FRUIT FLIES (DIPTERA, TEPHRITIDAE) OF KAKAMEGA FOREST,
KENYA
Author(s): Robert S. Copeland, Wilberforce Okeka, Amnon Freidberg, Bernhard Merz, Ian M. White,
Marc De Meyer, and Quentin Luke
Source: Journal of East African Natural History, 94(2):247-278.
Published By: Nature Kenya/East African Natural History Society
DOI: http://dx.doi.org/10.2982/0012-8317(2005)94[247:FFDTOK]2.0.CO;2
URL: http://www.bioone.org/doi/full/10.2982/0012-8317%282005%2994%5B247%3AFFDTOK
%5D2.0.CO%3B2
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Journal of East African Natural History 94(2): 247–278 (2005)
FRUIT FLIES (DIPTERA, TEPHRITIDAE) OF KAKAMEGA
FOREST, KENYA
Robert S. Copeland
Department of Entomology, Texas A&M University, College Station, TX, 77843, USA
and
International Centre of Insect Physiology and Ecology, P.O. Box 30772, Nairobi, Kenya
hunteriazeylanica@yahoo.com
Wilberforce Okeka
Forest Department, Kakamega Forest Station, Kakamega, Kenya
wokekasan@yahoo.co.uk
Amnon Freidberg
Department of Zoology, The George S. Wise Faculty of Life Sciences,
Tel Aviv University, Tel Aviv 69978, Israel
afdipter@post.tau.ac.il
Bernhard Merz
Muséum d’Histoire Naturelle, Département d’Entomologie
C.P. 6434, CH–1211 Genève 6, Switzerland
bernhard.merz@mhn.ville-ge.ch
Ian M. White
Department of Entomology, The Natural History Museum, Cromwell Road,
London, SW7 5BD, UK
imw@nhm.ac.uk
Marc De Meyer
Entomology Section, Royal Museum for Central Africa, Leuvensesteenweg 13,
B-3080 Tervuren, Belgium
marc.de.meyer@africamuseum.be
Quentin Luke
East African Herbarium, National Museums of Kenya, Box 45166, Nairobi, Kenya
quentin.luke@swiftkenya.com
ABSTRACT
A list of the Tephritidae from Kakamega Forest and it environs was compiled from
published records, Malaise trap, baited trap, and sweep-net collections, rearings from
flowers and fruits, and examination of museum specimens. One hundred and thirtyfive species are recorded, and collection information is provided for all of them.
248
R.S. Copeland et al.
Twenty-five of these species are undescribed, or were so at the time our study began.
Frugivorous Tephritidae were reared from 134 of 378 (35%) fruit collections made
within the forest. Fruits of 17 of 47 plant families (36%), and 37 of the 123 plant
species (30%) yielded Tephritidae. The plant families of greatest importance in the
life histories of frugivorous species were the Sapotaceae, Rubiaceae, Flacourtiaceae,
Rosaceae and Solanaceae, while the Asteraceae provided hosts for most of the flowerfeeding species. The affinities of the Kakamega forest tephritid fauna with those of
other regions, in particular the main central and western African rain forests and the
East African coastal forests, are discussed.
INTRODUCTION
Fruit flies (Diptera: Tephritidae) are a biologically diverse family of flies whose larvae are
mainly phytophagous, infesting fruits, flower heads or stems of a wide range of host plants.
More than 4000 species are known worldwide (Thompson, 1999), and they comprise a
significant part of regional biotas. Tephritids are also among the most economically
important groups of insects in the Afrotropical Region, being a major constraint to
commercial and subsistence farming in the region. The destructive association of several
species with commercially grown fruit and vegetable crops makes them the subject of
intensive agricultural research.
In the Pleistocene, Kakamega Forest was part of a rain forest belt connected to the
lowland rain forests of central and western Africa (Round-Turner, 1994), and covering large
parts of Kenya and Uganda. Approximately 10 000 years ago, the forest was reduced to a
number of isolated patches in East Africa, with Kakamega the easternmost relict. Human
encroachment and activities have further reduced the size of this unique Guineo-Congolian
remnant in Kenya (Kokwaro, 1988), and more than half of the forested area is considered
disturbed (Beentje, 1990).
Kakamega Forest (0°10’–0°21’N, 34°47’–34°58’E, altitude 1500–1700 m) is considered
an example of dry Guineo-Congolian forest (figure 1). It has a slightly cooler and less humid
climate than the equatorial rain forests of West and Central Africa (Kokwaro, 1988), with a
mean monthly minimum temperature of 14.8–15.2°C, mean monthly maximum of 26.4–
31.1°C, and an average yearly rainfall just below 2000 mm (Ojany & Ogendo, 1988). The
forest reserve area covers slightly less than 20 000 hectares (Wass, 1995) and has a mosaic
structure, with disturbed and undisturbed forest patches, open glades, and plantations of
exotic and indigenous trees. For a complete vegetation survey and history of the use of the
forest, see Kokwaro (1988) and Mutangah et al. (1992).
Kakamega Forest is an area with high conservation value, harbouring many plant and
animal species found nowhere else in Kenya. With regard to biodiversity and its threatened
status, it is listed among the top five forest areas in the country (Wass, 1995). However, the
invertebrate fauna is still poorly known with only a few groups, such as dragonflies
(Clausnitzer, 1999, Clausnitzer, 2005) and hover flies (De Meyer, 2001a) studied in any
detail. As part of an effort to increase knowledge of the biodiversity of this rich area, we
present here all the published and unpublished records, known to us, of tephritids from the
Kakamega Forest area.
Fruit Flies of Kakamega Forest, Kenya
249
Figure 1: The gazetted boundaries of tropical rain forests of Kakamega District and their allies,
the Nandi Forests (taken, with permission, from Kokwaro (1988)).
MATERIAL AND METHODS
Many of the fruit fly records are the result of a Kenya-wide sampling programme conducted
to determine the wild-fruit hosts of tephritids, particularly for species of economic
importance. The survey was a joint project of Texas A&M University and the International
Centre of Insect Physiology and Ecology (TAMU/ICIPE). Sampling and rearing methods for
that programme are described in Copeland et al. (2002, 2004). Fruit collections in Kakamega
Forest were made at approximately monthly intervals from March 1999 to January 2002.
Three spot-samples were collected in 2005. Plant specimens were pressed in the field and
identified later by one of us (QL).
250
R.S. Copeland et al.
Although progress has been made in resolving the higher classification of the Tephritidae
(Korneyev, 1999, Hancock, 2003, Hancock et al., 2003), no lasting consensus has been
reached, and the matter will likely remain unsettled until a molecular phylogeny is
constructed. In general, we list tephritid species according to the recent synopsis of the
higher classification of the Tephritidae (Norrbom et al., 1999). However, we accept
Korneyev’s (1999) addition of the Blepharoneurinae to the three subfamilies (Phytalmiinae,
Trypetinae, and Tephritinae) recognized by Norrbom et al. (1999). Additionally, we follow
the changes made by Hancock (2003) and Hancock et al. (2003) to the classification of the
Tephritinae, the most important of which were the establishment of the new tribes
Axiothaumini and Pliomelaenini.
Under each taxon the collecting date, method, collector, number of specimens, and the
collection depository are given. When exact geographical co-ordinates are known (e.g. from
GPS readings), these are included. In the list, [H] indicates that at least one new host plant
record is presented in the collection data immediately following a species name and, to make
it easy to locate, that record is underlined where it first appears in the text. If the fruit
represents the first host record for a particular tephritid species, the [H] is indicated in bold,
as is the first mention of that host in the text. A new country record for Kenya is indicated by
[K]. A previously undescribed species is indicated with a bold [S].
Not all material could be identified to species (or sometimes even genus) level. Up to date
revisions or identification keys are lacking for several groups, especially for those within the
Schistopterini and Myopitini. For some collections of fruits that yielded both Ceratitis
anonae and Ceratitis fasciventris, a portion of the females has not yet been assigned to
species. These are indicated under the listings for C. anonae.
Many records included in the checklist represent species collected near, but outside of,
Kakamega Forest (e.g. Kaimosi Forest, Malava, and Sabatia). Specimens collected from
these areas are identified as such in the text. We have included these records because these
species are likely to be found in, or on the fringes of, Kakamega Forest. Records that are
unaccompanied by trapping or rearing information were captured by sweep net.
Other abbreviations used in the text are:
AF = A. Freidberg
RSC = R.S. Copeland
BM = B. Merz
TAU = Tel Aviv University, Department of Zoology, Tel Aviv
ICIPE = International Centre of Insect Physiology and Ecology, Nairobi
NMK = National Museums of Kenya, Department of Invertebrate Zoology, Nairobi
MHNG = Natural History Museum, Geneva
NHM = Natural History Museum, London
TAMU/ICIPE = Texas A&M University and International Centre of Insect Physiology
and Ecology tephritid survey
cf. = confer [L.] compare (in the sense of “probably equals” when referring to a species)
m = male
f =female
CHECKLIST
Blepharoneurinae
Baryglossa sp.
11.v.1991, AF & F. Kaplan (1f, TAU); 14.i.1996, I. Yarom & AF (1m, TAU).
Fruit Flies of Kakamega Forest, Kenya
251
Phytalmiinae
Acanthonevrini
Aethiothemara trigona Hendel
8–9.xi.1983, AF (1f, TAU); 20–21.xi.1986, AF (1m, TAU); 11.v.1991, AF & F. Kaplan
(2m, TAU); 0o14.13’N, 34o51.87’E, 3–10.vii.1999, Malaise trap, RSC (1f, RSC);
0o14.13’N, 34o51.87’E, 11–18.ix.1999, Malaise trap, RSC (1, RSC); 0o14.13’N,
34o51.87’E, 16–23.viii.2000, Malaise trap, RSC (1, RSC); 0o14.13’N, 34o51.87’E, 12–
19.ix.2000, Malaise trap, RSC (1, RSC).
Ptiliniola edwardsi Munro
20–21.xi.1986, AF (1f, TAU); 22.xi.1989, AF & F. Kaplan (1m, TAU); 0o14.13’N,
34o51.87’E, 26.iii–2.iv.2000, Malaise trap, RSC (1f, RSC).
Phytalmiini
Sosiopsila sp. cf. metadacus (Speiser)
0o14.13’N, 34o51.87’E, 13–20.ii.2000, Malaise trap, RSC (1m, RSC).
Trypetinae
Adramini
Celidodacus obnubilus (Karsch) (figure 2)
0o14.13’N, 34o51.87’E, 20–27.xi.1999, Malaise trap, RSC (1m, RSC).
Coelopacidia sp.
0o14.13’N, 34o51.87’E, 4–11.xii.1999, Malaise trap, RSC (1, RSC); 0o14.13’N, 34o51.87’E,
30.viii.–5.ix.2000, Malaise trap, RSC (1, RSC).
Coelopacidia sp. cf. apicalis Hendel
20–21.xi.1986, AF (1f, TAU); 30 km N of Kakamega (forest), 23.xi.1986, AF (1m, 1f,
TAU); Sabatia, 12.iii.1993, BM (1m, MHNG).
Coelopacidia vivax (Munro)
14.i.1996, I. Yarom & AF (1f, TAU).
Coelotrypes pulchellus (Bezzi)
0o14.13’N, 34o51.87’E, 13–20.xi.1999, Malaise trap, RSC (1, RSC).
Coelotrypes sp. nr. vittatus (Bezzi)
Malava, 25 km N of Kakamega, 23.xi.1989, AF & F. Kaplan (6m, 2f, TAU); same
collecting data, ex flowerbud Astripomoea malvacea (Klotzsch) A.Meeuse (Convolvulaceae),
9.xii.1989 (pupa: 24.xi.1989) (21m, 8f, TAU).
Coelotrypes sp. cf. vittatus (Bezzi)
Malava, 25 km N of Kakamega, 23.xi.1989, AF & F. Kaplan, ex flowerbud Ipomoea batatas
(L.) Lam. (Convolvulaceae), xii.1989 (1m, TAU).
Coelotrypes sp. cf. inumbratus (Munro)
0o14.13’N, 34o51.87’E, 12–19.iii.2000, Malaise trap, RSC (1f, RSC); same collecting data,
18–25.vi.1999 (1f, RSC).
252
R.S. Copeland et al.
Coelotrypes sp. nov. A [S]
22.xi.1989, AF & F. Kaplan (1m, TAU); 8.xii.1989, ex flowerbud Ipomoea tenuirostris
Choisi ssp. tenuirostris (Convolvulaceae), AF & F. Kaplan (pupa: 23.xi.1989) (1f, TAU);
0o14.13’N, 34o51.87’E, 9–16.iv.2000, Malaise trap, RSC (1f, RSC).
Coelotrypes sp. nov. B [S]
0o14.13’N, 34o51.87’E, 19–26.vi.2000, Malaise trap, RSC (1f, RSC).
Meracanthomyia antennata Hendel
0o14.13’N, 34o51.87’E, 18–25.vii.2000, Malaise trap, RSC (1, RSC).
Trypanophion sp. cf. gigas Bezzi
0o14.13’N, 34o51.87’E, 25.ix–2.x.1999, Malaise trap, RSC (1, RSC).
Carpomyini
Notomma sp. (mutilum group)
10.x.1998, F. Kaplan & AF (1f, TAU).
Taomyia marshalli Bezzi [H] (figure 4)
14.i.1996, I. Yarom & AF (4f, TAU); 4.iii.1999, 0o14.54’N, 34o51.94’E, ex fruits
Dracaena fragrans (L.) Ker Gawl. (Dracaenaceae) (figure 5), ICIPE/USAID coll. no. 31,
RSC (39m, 48f, ICIPE, NMK); 13.iv.1999, 0o14.54’N, 34o51.94’E, ex fruits Dracaena
fragrans, ICIPE/USAID coll. no. 69, RSC (many, not counted, ICIPE, NMK); 25.i.2000,
0o13.44’N, 34o53.44’E, ex fruits Dracaena fragrans, ICIPE/USAID coll. no. 446, RSC
(76m, 74f, ICIPE, NMK); 30.iii.2000, 0o14.16’N, 34o51.82’E, ex fruits Dracaena fragrans,
ICIPE/USAID coll. no. 597, RSC (120m, 140f, ICIPE, NMK); 1.v.2000, 0o14.13’N,
34o51.87’E, ex fruits Dracaena fragrans, ICIPE/USAID coll. no. 647, RSC (19m, 26f,
ICIPE, NMK); 2.vi.2000, 0o14.16’N, 34o51.82’E, ex fruits Dracaena fragrans,
ICIPE/USAID coll. no. 714, RSC (16m, 14f, ICIPE, NMK); 6.vi.2001, 0o13.44’N,
34o53.44’E, ex fruits Dracaena fragrans, ICIPE/USDA coll. no. 1302, RSC (not counted,
ICIPE, NMK); 29.1.2002, 0o13.44’N, 34o53.44’E, ex fruits Dracaena fragrans,
ICIPE/USDA coll. no. 1728, RSC (not counted, ICIPE, NMK).
Dacini
Ceratitidina
Carpophthoromyia amoena Enderlein
8–9.xi.1983, I. Yarom (1m, TAU); 14.i.1996, I. Yarom & AF (1f, TAU).
Carpophthoromyia dimidiata Bezzi (figure 6)
2.v.2000, 0o13.44’N, 34o53.44’E, ex fruits Drypetes gerrardii Hutch. (Euphorbiaceae)
(figure 7), ICIPE/USAID coll. no. 638, RSC (1m, 1f, NMK); 2.vi.2000, 0o13.66’N,
34o53.12’E, ex fruits Drypetes gerrardii, ICIPE/USAID coll. no. 712, RSC (16m, 13f,
ICIPE, NMK).
Ceratitis (Ceratalaspis) argenteobrunnea Munro [H,K]
20.ix.2000, 0o13.06’N, 34o53.98’E, ex fruits Vepris nobilis (Delile) Mziray (Rutaceae),
ICIPE/USAID coll. no. 873, RSC (3m, 8f, ICIPE, NMK).
Fruit Flies of Kakamega Forest, Kenya
253
Ceratitis (Ceratalaspis) connexa (Bezzi) [H,K]
26.ii.2000, 0o15.16’N, 34o51.68’E, ex fruits Synsepalum brevipes (Baker) T.D.Penn.
(Sapotaceae), ICIPE/USAID coll. no. 540, RSC (1f, NMK).
Ceratitis (Ceratalaspis) stictica Bezzi [H,K]
3.vi.2000, 0o13.44’N, 34o53.44’E, ex fruits Craterispermum schweinfurthii Hiern
(Rubiaceae), ICIPE/USAID coll. no. 709, RSC (7m, 3f, ICIPE, NMK); 13.vii.2000,
0o13.44’N, 34o53.44’E, ex fruits Craterispermum schweinfurthii, ICIPE/USAID coll. no.
744, RSC (3m, 4f, ICIPE, NMK); 31.vii.2001, 0o12.35’N, 34o52.88’E, ex fruits
Craterispermum schweinfurthii, ICIPE/USDA coll. no. 1428, RSC (1m, 1f, NMK).
Ceratitis (Ceratalaspis) venusta (Munro) [H]
21.9.2000, 0o13.66’N, 34o53.12’E, ex fruits Solanum anguivi Lam. (Solanaceae),
ICIPE/USAID coll. no. 883, RSC (13m, 10f, ICIPE, NMK).
Ceratitis (Ceratitis) caetrata Munro [H] (figure 8)
31.viii.1999, 0o14.38’N, 34o51.86’E, ex fruits Manilkara butugi Chiov. (Sapotaceae),
ICIPE/USAID coll. no. 273, RSC (3m, 1f, ICIPE, NMK); 14.x.1999, 0o14.20’N,
34o51.82’E, ex fruits Manilkara butugi, ICIPE/USAID coll. no. 309, RSC (2m, 1f, NMK);
14.x.1999, 0o13.14’N, 34o53.76’E, ex fruits Manilkara butugi, ICIPE/USAID coll. no. 311,
RSC (26m, 25f, ICIPE, NMK); 22.xi.1999, 0o14.26’N, 34o51.82’E, ex fruits Manilkara
butugi, ICIPE/USAID coll. no. 360, RSC (1m, 2f, ICIPE, NMK); 23.xi.1999, 0o13.14’N,
34o53.76’E, ex fruits Manilkara butugi, ICIPE/USAID coll. no. 371, RSC (8m, 6f, ICIPE,
NMK); 20.ix.2000, 0o13.14’N, 34o53.76’E, ex fruits Mimusops bagshawei S.Moore
(Sapotaceae) (figure 9), ICIPE/USAID coll. no. 879, RSC (72m, 92f, ICIPE, NMK);
11.x.2000, 0o13.14’N 34o53.76’E, ex fruits Mimusops bagshawei, ICIPE/USAID coll. no.
914, RSC (49m, 39f, ICIPE, NMK); 29.i.2002, 0o14.13’N, 34o51.87’E, ex fruits Manilkara
butugi, ICIPE/USDA coll. no. 1732, RSC (uncounted, NMK).
Ceratitis (Ceratitis) capitata (Wiedemann) [H]
14.x.1999, 0o14.20’N, 34o51.82’E, ex fruits Manilkara butugi, ICIPE/USAID coll. no. 309, RSC
(uncounted, NMK); 14.x.1999, 0o13.14’N, 34o53.76’E, ex fruits Manilkara butugi,
ICIPE/USAID coll. no. 311, RSC (3m, 3f, NMK); 23.xi.1999, 0o13.14’N, 34o53.76’E, ex fruits
Manilkara butugi, ICIPE/USAID coll. no. 371, RSC (uncounted, NMK); 26.i.2000,
0o14.16’N, 34o51.82’E, ex fruits Prunus africana (Hook.f.) Kalkman (Rosaceae),
ICIPE/USAID coll. no. 465, RSC (1m, NMK); 11.x.2000, 0o13.14’N 34o53.76’E, ex fruits
Mimusops bagshawei, ICIPE/USAID coll. no. 914, RSC (1m, 3f, ICIPE, NMK).
Ceratitis (Hoplolophomyia) cristata (Bezzi) [H,K]
15.viii.2000, 0o13.06’N, 34o53.98’E, ex fruits Vepris nobilis, ICIPE/USAID coll. no. 807,
RSC (not counted, NMK); 20.ix.2000, 0o13.06’N, 34 o53.98’E, ex fruits Vepris nobilis,
ICIPE/USAID coll. no. 873, RSC (15m, 14f, ICIPE, NMK); 11.x.2000, 0o13.06’N,
34o53.98’E, ex fruits Vepris nobilis, ICIPE/USAID coll. no. 904, RSC (2f, NMK).
Ceratitis (Pardalaspis) bremii Guérin-Meneville [H,K]
8–9.xi.1983, I. Yarom and AF (2m, TAU); 1.v.2000, 0o14.13’N, 34o51.87’E, ex fruits
Pouteria altissima (A.Chev.) Baehni (Sapotaceae), ICIPE/USAID coll. no. 648, RSC (1m,
2f, ICIPE, NMK).
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R.S. Copeland et al.
Ceratitis (Pardalaspis) ditissima (Munro) [K]
25.ii.2000, 0o14.13’N, 34o51.87’E, ex fruits Chrysophyllum albidum G.Don. (Sapotaceae),
ICIPE/USAID coll. no. 548, RSC (194m, 192f, ICIPE, NMK); 29.iii.2000, 0o14.16’N,
34o51.82’E, ex fruits Chrysophylium albidum, ICIPE/USAID coll. no. 573, RSC (36m, 31f,
ICIPE, NMK); 29.iii.2000, 0o13.14’N, 34o53.76’E, ex fruits Chrysophyllum albidum,
ICIPE/USAID coll. no. 595, RSC (30m, 23f, ICIPE, NMK); 1.v.2000, 0o13.06’N,
34o53.98’E, ex fruits Chrysophyllum albidum, ICIPE/USAID coll. no. 646, RSC (2f, NMK);
7.vi.2000, 0o12.590’N, 34o55.302’E, ex fruits Chrysophyllum albidum, ICIPE/USAID coll.
no. 548, RSC (not counted, NMK).
Ceratitis (Pardalaspis) hamata De Meyer [H,K]
25.i.2000, 0o11.90’N, 34o11.90’E, ex fruits Synsepalum cerasiferum (Welw.) T.D.Penn.
(Sapotaceae), ICIPE/USAID coll. no. 435, RSC (16m, 6f, ICIPE, NMK); 25.ii.2000,
0o11.90’N, 34o11.90’E, ex fruits Synsepalum cerasiferum, ICIPE/USAID coll. no. 534, RSC
(69m, 29f, ICIPE, NMK); 30.iii.2000, 0o11.90’N, 34o11.90’E, ex fruits Synsepalum
cerasiferum, ICIPE/USAID coll. no. 575, RSC (1m, 1f, NMK).
Ceratitis (Pterandrus) anonae Graham [H]
4.iii.1999, 0o14.54’N, 34o51.94’E, ex fruits Antiaris toxicaria (Pers.) Lesch. (Moraceae),
ICIPE/USAID coll. no. 26, RSC (1m, NMK); 3.viii.1999, 0o13.66’N, 34o53.13’E, ex fruits
Solanum mauritianum Scop. (Solanaceae), ICIPE/USAID coll. no. 213, RSC (8m, 2f, 5f
undetermined, ICIPE, NMK); 20.v.1999, 0o14.54’N, 34o51.94’E, ex fruits Pouteria
altissima, ICIPE/USAID coll. no. 133, RSC (10m, 4f, 3f undetermined, ICIPE, NMK);
20.v.1999, 0o14.54’N, 34o51.94’E, ex fruits Englerophytum oblanceolatum (S.Moore)
T.D.Penn. (Sapotaceae), ICIPE/USAID coll. no. 126, RSC (1m, NMK); 19.vi.1999,
0o13.10’N, 34o54.06’E, ex fruits Rawsonia lucida Harv. & Sond. (Flacourtiaceae),
ICIPE/USAID coll. no. 170, RSC (3m, 1f, NMK); 31.viii.1999, 0o14.090’N, 34o52.067’E,
ex fruits Solanum mauritianum, ICIPE/USAID coll. no. 265, RSC (1m, 1f, NMK).
22.xi.1999, 0o15.28’N, 34o51.36’E, ex fruits Leptactina platyphylla (Hiern) Wernham
(Rubiaceae), ICIPE/USAID coll. no. 350, RSC (35m, 41f, ICIPE, NMK); 23.xi.1999,
0o13.14’N, 34o54.14’E, ex fruits Solanum mauritianum, ICIPE/USAID coll. no. 351, RSC
(3m, 1f, 2f undetermined, ICIPE, NMK); 16.xii.1999, 0o14.15.28’N, 34o51.36’E, ex fruits
Solanum mauritianum, ICIPE/USAID coll. no. 396, RSC (3m, 4f, 3f undetermined,
ICIPE,NMK); 16.xii.1999, 0o15.28’N, 34o51.36’E, ex fruits Leptactina platyphylla,
ICIPE/USAID coll. no. 398, RSC (8m, 7f, ICIPE, NMK); 25.i.2000, 0o13.14’N,
34o53.76’E, ex fruits Antiaris toxicaria, ICIPE/USAID coll. no. 450, RSC (16m, 20f,
ICIPE, NMK); 25.i.2000, 0o13.67’N, 34o53.76’E, ex fruits Antiaris toxicaria,
ICIPE/USAID coll. no. 437, RSC (41m, 41f, ICIPE, NMK); 25.i.2000, 0o14.16’N,
34o51.82’E, ex fruits Solanum mauritianum, ICIPE/USAID coll. no. 445, RSC (20m, 14f,
ICIPE, NMK); 26.i.2000, 0o14.54’N, 34o51.94’E, ex fruits Antiaris toxicaria,
ICIPE/USAID coll. no. 439, RSC (33m, 30f, ICIPE, NMK); 25.ii.2000, 0o11.895’N,
34o52.680’E, ex fruits Solanum mauritianum, ICIPE/USAID coll. no. 542, RSC (1m, 5f
undetermined, ICIPE, NMK); 25.ii.2000, 0o11.90’N, 34o52.68’E, ex fruits Antiaris
toxicaria, ICIPE/USAID coll. no. 530, RSC (1m, 1f, NMK); 26.ii.2000, 0o15.16’N,
34o51.68’E, ex fruits Synsepalum brevipes, ICIPE/USAID coll. no. 540, RSC (4m, 5f, 5f
undetermined, ICIPE, NMK); 29.iii.2000, 0o15.16’N, 34o51.68’E, ex fruits Synsepalum
brevipes, ICIPE/USAID coll. no. 577, RSC (5m, 15f undetermined, ICIPE, NMK);
30.iii.2000, 0o14.16’N, 34o51.82’E, ex fruits Antiaris toxicaria, ICIPE/USAID coll. no.
Fruit Flies of Kakamega Forest, Kenya
255
576, RSC (8m, ICIPE, NMK); 30.iii.2000, 0o11.895’N, 34o52.680’E, ex fruits Solanum
mauritianum, ICIPE/USAID coll. no. 585, RSC (18m, 21f undetermined, ICIPE, NMK);
30.iv.2000, 0o14.16’N, 34o51.82’E, ex fruits Solanum mauritianum, ICIPE/USAID coll. no.
640, RSC (4m, 2f undetermined, ICIPE, NMK); 30.iv.2000, 0o14.16’N, 34o51.82’E, ex
fruits Strombosia scheffleri Engl. (Olacaceae), ICIPE/USAID coll. no. 661, RSC (12m, 24f
undetermined, ICIPE, NMK); 1.v.2000, 0o14.13’N, 34o51.87’E, ex fruits Englerophytum
oblanceolatum, ICIPE/USAID coll. no. 641, RSC (12m, 31f undetermined, ICIPE, NMK);
28.v.2000, 0o14.16’N, 34o51.82’E, ex fruits Solanum mauritianum, ICIPE/USAID coll. no.
690, RSC (2m, 3f undetermined, ICIPE, NMK); 2.vi.2000, 0o14.13’N, 34o51.87’E, ex fruits
Englerophytum oblanceolatum, ICIPE/USAID coll. no. 706, RSC (76m, 93f undetermined,
ICIPE, USAID); 2.vi.2000, 0o14.13’N, 34o51.87’E, ex fruits Pouteria altissima,
ICIPE/USAID coll. no. 689, RSC (1m, 1f, 2f undetermined, ICIPE, USAID); 2.vi.2000,
0o12.34’N, 34o53.57’E, ex fruits Tiliacora funifera (Miers.) Oliv. (Menispermaceae),
ICIPE/USAID coll. no. 713, RSC (2m, 1f, NMK); 13.vii.2000, 0o12.34’N, 34o53.57’E, ex
fruits Solanum mauritianum, ICIPE/USAID coll. no. 743, RSC (4m, 3f undetermined,
ICIPE, NMK); 13.vii.2000, 0o12.34’N, 34o53.57’E, ex fruits Psidium guajava L.
(Myrtaceae), ICIPE/USAID coll. no. 750, RSC (41m, 49f undetermined, ICIPE, NMK);
15.viii.2000, 0o13.69’N, 34o53.12’E, ex fruits Solanum mauritianum, ICIPE/USAID coll.
no. 816, RSC (3m, 2f, 7f undetermined, ICIPE, NMK); 16.viii.2000, 0o14.16’N,
34o51.82’E, ex fruits Solanum mauritianum, ICIPE/USAID coll. no. 815, RSC (1m, 2f,
ICIPE, NMK); 12.ix.2000, 0o12.34’N, 34o53.57’E, ex fruits Solanum mauritianum,
ICIPE/USAID coll. no. 895, RSC (3m, 1f undetermined, ICIPE, NMK); 20.ix.2000,
0o14.16’N, 34o51.82’E, ex fruits Solanum mauritianum, ICIPE/USAID coll. no. 889, RSC
(17m, 16f undetermined, ICIPE, NMK); 18.ii.2001, 0o14.16’N, 34o51.82’E, ex fruits
Antiaris toxicaria, ICIPE/USDA coll. no. 1056, RSC (2f, NMK); 19.ii.2001, 0o13.66’N,
34o53.12’E, ex fruits Antiaris toxicaria, ICIPE/USDA coll. no. 1057, RSC (69m, 69f,
ICIPE, NMK); 26.iv.2001, 0° 14.16’N, 34o51.82’N, ex fruits Morus mesozygia Stapf. ex
A.Chev. (Moraceae), ICIPE/USDA coll. no. 1199, RSC (1m, NMK); 6.vi.2001, 0o14.13’N,
34o51.87’E, ex fruits Antiaris toxicaria, ICIPE/USDA coll. no. 1312, RSC (3m, 2f, ICIPE,
NMK).
Ceratitis (Pterandrus) fasciventris (Bezzi) [H]
3.viii.1999, 0o13.66’N, 34o53.13’E, ex fruits Solanum mauritianum, ICIPE/USAID coll. no.
213, RSC (1m, NMK); 20.v.1999, 0o14.54’N, 34o51.94’E, ex fruits Pouteria altíssima,
ICIPE/USAID coll. no. 133, RSC (1f, NMK); 19.vi.1999, 0o13.10’N, 34o54.06’E, ex fruits
Rawsonia lucida, ICIPE/USAID coll. no. 170, RSC (4m, 1f, NMK); 31.viii.1999,
0o14.090’N, 34o52.067’E, ex fruits Solanum mauritianum, ICIPE/USAID coll. no. 265, RSC
(2f, NMK); 23.xi.1999, 0o13.14’N, 34o54.14’E, ex fruits Solanum mauritianum,
ICIPE/USAID coll. no. 351, RSC (8m, 11f, ICIPE, NMK); 16.xii.1999, 0o14.15.28’N,
34o51.36’E, ex fruits Solanum mauritianum, ICIPE/USAID coll. no. 396, RSC (6m, 6f,
ICIPE, NMK); 25.i.2000, 0o14.16’N, 34o51.82’E, ex fruits Solanum mauritianum,
ICIPE/USAID coll. no. 445, RSC (27m, 31f, ICIPE, NMK); 26.i.2000, 0o14.16’N,
34o51.82’E, ex fruits Prunus africana, ICIPE/USAID coll. no. 436, RSC (1m, NMK);
25.ii.2000, 0o11.895’N, 34o52.680’E, ex fruits Solanum mauritianum, ICIPE/USAID coll.
no. 542, RSC (not counted, ICIPE, NMK); 26.ii.2000, 0o15.16’N, 34o51.68’E, ex fruits
Synsepalum brevipes, ICIPE/USAID coll. no. 540, RSC (20m, 13f, ICIPE, NMK);
29.iii.2000, 0o15.16’N, 34o51.68’E, ex fruits Synsepalum brevipes, ICIPE/USAID coll. no.
577, RSC (6m, ICIPE, NMK); 30.iii.2000, 0o11.895’N, 34o52.680’E, ex fruits Solanum
256
R.S. Copeland et al.
mauritianum, ICIPE/USAID coll. no. 585, RSC (1m, NMK); 30.iv.2000, 0o14.16’N,
34o51.82’E, ex fruits Solanum mauritianum, ICIPE/USAID coll. no. 640, RSC (not counted,
ICIPE, NMK). 30.iv.2000, 0o14.16’N, 34o51.82’E, ex fruits Strombosia scheffleri,
ICIPE/USAID coll. no. 661, RSC (14m, ICIPE, NMK); 1.v.2000, 0o14.13’N, 34o51.87’E,
ex fruits Englerophytum oblanceolatum, ICIPE/USAID coll. no. 641, RSC (9m, ICIPE,
NMK); 28.v.2000, 0o14.16’N, 34o51.82’E, ex fruits Solanum mauritianum, ICIPE/USAID
coll. no. 690, RSC (not counted, ICIPE, NMK); 2.vi.2000, 0o14.13’N, 34o51.87’E, ex fruits
Englerophytum oblanceolatum, ICIPE/USAID coll. no. 706, RSC (15m, ICIPE, USAID);
2.vi.2000, 0o14.13’N, 34o51.87’E, ex fruits Pouteria altissima, ICIPE/USAID coll. no. 689,
RSC (1m, USAID); 13.vii.2000, 0o12.34’N, 34o53.57’E, ex fruits Solanum mauritianum,
ICIPE/USAID coll. no. 743, RSC (not counted, ICIPE, NMK); 13.vii.2000, 0o12.34’N,
34o53.57’E, ex fruits Psidium guajava, ICIPE/USAID coll. no. 750, RSC (not counted,
ICIPE, NMK); 15.viii.2000, 0o13.69’N, 34o53.12’E, ex fruits Solanum mauritianum,
ICIPE/USAID coll. no. 816, RSC (not counted, ICIPE, NMK); 16.viii.2000, 0o14.16’N,
34o51.82’E, ex fruits Solanum mauritianum, ICIPE/USAID coll. no. 815, RSC (not counted,
ICIPE, NMK); 17.viii.2000, 0o13.66’N, 34o52.10’E, ex fruits Solanum mauritianum,
ICIPE/USAID coll. no. 814, RSC (1m, NMK); 12.ix.2000, 0o12.34’N, 34o53.57’E, ex fruits
Solanum mauritianum, ICIPE/USAID coll. no. 895, RSC (not counted, ICIPE, NMK);
20.ix.2000, 0o14.16’N, 34o51.82’E, ex fruits Solanum mauritianum, ICIPE/USAID coll. no.
889, RSC (3m, NMK); 1.xi.2000, 0o12.990’N, 34o53.080’E, ex fruits Psidium guajava,
ICIPE/USAID coll. no. 1501, RSC (not counted, ICIPE, NMK).
Ceratitis (Pterandrus) flexuosa (Walker) [H,K]
4.iii.1999, 0o14.54’N, 34o51.94’E, ex fruits Antiaris toxicaria, ICIPE/USAID coll. no. 26,
RSC (1m, 3f, ICIPE, NMK); 25.i.2000, 0o13.67’N, 34o53.76’E, ex fruits Antiaris toxicaria,
ICIPE/USAID coll. no. 437, RSC (141m, 121f, ICIPE, NMK); 25.i.2000, 0o13.14’N,
34o53.76’E, ex fruits Antiaris toxicaria, ICIPE/USAID coll. no. 450, RSC (71m, 63f,
ICIPE, NMK); 26.i.2000, 0o14.54’N, 34o51.94’E, ex fruits Antiaris toxicaria,
ICIPE/USAID coll. no. 439, RSC (44m, 56f, ICIPE, NMK); 30.iii.2000, 0o14.16’n,
34o51.82’E, ex fruits Antiaris toxicaria, ICIPE/USAID coll. no. 576, RSC (151m, 168f,
ICIPE, NMK); 18.ii.2001, 0o14.16’N, 34o51.82’E, ex fruits Antiaris toxicaria,
ICIPE/USDA coll. no. 1056, RSC (23m, 15f, ICIPE, NMK); 19.ii.2001, 0o13.66’N,
34o53.12’E, ex fruits Antiaris toxicaria, ICIPE/USDA coll. no. 1057, RSC (400m, 466f,
ICIPE, NMK); 6.vi.2001, 0o14.13’N, 34o51.87’E, ex fruits Antiaris toxicaria, ICIPE/USDA
coll. no. 1312, RSC (1m, NMK).
Ceratitis (Pterandrus) rubivora Coquillett [H]
Sabatia, 12.iii.1993, BM (1m, MHNG), 13.x.1999, 0o15.275’N, 34o51.358’E, ex fruits
Rubus niveus Thunb. (Rosaceae), ICIPE/USAID coll. no. 320, RSC (1m, NMK);
16.xii.1999, 0o15.28’N, 34o51.36’E, ex fruits Rubus pinnatus Willd. (Rosaceae),
ICIPE/USDA coll. no. 386, RSC (1m, NMK); 24.i.2000, 0o15.28’N, 34o51.36’E, ex fruits
Rubus pinnatus, ICIPE/USDA coll. no. 442, RSC (4m, 11f, ICIPE, NMK); 2.vi.2000,
0o12.34’N, 34o53.57’E, ex fruits Rubus pinnatus, ICIPE/USAID coll. no. 696, RSC (1f,
NMK); 13.vii.2000, 0o12.34’N, 34o53.57’E, ex fruits Rubus pinnatus, ICIPE/USAID coll.
no. 755, RSC (4m, 5f, ICIPE, NMK); 31.x.2001, 0o13.045’N, 34o54.077’E, ex fruits
Rubus apetalus Poir. (Rosaceae), ICIPE/USDA coll. no. 1482, RSC (not counted, ICIPE,
NMK).
Fruit Flies of Kakamega Forest, Kenya
257
Ceratitis (Pterandrus) sp. nov. “CO” [S,H,K]
26.ii.2000, 0o15.16’N, 34o51.68’E, ex fruits Synsepalum brevipes, ICIPE/USAID coll. no.
540, RSC (2m, ICIPE, NMK).
This species, new to science, will be formally described in a paper dealing with the subgenus
Pterandrus (De Meyer & Freidberg, in press).
Ceratitoides nigromaculatus Hendel [K]
23–30.iv.2000, Malaise trap, RSC (1f, NMK).
Perilampsis sp.
20–21.xi.1986, AF (1f, TAU); 30 km N of Kakamega (forest), 23.xi.1986, AF (1m, TAU).
Nippia sp. nov. nr. reticulata Hancock [S]
25 km S of Kakamega, 19.xi.1986, AF (2m,1f, TAU); Rt. A1, 15 km S of Kakamega,
10.v.1991, AF & F. Kaplan (1m, TAU).
Trirhithrum albomaculatum (Röder) [H]
13.iv.1999, 0o13.14’N, 34o54.14’E, ex fruits Dovyalis macrocalyx (Oliv.) Warb.
(Flacourtiaceae), ICIPE/USAID coll. no. 66, RSC (5m, 5f, ICIPE, NMK); 1.v.2000,
0o14.13’N, 34o51.87’E, ex fruits Dovyalis macrocalyx, ICIPE/USAID coll. no. 659, RSC
(9m, 4f, ICIPE, NMK); 26.iv.2001, 0o14.16’N, 34o51.82’E, ex fruits Dovyalis macrocalyx,
ICIPE/USDA coll. no. 1207, RSC (7m, 7f, ICIPE, NMK).
Trirhithrum coffeae Bezzi [H]
14.i.1996, I. Yarom & AF (1m, TAU). 23.xi.1999. 0o13.14’N, 34o53.76’E, ex fruits Coffea
eugenioides S.Moore (Rubiaceae), ICIPE/USAID coll. no. 342, RSC (3m, 3f, ICIPE,
NMK); 9.xii.1999, 0o13.14’N, 34o53.76’E, ex fruits Coffea eugenioides, ICIPE/USAID coll.
no. 383, RSC (1m, 1f, ICIPE, NMK); 15.xii.1999, 0o13.14’N, 34o53.76’E, ex fruits Coffea
eugenioides, ICIPE/USAID coll. no. 401, RSC (80m, 83f, ICIPE, NMK); 25.i.2000,
0o13.14’N, 34o53.76’E, ex fruits Coffea eugenioides, ICIPE/USAID coll. no. 470, RSC (1m,
3f, ICIPE, NMK).
Trirhithrum demeyeri White [S,H,K]
13.iv.1999, 0o14.54’N, 34º 54.14’E, ex fruits Rawsonia lucida, ICIPE/USAID coll. no. 73,
RSC (2m, 1f, ICIPE, NMK); 19.vi.1999, 0o13.10’N, 34o54.06’E, ex fruits Rawsonia lucida,
ICIPE/USAID coll. no. 170, RSC (4m, 1f, ICIPE, NMK); 3.viii.1999, 0o13.10’N,
34o54.06’E, ex fruits Rawsonia lucida, ICIPE/USAID coll. no. 217, RSC (2m, 4f, ICIPE,
NMK); 7.vi.2001, 0o13.14’N, 34o53.76’E, ex fruits Rawsonia lucida, ICIPE/USDA coll. no.
1303, RSC (many, uncounted, ICIPE, NMK).
Trirhithrum fraternum Munro [H,K]
3.vi.2000, 0o13.44’N, 34o53.44’E, ex fruits Craterispermum schweinfurthii, ICIPE/USAID
coll. no. 709, RSC (4m, 1f, ICIPE, NMK); 13.vii.2000, 0o13.44’N, 34o53.44’E, ex fruits
Craterispermum schweinfurthii, ICIPE/USAID coll. no. 744, RSC (9m, 7f, ICIPE, NMK).
Trirhithrum inscriptum (Graham) [H,K] (figure 10)
1.v.2000, 0o12.34’N, 34o53.57’E, ex fruits Culcasia falcifolia Engl. (Araceae) (figure 11),
ICIPE/USAID coll. no. 639, RSC (6m, 8f, ICIPE, NMK); 2.vi.2000, 0o12.34’N,
258
R.S. Copeland et al.
34o53.57’E, ex fruits Culcasia falcifolia, ICIPE/USAID coll. no. 702, RSC (2m, 2f,
ICIPE,NMK); 13.vii.2000, 0o12.34’N, 34o53.57’E, ex fruits Culcasia falcifolia,
ICIPE/USAID coll. no. 753, RSC (1f, NMK); 26.iv.2001, 0o14.16’N, 34o51.82’E, ex fruits
Culcasia falcifolia, ICIPE/USDA coll. no. 1201, RSC (11m, 19f, ICIPE, NMK); 6.vi.2001,
0o14.13’N, 34o51.87’E, ex fruits Culcasia falcifolia, ICIPE/USDA coll. no. 1304, RSC (not
counted, ICIPE, NMK).
Trirhithrum meladiscum Munro [H]
22.xi.1999, 0o15.28’N, 34o51.36’E, ex fruits Psychotria peduncularis (Salisb.) Steyerm.
(Rubiaceae), ICIPE/USAID coll. no. 359, RSC (7m, 6f, ICIPE, NMK); 16.xii.1999,
0o15.28’N, 34o51.36’E, ex fruits Psychotria peduncularis, ICIPE/USAID coll. no. 387, RSC
(34m, 31f, ICIPE, NMK); 24.i.2000, 0o15.28’N, 34o51.36’E, ex fruits Psychotria
peduncularis, ICIPE/USAID coll. no. 473, RSC (7m, 17f, ICIPE, NMK); 20.ii.2001,
0o15.28’N, 34o51.36’E, ex fruits Psychotria peduncularis, ICIPE/USDA coll. no. 1060, RSC
(2m, 6f, ICIPE, NMK).
Trirhithrum micans Munro [K]
14.i.1996, I. Yarom & AF (1f, TAU).
Trirhithrum nigrum (Graham) [H,K]
11.v.1991, AF & F. Kaplan (1f, TAU). 1.v.2000, 0o12.34’N, 34o53.57’E, ex fruits Culcasia
falcifolia, ICIPE/USAID coll. no. 639, RSC (24m, 22f, ICIPE, NMK); 2.vi.2000,
0o12.34’N, 34o53.57’E, ex fruits Culcasia falcifolia, ICIPE/USAID coll. no. 702, RSC (1m,
2f, NMK); 13.vii.2000, 0o12.34’N, 34o53.57’E, ex fruits Culcasia falcifolia, ICIPE/USAID
coll. no. 753, RSC (2f, NMK); 26.iv.2001, 0o14.16’N, 34o51.82’E, ex fruits Culcasia
falcifolia, ICIPE/USDA coll. no. 1201, RSC (11m, 10f, ICIPE, NMK); 6.vi.2001,
0o14.13’N, 34o51.87’E, ex fruits Culcasia falcifolia, ICIPE/USDA coll. no. 1304, RSC (not
counted, ICIPE, NMK).
Trirhithrum occipitale Bezzi
22.xi.1989. AF & F. Kaplan (2m, 1f, TAU). 14.i.1999. I. Yarom & AF (5m, 6f, TAU).
Sabatia, 12.iii.1993. B. Merz (1m, MHNG).
Trirhithrum overlaeti Munro [H,K]
31.x.2001, 0o13.806’N, 34o51.983’E, ex fruits Solanum terminale Forssk. (Solanaceae),
ICIPE/USDA coll. no. 1486, RSC (1f, NMK).
Dacina
Bactrocera amplexa (Munro) [H] (figure 12)
30.iv.2000, 0o14.16’N, 34o51.82’E, ex fruits Strombosia scheffleri (figure 13), ICIPE/USAID
coll. no. 661, RSC (88m, 65f, 91 sex undet., ICIPE, NMK); 2.vi.2000, 0o14.13’N, 34o51.87’E,
ex fruits Strombosia scheffleri, ICIPE/USAID coll. no. 699, RSC (2m, 1f, ICIPE, NMK).
Bactrocera munroi White [S,H,K] (figure 14)
13.iv.1999, 0o13.14’N, 34o54.14’E, ex fruits Prunus africana, ICIPE/USAID coll. no. 67,
RSC (3m, 1f, ICIPE/NMK); 29.iii.2000, 0o14.13’N, 34o51.87’E, ex fruits Prunus africana,
ICIPE/USAID coll. no. 574, RSC (1f, NMK). 14.viii.2000, 0o14.13’N, 34o51.87’E, ex
fruits Olea welwitschii (Knobl.) Gilg & G.Schellenb (figure 15). (Oleaceae), ICIPE/USAID
Fruit Flies of Kakamega Forest, Kenya
259
coll. no. 821, RSC (1f, NMK); 15.viii.2000, 0o13.66’N, 34o53.13’E, ex fruits Olea
welwitschii, ICIPE/USAID coll. no. 817, RSC (1f, NMK); 12.ix.2000, 0o14.13’N,
34o51.87’E, ex fruits Olea welwitschii, ICIPE/USAID coll. no. 884, RSC (1m, 2f, NMK).
Dacus arcuatus Munro
19.ii.2001, 0o13.175’N, 34o53.377’E, ex fruits Pergularia daemia (Forssk.) Chiov.
(Apocynaceae), ICIPE/USDA coll. no. 1064, RSC (5m, 5f, ICIPE, NMK).
Dacus armatus Fabricius
13.iii.1993, BM, cue lure trap (34m, MHNG).
Dacus bivittatus (Bigot)
5 km E of Kaimosi, 12.iii.1993, BM (1f, MHNG); 13.iii.1993, BM, cue lure trap (7m,
MHNG); 6.vi.2001, 0o12.34’N, 34o53.57’E, ex fruits Peponium vogelii (Hook.f.) Engl.
(Cucurbitaceae), ICIPE/USDA coll. no. 1292, RSC (2m, NMK); 1.xi.2001, 0o13.09’N,
34o53.28’E, ex fruits Peponium vogelii, ICIPE/USDA coll. no. 1479, RSC (1m, NMK); 16–
23.i.2000, 0o13.14’N, 34o54.14’E, RSC, Malaise trap (1m, NHM).
Dacus croceus Munro [K]
14.i.1996, I. Yarom & AF (1m, TAU).
Dacus fasciolatus Collart
11–18.ix.1999, 0o13.14’N, 34o54.14’E, RSC, Malaise trap (1m, NHM).
Dacus humeralis (Bezzi) [K]
13.iii.1993, BM, cue lure trap (4m, MHNG).
Dacus limbipennis Macquart (figure 16)
3.viii.1999, 0o12.791’N, 34o55.391’E, ex fruits Momordica foetida Schumach.
(Cucurbitaceae), ICIPE/USAID coll. no. 215, RSC (1m, 1f, ICIPE, NMK);
31.viii.1999, 0o13.66’N, 34º 53.12’E, ex fruits Momordica cissoides Benth.
(Cucurbitaceae) (figure 17), ICIPE/USAID coll. no. 260, RSC (12m, 2f, ICIPE, NMK);
25.ii.2000, 0o15.04’N, 34o51.88’E, ex fruits Momordica cissoides, ICIPE/USAID coll.
no. 525, RSC (3m, NMK); 19.ii.2001, 0o12.371’N, 34o53.621’E, ex fruits Momordica
foetida, ICIPE/USDA coll. no. 1059, RSC (8m, 10f, ICIPE, NMK); 6.vii.2001,
0o12.755’N, 34o55.430’E, ex fruits Momordica foetida, ICIPE/USDA coll. no. 1294,
RSC (4m, 4f, ICIPE, NMK);26.iii–3.iv.2000, 0o13.14’N, 34o54.14’E, RSC, Malaise
trap (1f, NHM).
Dacus macer Bezzi [K]
Kakamega, 18.xii.1970, “5200 ft”, A.E. Stubbs (1m, NHM); 18–22.i.1972, 0o15’N,
34o52’E, “5100ft”, C.F. Huggins (1f, NHM).
Dacus maynei Bezzi
12.iii.1993, Sabatia, SE of Kakamega, BM (3m, 2f, MHNG); 12.iii.1993, Kaimosi, 5 km
SE of Kakamega, BM, (1m, MHNG); 23–30.x.1999, 0o13.14’N, 34o54.14’E, RSC, Malaise
trap (1f, NHM).
260
R.S. Copeland et al.
Dacus notalaxus Munro
13.iii.1993, BM, cue lure trap (1m, MHNG).
Dacus punctatifrons Karsch
13.iii.1993, BM, cue lure trap (23m, MHNG); 26.iii–3.iv.2000, RSC, Malaise trap (1f,
NHM); 6.vi.2001, 0o12.34’N, 34o53.57’E, ex fruits Peponium vogelii, ICIPE/USDA coll. no.
1292, RSC (2m, 2f, ICIPE, NMK); 31.x.2001, 0o13.626’N, 34o52.822’E, ex fruits Zehneria
scabra (L.f.) Sond. (Cucurbitaceae), ICIPE/USDA coll. no. 1504, RSC (4f, ICIPE, NMK).
Dacus sphaeristicus Speiser
5 km E of Kaimosi, 12.iii.1993, BM (1m, MHNG); 0o14.13’N, 34o51.87’E, 11–18.ix.1999,
Malaise trap, RSC (1, RSC).
Dacus telfaireae (Bezzi)
Kakamega, December 1956, R. Carcasson (not counted, NMK).
Tephritinae
Axiothaumini
Cryptophorellia flava Freidberg & Hancock
20–21.xi.1986, AF (1m, TAU); 5 km E of Kaimosi, BM (4f, MHNG).
Cryptophorellia longicauda Freidberg & Hancock
20–21.xi.1986, AF (2m, TAU).
Dithrycini
Bezzina margaritifera (Bezzi)
10 km N of Kakamega, 22.xi.1986, AF (3m, TAU); 30 km N of Kakamega (forest),
23.xi.1986, I. Susman (1m, TAU); 5 km E of Kaimosi, 12.iii.1993, BM (1m, MHNG);
13.iii.1993, BM (1, MHNG).
Myopitini
Stamnophora vernoniicola (Bezzi)
Malava, 25 km N of Kakamega, 23.xi.1989, AF & F. Kaplan (13m, 10f, TAU), ex
flowerhead Vernonia adoensis Walp. (Asteraceae), 28.xi–18.xii.1989; 5 km E of Kaimosi,
12.iii.1993, BM (4f, 1 specimen lacking abdomen, MHNG).
Stamnophora sp. nov. [S]
20–21.xi.1986, AF (225, TAU), ex stem gall Vernonia auriculifera Hiern
(Asteraceae); xi–xii.1986; 10 km N of Kakamega, 22.xi.1986, AF (1m, 2f, TAU);
22.xi.1986, AF (5m, 4f, TAU); 15 km N of Kakamega, 22.xi.1986, I. Susman (2f,
TAU); 5 km E of Kaimosi, 12.iii.1993, BM (1m, 1f, MHNG); Sabatia, 12.iii.1993,
BM (2f, MHNG).
Spinicosta sp. nov. A [S]
Malava, 25 km N of Kakamega, 23.xi.1989, AF & F. Kaplan (4m, 1f, TAU); same
collection data, ex flowerhead Vernonia adoensis, 26.xi–18.xii.1989 (2m, TAU); 10 km N of
Kakamega, 22.xi.1986, AF (1m, TAU).
Fruit Flies of Kakamega Forest, Kenya
261
Spinicosta sp. nov. B [S]
Near Kakamega town, 10.x.1986, F. Kaplan & AF (1m, TAU). Ex Vernonia amygdalina
Delile (Asteraceae).
Pliomelaenini
Elaphromyia adatha (Walker)
20–21.xi.1986, AF (4m, TAU); 8–9.xi.1983, AF (1m, TAU); 14.i.1996, I. Yarom & AF
(2f, TAU); 22.xi.1989, AF & F. Kaplan (4f, TAU); 30 km N of Kakamega (forest),
23.xi.1986, AF (3f, TAU); 25 km S of Kakamega, 19.xi.1986, AF (1m, 1f, TAU);
13.iii.1993, BM (1f, MHNG).
Elaphromyia pallida Bezzi
20–21.xi.1986, AF (3m, TAU); 5 km E of Kaimosi, 12.iii.1993, BM (1m, MHNG).
Hyaloctoides semiater (Loew)
5 km E of Kaimosi, 12.iii.1993, BM (1f, MHNG); Sabatia, 12.iii.1993, BM (1f, MHNG).
Platomma nigrantior Munro
Malava, 25 km N of Kakamega, 23.xi.1989, AF & F. Kaplan (1f, TAU).
Pliomelaena brevifrons (Bezzi)
20–21.xi.1986, AF (1f, TAU); cf. P. brevifrons, 11.iii.1993, BM (1f, MHNG); cf.
P. brevifrons, Sabatia, 12.iii.1993, BM (1f, MHNG); 0o14.13’N, 34o51.87’E, 20–
27.xi.1999, Malaise trap, RSC (1, RSC); 0o14.13’N, 34o51.87’E, 4–11.xii.1999, Malaise
trap, RSC (2, RSC).
Pliomelaena brevifrons (Bezzi) var. xyphosiina Bezzi (det. D.L. Hancock)
14.i.1996, I. Yarom & AF (2f, TAU).
Pliomelaena sp. cf. discosa Munro
20–21.xi.1986, AF and I. Susman (2f, TAU); 30 km N of Kakamega (forest), 23.xi.1986
(1f, TAU).
Pliomelaena sp. B.
15 km S of Kakamega, Rt. A1, 10.v.1991, AF & F. Kaplan (1f, TAU).
Pseudafreutreta biseriata (Bezzi)
8–9.xi.1983, AF and I.Yarom (71m, 33f, TAU); 20–21.xi.1986, AF (1f, TAU); 11.v.1991,
AF & F. Kaplan (10m, 11f, TAU); 5 km E of Kaimosi, 12.iii.1993, BM (1m, MHNG).
Pseudafreutreta katangana (Munro)
20–21.xi.1986, AF (4f, TAU), ex inflorescence (covered by froth) Pseuderanthemum
ludovicianum (Büttner) Lindau (Acanthaceae), 5.xii.1986; 0o14.13’N, 34o51.87’E, 20–
27.xi.1999, Malaise trap, RSC (1, RSC).
Pseudafreutreta sp. nov. [S]
11.v.1991, AF & F. Kaplan (14m, 17f, TAU); 14.i.1996, I. Yarom & AF (20m, 19f, TAU).
262
R.S. Copeland et al.
Stephanotrypeta taeniaptera (Bezzi)
10 and 15 km N of Kakamega, 22.xi.1986, AF and I. Susman (4m, 6f, TAU); Sabatia,
12.iii.1993, BM (1f, MHNG); 0o14.13’N, 34o51.87’E, 16–23.i.2000, Malaise trap, RSC (2,
RSC).
Schistopterini
Clematochaeta sp. nov. A [S]
25 km S of Kakamega, 19.xi.1986, AF (1f, TAU).
Clematochaeta sp. nov. B [S]
8–9.xi.1983, AF (21m, 15f, TAU).
Cordylopteryx sp. nov. A [S]
Malava, 25 km N of Kakamega, 23.xi.1989, AF & F. Kaplan (4m, 2f, TAU);
20–21.xi.1986, AF (1m, TAU). Ex Vernonia adoensis.
Cordylopteryx sp. nov. B [S]
8–9.xi.1983, AF (3m, 13f, TAU). Ex Vernonia calvoana (Hook.f.) Hook.f. (Asteraceae).
Eutretosoma sp. nov. A [S]
Near Kakamega town, 10.x.1998, F. Kaplan & AF (2m, 2f, TAU).
Eutretosoma sp. nov. B [S]
Near Kakamega town, 10.x.1998, F. Kaplan & AF (5f, TAU).
Nigroptera nigra (Bezzi)
8–9.xi.1983, AF (1f, TAU); near Kakamega town, 10.x.1998, F. Kaplan & AF (4m, 5f,
TAU); 10 km N of Kakamega, 22.xi.1986, AF (1m, 3f, TAU); 25 km S of Kakamega,
19.xi.1986, AF (2m, 2f, TAU). Ex Aspilia sp. (Asteraceae).
Nigroptera sp. nov. A [S]
20–21.xi.1986, AF (3m, 1f, TAU); 15 km N of Kakamega, 22.xi.1986, I. Susman (1m,
TAU); 25 km S of Kakamega, 19.xi.1986, AF (1m, TAU); 22.xi.1986, AF (2m, 1f, TAU).
Ex Vernonia amygdalina.
Nigroptera sp. nov. B [S]
20–21.xi.1986, AF (2m, 3f, TAU); 22.xi.1986, AF (2f, TAU). Ex Vernonia auriculifera.
Rhochmopterum lutescens (Bezzi)
Near Kakamega town, 10.x.1998, F. Kaplan & AF (1m, TAU), 25 km S of Kakamega,
19.xi.1986, AF (1f, TAU). Hosts various Vernonieae, particularly Bothriocline tomentosa
(Oliv. & Hiern) H.Wild & G.V.Pope (Asteraceae).
Rhochmopterum sp. nov. [S]
25 km S of Kakamega, 19.xi.1986, AF (1m, TAU). Ex Gutenbergia cordifolia Oliv.
(Asteraceae).
Fruit Flies of Kakamega Forest, Kenya
263
In addition, several schistopterine specimens were collected that could not be identified to
genus level. This material is currently under study and seems to harbour several taxa new to
science.
Tephrellini
Afraciura quaternaria (Bezzi)
22.xi.1986, AF (1m, TAU);10 km N of Kakamega, 22.xi.1986, AF (1m, TAU); Sabatia,
12.iii.1993, BM (1m, MHNG).
Brachyaciura sp. (figure 18)
20–21.xi.1986, AF and I. Susman (1m, 1f, TAU); 10 & 11.v.1991, AF & F. Kaplan (4m,
TAU); 25 km S of Kakamega, 19.xi.1986, AF and I. Susman (15m, 8f, TAU); 15 km S of
Kakamega, Rt. A1, 22.xi.1986, AF and I. Susman (10m, 3f, TAU); 10.v.1991, AF and F.
Kaplan (8m, 3f, TAU); 22.xi.1989, AF & F. Kaplan (1m, TAU); Malava, 25 km N of
Kakamega, 23.xi.1989, AF & F. Kaplan (1m, 1f, TAU); 22.xi.1986, AF (2m, TAU) ex
flower Acanthus pubescens (Oliv.) Engl. (Acanthaceae) (figure 19); 11 & 13.iii.1993, BM
(7m, 4f, MHNG); 16 km N of Kakamega, 11.iii.1993, BM (5m, 1f, MHNG); 5 km E of
Kaimosi, 12.iii.1993, BM (11m, 8f, MHNG); Sabatia, 12.iii.1993, BM (6m, 3f, MHNG);
0o14.13’N, 34o51.87’E, 26.iii–2.iv.2000, Malaise trap, RSC (1, RSC); 19.ii.2001,
0o11.90’N, 34o52.68’E, ex fruits Acanthus eminens C.B.Clarke (Acanthaceae), USDA/ICIPE
coll. no. 1051, RSC (many, uncounted, ICIPE, NMK); 0o14.602’N, 34o51.235’E,
20.ii.2001, ex fruits Acanthus pubescens, USDA/ICIPE coll. no. 1068, RSC (61m, 51f,
ICIPE, NMK).
This material represents at least two, and perhaps three, species, one of which might be
B. limbata. They differ in wing pattern and aculeus shape. All are associated with flowers or
fruits of species of Acanthus (Acanthaceae).
Dicheniotes katonae (Bezzi)
11 & 13.iii.1993, BM (2f, MHNG).
Dicheniotes turgens Munro
10.x.1998, F. Kaplan & AF (1m, TAU).
Gymnaciura austeni (Munro)
30 km N of Kakamega (forest), 23.xi.1986, AF (1f, TAU); cf. G. austeni, Sabatia,
12.iii.1993, BM (1f, MHNG).
Gymnaciura sp. nov. [S]
10 km N of Kakamega, 22.xi.1986, AF (1m, TAU).
Metasphenisca sp.
0o14.13’N, 34o51.87’E, 29.iii–2.iv.1999, Malaise trap, RSC (1, RSC); 0o14.13’N,
34o51.87’E, 4–11.xii.1999, Malaise trap, RSC (1, RSC).
Metasphenisca sp. cf. frondifer (Munro)
30 km N of Kakamega (forest), 23.xi.1986, I. Susman (1m, TAU);13.iii.1993, BM (1m, 2f,
MHNG); 5 km E of Kaimosi, 12.iii.1993, BM (1f, MHNG).
264
R.S. Copeland et al.
Ocnerioxyna sp. cf. hemilea Séguy.
0o14.13’N, 34o51.87’E, 4–11.xii.1999, Malaise trap, RSC (1, RSC).
Ocnerioxyna maripilosa (Munro)
22.xi.1986, AF (2m, TAU); 16 km N of Kakamega, 11.iii.1993, BM (1f, MHNG).
Paraciura perpicillaris (Bezzi)
22.xi.1989, AF & F. Kaplan (1m, TAU).
Paraciura sp. [S]
22.xi.1989, AF & F. Kaplan (1f, TAU); 20–21.xi.1986, AF (3m,1f, TAU); 8–9.xi.1983, I.
Yarom (1m, TAU); 11.v.1991, AF & F. Kaplan (1m, TAU); 30 km N of Kakamega
(forest), 23.xi.1986, AF and I. Susman (1m, 1f, TAU); Sabatia, 12.iii.1993, BM (1f,
MHNG).
The specimens at hand probably represent at least two species. Pending a revision of the
genus they are included here, unnamed.
Paraspheniscoides binarius (Loew) var. adeptus (Munro)
25 km S of Kakamega, 19.xi.1986, AF (3m, 2f, TAU); 10 km N of Kakamega, 22.xi.1986,
AF (1m, 1f, TAU); Rt. A1, 15 km S of Kakamega, 10.v.1991, AF & F. Kaplan (3f, TAU);
11.v.1991, AF & F. Kaplan (1m, TAU); 5 km E of Kaimosi, BM (2m, MHNG).
Pediapelta sp. nov. nr. alexina Munro [S]
8–9.xi.1983, AF (4m, TAU).
Pediapelta ternaria (Loew)
16 km N of Kakamega, 11.iii.1993, BM (1m, MHNG).
Tephraciura sphenoptera (Bezzi)
13.iii.1993, BM (1f, MHNG).
Ypsilomena compacta (Bezzi)
10 km N of Kakamega, 22.xi.1986, AF (1m, TAU); Malava, 25 km N of Kakamega,
23.xi.1989, AF & F. Kaplan (1f, TAU).
Tephritini
Actinoptera sp. nov. nr. contacta Munro [S]
11.iii.1993, BM (5m, MHNG).
Antoxya oxynoides (Bezzi)
8–9.xi.1983, AF (1m, TAU); 13.iii.1993, BM (1f, MHNG); Sabatia, 12.iii.1993, BM (1m,
MHNG); 0o14.13’N, 34o51.87’E, 6–13.xi.1999, Malaise trap, RSC (1, RSC).
Campiglossa sp.
0o14.13’N, 34o51.87’E, 20–27.xi.1999, Malaise trap, RSC (1, RSC).
Campiglossa ignobilis (Loew)
5 km E of Kaimosi, 12.iii.1993, BM (1f, MHNG).
Fruit Flies of Kakamega Forest, Kenya
265
Dectodesis sp. nov. [S]
11.iii.1993, BM (1m, 1f, MHNG); Kaimosi, 11.iii.1993 (1f, MHNG); 5 km E of Kaimosi,
12.iii.1993 (1m, MHNG).
Dectodesis sp.
Near Kakamega town, 10.x.1998, F. Kaplan & AF (1m, TAU).
Dectodesis sp. cf. auguralis (Bezzi)
Kaimosi, 11.iii.1993, BM (1m, MHNG).
Dectodesis sp. cf. hexapoda (Bezzi)
11.v.1991, AF & F. Kaplan (6m, 7f, TAU).
Dioxyna sororcula (Wiedemann)
Kaimosi, 11.iii.1993, BM (many, MHNG); 5 km E of Kaimosi, 12.iii.1993, BM (many,
MHNG); Sabatia, 12.iii.1993, BM (many, MHNG).
Gymnosagena sp. nov. [S]
20–21.xi.1986, AF (1, TAU).
Mastigolina sp. nov. nr. bequaerti (Munro) [S]
Kaimosi, 11.iii.1993, BM (1f, MHNG).
Scedella dissoluta (Loew)
5 km E of Kaimosi, 12.iii.1993, BM (1m, MHNG); Sabatia, 12.iii.1993, BM (2f, MHNG).
Scedella sp. cf. cyana (Walker)
Near Kakamega town, 10.x.1998, F. Kaplan & AF (1m, 2f, TAU); Malava, 25 km N of
Kakamega, 23.xi.1989, AF & F. Kaplan (2m, TAU), ex Vernonia sp. (Asteraceae).
Scedella glebosa Munro
Kaimosi, 11.iii.1993, BM (1f, MHNG); 5 km E of Kaimosi, 12.iii.1993, BM (3m, 6f,
MHNG); Sabatia, 12.iii.1993, BM (5m, 1f, MHNG).
Scedella incurva Munro
10 km N of Kakamega, 22.xi.1986, AF (4m, TAU); 25 km S of Kakamega, 19.xi.1986, AF
(1m, TAU); 11.iii.1993, BM (2m, MHNG); 5 km E of Kaimosi, 12.iii.1993, BM (1,
MHNG); Sabatia, 12.iii.1993, BM (3m, 1f, MHNG).
Scedella praetexta (Loew)
Near Kakamega town, 10.x.1998, F. Kaplan & AF (1m, TAU); 25 km S of Kakamega,
19.xi.1986, AF (2m, TAU); 10 km N of Kakamega, 22.xi.1986, AF (1f, TAU); 11 &
13.iii.1993, BM (4m, 1f, MHNG); 5 km E of Kaimosi, 12.iii.1993, BM (2m, 1f, MHNG);
Sabatia, 12.iii.1993, BM (1m, MHNG).
Scedella sandoana Munro
11.iii.1993, BM (1f, MHNG); 5 km E of Kaimosi, 12.iii.1993, BM (1m, MHNG); Sabatia,
12.iii.1993, BM (1m, MHNG).
266
R.S. Copeland et al.
Scedella spp.
Additional unidentified material, probably including additional spp.
Spathulina acroleuca (Schiner)
30 km N of Kakamega (forest), 23.xi.1986, AF (1f, TAU); 5 km E of Kaimosi, 12.iii.1993,
BM (1m, MHNG).
Stelladesis lamborni (Munro)
Kaimosi, 11.iii.1993, BM (1m, MHNG); 5 km E of Kaimosi, 12.iii.1993, BM (5m, 6f,
MHNG); Sabatia, 12.iii.1993, BM (3m, 1f, MHNG).
Stelladesis sp. nov. nr. lutescens (Bezzi) [S]
5 km E of Kaimosi, 12.iii.1993, BM (1m, 1f, MHNG); Sabatia, 12.iii.1993, BM (1f,
MHNG).
Stelladesis woodi (Bezzi)
5 km E of Kaimosi, 12.iii.1993, BM (1m, MHNG); Sabatia, 12.iii.1993, BM (1m, 1f,
MHNG).
In addition many unidentified specimens are housed in the collections of the MHNG, pending
a revision of the genus.
Telaletes sp.
8–9.xi.1983, AF (1m, 2f, TAU); 5 km E of Kaimosi, 12.iii.1993, BM (1f, MHNG);
Sabatia, 12.iii.1993, BM (1m, MHNG); 14.i.1996, I. Yarom & AF (1f, TAU).
Trupanea sp. cf. tubulata Munro
0o14.13’N, 34o51.87’E, 11–18.ix.1999, Malaise trap, RSC (1, RSC).
Trupanodesis aurea (Bezzi)
8–9.xi.1983, AF (1f, TAU); 20–21.xi.1986, AF (1m, TAU); 11.v.1991, AF & F. Kaplan
(1f, TAU); 14.i.1996, I. Yarom & AF (1m, 1f, TAU); 22.xi.1986, AF (1m, TAU); 5 km
E of Kaimosi, 12.iii.1993, BM (4m, 2f, MHNG); Sabatia, 12.iii.1993, BM (1m, 2f,
MHNG).
Terelliini
Craspedoxantha polyspila Bezzi
5 km E of Kaimosi, 12.iii.1993, BM (1m, MHNG).
Craspedoxantha vernoniae Freidberg
Malava, 25 km N of Kakamega, 23.xi.1989, AF & F. Kaplan (1f, TAU); 5 km E of
Kaimosi, 12.iii.1993, BM (2m, 1f, MHNG); Sabatia, 12.iii.1993, BM (1f, MHNG).
Craspedoxantha yaromi Freidberg
5 km E of Kaimosi, 12.iii.1993, BM (13m, 6f, MHNG); Sabatia, 12.iii.1993, BM (3m, 2f,
MHNG).
Fruit Flies of Kakamega Forest, Kenya
267
RESULTS AND DISCUSSION
Tephritidae from the Kakamega area
A total of 135 species of Tephritidae are recognized from Kakamega Forest and the
surrounding area of which at least 25 (18%) are undescribed, or were so when this study
began (table 1). These species represent 14 tribes and 59 genera from the four subfamilies of
Tephritidae. Over half of these species are Tephritinae, most of which breed in flowers.
These latter records reflect the extensive collections made in open areas near Kakamega
Forest by AF and BM.
Table 1. Tephritidae recorded from the Kakamega area.
Subfamily
Blepharoneurinae
Phytalmiinae
Tribe
Blepharoneurini
Acanthonevrini
Trypetinae
Phytalmiini
Adramini
Subtribe
Carpomyini
Dacini
Ceratitidina
Dacina
Tephritinae
Axiothaumini
Dithrycini
Myopitini
Pliomelaenini
Schistopterini
Tephrellini
Genus
Baryglossa
Aethiothemara
Ptiliniola
Sosiopsila
Celidodacus
Coelopacidia
Coelotrypes
Meracanthomyia
cf Trypanophion
Notomma
Taomyia
Carpophthoromyia
Ceratitis
Ceratitoides
Perilampsis
Nippia
Trirhithrum
Bactrocera
Dacus
Cryptophorellia
Bezzina
Stamnophora
Spinicosta
Elaphromyia
Hyaloctoides
Platomma
Pliomelaena
Pseudafreutreta
Stephanotrypeta
Clematochaeta
Cordylopteryx
Eutretosoma
Nigroptera
Rhochmopterum
Afraciura
No. of
species
1
1
1
1
1
3
6
1
1
1
1
2
15
1
1
1
10
2
12
2
1
2
2
2
1
1
4
3
1
2
2
2
3
2
1
No. of
undescribed
species
0
0
0
0
0
0
2
0
0
0
0
0
1
0
0
1
1
1
0
0
0
1
2
0
0
0
0
1
0
2
2
2
2
1
0
268
Subfamily
Tephritinae
R.S. Copeland et al.
Tribe
Tephrellini
Tephritini
Terelliini
Subtribe
Genus
Brachyaciura
Dicheniotes
Gymnaciura
Metasphenisca
Ocnerioxyna
Paraciura
Paraspheniscoides
Pediapelta
Tephraciura
Ypsilomena
Actinoptera
Antoxya
Campiglossa
Dectodesis
Dioxyna
Gymnosagena
Mastigolina
Scedella
Spathulina
Stelladesis
Telaletes
Trupanea
Trupanodesis
Craspedoxantha
No. of
species
2
2
2
2
2
2
1
2
1
1
1
1
2
4
1
1
1
6
1
3
1
1
1
3
No. of
undescribed
species
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
1
1
0
0
1
0
0
0
0
Most of the classes of larval feeding behaviour among Tephritidae are represented in
the species associated with Kakamega Forest, including fruit-pulp feeders (most
Ceratitidina and Dacina species; White & Elson-Harris, 1992), seed predators
(Taomyia), stem borers (Coelopacidia; Hancock, 1986), stem-gall formers (Notomma;
Munro, 1952), specialists on the fruit capsules or flowers of the Acanthaceae, Lamiaceae
and Verbenaceae (Pliomelaenini and Tephrellini; Munro, 1947), and strict flower
feeders. Among the latter are species specializing on cucurbitaceous flowers
(Baryglossa; Norrbom & Condon, 2000), those feeding on buds of the Convolvulaceae
(Coelotrypes; Hancock, 1986), and the many species that exploit the great diversity of
afrotropical Asteraceae, including flower-head feeders that induce gall formation
(Stamnophora; Freidberg & Norrbom, 2000) and those that do not (e.g. Craspedoxantha;
Freidberg, 1985). Table 2 lists the known hosts of flower-feeding Tephritidae in the
Kakamega area.
Among the species absent from collections made in Kakamega Forest were the
grass borers (Trypetinae, Dacini, Gastrozonina), diverse in Asia but represented in
Africa by only a single genus, Bistrispinaria Speiser (Hancock, 1999). There are
numerous natural, open glades within the forest (apparently a result of suppresion of
tree growth by subsurface rock (Kokwaro, 1988)) that support grass populations used
primarily as thatching material by the local people. Additionally, wild grasses are
found thoughout the local area on farms, particularly in fallow fields. In western
Kenya, Bistrispinaria magniceps (Bezzi) and Bistrispinaria fortis (Speiser) have
recently been reared from the wild grasses Panicum maximum Jacq. and Pennisetum
Fruit Flies of Kakamega Forest, Kenya
269
polystachion (L.) Schult., respectively (RSC, unpubl. data). Both grass species occur
around Kakamega, and further searches may reveal the presence of Bistrispinaria
species within or near the forest. It should be added that Ceratitoides, which is found
in Kakamega, but for which hosts are unknown, may prove to be a grass-breeder (D.L.
Hancock, pers. comm.).
Table 2. Summary of known flower hosts of Tephritidae from the Kakamega Forest area.1
Plant family
Acanthaceae
Asteraceae
Plant species
Acanthus pubescens
Pseuderanthemum ludovicianum
Aspilia sp.
Bothriocline tomentosa
Gutenbergia cordifolia
Vernonia adoensis
Vernonia amygdalina
Tephritid species
Brachyaciura sp.
Pseudafreutreta katangana
Nigroptera nigra
Rhochmopterum lutescens
Rhochmopterum sp. nov.
Stamnophora vernoniicola
Spinicosta sp. nov. B
Nigroptera sp. nov. A
Cordylopteryx sp. nov. B
Scedella sp. cf. cyana
Coelotrypes sp. nr. vittatus
Coelotrypes sp. cf. vittatus
Coelotrypes sp. nov. A
Vernonia calvoana
Vernonia sp.
Convolvulaceae
Astripomoea malvacea
Ipomoea batatas
Ipomoea tenuirostris ssp.
tenuirostris
1
Most flowers collected in open areas within 25 km of the forest
Tephritidae reared from fruits
A total of 378 collections comprising 97 001 fruits (112.5 kg) were made in Kakamega Forest.
Of these collections, the plant species was identified for all but 11 samples, and no tephritids
were reared from the few unidentified fruits. Tephritids were reared from 134 of the 378
samples (35%), from fruits of 17 of 47 plant families (36%), and 37 of the 123 plant species
(30%) collected in the forest. The percentages of families and species acting as hosts of
tephritids in Kakamega Forest were very similar to those from wet, highland forests in central
Kenya (37.5% and 30.2%, respectively; number of samples=476) and from the Shimba Hills,
a wet coastal forest (34.8% and 24.4% respectively; number of samples=465) (RSC, unpubl.
data). Table 3 summarizes the known hosts of frugivorous Tephritidae in Kakamega Forest.
Among these hosts, species of Sapotaceae stand out as particularly important for the diversity
of Ceratitis species in Kakamega Forest. Sapotaceous fruits were hosts to nine Ceratitis species,
among them three species (C. anonae, C. capitata, and C. fasciventris) that are pests of
commercial fruits. A single collection of Synsepalum brevipes in Kakamega Forest produced
four tephritid species, the highest number from any fruit collection (n=3701) made during five
years of sampling throughout Kenya (RSC, unpubl. data). All seven species of Sapotaceae that
occur in Kakamega Forest were tephritid hosts, including fruits of both canopy
(Chrysophyllum, Manilkara, Mimusops, Pouteria, Synsepalum) and understory
(Englerophytum) tree species.
Table 4 provides more detail of the relative importance of plant families as hosts of fruitfeeding tephritids. Although sampling effort was not equal among families, the data
provide strong evidence for the importance of several plant families (e.g. Sapotaceae,
Rubiaceae, Solanaceae, Rosaceae, Flacourtiaceae, Moraceae) in fruit fly production.
270
R.S. Copeland et al.
Table 3. Summary of known fruit hosts of Tephritidae from Kakamega Forest.
Plant family
Acanthaceae
Apocynaceae
Araceae
Cucurbitaceae
Plant species
Acanthus eminens
Acanthus pubescens
Pergularia daemia
Culcasia falcifolia
Momordica cissoides
Momordica foetida
Peponium vogelii
Dracaenaceae
Euphorbiaceae
Flacourtiaceae
Zehneria scabra
Dracaena fragrans
Drypetes gerrardii
Dovyalis macrocalyx
Rawsonia lucida
Menispermaceae
Moraceae
Tiliacora funifera
Antiaris toxicaria
Myrtaceae
Morus mesozygia
Psidium guajava
Olacaceae
Strombosia scheffleri
Oleaceae
Rosaceae
Olea welwitschii
Prunus africana
Rubiaceae
Rubiaceae
Rutaceae
Sapotaceae
Rubus apetalus
Rubus niveus
Rubus pinnatus
Coffea eugenioides
Craterispermum schweinfurthii
Leptactina platyphylla
Psychotria peduncularis
Vepris nobilis
Chrysophyllum albidum
Englerophytum oblanceolatum
Manilkara butugi
Mimusops bagshawei
Pouteria altissima
Tephritid species
Brachyaciura sp.
Brachyaciura sp.
Dacus arcuatus
Trirhithrum nigrum
Trirhithrum inscriptum
Dacus limbipennis
Dacus limbipennis
Dacus bivittatus
Dacus punctatifrons
Dacus punctatifrons
Taomyia marshalli
Carpophthoromyia dimidiata
Trirhithrum albomaculatum
Ceratitis anonae
Ceratitis fasciventris
Trirhithrum demeyeri
Ceratitis anonae
Ceratitis anonae
Ceratitis fasciventris
Ceratitis flexuosa
Ceratitis anonae
Ceratitis anonae
Ceratitis fasciventris
Bactrocera amplexa
Ceratitis anonae
Ceratitis fasciventris
Bactrocera munroi
Bactrocera munroi
Ceratitis capitata
Ceratitis fasciventris
Ceratitis rubivora
Ceratitis rubivora
Ceratitis rubivora
Trirhithrum coffeae
Ceratitis stictica
Trirhithrum fraternum
Ceratitis anonae
Trirhithrum meladiscum
Ceratitis argenteobrunnea
Ceratitis cristata
Ceratitis ditissima
Ceratitis anonae
Ceratitis fasciventris
Ceratitis caetrata
Ceratitis capitata
Ceratitis caetrata
Ceratitis capitata
Ceratitis anonae
Fruit Flies of Kakamega Forest, Kenya
Plant family
Sapotaceae
271
Plant species
Pouteria altissima
Synsepalum brevipes
Solanaceae
Synsepalum cerasiferum
Solanum anguivi
Solanum mauritianum
Solanum terminale
Tephritid species
Ceratitis bremii
Ceratitis fasciventris
Ceratitis anonae
Ceratitis connexa
Ceratitis fasciventris
Ceratitis sp. nov. “CO”
Ceratitis hamata
Ceratitis venusta
Ceratitis anonae
Ceratitis fasciventris
Trirhithrum overlaeti
Families producing both large fruits (seeds primarily dispersed by mammals; e.g.
Sapotaceae), and small ones (seeds primarily dispersed by birds; e.g. Rubiaceae) were
exploited by tephritids.
Table 4. Relative importance of plant families to frugivorous Tephritidae in Kakamega Forest1.
Plant family
Sapotaceae
Rubiaceae
Flacourtiaceae
Rosaceae
Solanaceae
Cucurbitaceae
Moraceae
Olacaceae
Araceae
Myrtaceae
Rutaceae
Acanthaceae
Apocynaceae
Dracaenaceae
Euphorbiaceae
Menispermaceae
Oleaceae
Alangiaceae
Annonaceae
Araliaceae
Boraginaceae
Capparaceae
Clusiaceae
Colchicaceae
Dioscoreaceae
Ebenaceae
Fabaceae
Hippocrateaceae
Icacinaceae
Lamiaceae
Lauraceae
Loganiaceae
Meliaceae
% of plant species
producing tephritids (n)
100 (7)
23.5 (17)
40 (5)
57.1 (7)
37.5 (8)
80 (5)
66.7 (3)
100 (1)
100 (1)
50 (2)
33.3 (1)
100 (2)
20 (5)
100 (1)
12.5 (8)
25 (4)
100 (1)
0 (1)
0 (3)
0 (2)
0 (1)
0 (1)
0 (1)
0 (1)
0 (1)
0 (1)
0 (2)
0 (1)
0 (1)
0 (1)
0 (1)
0 (1)
0 (1)
% of samples
producing tephritids (n)
59.2 (49)
24.5 (53)
35 (20)
37 (27)
77.8 (27)
56.3 (16)
64.7 (17)
66.7 (3)
62.5 (8)
75 (4)
37.5 (8)
100 (2)
16.7 (6)
90.9 (11)
21.4 (14)
10 (10)
50 (6)
0 (1)
0 (3)
0 (3)
0 (5)
0 (2)
0 (3)
0 (1)
0 (1)
0 (1)
0 (2)
0 (3)
0 (3)
0 (1)
0 (1)
0 (4)
0 (1)
No. of tephritid species
produced by family
9
5
4
4
4
3
3
3
2
2
2
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
272
R.S. Copeland et al.
Plant family
% of plant species
% of samples
No. of tephritid species
producing tephritids (n)
producing tephritids (n) produced by family
Melianthaceae
0 (1)
0 (1)
0
Myrsinaceae
0 (1)
0 (2)
0
Orchidaceae
0 (1)
0 (1)
0
Passifloraceae
0 (1)
0 (1)
0
Phytolaccaceae
0 (1)
0 (1)
0
Piperaceae
0 (1)
0 (8)
0
Rhamnaceae
0 (3)
0 (8)
0
Sapindaceae
0 (3)
0 (8)
0
Ulmaceae
0 (4)
0 (5)
0
Urticaceae
0 (1)
0 (1)
0
Verbenaceae
0 (1)
0 (1)
0
Violaceae
0 (1)
0 (2)
0
Vitaceae
0 (3)
0 (10)
0
Zingiberaceae
0 (1)
0 (2)
0
1
Plant families ranked in descending order by number of tephritid species they produced
Notes on tephritid groups
Trypetinae
The Trypetinae include many genuine forest species, predominately associated with shrubs
and trees. In Africa, the best known of these are species of the genus Ceratitis. In total,
46 Ceratitis species are reported from Kenya out of a total of 94 for the entire Afrotropical
region. Of these 46, 15 are found in Kakamega Forest. This is a fairly high proportion
compared to other localities intensively collected within the country. To date, the only area
reported with a higher diversity of Ceratitis species is the central highland region around
Nairobi.
In Kenya, Kakamega is the only known locality for three Ceratitis species; C. ditissima,
C. hamata, and C. stictica. These species are representative of the floral and faunal affinities
of Kakamega Forest with the Guineo-Congolian forest belt of central and western Africa
(Kokwaro, 1988; Wass, 1995), and are most commonly found in these latter regions.
However, most of these species are also reported from isolated localities in other eastern or
southern African countries. Recently, C. bremii, previously known in Kenya only from
Kakamega Forest, was reared from fruits collected in a Kenyan coastal forest (Shimba Hills;
RSC, unpublished data). Like Kakamega Forest, East African coastal forests have biotic
affinities with the Guineo-Congolian flora, at least at the generic level (Clarke, 2000). The
only host records for C. bremii are from Kenya. Interestingly, both hosts are of the same
genus (Pouteria altissima in Kakamega Forest and Pouteria alnifolia Baker (Roberty) in
Shimba Hills), suggesting that C. bremii may be exceedingly stenophagous (sensu De Meyer,
2001b). Another species, C. flexuosa, is also primarily a West to Central African species
that, in Kenya, like C. bremii, is found only in Kakamega and coastal forests. Throughout its
range, C. flexuosa has been reared nearly exclusively from Antiaris species (there is a single
record from Dorstenia sp., also a Moraceae), and the known distributions of Antiaris and C.
flexuosa are essentially identical. It is likely that C. flexuosa is a forest-dependent
stenophagous species.
The other Ceratitis species known from Kakamega Forest fit the general pattern of
distribution that has been outlined for the genus (De Meyer, 2001b). Ceratitis anonae, for
example, has a largely equatorial distribution with an eastern boundary in western Kenya, but
stretching beyond Kakamega. Other species, like C. rubivora, are found only in southern to
eastern Africa and often reach their northernmost distribution in Kenya.
Fruit Flies of Kakamega Forest, Kenya
273
A recent revision of the genus Trirhithrum (White et al., 2003) listed 10 species from
Kakamega, largely based on 311 reared samples provided by the TAMU/ICIPE sampling
programme. Three of these species appear to reach the eastern limit of their distribution at
Kakamega: T. inscriptum and T. nigrum (mostly found in western and central Africa), and T.
stubbsi (an isolated record from Cameroon, in addition to records from Kakamega). Another
species worthy of special mention is T. micans, which is only known from the Ruwenzori
area of Uganda and Kakamega. Other species are widespread in at least eastern Africa.
Previously T. coffeae was only known from cultivated Coffea arabica L. Our rearing of T.
coffeae from Coffea eugenoides is the first record of a wild-fruit host for this tephritid.
A revision of the Dacina (as Dacidae) of Africa was published as recently as 1984 (Munro,
1984) but was largely based on much earlier work by the author (1926–1969). Prior to the
TAMU/ICIPE survey, the only organized fruit rearing carried out in Kenya was the unpublished
work of V.G.L. van Someren in the 1930s and 1940s, mainly around Nairobi. It is therefore
difficult to comment on the affinities of the Dacina of Kakamega with any accuracy. However,
four species do appear to have a primarily western distribution, as follows: D. armatus (also has
isolated records from Uganda), D. maynei (also has records from Uganda and Rwanda), D.
humeralis (a West African species for which Kakamega is the easternmost locality record), and
D. limbipennis (also known from Uganda). A few very rare species have been found at
Kakamega, including Bactrocera amplexa (previously known only from Meru, Kenya and
Amani, Tanzania) and a possible find of D. notalaxus (only known from Tora, Uganda).
Of the stem-boring genus Coelopacidia, hosts are known only from the Asteraceae
(Senecio) and the Apiaceae (=Umbelliferae) (Polemannia) (Hancock, 1986). Neither genus
is represented in the flora of Kakamega Forest (Kokwaro, 1988). However several species of
both Asteraceae and Umbelliferae are present and one or more may be hosts of the 3
Coelopacidia species recorded from the forest. Celidodacus species are known to use the
large, turgid fruits of species of the forest-associated climber Canavalia DC, a member of
the Fabaceae. In western Kenya, C. obnubilus has been reared from Canavalia africana
Dunn (figure 3; RSC, unpbl. data).
Non-Trypetinae taxa
Among the Tephritinae, the main tribes represented in and around Kakamega Forest are the
Pliomelaenini, Schistopterini, Tephrellini and Tephritini. Several smaller tribes are also
represented. The Tephritinae are usually poorly represented in forests and most of the species
of this subfamily listed above are actually common species of the savannah or other open
biotopes, penetrating into forest primarily along roads, following their ruderal hosts. Genera
such as Afraciura, Eutretosoma and Scedella serve as good examples. Conversely, genera
and species such as Brachyaciura (infesting Acanthus species, a typical forest shrub (Beentje,
1994)), Elaphromyia, Paraciura, Pseudafreutreta katangana (infesting another forest shrub,
Pseuderanthemum ludovicianum), Antoxya oxynoides, and a few species of Schistopterini
(e.g. Nigroptera) are typical forest-dwelling species.
Other interesting taxa are species of the Blepharoneurinae and Phytalmiinae genera
Baryglossa, Aethiothemara, and Ptiliniola. These genera are typical of forests in Africa, and
Aethiothemara trigona is the first representative of this genus found east of Uganda.
Baryglossa species are known only from flowers of the Cucurbitaceae, and several climbers
of this family (e.g. Coccinia barteri (Hook.f.) Keay, Momordica cissoides, Momordica
foetida, and Peponium vogelii are known from Kakamega Forest.
274
R.S. Copeland et al.
ACKNOWLEDGEMENTS
Millicent Okumu and Peris Machera provided excellent technical help in overseeing the
rearing of tephritids from all the fruits collected in Kakamega Forest. We thank the Director
General, ICIPE, for providing office and laboratory space and a field vehicle for this project.
This work was supported in part by USAID grant no. PCE-G-00-98-0048-00 and in part by
USDA/CSREES/IFAFS grant no. 00-52103-9651, (both grants in collaboration with the
International Centre of Insect Physiology an Ecology and its African Fruit Fly Initiative),
both to R. Wharton, Texas A&M University.
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Figure 2: Celidodacus obnubilus female.
Figure 3: Pod of Canavalia africana
(climber), host of Celidodacus obnubilus.
Figure 4: Taomyia marshalli female.
Figure 5: Fleshy fruit of Dracaena fragrans
(shrub/tree), host of Taomyia marshalli.
Figure 6: Carpophthoromyia dimidiata female.
Figure 7: Fleshy fruit of Drypetes gerrardii
(tree), host of Carpophthoromyia dimidiata.
Fruit Flies of Kakamega Forest, Kenya
277
Figure 8: Ceratitis caetrata male (note long,
capitate setae).
Figure 9: Fleshy fruit of Mimusops
bagshawei (tree), host of Ceratitis caetrata.
Figure 10: Trirhithrum inscriptum female.
Figure 11: Fleshy fruit of Culcasia falcifolia
(climber), host of Trirhithrum inscriptum.
Figure 12: Bactrocera amplexa male.
Figure 13: Fleshy fruit of Strombosia scheffleri
(tree), host of Bactrocera amplexa.
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R.S. Copeland et al.
Figure 14. Bactrocera munroi female.
Figure 15. Fleshy fruit of Olea welwitschii
(tree), host of Bactrocera munroi.
Figure 16. Dacus limbipennis male
Figure 17. Fleshy fruit of Momordica cissoides
(climber), host of Dacus limbipennis.
Figure 18. Brachyaciura sp. female.
Figure 19. Capsule of Acanthus pubescens
(shrub), host of Brachyaciura sp.
Quentin Luke