structure in African eco-
systems. In: Huntley BJ, Walker BH (eds) Ecology of tropical savannas. Springer, Heidelberg,
pp 193–216
Bergman B, Verissimo L (2008) Avaliação do Estatuto de Áreas Protegidas do Sudeste do Kuando
Kubango,- Projecto Integrado de Gestão da Bacia Hidrográfica do rio Okavango. OKACOM-
USAID, 48 pp
Bersacola E, Svensson M, Bearder S et al (2014) Hunted in Angola. Surveying the Bushmeat trade.
SWARA, January–March 2014:31–36
Billes A, Fretey J, Verhage B et al (2006) First evidence of leatherback movement from Africa to
South America. Mar Turt Newsl 111:13–14
Science & conservation: A modern synthesis. Springer, Cham
Breytenbach J (2015) Eden’s Exiles. Protea Boekhuis, Pretoria, 306 pp
Carruthers J (2017) National park science. A century of research in South Africa. Cambridge
University Press, Cambridge, 512 pp
CBD (2010) Strategic plan for biodiversity 2011–2020 and the Aichi targets. Secretariat for the
Convention on Biodiversity, Montreal
Crawford-Cabral JC (1970) Alguns aspectos da ecologia da Palanca real. Bol Instituto de
Investigação Cientifica de Angola 7:7–42
Crawford-Cabral JC (1971) A Suricata do Iona, subspécie nova. Bol Instituto de Investigação
Cientifica de Angola 8:65–83
Dean WRJ (2000) The birds of Angola. An annotated checklist. British Ornithologists Union,
Tring, 433 pp
tion: A modern synthesis. Springer, Cham
Driver A, Sink KJ, Nel JL et al (2012) National Biodiversity Assessment 2011: an assessment
of South Africa’s biodiversity and ecosystems. Synthesis Report. South African National
Biodiversity Institute and Department of Environmental Affairs, Pretoria
Estes RD, Estes RK (1974) The biology and conservation of the Giant Sable Antelope Hippotragus
niger variani Thomas, 1916. Proc Acad Natl Sci Phila 126(7):73–104
Figueiredo E, Smith GF (2008) Plants of Angola/Plantas de Angola. Strelitzia 22:1–279
Frade F (1956) Reservas naturais de Angola – I. Alguns mamíferos da Reserva da Quiçama. Anais
Junta Invest Ultram 11(3):228–245
Frade F (1960) Os animais na etnologia ultramarina. Estudos, Ensaios e Documentos 84:211–240
GoA (Government of Angola) (1998) Lei de Bases do Ambiente. Ministry of Fisheries and
Environment, Luanda
GoA (Government of Angola) (2006) National biodiversity strategy and action plan. Ministry of
Urban Affairs and Environment, Luanda, 54 pp
de Angola (PLENARCA). Ministry of Environment, Luanda
GoA (Government of Angola) (2018) Plano Estratégico para o Sistema de Áreas de Conservação
de Angola (PESAC). Ministry of Environment, Luanda
Hall BP, Moreau RE (1962) The rare birds of Africa. Bull Brit Mus Nat Hist (Zool) 8:315–381
Holness S, Kirkman S, Samaai T et al (2014) Spatial biodiversity assessment and spatial man-
agement, including marine protected areas. Final report for the Benguela current commission
project BEH 09-01
Humbert H (1940) Zones e Étages de Végétation dans le Sud-Oest de l’Angola. C.R. somm. Scéanc
Soc Biogèogra 17:47–57
Huntley BJ (1971) Preliminary guide to the National parks and reserves of Angola. Report 3.
Repartição Tecnica da Fauna, Direcção Provincial dos Serviços de Veterinária, Luanda,
Mimeograph report, 17 pp
Huntley BJ (1972) A Plan for the future of the Giant Sable of Angola. Report 11. Repartição
Tecnica da Fauna, Direcção Provincial dos Serviços de Veterinária, Luanda, Mimeograph
report, 9 pp
Huntley BJ (1973) Distribution and status of the larger mammals of Angola, with particular ref-
erence to rare and endangered species. Report 21. Repartição Tecnica da Fauna, Direcção
Provincial dos Serviços de Veterinária, Luanda, Mimeograph report, 14 pp
Huntley BJ (1974a) Vegetation and Flora Conservation in Angola. Report 22. Repartição Tecnica
da Fauna, Direcção Provincial dos Serviços de Veterinária, Luanda, Mimeograph report, 13 pp
Huntley BJ (1974b) Iona national park: administration, management, research and tourism. Report
23. Repartição Tecnica da Fauna, Direcção Provincial dos Serviços de Veterinária, Luanda,
Mimeograph report, 19 pp, 5 Figs
Huntley BJ (1974c) Ecosystem conservation priorities in Angola. Report 28. Repartição Tecnica
da Fauna, Direcção Provincial dos Serviços de Veterinária, Luanda, Mimeograph report, 22 pp
Huntley BJ (1974d) Outlines of wildlife conservation in Angola. J S Afr Wildl Manag Assoc
4:157–166
Huntley BJ (2003) Quiçama national park. Integrated conservation management plan. Report to
Ministry of Urban Affairs and Environment, Luanda, 20 pp
Huntley BJ (2010) Estratégia de Expansão de Rede da Áreas Protegidas da Angola/proposals for
an Angolan protected area expansion strategy (APAES). Unpublished report to the Ministry of
Environment, Luanda, 28 pp, map
Huntley BJ (2017) Wildlife at war in Angola. The rise and fall of an African Eden. Protea Book
IUCN (1992) Environment status quo assessment report. IUCN Regional Office for Southern
Africa, Harare, 255 pp
IUCN (2018) The IUCN Red List of threatened species. Ver. 2017–3. http://www.iucnredlist.org
conservation: A modern synthesis. Springer, Cham
Kirkman SP, Yemane D, Kathena J et al (2013) Identifying and characterizing demersal fish biodi-
versity hotspots in the Benguela current large marine ecosystem: relevance in the light of global
changes. ICES J Mar Sci 70:943–954
Kirkman SP, Blamey L, Lamont T et al (2016) Spatial characterization of the Benguela ecosystem
for ecosystem-based management. Afr J Mar Sci 38:7–22
Margules CR, Pressey RI (2000) Systematic conservation planning. Nature 405:243–253
Martin E, Vigne L (2014) Luanda – the largest illegal ivory market in Africa. Pachyderm 55:30–37
N (eds) Biodiversity of Angola. Science & conservation: A modern synthesis. Springer, Cham
Mendelsohn JM, Mendelsohn S (2018) Sudoeste de Angola: um retrato da terra e da vida. South
West Angola: a portrait of land and life. Raison, Windhoek
Mendes L, Bivar-De-Sousa A, Williams M (2019) The butterflies and skippers of Angola. In:
Milliken T, Pole A, Huongo A (2006) No peace for elephants: unregulated domestic ivory mar-
kets in Angola and Mozambique. TRAFFIC online report series no. 11. Traffic East/Southern
Africa, Harare, Zimbabwe, 46 pp
Morais M (2008) Tartarugas Marinhas na Costa de Cabinda. Plano de conservação e gestão para a
implementação do projecto de prospecção sísmica “on shore”. Holisticos/Chevron, 67p
Morais M, Torres MOF, Martins MJ (2005) Análise da Biodiversidade Marinha e Costeira e
Identificação das Pressões de Origem Humana sobre os Ecossistemas Marinhos e Costeiros.
Ministerio do Urbanismo e Ambiente, Luanda, 140 pp
Mucina L, Rutherford MC (2006) The vegetation of South Africa, Lesotho and Swaziland.
Strelitizia 19:1–807
Myers N (1988) Threatened biotas: ‘hotspots’ in tropical forests. Environmentalist 8:120
Myers N, Mittermeier RA, Mittermeier CG et al (2000) Biodiversity hotspots for conservation
priorities. Nature 403:853–858
Newton da Silva S (1952) Wild life and its protection in Angola. Oryx 1:7
Newton da Silva S (1970) A Grande Fauna Selvagem de Angola. Serviços de Veterinária, Luanda,
151 pp
NGOWP (2018) National Geographic Okavango wilderness project. Initial findings from explora-
tion of the upper catchments of the Cuito, Cuanavale and Cuando Rivers in Central and South-
Eastern Angola (May 2015 to December 2016). National Geographic Okavango Wilderness
Project, 352 pp
Peace Parks Foundation (2016) Annual review 2016. Peace Parks Foundation, Stellenbosch
Pinto AA d R (1983) Ornitologia de Angola, vol 1. Instituto de Investigação Científica Tropical,
Lisbon, 696 pp
Projecto Kitabanga (2017) Projecto Kitabanga – Conservação de tartarugas marinhas – Folhetos
2017. Universidade Agostinho Neto/Faculdade de Ciências, Luanda
Raimondo D, von Staden L, Foden W et al (2009) Red list of South African plants 2009. Strelitzia
25:1–668
Rejmánek M, Huntley BJ, le Roux JJ et al (2017) A rapid survey of the invasive plant species in
Russo V, Huntley BJ, Lages F, Ferrand N (2019) Conclusions: biodiversity research and conserva-
tion opportunities. In: Huntley BJ, Russo V, Lages F, Ferrand N (eds) Biodiversity of Angola.
Science & Conservation: a modern synthesis. Springer, Cham
Schlossberg S, Chase MJ, Griffin CR (2018) Poaching and human encroachment reverse recov-
ery of African savannah elephants in south-East Angola despite 14 years of peace. PLoS One
13(3):e0193469. https://doi.org/10.1371/journal.pone.0193469
(eds) Biodiversity of Angola. Science & conservation: A modern synthesis. Springer, Cham
Stassen N (ed) (2010) Reminiscences concerning the life of William James Bushnell Chapman &
An account of the entry of the Trek Boers into Angola and of their sojourn during the forty-
eight years they struggled in that country under Portuguese rule. Protea Book House, Pretoria,
476 pp
Stassen N (2016) The Thirstland Trek, 1874–1881. Protea Book House, Pretoria, 73 pp
Svensson MS, Bersacola E, Bearder SK et al (2014) Open sale of elephant ivory in Luanda,
Angola. Oryx 48:13–14
Teixeira JB (1968) Parque Nacional do Bicuar. Carta da vegetação (1a aproximação) e memória
descritiva. Instituto de Investigação Agronómica de Angola, Nova Lisboa
Teixeira JB, Matos GC, Sousa JNB (1967) Parque Nacional da Quiçama. Carta da vegetação e
memória descritiva. Instituto de Investigação Agronómica de Angola, Nova Lisboa
Varian HF (1953) Some African milestones. Books of Rhodesia, Bulawayo, 78 pp
and conservation. PhD thesis. University of Porto, Porto
Vaz Pinto P (2019) The Giant Sable Antelope: Angola’s national icon. In: Huntley BJ, Russo V,
Lages F, Ferrand N (eds) Biodiversity of Angola. Science & conservation: A modern synthesis.
Springer, Cham
Walker JF (2004) A certain curve of horn. The hundred-year quest for the Giant Sable Antelope of
Angola. Grove/Atlantic Inc., New York, 514 pp
Walker JF (2009) Ivory’s ghosts. The white gold of history and the fate of elephants. Atlantic
Monthly Press, New York, 296 pp
Weir CR, Ron T, Morais M (2007) Nesting and at-sea distribution of marine turtles in Angola, West
Africa, 2000–2006: occurrence, threats and conservation implications. Oryx 41(2):224–231
Chapter 19
Museum and Herbarium Collections
for Biodiversity Research in Angola
Rui Figueira and Fernanda Lages
Abstract The importance of museum and herbarium collections is especially great

in biodiverse countries such as Angola, an importance as great as the challenges
facing the effective and sustained management of such facilities. The interface that
Angola represents between tropical humid climates and semi-desert and desert
regions creates conditions for diverse habitats with many rare and endemic species.
Museum and herbarium collections are essential foundations for scientific studies,
providing references for identifying the components of this diversity, as well as
serving as repositories of material for future study. In this review we summarise the
history and current status of museum and herbarium collections in Angola and of
information on the specimens from Angola in foreign collections. Finally, we pro-
vide examples of the uses of museum and herbarium collections, as well as a road-
map towards strengthening the role of collections in biodiversity knowledge
generation.
Keywords Bioinformatics · Catalogue of life · Checklists · Conservation ·

Expeditions · GBIF · Natural history collections
R. Figueira (*)
e-mail: ruifigueira@isa.ulisboa.pt
F. Lages
ISCED – Instituto Superior de Ciências da Educação da Huíla, Lubango, Angola
e-mail: f_lages@yahoo.com.br

https://doi.org/10.1007/978-3-030-03083-4_19
514 R. Figueira and F. Lages
Introduction
Natural History Collections (NHCs) are the basic building blocks for the discovery
and understanding of the diversity of life. Scientific names are currently available
for about two million of the eight million species estimated to live on Earth (Mora
et al. 2011). In museum and herbarium collections, researchers try to compile and
organise the most complete representation of biological diversity. This is motivated
not (only) because some people have a:
metaphysical angst, perhaps because they cannot bear the idea of chaos being the one ruler
of the universe, which is why (…) they attempt to impose some order on the world…
as surmised by the Nobel literature prize winner José Saramago in his book ‘All the
Names’ (Saramago 2000) but also by the need to have reference samples to identify
species, know how and where they live, their biotic and abiotic interactions, their
links to communities and ecosystems, and finally, all subjects that define natural
history (Tewksbury et al. 2014).
These needs are met by specimens serving as vouchers of a species’ occurrence,
collected in a specific habitat and in certain circumstances of time, space, traits and
sampling methods. Globally, it is estimated that biological collections contain three
billion preserved specimens (Brooke 2000; Wheeler et al. 2012). Specimens main-
tained in biological collections include the material samples on which new species
are described – the type specimens – but also additional specimens that represent
the variety and variability that a biologist needs to recognise to become a good prac-
titioner of species identification. Collections are essential for taxonomic and sys-
tematic research, but also for studies in ecology, evolution, biogeography,
conservation, climate change effects, and other fields, as will be discussed later in
this chapter.
Building an inventory of the biodiversity of Angola, as a national checklist,
begins with an initial register of species present in biological collections in Angola
and abroad. With fifteen ecoregions (Olson et al. 2001; Burgess et al. 2004; Huntley
2019), Angola is one of the most biodiverse countries of the world, so biological
collections will or should reflect that diversity. The number of endemic species is
recognised to be high in several groups, for example in birds (Mills and Melo 2013;
Dean et al. 2019) and plants (Figueiredo et al. 2009a, b; Goyder and Gonçalves
2019). However, despite this richness, the Angolan Escarpment could not be recog-
nised as one of the biodiversity hotspots of the world due to the lack of information
on its species diversity (Myers et al. 2000). This might be a consequence of the
under-representation of Angola’s biological diversity in natural history collections.
This situation results, in part at least, from a combination of factors like the restric-
tions placed on field explorations due to the war situation in the country over several
decades, and to limited access to the country’s natural history collections such as
those of the Museu do Dundo.
It is possible, nevertheless, to redress this situation. The pressure on biological
collections has been very high for the last two decades due to cuts in budgetary
19 Museum and Herbarium Collections for Biodiversity Research in Angola 515
support. For example, in the USA, 100 herbaria have closed since 1997 (Deng
2015). Paradoxically, the use of such collections has increased at an exponential rate
in the same period (Pyke and Ehrlich 2010; Lavoie 2013). This increase in use
might be related to the fact that biodiversity informatics and cyber-infrastructure
developments now contribute to compress time and space, facilitating virtual access
to specimens, data and literature. A researcher of Angolan biodiversity, working in
Angola or working abroad, can now access hundreds of thousands of biodiversity
records online. Such databases include images of the specimens in collections
hosted elsewhere in the world, and species descriptions in old or inaccessible jour-
nals. In parallel, molecular tools have recently seen great advances, with the arrival
of next-generation sequencing tools that promise to overcome limitations of DNA
fragmentation caused by certain preservation methods used in collections (Yeates
et al. 2016). This will facilitate attaching genetic sequences to specimens, and sup-
port biodiversity field surveys to achieve faster results. An optimistic view is held by
a group of experts who suggest that it is possible to describe ten million species in
50 years, virtually describing all species that currently remain to be discovered
(Wheeler et al. 2012). The authors conclude that this goal might be reached by inter-
disciplinary partnerships using and developing cyber technologies.
This background creates a favourable environment for the development and the
increase of the role of museum and herbarium collections to support the advance of
biodiversity knowledge globally, but also in Angola. In this chapter, we will provide
a short review of the current status and knowledge of Angolan biodiversity based on
these collections, their importance for biodiversity research, and some indications
on how biodiversity informatics and cyber-infrastructure could facilitate their use in
biodiversity knowledge generation.
Museum and Herbarium Collections from Angola
Internet access to information about species from Angola represented in museum

and herbarium collections around the world is now possible. This has been facili-
tated by global networks and infrastructure resulting from several initiatives, based
on biodiversity informatics standards, protocols, tools, manuals and quality control
procedures that, being interoperable, create a digital global biodiversity observa-
tory. The most visible facility is probably the intergovernmental Global Biodiversity
Information Facility (GBIF), created in 2001, through which more than one thou-
sand institutions share, in 2018, over one billion records, including 145 million
records based on specimens preserved in collections. These records are freely and
openly accessible to all using an Internet browser, at www.gbif.org. However, we
are still far from having all specimens from collections catalogued in databases. In
Europe, for example, only 10% of NHC specimens are digitally catalogued. But
these inventories provide a good basis for knowing what expeditions and studies
have contributed specimens of the different biological groups to museum and her-
barium collections.
Most of the world’s’ biodiversity is located in the tropics. But the NHCs of
Europe and North America hold the largest collections of material from the tropics,
not the NHCs of the countries of origin (Peterson et al. 2016). For historical reasons,
many of the best collections of African countries are not located in the country of
origin, but in the former colonising countries, or in other countries that conducted
field expeditions in Africa. This is the case of Angola, where the highest representa-
tion of Angolan biodiversity in collections is in Portuguese or other European or
North American NHCs. For this reason, the repatriation of information that can be
achieved through GBIF, where data is mobilised and made available to the country
of origin, represents an important asset to assist biodiversity research and conserva-
tion in Angola.
The representation in NHCs of Angola’s biodiversity varies between taxonomic
groups, regions and time periods, and depends on the history of expeditions and
studies conducted in the country through the last 150 years. To provide an overview
of the museum and herbarium specimens collected in Angola, we compiled a data-
set with information from several sources. We used the full dataset available through
GBIF as on May 25, 2018 (GBIF.org 2018), containing 149,701 records for all
groups. This was merged with other sources for specific groups, information that is
not yet published by GBIF. These datasets include the database of the bird collec-
tions of the Herbarium and Museum of Ornithology and Mammalogy of Lubango
(Lages 2016, pers. com.) containing 34,471 records, and the Herbarium of the
University of Coimbra (Santos and Sales 2018), with 7864 herbarium records. Both
resources are soon to be published through GBIF. Finally, we accessed RAINBIO
(Gilles et al. 2016) that holds 1884 herbarium records from Angola. In the aggre-
gated dataset, the possible duplication of records between RAINBIO and GBIF was
checked and removed, as well as a careful check of the information about province,
which was standardised or completed whenever possible. The total aggregated data-
set for this analysis contains 193,839 records, of which 158,185 records contain
information about the province and 154,631 records with information about the
sampling year (Table 19.1). These records are published by more than 200 institu-
tions from 28 countries (Fig. 19.1), and should be considered a partial view of the
complete holding of specimens from Angola in collections worldwide.
The oldest specimens from Angola known in museums date from the late seven-
teenth and beginning of the eighteenth centuries. These are of plants included in the
herbarium collection of the Natural History Museum, London and include 36 speci-
mens collected in the region of Luanda by Mason in 1669 (Romeiras 1999; Goyder
and Gonçalves 2019), followed by samples collected by John Kirckwood in
Cabinda. The first records available through GBIF for Angola are from 1758, of
mussels from the Malacology collection of the Balley-Matthews National Shell
Museum, in the USA, which includes 70 specimens collected up to the end of the
eighteenth century.
The time profile of the specimens collected in Angola (Fig. 19.2) only shows
three mussel records from before 1800, presented as small peaks. The first records
of the nineteenth century also create a small peak in 1804, based on material col-
lected during the Viagens Philosophicas. These expeditions were organised by
Table 19.1 Data records of specimens from Angola in collections available in online resources, as
of May 2018
Source Reference Collection type N° of records N° of types
GBIFa GBIF.org (2018) Herbarium 85,360 8877
Fungi 601 283
Mammals 4641 39
Birds 58,821 187
Herpetological 7269 269
Fish 9227 569
Arthropods 11,480 1518
Invertebrates 5425 657
Microorganisms 220 10
Not classified 1046
TOTAL 148,573 12,409
ISCEDb Lages (2016, pers. com.) Bird 34,471
Coimbrac Santos and Sales (2018) Herbarium 7864 634
RAINBIO Gilles et al. (2016) Herbarium 1884 Not available
a
Global Biodiversity Information Facility
b
Museum of Ornithology and Mammalogy, ISCED-Huíla, Lubango
c
Herbarium of the University of Coimbra
Fig. 19.1 Hosting country of specimens from Angola in collections available in online resources,
as of May 2018. The size of the pie relates to the number of records published by each country,
decreasing in logarithmic basis. PRT Portugal, AGO Angola, USA United States, GRB United
Kingdom, ZAF South Africa, BEL Belgium, FRA France, NLD The Netherlands, SWE Sweden,
DEU Germany
Portugal to explore the former Portuguese overseas territories of Brazil, Goa, Cape
Verde, Mozambique and Angola. The naturalist Joaquim José da Silva was in charge
of sampling plants and animals from Angola, and stayed in the country between
1783 and 1808. The materials collected were sent to Lisbon. However, during the
Fig. 19.2 The time profile of the specimens collected in Angola. The horizontal bars for each col-
lection type, indicated on the right axis, indicates presence of specimens of that collection type in
the corresponding period
French invasion of Portugal, these specimens were removed to the MNHN in Paris
by Saint-Hillaire in 1808 (Barbosa du Bocage 1862 in Alves et al. 2014).
In the time profile (Fig. 19.2), the first significantly high value is visible in the
beginning of the second half of the nineteenth century, when the Austrian botanist
Friedrich Welwitsch was commissioned by the Portuguese government to explore
the flora of Angola. In the expedition named Iter Angolense, he sampled more than
10,000 specimens (Albuquerque et al. 2009), between 1853 and 1860, of which
more than 1000 were used to describe new species. Several sets of the collection
were made by Welwitsch and distributed to several herbaria, but the most complete
sets are located in Lisbon, at the LISU herbarium, and in the BM herbarium, in
London. In total, more than 20,000 duplicates were sent to the major herbaria in
Europe (Albuquerque and Correia 2010). Welwitsch sampled not only vascular
plants but also cryptogams, including 350 lichen specimens with 50 type speci-
mens, and also mammals. The most well-known and notable species he found is
Welwitschia mirabilis from the Namib Desert, in southwest Angola. The genus was
named in Welwitsch’s honour by Sir Joseph Hooker, and is the most iconic plant
species of Angola.
Other expeditions contributed to diversify, in terms of biological groups, the
addition of specimens to collections. These collections are detailed in other chapters
of this volume (Baptista et al. 2019; Beja et al. 2019; Branch et al. 2019; Dean et al.
2019; Kipping et al. 2019; Mendes et al. 2019; Skelton 2019). The institutions to
which foreign collectors sent material are listed in Appendix. The Portuguese natu-
ralist José Anchieta collected plants and animals of several groups (birds, reptiles
and amphibians, mammals, fishes) between 1850 and 1897 (Albuquerque and
Correia 2010; Mills et al. 2010; Alves et al. 2014; Ceríaco 2014). Other main
collectors in that period are the German botanist Hugo Baum, with specimens
deposited at the Berlin Herbarium, and which expedition was reviewed by Figueiredo
et al. (2009a, b). The Portuguese explorers Hermenegildo Capelo and Roberto Ivens
contributed with plants specimens to the LISU herbarium (University of Lisbon),
the German botanist Alexandre von Mechow to the Berlin herbarium, the American
naturalist William H Brown, with birds, mammals and fishes to the collections of
the NMNH, Smithsonian Institute. Several bird collectors also promoted expedi-
tions, like Axel W Eriksson (Vänersborg Museum, Sweden), the French ornitholo-
gists Albert Lucan and Louis Petit (NHM, London), and P van Kellen (Naturalis,
The Netherlands). These naturalists also collected specimens of biological groups
other than their main field of interest, like butterflies, bees, hemipterans, etc.
During the twentieth century, a crescendo of the number of specimens added to
collections was observed continuously until 1957 and remained high until a drop in
1975. After this year, with the start of the civil war that lasted for 27 years, very few
specimens were added to collections. Finally, after the year 2000, with the end of
the civil war (in 2002), there is a recovery in the deposition of specimens into col-
lections, but not to the levels observed before 1975. However, for the recent period
we need to consider the time lags between the end of expeditions, the deposition of
specimens in collections and the making of data accessible through databases.
Currently, the average time gap between specimens being collected and identified is
21 years (Fontaine et al. 2012). Simultaneously, a change in sampling ethics and
tight permit issuing control by national authorities might also explain lower sam-
pling rates per expedition (Prathapan et al. 2018).
In the twentieth century, a series of large expeditions to Angola increased knowl-
edge about the flora and fauna of the country. In terms of plants, the largest plant
collections were those of the botanist John Gossweiler, sampling in all provinces of
the country, with a total of 14,600 numbers, between years 1900 and 1950. The most
complete set of this collector is deposited in the LISC herbarium, at the University
of Lisbon, but many duplicate specimens were distributed to other herbaria, namely
COI, BM, LISU, P, K, LUA. All herbarium acronyms are according to the Index
Herbariorum (Thiers 2018). The second most prolific collector was JM Brito
Teixeira, a follower of Gossweiler who collected about 13,000 numbers in all prov-
inces between 1949 and 1969. Several botanical expeditions were organised from
Portugal, either with a focus in botany, agronomy or forestry, while others were
promoted by institutions newly created in Angola. The Instituto de Investigação
Científica de Angola (IICA) was created in 1958, with herbarium and zoological
collections based in Lubango. Another research institution was the Instituto de
Investigação Agronómica, established in 1961, based in Huambo. Frequently, staff
from institutions from both Portugal and Angola worked together in field expedi-
tions, because, formally, the new institutions in Angola were dependencies of the
equivalent Portuguese institutions. Therefore, duplicate samples were distributed to
herbaria in Angola (LUBA or LUAI).
The main collectors were Luiz Carrisso (based in COI), Francisco de Ascensão
Mendonça (LISC), Francisco de Sousa (LISC), Eduardo Mendes (LISC), Romeu
Santos (LUBA), Óscar Azancot de Menezes, Carlos Henriques, Luís
randvaux-Barbosa (LISC) and the British botanist Arthur W Exell (BM). A con-
G
siderable set of specimens without the individual collector being indicated were
collected in the scope of the Missão de Estudos Florestais a Angola (MEFA),
between 1957 and 1960. The last large plant collectors in the twentieth century
sampled between 1970 and 1974 were António RF Raimundo, Gilberto Cardoso de
Matos, Paul Bamps, Roger Dechamps and Eurico S. Martins. A detailed list of col-
lectors, including the time range of the collections and provinces is available in
Figueiredo and Smith (2008).
Concerning animal collections, the largest collections created or with the largest
growth in the twentieth century are of birds. In this case, the ornithological collec-
tion of the Museum and Herbarium of Lubango is by far the largest and most repre-
sentative of Angola, with circa 40,000 specimens, and probably one of the largest
collections of birds based in Africa. The oldest specimens in the collection are from
1948, but the relevant sampling started in 1958, the year of the creation of IICA. The
collection was established by António da Rosa Pinto, with many staff of IICA con-
tributing significantly with specimens: at least 13 people each added more than 500
bird specimens. The collection contains specimens from throughout the country, but
75% of the specimens are from the western and southern provinces, with 25% from
Huíla where the collection is based (Lubango).
Dean et al. (2019) presents details on the ornithological collectors of Angola. In
the first half of the century, about 13,000 specimen records can be found through
GBIF, the main publishing institutions being AMNH, NHMUK, CM, FMNH and
GNM. The main collectors are WJ Ansorge, R Boulton, H Lynes, CH Pemberton
and the main provinces with records are Cuanza-Norte, Bengo, Malanje, Benguela,
Namibe, and Bié. Between 1950 and 1974, apart from IICA staff, the main collec-
tors were Gerd H Heinrich and T Archer, which collected about 900 specimens held
by YPM and USNM collections.
The history of mammal collecting in Angola is presented by Beja et al. (2019).
In the twentieth century, the year 1925 presents an exceptionally high number of
records of about 1400 specimens in mammal collections. This corresponds to the
Arthur Vernay expedition to Angola, which specimens are in the AMNH collection.
The following years with high values are 1932 and 1933, corresponding to the
Phipps-Bradley Expedition, specimens also at the AMNH collection. In 1936, KH
Prior sampled in Benguela, which specimens are located at the collection MVZ. In
1954 and 1955, GH Heinrich sampled in several provinces, which materials are held
at FMNH. Considering the collections with the highest number of records available
through GBIF, in general, the order with the highest number of specimens is
Rodentia.
The number of records for mammals is in general low in the dataset compiled. A
reason for this might be the incomplete status of digitisation of mammal collections
in databases. This means that the values mentioned might not be representative of
the total holdings of mammals from Angola. For example, the mammal collection
from Lubango was not taken into account in this analysis, because the digitising
process is still ongoing, in terms of data quality verification and import to a data
management system. For the same reason, it is possible that in other collection
types, the number of specimen records available online is also not representative of
the true sampling effort for these collections. For example, in the dataset used in this
chapter, no records are included from collections based in Portuguese institutions
for fish or butterflies, although several zoological expeditions were organised by the
Instituto de Investigação Científica Tropical (IICT) to Angola, since 1950. This is
the case of ichthyological expeditions to lakes Cameia and Dilolo, by Fernando
Frade and Teixeira Pinto, in 1958. Another example of a collection that still needs to
be mobilised (although already studied) is the Lepidoptera order of the Entomological
collection of IICT. This collection was extensively studied in the preparation of the
book Butterflies of Angola (Mendes et al. 2013), with more than 15,000 specimens
reviewed. However, the records of these specimens are not yet accessible.
In a related subject on collection accessibility, it should be noted that the Instituto
de Investigação Científica Tropical (IICT) was integrated in the University of
Lisbon in July 2015, as a special unit. This will not, however, change the possibility
of accession to collections, except for the period while the collections are being
moved. The new unit in the University shares the Director with the National Museum
of Natural History and Science of the University of Lisbon, but all the zoological
and herbarium collections of IICT will be retained as distinct collections. This is
relevant for the study of the biodiversity of Angola because these collections are
important, not only because they host many type specimens, but also because some
are the most representative worldwide of the biodiversity of Angola. For example,
the LISC herbarium of IICT has ca. 70,000 specimens, which is the largest world-
wide for Angola, because it merged duplicate specimens from several expeditions,
while in Angola these were hosted in separate herbaria (LUBA, LUAI, LUA).
The specimen-based atlas of the butterflies of Angola highlighted the importance
of access to valuable but privately owned collections. In that example, four private
collections were consulted.
The specimens collected in Angola are not evenly distributed across the country,
as it is often observed in natural history collections (Lavoie 2013; see also Dean
et al. 2019; Mendes et al. 2019; Beja et al. 2019). The bias is reflected both in the
spatial coverage of the collections, as well as in the groups’ representativeness
across the country (Fig. 19.3). Some regions of Angola are clearly under-represented
in collections, as is the case of the provinces of Zaire and Uíge, in the northwest, and
most of the eastern provinces, including Lunda-Norte, Lunda-Sul, Moxico and
Cuando Cubango. According to Crawford-Cabral (2010), there is a triangle that
extends from Bié northeast to Lunda-Sul and southeast to the Cuando River, where
there is a serious lack of knowledge about the fauna. This includes the interesting
areas of the Upper Zambezi. The province of Huíla stands out has having almost
twice the number of specimens compared with the second province, Namibe. This
is possibly a result of the establishment, in Huíla, of the collections and research
staff of IICA, which impact is also noted in its neighbouring provinces.
In most provinces, plant collections outnumber other taxa, but in four – Benguela,
Cuanza-Sul, Malanje, Namibe – the number of birds in collections exceeds plants.
The number of mammals in collections has some expression in Bié, Benguela,
Cuanza-Sul, Huíla and Malanje. As for fish collections, these are more present for
Fig. 19.3 Number of records per province, with darker colours corresponding to higher numbers.
Each pie chart depicts the breakdown per collection type, and displays the number of records for
the province. The records of Luanda and Bengo were aggregated in one chart. Province names: Bo
Bengo, BE Benguela, BI Bié, CA Cabinda, CC Cuando Cubango, CN Cuanza-Norte, CS Cuanza-
Sul, Cu Cunene, HA Huambo, HI Huíla, LA Luanda, LN Lunda-Norte, LS Lunda-Sul, MA Malanje,
MO Moxico, NA Namibe, UI Uíge, ZA Zaire
the areas of Zambezi River, in Moxico, and in the Lundas. In this last region, we find
an important representation of arthropods in collections, which might be a result of
the activity of the Museu do Dundo, created in 1942. The museum included the
Laboratory of Biology, where A Barros de Machado and E Luna de Carvalho
established numerous international connections with specialists, with exchange of
specimens with other collections.
Current Status of the Natural History Collections in Angola
The landscape of museum and herbarium collections in Angola is considerably

diverse in terms of institutional governance. Although all the hosting institutions are
public entities, they depend on different government ministries, which implies dif-
ferent priorities and funding programmes. As a result, it has been difficult to develop
a common strategy for the development and use of NHCs in Angola. Currently, the
different institutions have different capacities and dimensions. Most are still inac-
tive or starting their activities, mainly by performing inventories, digitising and sys-
tematising information on species.
Research activities are also recent and have relied on international collaboration
projects such as the Southern African Botanical Diversity Network (SABONET),
the Angolan Biodiversity Assessment and Capacity Building Project, the African
Plants Initiative (API), the Southern African Science Service Centre for Climate
Change (SASSCAL), the Future Okavango (TFO), and the National Geographic
Okavango Wilderness Project, among others. These funding opportunities allowed
the support of the cooperation with relevant international institutions (Royal Botanic
Gardens, Kew and the South African National Biodiversity Institute – SANBI),
enabling the rehabilitation of some of the country’s collections and the training of
qualified personnel for their management, expansion and valorisation. In this con-
text, since 2013, Angola has benefited from the GBIF training initiatives by the
Portuguese node, for computerisation and publication of biodiversity data.
Another cooperation programme has also recently contributed to the advance of
data mobilisation and capacity enhancement in Angola. Within the scope of the
program Biodiversity Information for Development (BID), managed by GBIF with
funds from the European Commission, Angola was granted a national project, led
by SASSCAL, which started in 2016. Apart from the data mobilisation activities,
some of which are based on collections, the project will enable Angola to participate
in training workshops promoted by GBIF and other partners on biodiversity data
publication, data quality, and data use.
Active Collections
In the scope of this chapter, we consider active collections those that are supporting
or developing research activities, contributing to the increase of the value of the col-
lection, by means of new additions of biological specimens, or valorisation through
taxonomic revision, digitisation, and use by researchers on-site and online.
Herbarium LUBA, Instituto Superior de Ciências de Educação da Huíla

(ISCED), Lubango
Founded in 1958 with the creation of the Instituto de Investigação Científica de

Angola (IICA), the collection currently houses 15,902 plant specimens belonging to
202 families and 3520 species. The main collectors were G Barbosa, A de Menezes,
R Santos, R Correia and JM Daniel. After independence, approximately half of the
collection was transferred to Luanda, becoming part of the LUAI herbarium.
Currently, the database of the collection is being prepared, and 200 specimens are
available in high resolution through the Global Plants repository at http://plants.
jstor.org.
Herbarium LUAI, Universidade Agostinho Neto, Luanda
The Luanda Herbarium incorporated part of the LUBA Herbarium, which was par-
tially transferred to Luanda to be hosted at the National Centre for Scientific
Research of the University of Agostinho Neto (Martins and Martins 2002). LUAI
currently houses about 35,000 botanical specimens representing approximately
5000 species. The main collectors were A de Menezes, M Batalha, JM Daniel, M
Lopes, R Santos, B Sousa and F Sousa. From 1995 until 2007, this herbarium
housed 45,000 botanical samples of the LUA Herbarium. The herbarium currently
has the digital infrastructure for databasing, but the pace of cataloguing has been
slow.
Herbarium LUA, Instituto de Investigação Agrária (IIA), Ministry

of Agriculture, Huambo
LUA was the first herbarium created in Angola, in Huambo Province, in 1941
(Martins and Martins 2002). Its collection includes about 40,000 specimens. The
main contributors were G Barbosa, J Gossweiler, C de Matos, OA Leistner, EJ
Mendes, FA Mendonça, R Monteiro and F Murta. There is a digitisation programme
in preparation, within the scope of the National Project of the BID programme.
ird and Mammal Collections, Instituto Superior de Ciências de Educação

B
da Huíla (ISCED), Lubango
These collections were created as an IICA section installed in Lubango in the late
1950s. The first record dates from 1958, and until 1975 about 40,000 specimens
were incorporated. The bird collection consists of 34,471 skins, as well as eggs, nests
and embryos, distributed across 26 orders, 84 families and 305 genera. It is worth
noting the contribution of António Rosa Pinto, representing 21% of the specimens of
the collection sampled between 1958 and 1972 throughout the country (see map in
Dean et al. 2019). The collection of mammals consists of 4299 skins of 157 species
distributed by 11 Orders, 56 Families and 103 Genera, and an unregistered number
of skeletons and skulls. The temporal coverage is between 1960 and 1978, and the
main collectors were J Crawford Cabral, AP Simões, C Simões and E Epalanga. The
full bird collection and the group of chiroptera within mammals, with about 300
specimens, are on database, and will be published through GBIF.
Herpetological Collection, Instituto Superior de Ciências de Educação da

Huíla (ISCED), Lubango
This is the first herpetological collection created after 1975, as an output of a study
included in the SASSCAL project, under the responsibility of Ninda Baptista. It
houses 1081 specimens of reptiles and amphibians (approx. 30–70% respectively),
preserved in alcohol, as well tadpoles, eggs and tissue samples. Two collectors –
Ninda Baptista and Pedro Vaz Pinto are important contributors to this collection.
Entomological Collection, Instituto de Investigação Agrária (IIA), Ministry

of Agriculture, Huambo
Based on registry books, the collection contains the 44,884 specimens. There is also
a digitisation programme for this collection, with the Odonata (1006 records)
already digitised and published through GBIF (Cassinda et al. 2018), and with other
orders to follow.
nake Collection of the Research and Information for Drug and Toxicology

S
Center (CIMETOX), Malanje
A collection of snakes was recently created in Malanje, at the Research and

Information for Drug and Toxicology Center of the Medical Faculty, in Malanje
(Oliveira et al. 2016). The number of specimens is not available at this stage.
Inactive Collections
oological Collection of Dundo, Museu do Dundo, Ministry of Culture,

Z
Dundo
Although the creation of the Museu do Dundo dates from 1942, the zoological col-
lection began in 1936 (Machado 1952). This museum is best known for its valuable
ethnographic collection, so at the time of its restoration at the beginning of this
century, biological collections were not covered and some of them are in danger of
deterioration. According to EC Afonso, curator of the Biological section in the

1980s, the Museum houses about 50,000 specimens of mammals, fishes, reptiles,
amphibians and insects, the latter being the largest collection with about 30,000
specimens. Due to its value, we expect that in the future this collection can be
studied and restored. The museum originally also had a herbarium (DIA), that no
longer exists.
Museu Nacional de História Natural, Ministry of Culture, Luanda
The museum was created in 1938, and moved to the current location in 1956. The
museum holds mounted specimens of mammals, fishes, birds, reptiles and insects.
However, it was not possible to determine if there is an active collection, the number
of specimens, and their value for scientific research. There is no insect collection.
This museum is currently responsible for the management of Museu do Dundo.
Current and Potential Uses of Biological Collections
Museum and herbarium collections are examples where the whole is greater than
the sum of the parts. Each specimen, as a voucher of a species found in nature, car-
ries biological data (in its genes, tissues, traits, biochemistry) and metadata (on its
label or attached field notes) providing contextual information on the location, date,
habitat and ecology of that specimen. But from a set of specimens that make a col-
lection, it is possible to do comparisons, grouping and separating them by their
features, in what, ultimately, leads to the description or identification of a species.
The possibility of making and analysing comparisons between specimens is funda-
mental to our developing the knowledge of what the species is, and therefore, our
understanding of biodiversity. There is great value in having a collection. Because
of this, we need to resist the fate predicted for collections in the previous quote from
Saramago, that in the continuation of the text, says:
… and for a short while they manage [to impose some order on the world], but only as long
as they are there to defend their collection, because when the day comes when it must be
dispersed, and that day always comes, either with their death or when the collector grows
weary, everything goes back to its beginnings, everything returns to chaos.
We would lose too much in letting everything return to chaos, we cannot afford it.
Without a doubt, there is a cost in maintaining a collection. Several reports have
been published about the termination of collections due to budget restrictions, either
by closing doors, restricting or diverting staff to other tasks, or aggregating collec-
tions in large facilities (Gropp 2003; Deng 2015; Kemp 2015). For example, in Italy,
by 2014, it was estimated that one-third of the biological specimens were lost
through lack of preservation or bad practice (Nature Editorial 2014), and in the USA
100 herbaria have been closed since 1997 (Deng 2015). Discarding a collection
brings with it the loss of investments in field expeditions, the costs of subsequent
preservation of specimens across decades or centuries, and even the fact that many
species may not be found anymore in the original collecting sites, due to the loss of
habitats or restrictions in sampling because of conservation or ethical reasons. As an
indicator, insurance companies in Norway value herbarium specimens at €21 each
(Hannu Saarenmaa 2017, pers. com.). Here we will focus on the uses of collections
and how they underpin scientific research, biodiversity conservation, food security,
and other societal and economic benefits. Several reviews have discussed these
uses, providing examples (Brooke 2000; Suarez and Tsutsui 2004; Tewksbury et al.
2014; Rocha et al. 2014). It is worth mentioning that, with the development of labo-
ratory methods, technology and other tools still to be invented, there are potential
applications of the collections in the future that we cannot foresee.
Preserving and Documenting Biodiversity
The most fundamental use of museum and herbarium collections is to support tax-
onomy and systematics, serving as references for species description, identification,
and naming of species. One of the most important roles of collections is to preserve
the physical specimens that served as samples for the formal scientific description
of a new species for science. These specimens, often more than one, are called
types. One of these specimens is usually designated as the holotype by the author of
the species – the one chosen to be most representative of the characteristics of the
species – but duplicates can also be mentioned in the publication of the species, and
distributed to other collections. This distribution is important for security reasons,
to ensure that if the holotype is lost because of an unfortunate event, other speci-
mens that were used the initial description of the species are preserved. In 2017, a
package with specimens sent by the National Museum of Natural History in Paris to
the Queensland Herbarium in Australia, including some type specimens, was incin-
erated by Australian customs officers (Stokstad 2017).
Types are, therefore, special specimens, so their management is undertaken with
extreme care. Digitisation programmes normally prioritise these specimens, to pro-
vide digital preservation and alternative access to the specimens via the internet. In
the case of plant type specimens, the Global Plants Initiative framed this task, with
support of the Andrew W Mellon Foundation. The repository Global Plants (http://
plants.jstor.org) aggregates and provides access to more than two million high reso-
lution images of types, including 3461 images of type specimens collected in
Angola. The total number of types from Angola, from all groups, reported through
GBIF and Global Plants is 6983 (Table 19.2).
Museum and herbarium collections serve not only to preserve types and other
specimens already identified, but also organisms yet to be identified. In fact, many
specimens remain unidentified for several years, either because there is no capacity
for their immediate processing after being collected, or its identification represents
taxonomic challenges, sometimes at the level of the description of a new species for
Table 19.2 Number of type specimens from Angola in NHC collections worldwide
Collection type Holotypes Other types Total per collection
Plants 1236 2225 3461
Fungi 24 259 283
Mammals 25 14 39
Birds 108 79 187
Fish 82 487 569
Herpetological 40 229 269
Arthropods 300 1218 1518
Invertebrates 146 511 657
Microorganisms 10 10
Total 1961 5022 6983
The sources of the data are Global Plants (http://plants.jstor.org), for plant specimens, and GBIF
(GBIF.org 2018), for other collection types
science. In plants, only 16% of newly collected species are described within the first
5 years after being collected, and approximately 25% of new species are described
using specimens more than 50 years after their collection (Bebber et al. 2010), and
in animals, the situation is likely to be similar (Kemp 2015).
Collections are the main source for documenting diversity not only between spe-
cies but also within species. The majority of specimens in collections are not types,
but regular specimens sampled at a certain date and location, by one or more collec-
tors, and representing a species. These specimens and their associated information
represent what we call primary biodiversity data, supporting different types of stud-
ies and applications. The set of specimens of a species, from one or more collec-
tions, allow identifying the range of natural variation of the several traits that are
analyzed in the process of defining the species. Often, labels attached to the speci-
mens, or registry books associated with the collections include information on traits
that are recorded at the time of collection or when the specimen is added to the
collection (e.g. size, weight, length, maturity stage, colour of the flower in plants,
the presence of fruits, etc.). Additionally, the habitat, interactions with other species
(e.g. parasite of, epiphyte on), its use by local populations in traditional medicine,
food, and construction (mainly plants) is recorded.
Most taxonomic studies will require access to more than one collection, to allow
a comprehensive analysis of the variability of the species of interest. Finding which
collections have specimens important to the study might be demanding, but current
digitisation projects underway in many collections do facilitate the task enormously.
This is the case of some collections based in Angola at the Museum and Herbarium
of Lubango, the entomological collections of Instituto de Investigação Agronómica,
in Huambo, and the Herbarium of the Centro de Botânica da Universidade Agostinho
Neto, in Luanda. The first two initiatives are currently preparing the publication of
their databases through GBIF, for global and open access to data, which is possible
even though Angola is not yet a member of the organisation. Through GBIF,
researchers can have access to recorded information including most of the details
included in the specimen’s label, and if available, an image of the specimen, a sound
recording or a video attached to the specimen record. There are currently approxi-
mately 49,000 specimens from Angola with media attachments.
Other mega-science digital platforms also support capacity development in taxo-
nomic studies (Triebel et al. 2012). These include the Catalogue of Life, a global
checklist for all groups, that integrates more than 168 international or group-specific
checklists or taxonomic databases (Catalogue of Life 2018); the Biodiversity
Heritage Library (2018) which provides digital access to legacy literature, including
many publications with the original descriptions of the species, and automatic func-
tions for searching scientific names; the Encyclopedia of Life (2018), also an inte-
grative portal to information about species description, classification, multimedia,
and distribution maps of; the Barcode of Life (Ratnasingham and Hebert 2007)
which provides access to barcode sequence data; the IUCN Red List of Threatened
Species (2018), that promotes global and regional assessments of species conserva-
tion status; and the previously mentioned Global Plants Initiative. All these initia-
tives agree on common biodiversity informatics data standards, protocols and tools
that ensure the inter-operability across platforms under a common framework
(Hobern et al. 2012). This means that when researchers and institutions contribute
to or use one of these initiatives, they are reaching a global and transversal set of
resources covering several biodiversity dimensions which, although global, provide
detailed data applicable at the local or regional level.
These combinations of data are instrumental to prepare, for example, a national
checklist, like the national vascular plant list (Figueiredo and Smith 2008, Figueiredo
et al. 2009a, b). This task requires not only the compilation of information about
species and their distribution, but also synonyms, and sometimes helping to solve
taxonomic problems, species distribution ranges, and dealing with the scarcity of
information. Internet access to these and similar platforms is fundamental and an
important factor to reducing total costs of biodiversity research (Smith and
Figueiredo 2010).
An increase of the role of collections in preserving reference material on biodi-
versity is likely to occur with the addition of new methods for biodiversity identifi-
cation. Barcoding is one of the methods that determines DNA sequences that are
species-specific (Gross 2012), and it can speed up the identification of new species.
These sequences are stored in gene bank repositories, while the related physical
specimens, the source of the sequences, are stored as vouchers in collections.
Sometimes, specimens already represented in collections are found by barcoding to
belong to different species, therefore new arrangements are needed within collec-
tions. Museums are currently adopting new workflows with duplicate specimen pro-
cessing for traditional and molecular taxonomy (Gross 2012), in an articulated new
way of producing natural history knowledge (Strasser 2011).
etecting Changes in Species Distribution and in the

D
Environment
Understanding species distribution is usually a multidimensional problem that

involves information about species occurrences, climatic information, species
migrations and the availability of resources such as food and water. Primary biodi-
versity data from collections is often the only data resource to document the pres-
ence of species, either because of loss of habitats due to change of land use or
because of the local extinction of the species. This primary data, when combined
with environmental data, can be used to model the species distribution by numerical
tools that identify the environmental factors that are most closely associated with
occurrence of the species. This is, in turn, converted to a species distribution model
(SDM), expressed as a spatial map of probability of occurrence. Although subject to
problems associated with sample bias (Beck et al. 2014; Gomes et al. 2018), these
models can be an improved approach to gap analysis (Peterson and Kluza 2003), or
help to plan sampling effort to sites with a high potential of occurrence but that has
not yet been surveyed.
Distribution data from museums frequently supports assessments of the impact
of climate change on species distribution. By comparing two models, one for the
present distribution and another for a hypothesised future distribution, it is possible
to identify changes, including the expansion or reduction of the area of distribution.
The present distribution can be modeled from collection data, using a matching
period for climate date. A projection for a future state can be performed, using the
same distribution data, but with future climate scenarios data. Using this approach,
Warren et al. (2013) found that half of the plant species and one-third of animal spe-
cies can lose half of the suitable climate range by 2080. Another example with links
to human health is provided by Capinha et al. (2014), using as target species the
mosquito Aedes aegypti, the vector of dengue fever. These authors used collection
data combined with other data sources to determine the macroclimatic conditions
presently occupied by this mosquito, and the shifts in its distribution in the near
future (2010–2039), based on models of possible climate scenarios.
In assessing biological invasions, the use of NHCs is also essential. The histori-
cal record of an alien species needs to be determined so that its native distribution is
identified as well as its habitat and environmental requirements, life cycle, biotic
and abiotic interactions. Frequently, data and information on these parameters are
only available from museum or herbarium collections. These data will not only
allow to assess the invasion risk of a species, but also to predict its spread to new
regions, which can be done by projections of species distribution models, as for
example, the Giant African Snail (Sarma et al. 2015) or a result of climate change in
lantana (Taylor and Kumar 2014). Even if historical records were not available in
collections to support a study, these are essential as repositories of new records of
surveyed areas for future assessments (Rejmánek et al. 2017). Collections are also
important for the assessment of impacts in areas of invasion, in the determination of
species affected by the alien species.
Biodiversity Conservation
The IUCN Red List has become a standard with which to monitor a species’ conser-
vation status. Several of the criteria to determine the IUCN category of threat can be
obtained from natural history collections, such as features of life history, biology,
and geographical range. Williams and Crouch (2017) investigated whether herbar-
ium records could suffice for accurate estimation of the plant geographical range in
South Africa following IUCN Red List criteria. They concluded that results improve
when information from national herbaria is complemented by local or regional her-
barium datasets. For Cape Verde, herbarium data was also used in Red List assess-
ments of the endemic flora (Romeiras et al. 2016). However, the role of collections
in Red List assessments starts from the point of acertaining the correct identification
of each specimen in a survey, and before the application of any criteria, based on a
common taxonomy for the group of species under scrutiny (e.g. Grubb et al. 2003).
Another contribution of NHCs to biodiversity conservation is in reintroduction
programmes. When a local population of one species is extinct or threatened, the
reintroduction of new individuals can be done to increase population levels.
However, the genetic profile of the local population should be determined, in order
to ensure that new reintroduced individuals are the closest possible to the original
population, and thus well adapted to the environmental conditions of the new loca-
tion. If the local populations are extinct, museum or herbarium collections might be
the only resource to determine the genetic profile of the original populations, if
specimens are preserved in the NHCs from the original population. Collections are
also instrumental in determining other aspects of translocation planning, including
climate and habitat requirements (IUCN/SSC 2013).
A service provided by NHCs is also related to the trade of wild animals and
plants, within the scope of the CITES. Frequently, specimens of wild species are
seized by customs officials and the species identification is needed to check against
the species lists in the annexes of the convention. NHC taxonomists are frequently
asked to assist customs officials in identifying the species and the most likely source
of these organisms.
Museum and herbarium collections can also be used to verify if the network of
protected areas is effective in ensuring the protection of threatened species. Romeiras
et al. (2014) used collection data to make a biogeographic analysis of 18 high-value
timber trees from Angola. The authors concluded that these species could be
grouped within four regions, which had little correspondence to currently recog-
nised WWF ecoregions. They suggested that conservation plans based on WWF
ecoregions might provide the inappropriate basis for the conservation of these trees,
in which eight species were found to require high conservation priority because of
their very restricted distribution in Angola.
Supporting Sustainable Food Production
As in the definition of biodiversity adopted by the Convention of Biological

Diversity, the definition of agrobiodiversity by FAO highlights three levels of
diversity: diversity of genetic resources (varieties, breeds); diversity of species
used for food, fodder, fiber, fuel and pharmaceuticals; and the diversity of non-
harvested species that support production and diversity in the wider environment
that support agro-ecosystems (FAO 1999). Sustainable food production systems
demand attention to all these dimensions, which reinforces the role of NHCs. For
example, the conservation of crop wild relatives, which are wild species closely
related to crops, is important to ensure sources of genetic diversity useful to
develop more productive and resilient crops (Castañeda-Álvarez et al. 2016). A
priority in the conservation of such wild relatives is, therefore, the importance of
correct identification, based on herbarium data among other sources Castañeda-
Álvarez et al. 2016).
Natural history collections are also of importance in many other aspects of
agrosystems. This is the case of weed identification and the control of pests caused
by insects and fungi. Collections provide the resources for the identification of
these problem organisms, data for their first detection in a certain area, and infor-
mation about life history and distribution needed to determine the potential areas
of occurrence using bioclimatic and other modelling approaches. High biodiver-
sity in agrosystems can in some cases contribute to increasing productivity, by
promoting ecosystem services, e.g. through biological regulation of soil fertility
(Duru et al. 2015), for which NHCs are important to guide actions to increase
biodiversity in such systems, providing information on the original or potential
species native to the region.
Two further types of collections in support of food and forestry are also associ-
ated with herbarium collections. One of these is germplasm or seed bank collec-
tions. These seed collections are fundamental to the preservation of plant species,
by maintaining live and viable seeds for future use. There are more than 1300 seed
banks worldwide (Rajasekharan 2015), both for crop or wild species. These include
the Global Seed Vault, in Svalbard, Norway, which holds crop seeds for more than
5000 plant species, and the Millenium Seed Bank, at Kew and Wakehurst Place,
United Kingdom, which holds seeds for 10% of the world wild species. In Angola,
the Universidade Agostinho Neto hosts the only seed bank in the country as part of
the center for plant genetic resources. The other type of collection is the xylarium,
composed of wood samples, sometimes several pieces with different anatomical
sections of the wood of the same species. These samples are used, sometimes sup-
plemented by molecular genetics technologies, in identifying the products of illegal
activities in the timber trade (Yu et al. 2017).
Connecting Biodiversity to Society Through Education
Biodiversity is present, but rarely noticed, in the everyday life of humans. Natural
History Museums are important in bringing evidence of this to the population.
Through displays and exhibitions, it is possible to explain or demonstrate, in simple
terms, the value of biodiversity. In attractive displays, the direct value of biodiver-
sity can be shown in food, medicine, fuel, fiber, rubber, oils and building materials,
but also the indirect values through climate regulation, nutrient recycling, water and
air purification, pollination, and cultural, religious and aesthetic aspects. People
need to be informed by appropriate NHC displays in order to relate these values of
biodiversity to their daily life. These topics can also be explored more deeply to
explain biological concepts to students.
Specimens from collections facilitate the explanation of complex topics to visi-
tors. For example, concepts as life stages, evolution, adaptation to the environment,
species interaction and many others, are better explained using specimens as sup-
port. But these can also be used to provide insights to the science behind the scenes.
The causes of diversity, how genes express into forms and colors, the roles of micro-
scopic organisms that can move or destroy bodies a thousand times bigger, the work
of a taxonomist, a geneticists or a bioinformatician in understanding phylogenies,
the role of organisms as bioindicators of environmental changes, are all examples of
what might interest the visitor to a NHC. All these approaches can be comple-
mented by digital formats, through web pages that provide deeper coverage of the
topics displayed in the exhibitions. Via the Internet, it is also possible to use virtual
means to place species in their habitats and environments, or inform the visitor of
the species in one’s own neighbourhood. Other engagements with the public are
also possible, turning the visitor into a collaborator. For example, crowdsourcing
activities were implemented by some museums to make the databasing of speci-
mens’ labels, which is a time-consuming task in collection digitisation (Les
Herbonautes (2018); Notes from Nature (2018); DIGIVOL (2018).
Another example is citizen science participation, through which platforms citi-
zens can submit records, supported by images and other information, of species
occurrence, normally referring a date and a location (through GPS coordinates)
attached. Many projects of this type have emerged in recent years, the most visible
of the global scope being eBird (2018) for bird observations and iNaturalist.org
(2018) for any type of organism. Although subject to errors, these initiatives have
the enormous merit to expand the network of voluntary observers and are improving
their internal quality control mechanisms (using image analysis algorithms, for
example), to suggest or correct identifications. Records with attached images thus
become openly accessible to researchers, permitting validation of the records.
Museums can be windows to connect biodiversity science to society. Not only can
they contribute to educating people on concepts of biodiversity conservation and sus-
tainability but also to attracting new students and practitioners to biodiversity-related
topics. They can also be vibrant regional poles for research and natural history activi-
ties, interlinked with a global community of scientists and naturalists through digital
platforms. In some cases, their importance has also been recognised at an economic
level: the Natural History Museum in London has free entry to visitors, a benefit that
was earned after the demonstration of the economic benefit from attracting foreign
tourists to London.
oadmap for the Museum and Herbarium Collections

R
of Angola
The goal of developing knowledge about the biodiversity of Angola should be inti-
mately linked to strengthening the role of natural history collections as a reference
of biodiversity resources. That connection should be bi-directional, first to ensure
that natural history collections are used in studies about Angolan biodiversity, and
second that specimens documenting new distributional information are included in
collections for future reference. To ensure that the NHCs of Angola are prepared to
play this role, we propose the following roadmap for the museum and herbarium
collections of Angola.
Compile an Inventory of Collections from Angola
An inventory of the collections with specimens from Angola, either in Angolan

institutions or abroad, is important to produce an index of the available resources
and support a gap analysis of the biodiversity coverage of collections. This can be
done by a metadata description of the holdings of such collections, mentioning the
main taxonomic groups, time period, geographic area covered, main preservation
methods, total (estimated) number of specimens and number of species in the data-
base. An assessment of data needs (Asese and Schiwinger 2018) would provide
elements for future prioritisation of data mobilisation activities.
Identify Taxonomic Expertise and Promote Networking
A network of experts is essential to support NHC activities, in order to avoid, for

example, large time gaps between sampling and identification of specimens. These
can be taxonomists working in collections, but increasingly, ecologists, molecular
biologists and experts from other fields are performing taxonomic activities (Kemp
2015). This network of contacts should be developed to cover many biological
groups and be strongly tied to cooperation activities in training, study programmes

and projects focused on biodiversity. A route to promote the creation of this network
is the effective participation of Angola in the leading international networks, as is
the case of GBIF, or the Biodiversity Information Standards (TDWG) international
community, which leads and promotes developments in biodiversity informatics
worldwide.
Promote Data Repatriation Activities
Specimens from the country hosted in collections abroad contain important infor-
mation that should be available for studies and biodiversity management in Angola.
The repatriation of data can be promoted by a combination of initiatives that can
facilitate and speed up the access to it. For example, in the late 1990s, Mexico used
government funds to support visits of Mexican ornithologists to the largest bird col-
lection in the United States and in Europe, to catalog in a database bird specimens
from Mexico (Peterson et al. 2016). A similar approach could be done in the support
of visiting students or researchers from Angola to institutions hosting Angolan col-
lections. Furthermore, many collections have existing databasing projects, so coor-
dination with such activities could facilitate the prioritisation of data mobilisation.
The framework for these data mobilisation activities can be provided by the partici-
pation of countries and institutions in GBIF.
Include NHC Activities in University Curricula
In many graduate curricula in universities worldwide, a decrease in the importance

of natural history has been experienced through the last quarter of the twentieth
century. Fewer or no credits have been dedicated to traditional taxonomy, compared
to subjects in ecology, cellular and molecular biology, evolution and biotechnology.
However, natural history collections can now encompass these new methods,
remaining central to the goal of understanding the world’s biodiversity. We can see
NHCs as vibrant facilities that merge specimen and biomolecular preservation, and
biodiversity informatics infrastructure, being prepared to respond to societal chal-
lenges such as climate change, biodiversity loss and food security. Therefore, natu-
ral history can be attractive to teachers and graduate and postgraduate students in
universities, provided that its activities can be properly compensated. One way of
doing this is to reward data publication in career assessments of researchers and to
provide proper recognition through traceable citations to the use of collections in
scientific publications (Rouhan et al. 2017).
lign NHCs with National and International Agendas

A
on Biodiversity
The 2030 Agenda for Sustainable Development approved by the United Nations
includes several goals in which biodiversity and ecosystems take a central role. Goal
15 (Biodiversity, Forests, Desertification) is specifically targeted to halt biodiversity
loss, but biodiversity is also relevant for other Goals, as the Goal 2 (Hunger and
Food Security), Goal 12 (Sustainable Consumption and Production), Goal 13
(Climate Change) and Goal 14 (Oceans), if we consider ecosystem services or agro-
biodiversity. Angola will have the opportunity to participate in this agenda with
actions that fulfill international requirements, which in turn translate to national
priorities. Institutions with NHCs in Angola should be prepared to respond to the
needs that the implementation of this agenda requires, namely in providing the
essential information and expertise to support projects and reporting.
Conclusions
Museum and herbarium collections are restoring their paramount role in the study
of biodiversity, with the rapid developments seen recently in molecular biology and
in biodiversity informatics. These new tools contribute to speed up and add layers
of analysis to biodiversity assets represented in collections, not only for the materi-
als sampled in current projects but also for specimens collected through the history
of each collection. Many specimens have been kept hidden in collections for decades
before they were discovered as new species for science. Collections thus represent
an important asset by preserving the known (and unknown) biodiversity of a region
or a country, especially if they combine with these new approaches of analysis and
providing access to biodiversity information.
There is presently an under-representation of Angola’s biodiversity in NHCs.
The vastness of the country and the diversity of its ecoregions and habitats means
that this is a demanding task, but essential to support biodiversity knowledge and
conservation in the country. Obtaining a figure for the total number of specimens in
collections from Angola worldwide is difficult. However, from data available
through GBIF it is possible to obtain approximations. The current number of records
available online through the facility is circa 150,000, which is in the same order of
magnitude of other countries in southern Africa (except for South Africa, with 2,9
million and Democratic Republic of Congo, with 800,000). The situation is likely to
improve in the near future, with the start of participation by Angolan institutions in
GBIF (and the possible participation of the country), but the significance of these
numbers will need to be translated into effective access after a fitness for use and a
gap analysis of taxonomic and spatial coverage and biases of the data has been
conducted.
There are three herbaria and four zoological collections based in Angola, but not
all of them are currently active in the support of research or other biodiversity-
related activities. The three herbaria are or have plans to create a database for their
collections, and the hosting institution of two of them (ISCED and IIA) are already
registered as publishers of biodiversity data through GBIF, indicating that these
datasets will be openly available in the future. In terms of zoological collections, the
collections in ISCED and IIA also are developing databasing activities on their col-
lections, namely of birds and mammals, in the first case, and entomological, in the
second, with perspectives of online publishing through GBIF very soon. Some other
important collections remain, however, hidden or not easily accessible to research-
ers, such as the collections of Museu do Dundo, and the collections of the Museu
Nacional de História Natural. Little information is available for the current situa-
tion and accessibility of these collections, although an extensive literature is avail-
able about the activity of Museu do Dundo in a publication issued by that institution
in the 1950s and 1960s, with references to specimens in the collection (Machado
1995).
Online data availability is very important to attract national and international
researchers and specialists to use the collections in the country. This is important to
promote international collaboration and raise the capacity to use collections to
improve the knowledge of Angolan biodiversity, on topics related to ecology, evolu-
tion, and conservation. These collaborations are also important to promote data
mobilisation and quality improvement of collections based in Angola and abroad,
which is now supported by a framework of international digital platforms. But col-
lections in Angola need to be prepared to support new research activities in the field,
considering that the biodiversity of parts of the country is still relatively unknown
and in need of field surveys, as described in other chapters of this volume.
Furthermore, to face big environmental challenges like the loss of biodiversity, cli-
mate change, and invasive alien species, it is urgently necessary to provide more
information and knowledge about biodiversity, and collections are certainly the
most accessible way to begin.
Natural History Collections are also important to link biodiversity to society.
Many aspects of the importance of biodiversity to everyday life can be achieved
through attractive displays that link the natural curiosity of humans with features of
the structure and functioning of biodiversity, resulting in important impacts on the
education and awareness of communities. Stimulating displays and activities can
also contribute to attracting more young researchers to work in NHCs. Education is
one of the most important roles of collections, in association with other uses for
preservation, documentation and biodiversity conservation. Therefore, Natural
History Collections represent strategic infrastructures for a country: reason enough
to contradict the fate predicted by Saramago that all of them would return to chaos.
Appendix
Natural History Collections holding specimens from Angola
Acronym Institution
AMNH American Museum of Natural History (USA)
ARC Agricultural Research Council, Plant Protection Research Institute (South
Africa)
B Botanic Garden and Botanical Museum Berlin-Dahlem (Germany)
BMSM Bailey-Matthews National Shell Museum (USA)
BR Botanic Garden Meise (Belgium)
CAS California Academy of Sciences (USA)
CM Carnegie Museums (USA)
COI Herbarium of the Universidade de Coimbra
E Royal Botanic Garden Edinburgh (United Kingdom)
FCEyN, ArOBIS Centro Nacional Patagónico (Argentina)
UBA
FishBase FishBase
FMNH Field Museum (USA)
GNM Gothenburg Natural History Museum (Sweden)
Ifremer French Research Institute for Exploitation of the Sea (France)
IICT Instituto de Investigação Científica Tropical of the Universidade de Lisboa
ISCED Instituto Superior de Ciências de Educação da Huíla (Angola)
K Royal Botanic Gardens, Kew (United Kingdom)
KU University of Kansas Biodiversity Institute (USA)
LEGON-GC University of Ghana – Ghana Herbarium (Ghana)
MACN Museo Argentino de Ciencias Naturales (Argentina)
MHNG Muséum d’Histoire Naturelle de la Ville de Genève (Switzerland)
MNCN Spanish National Museum of Natural Sciences (Spain)
MNHN Museum National d’Histoire Naturelle (France)
MUHNAC Museu Nacional de História Natural e da Ciência da Universidade de Lisboa
(Portugal)
MVZ Museum of Vertebrate Zoology (USA)
NHMUK Natural History Museum (United Kingdom)
RBINS Royal Belgian Institute of Natural Sciences (Belgium)
RMCA Royal Museum for Central Africa (Belgium)
S Swedish Museum of Natural History (Sweden)
SAIAB South African Institute for Aquatic Biodiversity
SANBI South African National Biodiversity Institute
SMF Senckenberg (Germany)
SNSB-M Staatliche Naturwissenschaftliche Sammlungen Bayerns (Germany)
TM Ditsong National Museum of Natural History Collection (South Africa)
UPS Museum of Evolution in Uppsala (Sweden)
USNM National Museum of Natural History, Smithsonian Institution (USA)
VM Vänersborg Museum (Sweden)
(continued)
Acronym Institution
YPM Yale University Peabody Museum (USA)
ZMB Collection Crustacea, Senckenberg (Germany)
ZMUC Zoological Museum, Natural History Museum of Denmark (Denmark)
References
Albuquerque S, Correia AI (2010) The Welwitsch collections – Iter Angolense (1853–1860) at

LISU. In: Van Der Burgt X, Van Der Maesen J, Onana J-M (eds) Systematics and conservation
of African plants. Royal Botanic Gardens, Kew, pp 787–790
Albuquerque S, Brummitt RK, Figueiredo E (2009) Typification of names based on the Angolan
collections of Friedrich Welwitsch. Taxon 58:641–646
Alves MJ, Bastos-Silveira C, Cartaxana A et al (2014) As Coleções Zoológicas do Museu Nacional
de História Natural e da Ciência. In: Alves MJ, Cartaxana A, Correia AM et al (eds) Professor
Carlos Almaça (1934–2010) – Estado da Arte em Áreas Científicas do Seu Interesse. Museus
da Universidade de Lisboa, Lisboa, pp 289–301
Asase A, Schwinger GO (2018) Assessment of biodiversity data holdings and user data needs for
Ghana. Biodivers Inform 13:27–37
Bebber DP, Carine MA, Wood JRI et al (2010) Herbaria are a major frontier for species discovery.
Proc Natl Acad Sci 107:22169–22171
Beck J, Böller M, Erhardt A et al (2014) Spatial bias in the GBIF database and its effect on model-
ing species’ geographic distributions. Eco Inform 19:10–15
Biodiversity Heritage Library (2018) URL https://www.biodiversitylibrary.org/. Accessed 12 June
2018
Brooke ML (2000) Why museums matter. Trends Ecol Evol 15:136–137
Burgess N, Hales JD, Underwood E et al (2004) Terrestrial ecoregions of Africa and Madagascar –
a conservation assessment. Island Press, Washington, DC, p 499
Cabral JC (2010) João Crawford Cabral (depoimento, 2009). IICT, Lisboa, 16 pp
Capinha C, Rocha J, Sousa CA (2014) Macroclimate determines the global range limit of Aedes
aegypti. EcoHealth 11(3):420–428
Cassinda S, Fernandes Elizalde S, Bassimba D (2018) Colecção Entomológica IIA. Instituto
de Investigação Agronómica – IIA. Occurrence dataset. https://doi.org/10.15468/bhqdhp.
Accessed via GBIF.org on 18 June 2018
Castañeda-Álvarez NP, Khoury CK, Achicanoy HA et al (2016) Global conservation priorities for
crop wild relatives. Nat Plants 2:16022
Catalogue of Life (2018) URL http://www.catalogueoflife.org/. Accessed 12 June 2018
Ceríaco LMP (2014) O “Arquivo Histórico Museu Bocage” e a História da História Natural em
Portugal. In: Alves MJ, Cartaxana A, Correia AM et al (eds) Professor Carlos Almaça (1934–
2010) – Estado da Arte em Áreas Científicas do Seu Interesse. Museus da Universidade de
Lisboa, Lisboa, pp 329–358
Deng B (2015) Plant collections left in the cold by cuts. Nature 523:16–16
DIGIVOL (2018) URL https://digivol.ala.org.au/. Accessed 15 June 2018
Duru M, Therond O, Martin G et al (2015) How to implement biodiversity-based agriculture to
enhance ecosystem services: a review. Agron Sustain Dev 35:1259–1281
eBird (2018) URL https://ebird.org/home. Accessed 15 June 2018
Encyclopedia of Life (2018) URL http://eol.org/. Accessed 12 June 2018
FAO (1999) Agricultural biodiversity, multifunctional character of agriculture and land confer-
ence, background paper 1. FAO, Maastricht
Figueiredo E, Smith GF (2008) Plants of Angola/Plantas de Angola. Strelitzia 22:1–279
Figueiredo E, Smith GF, Cesar J (2009a) The flora of Angola: first record of diversity and ende-
Figueiredo E, Soares M, Seibert G et al (2009b) The botany of the Cunene-Zambezi Expedition
with notes on Hugo Baum (1867-1950). Bothalia 39:185–211
Fontaine B, Perrard A, Bouchet P (2012) 21 years of shelf life between discovery and description
of new species. Curr Biol 22:R943–R944
GBIF.org (25 May 2018) GBIF occurrence download. https://doi.org/10.15468/dl.urk4kx
Gilles D, Zaiss R, Blach-Overgaard A et al (2016) RAINBIO: a mega-database of tropical African
vascular plants distributions. PhytoKeys 74:1–18
Gomes VHF, Ijff SD, Raes N et al (2018) Species distribution modelling: contrasting presence-
only models with plot abundance data. Sci Rep 8:1003
Gropp RE (2003) Are university natural science collections going extinct? Bioscience 53:550–550
Gross M (2012) Barcoding biodiversity. Curr Biol 22:R73–R76
Grubb P, Butynski TM, Oates JF et al (2003) Assessment of the diversity of African primates. Int
J Primatol 24:1301–1357
Hobern D, Apostolico A, Arnaud E, et al (2012) Global biodiversity informatics outlook: deliver-
ing biodiversity knowledge in the information age. Global Biodiversity Information Facility,
Copenhagen. Available at: https://www.gbif.org/document/80859
iNaturalist.org (2018) URL https://www.inaturalist.org/. Accessed 15 June 2018
IUCN/SSC (2013) Guidelines for reintroductions and other conservation translocations. IUCN
Species Survival Commission, Gland
Kemp C (2015) Museums: the endangered dead. Nature 518:292–294
Lavoie C (2013) Biological collections in an ever changing world: herbaria as tools for biogeo-
graphical and environmental studies. Perspect Plant Ecol Evol Syst 15:68–76
Les herbonautes (2018) URL http://lesherbonautes.mnhn.fr/. Accessed 15 June 2018
Machado AB (1952) Generalidades acerca da Lunda e da sua exploração biológica. Companhia de
Diamantes de Angola, Publicações Culturais 12:1–111
Machado AB (1995) Notícia sumária sobre a acção cultural da Companhia de Diamantes de
Angola. In: Diamang – Estudo do património cultural da ex-Companhia dos Diamantes de
Angola. Publicações do Centro de Estudos Africanos, Museu Antroplógico da Universidade de
Coimbra, Coimbra, pp 11–24
Martins E, Martins T (2002) Herbários de Angola: que futuro? Garcia da Orta Sér Bot 16(1–2):1–4
Mendes LF, Bivar-de-Sousa A, Figueira R (2013) Butterflies of Angola. Lepidoptera. Papilionoidea.

I. Hesperiidae, Papilionidae. IICT and CIBIO, Lisboa and Porto
Mills M, Melo M (2013) The checklist of the birds of Angola 2013. Associação Angolana para
Aves e Natureza, Luanda, Angola
Mills M, Franke U, Joseph G, Miato F, Milton S, Monadjem A, Oschadleus D, Dean W (2010)
Cataloguing the Lubango Bird Skin Collection: towards an atlas of Angolan bird distributions.
Bull ABC 17:43–53
Mora C, Tittensor DP, Adl S et al (2011) How many species are there on Earth and in the Ocean?
PLoS Biol 9:e1001127
Myers N, Mittermeier RA, Mittermeier CG, da Fonseca GAB, Kent J (2000) Biodiversity hotspots
for conservation priorities. Nature 403:853–858
Nature Editorial (2014) Save the museums. Nature 515:311–312
Notes from Nature (2018) URL https://www.notesfromnature.org/. Accessed 15 June 2018
Oliveira PSD, Rocha MT, Castro AG et al (2016) New records of Gaboon viper (Bitis gabonica)
in Angola. Herpetol Bull 136:42–43
new map of life on earth. Bioscience 51:933–938
Peterson AT, Kluza DA (2003) New distributional modelling approaches for gap analysis. Anim
Conserv 6:47–54
Peterson AT, Navarro-Sigüenza AG, Gordillo-Martínez A (2016) The development of ornithology
in Mexico and the importance of access to scientific information. Arch Nat Hist 43:294–304
Prathapan KD, Pethiyagoda R, Bawa KS et al (2018) When the cure kills – CBD limits biodiversity
research. Science 360:1405–1406
Pyke GH, Ehrlich PR (2010) Biological collections and ecological/environmental research: a
review, some observations and a look to the future. Biol Rev 85:247–266
Rajasekharan PE (2015) Gene banking for ex situ conservation of plant genetic resources. In: Plant
biology and biotechnology. Springer, New Delhi, pp 445–459
Ratnasingham S, Hebert PDN (2007) BOLD: the barcode of life data system (www.barcodinglife.
org). Mol Ecol Notes 7:355–364
Rejmánek M, Huntley BJ, Roux JJL et al (2017) A rapid survey of the invasive plant species in
Rocha LA, Aleixo A, Allen G, Almeda F et al (2014) Specimen collection: an essential tool.
Science 344:814–815
Romeiras M (1999) Subsídio para o conhecimento dos colectores botânicos em Angola. Revista
de Ciências Agrárias 22:73–83
Romeiras MM, Figueira R, Duarte MC et al (2014) Documenting biogeographical patterns of
African timber species using herbarium records: a conservation perspective based on native
trees from Angola. PLoS ONE 9:e103403
Romeiras MM, Catarino S, Gomes I et al (2016) IUCN Red List assessment of the Cape Verde
endemic flora: towards a global strategy for plant conservation in Macaronesia. Bot J Linn Soc
180:413–425
Rouhan G, Dorr LJ, Gautier L et al (2017) The time has come for Natural History Collections to
claim co-authorship of research articles. Taxon 66:1014–1016
Santos J, Sales F (eds) (2018) Catalogue of the herbarium of the University of Coimbra (COI).
Department of Life Sciences, Faculty of Sciences and Technology of the University of Coimbra.
http://coicatalogue.uc.pt. Accessed on 5 May 2018
Saramago J (2000) All the names. The Harvill Press, London
Sarma RR, Munsi M, Ananthram AN (2015) Effect of climate change on invasion risk of giant
African snail (Achatina fulica Férussac, 1821: Achatinidae) in India. PLoS ONE 10:e0143724
(eds) Biodiversity of Angola. Science & conservation: a modern synthesis. Springer Nature,
Cham
Smith GF, Figueiredo E (2010) E-taxonomy: an affordable tool to fill the biodiversity knowledge
gap. Biodivers Conserv 19:829–836
Stokstad E (2017) Botanists fear research slowdown after priceless specimens destroyed at
Australian border. Science News 11-05-2017. https://doi.org/10.1126/science.aal1175.
Accessed 13 June 2018
Strasser BJ (2011) The experimenter’s museum: GenBank, natural history, and the moral econo-
mies of biomedicine. Isis 102:60–96
Suarez AV, Tsutsui ND (2004) The value of museum collections for research and society.
Bioscience 54:66–74
Taylor S, Kumar L (2014) Impacts of climate change on invasive Lantana camara L. distribution
in South Africa. Afr J Environ Sci Technol 8:391–400
Tewksbury JJ, Anderson JGT, Bakker JD et al (2014) Natural history’s place in science and society.
Bioscience 64:300–310
The IUCN Red List of Threatened Species (2018) URL http://www.iucnredlist.org/. Accessed 12
June 2018
Thiers B (2018) Index Herbariorum: a global directory of public herbaria and associated staff.
New York Botanical Garden’s Virtual Herbarium. http://sweetgum.nybg.org/science/ih/
Triebel D, Hagedorn G, Rambold G (2012) An appraisal of megascience platforms for biodiversity
information. MycoKeys 5:45–63
Warren R, VanDerWal J, Price J et al (2013) Quantifying the benefit of early climate change miti-
gation in avoiding biodiversity loss. Nat Clim Chang 3:678–682
Wheeler QD, Knapp S, Stevenson DW et al (2012) Mapping the biosphere: exploring species to
understand the origin, organization and sustainability of biodiversity. Syst Biodivers 10:1–20
Williams VL, Crouch NR (2017) Locating sufficient plant distribution data for accurate estimation
of geographic range: the relative value of herbaria and other sources. S Afr J Bot 109:116–127
Yeates DK, Zwick A, Mikheyev AS (2016) Museums are biobanks: unlocking the genetic poten-
tial of the three billion specimens in the world’s biological collections. Curr Opin Insect Sci
18:83–88
Yu M, Jiao L, Guo J et al (2017) DNA barcoding of vouchered xylarium wood specimens of nine
endangered Dalbergia species. Planta 246:1165–1176
Chapter 20
Conclusions: Biodiversity Research
and Conservation Opportunities
Vladimir Russo, Brian J. Huntley, Fernanda Lages, and Nuno Ferrand
Abstract Angola is a country full of opportunities. Few countries offer more excit-
ing prospects for young scientists to discover and document the rich biodiversity,
complex ecosystem processes and undescribed species of plants and animals that
are to be found in its amazing diversity of landscapes and seascapes. The current
expanding support of the Angolan government and of international partners is
unprecedented, and the positive response from young students ensures the growth of
a new generation of biodiversity researchers and conservation professionals. Based
on a synthesis of biodiversity research and conservation activities of the past cen-
tury, we outline opportunities, approaches and priorities for a strengthened collab-
orative research and conservation agenda.
Keywords Africa · Angola · Biological discovery · Checklists · Endemic species ·

Research priorities · Science collaboration · Socio-ecological systems · Threatened
species
V. Russo (*)
e-mail: vladyrusso@gmail.com; vladimir.russo@holisticos.co.ao
B. J. Huntley
F. Lages
ISCED – Instituto Superior de Ciências da Educação da Huíla, Lubango, Angola
e-mail: f_lages@yahoo.com.br
N. Ferrand
CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Laboratório
Associado, Campus de Vairão, Universidade do Porto, Vairão, Portugal
Departamento de Biologia, Faculdade de Ciências, Universidade do Porto,
Porto, Portugal
Department of Zoology, Auckland Park, University of Johannesburg,
Johannesburg, South Africa
e-mail: nferrand@cibio.up.pt

https://doi.org/10.1007/978-3-030-03083-4_20
544 V. Russo et al.
Context: Challenges and Opportunities
This book was conceived on the basis of three simplistic assumptions. First, that
very little information is available on the biodiversity of Angola. The second
assumption was that colonial governments had neglected, and post-independence
authorities have been unsupportive of research on the fauna, flora and ecosystems of
Angola. Thirdly, we assumed that existing biodiversity knowledge was mostly lost
in dusty government archives or inaccessible scientific journals. In truth, the rich
assemblage of information gathered together in this synthesis volume demonstrates
the error of these assumptions. The perceived challenges of the past have become a
mosaic of opportunities for the future.
The energy, knowledge and dedication of this book’s 46 contributing authors has
resulted in a comprehensive synopsis of the ‘state of the science’ on the evolution
and diversity of Angola’s landscapes, flora, vegetation, all vertebrates, two iconic
invertebrate taxa, and key ecosystems in both marine and terrestrial environments.
What is immediately obvious from each chapter is the wealth of Angola’s natural
heritage, and how fragile it is to anthropogenic impacts and the vicissitudes of cli-
mate change. The vulnerability of the remnant forests of Angolan Escarpment and
Afromontane ecosystems to these pressures places a tremendous responsibility on
Angolans to study and protect these fingerprints of the past. Angola’s universities,
scientific organisations and government research institutions can lead the way to
strengthening our understanding of the evolution, structure and functioning of these
and the many other special habitats that make Angola unique in Africa in terms of
the diversity of biomes and ecoregions found within its borders.
Each chapter of this volume reveals research and conservation opportunities rel-
evant to the environment or taxon under discussion and presents compelling argu-
ments for greater levels of investment in both research and conservation. Some
needs are very specific, such as the importance of biodiversity surveys, vegetation
maps and socio-economic assessments of the country’s many protected areas as a
basis to achieving their effective management and the delivery of long-term benefits
to society. Other priorities are more broadly based, aimed at developing tools for
wide-scale natural resource-use planning, such as a new and detailed vegetation
map for the entire country, building on the results of the current forest inventory, and
for a national biodiversity data portal. Yet others focus on selected taxa that can
help, through modern genomic studies, to explain the processes of speciation that
have led to the richness of the country’s flora and fauna. All have a common pur-
pose – to effectively inform decisions that will ensure increasingly sustainable
development for all Angolans and for humanity at large.
20 Conclusions: Biodiversity Research and Conservation Opportunities 545
Towards a Biodiversity Conservation Research Strategy
Over 40 research topics are identified in the chapters of this volume. Such compila-
tions of research opportunities need to be embraced within national strategies,
matching needs with resources and priorities within a research agenda, while under-
standing the country’s challenges. In recent decades, a first point of departure in the
design of conservation science programmes has been the frameworks provided by
international multilateral environmental conventions – most specifically the
Convention on Biological Diversity to which Angola has been a signatory since
1998. The development of Angola’s National Biodiversity Strategy and Action Plan
(NBSAP) brought a logical structure to both policy and planning (GoA 2006, 2018).
These strategies have been further developed through setting nationally relevant
implementation goals such as those detailed in the Aichi Targets (CBD 2010). An
early lesson learned in the implementation of such strategies was the fundamental
importance of cooperative partnerships between multiple institutions. The global
scarcity of taxonomists, for example, has meant that no single country has the
capacity to study, understand and document all taxa and ecosystems. The strength
of organisations such as IUCN, GBIF, IPBES, etc., is based on shared information
and skills. The benefits of the recent surge of interest in Angola’s biodiversity dem-
onstrated by foreign universities, museums, non-governmental organisations and
intergovernmental agreements have yet to be fully exploited. For this, a strategic
approach, implemented opportunistically, is advantageous.
Building a programme of work for conservation science needs to be mobilised at
several levels, often simultaneously, rather than sequentially. At a first level, biodi-
versity assessments are a priority. Biodiversity surveys, which provide both the
building blocks of knowledge and unique training and capacity building opportuni-
ties, must maintain the positive momentum of the past decade. The preliminary
checklists of species presented in this volume should be regularly updated and
expanded to integrate these and other taxonomic groups through an electronic bio-
diversity data portal. Ideally, international institutions, in liaison with national hosts,
should assist in the coordination and curation of checklists, atlases and field guide-
books such as those already available for the reptiles and amphibians of Cangandala
National Park (Ceríaco et al. 2016), the ‘special birds’ of Angola (Mills 2018) and
the atlas of Angolan reptiles and amphibians (Marques et al. 2018). Checklists and
the natural history collections on which they are based need integration within inter-
national databases such as those of GBIF and the Catalogue of Life. A direct out-
come of this component of research is the development of taxonomic skills and of
para-taxonomists in Angola.
Second, the socio-ecological systems within which research and conservation
play out should not be neglected. Biodiversity does not exist in a vacuum – it has a
human face that governs the success or failure of interventions. Biodiversity scien-
546 V. Russo et al.
tists who ignore the human dimensions of research and conservation do so at their
peril. This is especially true within and around formal protected areas, where studies
are needed on the socio-economic drivers of change (from a subsistence to a
cash-based economy in rural areas), and the impacts of the bushmeat trade, slash-
and-burn agricultural practices, charcoal production and nomadic pastoralism, on
biodiversity conservation across the country. Such studies should also draw on local
indigenous knowledge in developing management plans and research projects. At a
regional scale, the assessment of the role of ecosystem services for sustainable liv-
ing in rural and urban environments provides information essential to long-term
development planning, ideally guided by a new and detailed vegetation map of
Angola. The interdependence of basic research, biodiversity assessments and
applied studies is obvious.
Third, there is no questioning the paramount importance of strengthening the
capacity of young Angolan researchers and of research institutions. These actions
should be framed within a collaborative and mutually beneficial strategy.
International collaboration is already assisting in this, but needs further support.
Scholarships, internships and mentoring programmes are fundamental, but field
experience is critical for young biodiversity researchers, and the promotion of bush
camps for student training in protected areas and biodiversity hotspots is a highly
effective mechanism for inspiring the new generation. The establishment of a vir-
tual network of Angolan conservation professionals using social media such as the
Angola Ambiente Facebook site is a highly effective initiative. Angolan institutions
also need strengthening and closer cooperation across government departments, and
the integration of their research results through a formal information portal, is a key
opportunity.
Research Opportunities from Genes to Landscapes
While the above three pillars form the foundations of a nascent biodiversity conser-
vation research agenda, with an emphasis on immediate and practical needs, there
are many fascinating questions relating to the functioning of Angola’s diverse eco-
systems that challenge the minds of biologists as they explore the country. The
research strategy should be reinforced by studies on the evolutionary and ecological
processes that account for Angola’s biodiversity. An understanding of the evolution
of Angola’s biota will be strengthened by the development of modern phylogenies
of key taxonomic groups, as proposed in many to the chapters of this volume. An
understanding of the evolution of species assemblages and of individual species is
of great value to guiding conservation measures, as already demonstrated by the
Angolan studies of Vaz da Silva (2015) and Vaz Pinto (2018). Advanced molecular
technologies allow new insights into many patterns revealed by basic surveys and
assessments. The population genetics and hybridisation of Angola’s two elephant
species needs urgent study before the last remnants of Forest Elephant are driven to
extinction. Similarly, iconic plants such as the baobab Adansonia digitata that in
Angola manifests as widely divergent phenotypes, from dwarf baobabs in Namibe
to the obese giants of Cunene and the slender columns of Cuanza-Norte, merit stud-
ies on their genetic diversity, ecology and traditional uses. Angola’s once vast popu-
lations of Welwitschia mirabilis await research on their population dynamics and
potential resilience to overgrazing by cattle in Iona National Park. The bizarre pat-
terns of ‘fairy circles’ of the Namib, ‘fairy forests’ of the Lundas and Moxico, of
dwarf succulents along the desert margins of Benguela, and the ecological role of
fog along the entire coast, are all ecological puzzles that need elucidation.
Many of these topics might at first sight appear of little more than academic
interest, but every element of applied science and technology rests on the funda-
mentals of curiosity-driven enquiry. The baseline compilation of checklists leads to
the identification of patterns of endemism and rarity, to be understood through phy-
logeographic studies across the country. From such studies, increasingly robust sce-
narios of evolutionary processes may be built. It is these phenomena of the patterns
in nature, their ecological functioning and the interactions that drive large-scale
environmental dynamics that will ultimately guide sustainable landuse management
and inform responses to the impacts of climate change. It is at the level of land-
scapes and seascapes that the nation’s economy and progress is built, and such
wide-horizon visioning, underpinned by fundamental research, is needed for the
sustainable development agendas of the twenty-first century.
At a landscape scale, an early priority should be to update the vegetation map of
Angola to give a more balanced and objective delineation of the country’s major
vegetation units, for landuse planning and conservation purposes. This can best be
achieved with the input of a multi-national team of workers, using modern remote
sensing technologies for vegetation classification, mapping and monitoring.
Vegetation classification and mapping skills take many years, even decades to
develop, and the close collaboration, even leadership, of foreign experts would be
valuable. An improved vegetation map will help expedite assessments of ecosystem
conservation status, landuse potential, research priorities and opportunities, and
help develop a predictive understanding of ecosystem structure and function.
At ecosystem scale, the importance of securing the effective management of pro-
tected areas, large and small, terrestrial and marine, is a sine qua non to the future
of Angola’s biodiversity. The biological and cultural importance of relatively small
protected areas, such as Ilhéu dos Pássaros, and future protected areas such as Mount
Moco, Namba, Cumbira, Tundavala and the rainforests of Cuanza-Norte, Uíge and
Zaire, and of turtle nesting grounds along the coast, should not be overlooked dur-
ing the pursuit of mega-parks that might excite the public and motivate politicians.
Targeted studies of the existing protected areas and of the key biodiversity hotspots
identified by the Ministry of Environment in successive strategies (GoA 2006, 2011,
2018) should be given priority, as these protected areas and hotspots most probably
hold more than 80% of Angola’s floral and faunal diversity in less than 15% of the
548 V. Russo et al.
country’s land area. As training grounds for young biologists and conservation sci-
entists, protected areas have no equal. Furthermore, identifying and rigorously pro-
tecting near-pristine zones within otherwise threatened protected areas such as
Quiçama, Luando, Iona and Mupa, and of the country’s extensive coastline, should
be an integral part of any protected areas strategy. The marine environment is espe-
cially sensitive to the impacts of human activities and science-based marine spatial
planning is essential if long-term conflicts between humans and the marine
environment are to be avoided. The importance of a focus on the biodiversity
hotspots and on the existing protected areas of Angola – the repositories of the
country’s natural wealth – is self-evident.
Conclusions
These outlines are of necessity simplistic and preliminary. Each of the chapters in
this book identifies research questions that can be addressed by an emerging genera-
tion of Angolan biodiversity scientists and conservation professionals. The chal-
lenges are exciting and demanding – offering multiple opportunities for intellectual
stimulation, knowledge generation and international collaboration. Angola is truly
alive with research and conservation opportunities. The country is still blessed with
vast areas of rich wilderness and unique habitats, and has the opportunity to stimu-
late scientists, conservationists and the general public to participate in programmes
of research and effective biodiversity conservation management. As this synthesis
volume demonstrates, the limits are boundless. Carpe diem!
References
CBD (2010) Strategic plan for biodiversity 2011–2020 and the Aichii targets. Secretariat of the
Convention on Biological Diversity, Montreal
Ceríaco LMP, Marques MP, Bandeira SA et al (2016) Anfíbios e Répteis do Parque Nacional da
Cangandala. Instituto Nacional da Biodiversidade e Áreas de Conservação/Museu Nacional de
História Natural e da Ciência, Luanda/Lisboa, 96 pp
GoA (Government of Angola) (2006) National biodiversity strategy and action plan (2007–2012).
Ministry of Urban Affairs and the Environment, Luanda, 55 pp
de Angola (PLERNACA). Ministério do Ambiente, Luanda
GoA (Government of Angola) (2018) Plano Estratégico para o Sistema de Áreas de Conservação
de Angola (PESAC). Ministério do Ambiente, Luanda
Marques MP, Ceríaco LMP, Blackburn DC, Bauer AM (2018) Diversity and distribution of the
amphibians and terrestrial reptiles of Angola. Atlas of historical and bibliographic records
(1840–2017). Proceedings of the California Academy of Sciences Series 4, 65:1–501
Mills MSL (2018) The special birds of Angola/As Aves Especiais de Angola. Go-Away-Birding,
Cape Town
piece to understand the biogeography of the Afromontane forests. MSc thesis. University of
Porto, Porto
and conservation. PhD thesis. University of Porto, Porto

2019 Book BiodiversityOfAngola

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

2019 Book BiodiversityOfAngola

Uploaded by

Copyright:

Available Formats

Brian J.

Huntley · Vladimir Russo

United Nations UNESCO Chair on Life on Land

ISBN 978-3-030-03082-7 ISBN 978-3-030-03083-4 (eBook)

Library of Congress Control Number: 2018966550

President of the Republic of Angola João Manuel Gonçalves Lourenço

This book was conceived as a collaborative and voluntary endeavour shared by

Finally, our thanks go to Margaret Deignan, Esther Rentmeester, Malini

Part I Introduction: Setting the Scene

Part II Flora, Vegetation and Landscape Change

Part III Invertebrate Diversity: Environmental Indicators

Part IV Vertebrates: Distribution and Diversity

Part V Research and Conservation Opportunities

19 Museum and Herbarium Collections for Biodiversity

Ninda Baptista Instituto Superior de Ciências da Educaҫão da Huíla, Rua

Werner Conradie National Geographic Okavango Wilderness Project, Wild Bird

Pedro Monterroso CIBIO-InBIO, Centro de Investigação em Biodiversidade e

Abstract Angola possesses an unusually rich diversity of ecosystems and species,

Keywords Africa · Biomes · Collaborative research · Conservation · Ecoregions

Angola is a country of unusually rich physiographic, climatic and biological diver-

© The Author(s) 2019 3

biodiversity is extremely limited when compared with most African countries.

Approach and Purpose of This Synthesis

Biodiversity Surveys: Historical Synopsis

The history of scientific exploration and biological collection in Angola is relatively

The Swiss zoologist, Monard (1935), had been less sanguine:

Research Collaboration in the Twenty-First Century

Namibia and provided a strong impetus to future multi-national projects in the

Abstract Angola is a large country of 1,246,700 km2 on the southwest coast of

Keywords Afromontane forest · Biogeography · Biomes · Climate change ·

© The Author(s) 2019 15

synthesised the diverse drivers of Angola’s ecological systems and agricultural

As a large country of 1,246,700 km2 on the southwest coast of Africa, Angola is

Geomorphology and Landscape Evolution

The general topography of Angola is illustrated in Fig. 2.1. In summary, coastal

continued from the 1950s to 1970s by Portuguese researchers, including Marques

Upper Cuanza Luena

The conservation importance of the Angolan river systems is of great signifi-

Vertisols Podzols Luvisols Ferralsols

Fig. 2.7 Mean annual rainfall in Angola

Biogeography, Biomes and Ecoregions

Angola’s geographic location, geological history, climate and physiography account

Africa: Congolian, Zambezian, Southern African, Ethiopian, Somalian, Sudanian

• Tropical and subtropical grasslands, savannas, shrublands, and woodlands;

The Biological Importance of the Angolan Escarpment

Table 2.2 Representation of biomes and ecoregions in southern African countries

Fig. 2.8 Ecoregions of Angola

planning in Angola (Revermann and Finckh 2019). What is equally important in

Albuquerque S, Figueirôa S (2018) Depicting the invisible: Welwitsch’s map of travellers in

Feio M (1964) A evolução da escadaria de aplanações do sudoeste de Angola. Garcia da Orta

© The Author(s) 2019 43

Keywords Benguela current · Ecologically or biologically significant marine areas

Physical and Oceanographic Context

The coastline of Angola, which is approximately 1650 km long, consists of sandy

There is a high diversity of demersal species in Angola relative to the temperate

 arine and Coastal Biodiversity Hotspots, Threats

Octávio Mateus, Pedro M. Callapez, Michael J. Polcyn, Anne S. Schulp,

Abstract This chapter provides an overview of the alpha paleobiodiversity of

© The Author(s) 2019 53

Keywords Ammonites · Benguela Basin · Cenozoic · Cretaceous · Cuanza Basin ·

The study of paleobiodiversity, i.e., the development of biodiversity through geo-

2. Life remains that can leave traces and fossilize

3. Remains that actually fossilized

Part I Introduction: Setting the Scene

Part II Flora, Vegetation and Landscape Change

Part III Invertebrate Diversity: Environmental Indicators

Part IV Vertebrates: Distribution and Diversity

Part V Research and Conservation Opportunities

19 Museum and Herbarium Collections for Biodiversity

Approach and Purpose of This Synthesis

Biodiversity Surveys: Historical Synopsis

Research Collaboration in the Twenty-First Century

Geomorphology and Landscape Evolution

Biogeography, Biomes and Ecoregions

The Biological Importance of the Angolan Escarpment

Physical and Oceanographic Context

arine and Coastal Biodiversity Hotspots, Threats

A Brief Geological History and Context of Angola

The Precambrian – The First Fossils

The Triassic – Inner Basins

The Early Cretaceous – The Opening of South Atlantic

Plants, Protists and Invertebrates

The Late Cretaceous – Marine Reptiles Flourish

In contrast to the diversity of chondrichthyans, the Late Cretaceous bony ray-

The Paleogene – Mammals Take Over

The Neogene – The Founding of Modern Biodiversity

The Quaternary – The Dominance of Humans

Final Remarks on the Fossil Record and Paleobiodiversity

History of Botanical Exploration in Angola

Floristic Diversity and Endemism

Biogeography, Regional Centres of Endemism and Vegetation

Recent Botanical Survey Initiatives

Future Botanical Work

Historical Exploration of Vegetation Patterns in Angola

I ntegration of the Vegetation Map of Angola

Regional and Local Studies on Vegetation Composition

Modern Approaches to Vegetation Mapping and Classification

Outlook: A Call for a New Vegetation Survey of Angola

Exploration of Geoxylic Grasslands

Suffrutex Flora and Endemism

nvironmental Conditions of Suffrutex-Grasslands

The substrate strongly influences the species composition of the suffrutex-

nowledge Gaps on the Evolution of the Geoxylic Suffrutices