07_Wagner&Böhme-2.qxd 12.08.2007 14:01 Uhr Seite 123 Bonner zoologische Beiträge Band 55 (2006) Heft 2 Seiten 123–150 Bonn, Juli 2007 Herpetofauna Kakamegensis – The amphibians and reptiles of Kakamega Forest, western Kenya PhilippWAGNER & Wolfgang BÖHME Bonn, Germany Abstract. We present an annotated checklist of the herpetofauna of Kakamega Forest with comments on the biology and systematics of the taxa. Twenty-five amphibian, one turtle, 22 lizard and 36 snake species are recorded from within the forest and its immediate environment. We discuss the generalized zoogeography of the forest and distribution pattern of the taxa comment on the protection of the forest. Analysis of the reptile species composition shows Kakamega Forest to be similar to the Guinea-Congolian rainforest and is considered the easternmost remnant of this forest block. Kakamega forest has a high diversity value for Kenya and represents a diversity hotspot on a national scale. Two species, Lygodac- tylus gutturalisand Psammophis phillipsi, are recorded in Kenya for the first time. Several other first records and the description of a new species (Agamidae: Agama finchi) were published already separately. Keywords.East Africa, Kenya, Kakamega Forest, herpetological survey, checklist, national diversity hotspot. 1. INTRODUCTION In Africa tropical rainforests extend from southern Sene- covery and scientific descriptions. Uganda, for example gal in the west to the coastal forests of Kenya and Tanza- has lost 86 % of its original forest in the past two decades nia in the east (COLLINS1992). The East African rainforests and the remaining parts are isolated fragments (VONESH belong to different biogeographical clades. Apart from the 2001). mountain forests and inselbergs, which are not clearly as- signable, there are three important forest types: the coastal Kakamega Forest in western Kenya is a similar isolated forests and the Eastern Arc mountain forests, both with a fragment. This small forest is the easternmost fragment high degree of endemism, and the easternmost outliers of of the equatorial rain forest system (CLAUSNITZER 2005; the Guinea-Congolian rain forest. The Eastern Arc Moun- DREWES1976; HAMILTON1976; KÖHLER2004; KOKWARO tains were already the object of several biological stud- 1988; SCHIØTZ 1976; VONESH 2001; WAGNER et al. sub- ies and have recently been reviewed in respect to their her- mitted; ZIMMERMANN 1972) and has recently been sur- petofauna by HOWELL(1993). The value of the coastal for- veyed in regard to its faunal communities by the ‘BIOTA est biodiversity was underestimated for a long time. Re- East Africa Project’and especially to its amphibian fau- cently the herpetofauna of the Kenyan part of the coastal na by e. g. SCHICKet al. (2005) and LÖTTERSet al. (2006). forest was surveyed by the Kifcon project (KIFCON1995). The herpetogeographical relationships to other fragments Only the forests associated with the Guinea-Congolian for- of the equatorial rain forest and other tropical forests have est (e. g. Budongo, Bwamba, Kibale, Bwindi, Mbira and recently been discussed by WAGNERet al. (subm.). Despite Mt. Elgon in Uganda) are at least partly explored. Her- belonging to Guinea-Congolian forest, Kakamega Forest petological inventories have been made for the Bwindi contains also numerous Afromontane elements (MUTAN- (DREWES & VINDUM 1991) and Kibale Forests (VONESH GAHet al. 1992; BENNUN& NJOROGE1999) in its flora and 2001). All together these forests harbour an estimated fauna. Therefore, the forest has a very large diversity and number of 333 amphibian (DUELLMAN1993), 105 snake zoogeographical value which has been shown by several (HUGHES 1983), 95 lizard, 16 turtle and 3 crocodilian authors for the different species groups (e. g. Odonata: species (BAUER 1993). This total herpetofaunal species CLAUSNITZER1999 & 2005; Amphibia: SCHICKet al. 2005; richness is more than 550. Considering the fact that zoo- Reptilia: this paper; Aves: ZIMMERMANN1972) and a lot logical research in Africa is becoming more and more dif- of species are not found elsewhere in Kenya. Additional- ficult because of the political instabilities in numerous ly, most of the remaining closed canopy forest within the countries, this number is certainly too low. Because trop- country is found in western Kenya (WASS1995). On a na- ical forests in Africa are seriously threatened numerous tional scale, the forest can be considered as a diversity undescribed species may become extinct before their dis hotspot and needs efficient protection. On the pan-African 07_Wagner&Böhme-2.qxd 12.08.2007 14:01 Uhr Seite 124 124 PhilippWAGNER& WolfgangBÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya (1935, 1936) and in the main forest by DREWES (1976). Subsequently no further reports on Kakamega reptiles have been published apart from the mentioning of single voucher specimens as e. g. in SPAWLS et al. (2002) and publications arising from this study (BÖHMEet al. 2005; KÖHLERet al. 2004; WAGNER& SCHMITZ2006; WAGNER et al. subm.). The aim of the present paper is to provide an overview of the herpetofauna of this forest as basic information for conservationists and wildlife biologists and to highlight the importance of the Kakamega Forest because of its im- pact and value on the biodiversity of Kenya. 2. DESCRIPTION OFTHE STUDYAREA Kakamega Forest is situated in the Kakamega District near Kakamega town in the Western Province of Kenya. The forest extends from 0°10’ and 0°21 N to 34°47’ and 34°58’E, covering an area of 240 km2, of which only 44.55 km2are protected by law (MITCHELL2004). The forest altitude varies between 1500 and 1700 m a.s.l. (above sea level) averaging 1650 m a.s.l. The forest be- comes part of the stratified landscape of the East African Rift Valley, situated 150 km to the east. The annual pre- cipitation ranges from 1500 to 2300 mm. The annual av- erage temperature is 27 °C at daytime and 15 °C at night. Two important rivers traverse the forest: the Isiukhu Riv- er in the north and the Yala River in the south. Both have their source in the Nandi Escarpment and drain into the Fig. 1. The main areas of investigation of BIOTAEast Africa nearby Lake Victoria. The forest block itself is surround- in the Kakamega Forest in the background a subset of a Land- ed by several forest fragments (e. g. Kisere, Malaba, sat 7 (ETM+) scene from 5 Feb 2001, contrast-enhanced band combination 5/4/3 for within-forest differentiation but printed Kaimosi), which differ in size, in the degree of destruc- in black and white. (Courtesy to G. SCHAABof BIOTA– E02). tion and their conservation status. The most important of these are the Kisere in the north and the Kaimosi fragment scale, Kakamega forest is not considered as a hot-spot be- in the south. Kisere is protected as a National Reserve cause in comparison with e. g. the Mt. Nlonako in whereas Kaimosi is unprotected but well known histori- Cameroon (99 amphibian [HERRMANN et al. 2005a] and cally from several collections made by A. LOVERIDGEand 89 reptile species, [HERRMANNet al.2005b]) the diversi- from the three herpetological taxa described by him from ty is comparatively low. Kaimosi and named after this forest: Agamakaimosae,Ty- phlopskaimosaeand Dendroaspisjamesonikaimosae. Compared with other vertebrate groups, East African am- phibians and reptiles are rather poorly studied and insuf- The eastern border of the forest is the 2200 m high Nan- ficiently known. In order to provide conservationists da- di Escarpment with its Northern Nandi and Southern Nan- ta for defining priorities for conservation it is necessary di forest. Both are considered to be montane forests be- to obtain basic information on the diversity and commu- cause of the occurrence of the tree fern Cyatheamanni- nity of forest amphibians and reptiles. ana as an indicator species of this forest type. The forests were contiguous with the Kakamega forest system until Amphibians have been intensively studied by several au- recently. MITCHELL(2004) pointed out that the North Nan- thors within the ‘Biota East Africa Project’(e. g. SCHICK di forest was not connected with Kakamega forest in the th et al. 2005; LÖTTERSet al. 2004; LÖTTERSet al. 2006; KÖH- 20 century although there was “dense forest” in the LERet al. 2006) whereas reptiles have been surveyed on- 1960´s between South Nandi forest and Kakamega ly superficially in the Kaimosi fragment by LOVERIDGE Forest. 07_Wagner&Böhme-2.qxd 12.08.2007 14:01 Uhr Seite 125 Bonner zoologische Beiträge 55 (2006) 125 Collection sites mentioned in this paper are as follows: the Collection codens: BIOTA= Biomonitoring Transect ‘Buyangu area’is the name of the northern part of the Analysis in Africa; BMNH = The Natural History Muse- Kakamega National Reserve (‘primary-like’forest), with um (British Museum [Natural History]), London, England; the Buyangu Village on his northern margin. Salazar Cir- CAS= California Academy of Science, San Francisco, cuit is an old plantation within the National Reserve, USA; MCZ= Museum of comparative Zoology, Harvard which is now secondary forest dominated by guava. Udo´s University, Cambridge/Massachusetts, USA; MHNG= Campsite is located within the northern part of the Nation- Muséum d´histoire naturelle, Geneva, Switzerland; al Reserve and is the home of the BIOTA field camp. NHMW= Naturhistorisches Museum Wien, Vienna, Aus- Isecheno is the low protected southern part of the forest. tria; NMK= National Museums of Kenya, Nairobi, Rondo Retreat Centre is a small hotel within this area. Kenya; PW= field number of the senior author; USNM= National Museum of Natural History, Smithsonian Insti- tution, Washington D.C., USA; ZFMK= Zoologisches 3. MATERIALAND METHODS Forschungsmuseum Alexander Koenig, Bonn, Germany. The material presented in this paper was partly collected during a three-month herpetological mission by the sen- 4.RESULTS ior author, which was carried out between March and June 2003. It was completed by some older voucher material 4.1. Checklist of the herpetofauna of Kakamega Forest from Kakamega Forest in the ZFMK collection, collect- Amphibia ed by H.W. HERRMANN, D. MODRYand P. NECAS. Mate- rial from Kakamega Forest is also part of the collections Pipidae Gray, 1825 of CAS, MHNG, NHMW, NMK, USNM, but this mate- Xenopus victorianusAhl, 1924 rial was only partly analysed by the authors. Relevant lit- erature data was also evaluated. During the 2003 mission 1924 Xenopus victorianusAhl, Mitt. Zool. Mus. Berlin, 11: 270. 170 reptiles were collected. Amphibians were not the main Specimens examined. ZFMK 81733-735, 81940. emphasis of the study but also collected and compared with the species list presented by SCHICK et al. (2005). Additional specimens. NMK A/3874/1, A/3935, A/3944, Specimens of the study were fixed in 98 % ethanol and A/4025/1-2, A/4062/1-8, A/4163. subsequently transferred to 70 % ethanol. For final dep- osition, they were equally partitioned between the NMK Key references:SCHICKet al. 2005. and ZFMK collections. The main forest and the forest Remarks: This species was often found in drift fence fragments were walked during both day and night and buckets in the Buyangu area near a small pond within the specimens, mostly arboreal, were caught dominantly by forest (ZFMK 81940). It was also found at Rondo Retreat visual encounter surveys along transects and opportunis- in the southern part of the forest (ZFMK 81735). The tic searches. In addition, Y-shaped drift-fences with pit- vouchers are assigned to Xenopusvictorianus in SCHICK falls were used for terrestrial species. Catching success et al. (2005) and to Xenopussp. in LÖTTERSet al. (2006). was low; only Adolfus africanus(Lacertidae) and amphib- ians were caught using this method. The roads were pa- trolled for snakes and the human inhabitants of the sur- Bufonidae Gray, 1825 rounding villages were recruited to help with collecting. Amietophrynus kisoloensis(Loveridge, 1932) The individual species accounts include the following 1932 Bufo regularis kisoloensis Loveridge, Occas. Pap. Boston Soc. Nat. parts: Specimens examined: gives a list of the material Hist., 8: 52. from museums collections examined by the authors; Ad- ditional specimens: refer to material known from other col- Specimens examined. ZFMK 81727-730, 81943. lections and not examined by the authors; Key references: Additional specimens. NMK A/104-106, A/108, A/1072/1- lists publications with more detailed information on 2, A/1293/1-2, A/1318/1-2, A/1648/1-3, A/2046/1-4, species of Kakamega forest; Remarks: gives information A/3055/1-3, A/3104/1-3, A/3153/1-9, A/3563/1-2, about the collected specimens and taxonomic statements. A/3750/1-3, A/3813, A/3850/2, A/3851/1-3, A/3943. We follow mostly the classification and taxonomic con- Remarks: ZFMK 81727-728 were collected in the clusions of FROSTet al. (2006), however there are sever- Buyangu area. ZFMK 81943 was found in the Malaba al taxonomic conclusions in their amphibian tree of live fragment of the forest. ZFMK 81729 and A/3055/1-3 are tat we can not fathom and in our opinion require addition- from Rondo Retreat Centre, NMK A/3104/1-3 from al evidence and study. Isecheno forest camp both located in the southern part of 07_Wagner&Böhme-2.qxd 12.08.2007 14:01 Uhr Seite 126 126 PhilippWAGNER& WolfgangBÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya the forest. NMK A/3850/2 was collected at the Isiukhu Key references:LÖTTERSet al. 2006. falls in the Buyangu area. Remarks:The specimen NMK A/198 was collected on a bridge near Kakamega town. NMK A/3105/2 was col- lected at the Kalunga glade at the Kakamega Forest. Amietophrynus maculatus(Hallowell, 1854) 1854 Bufo maculatusHallowell, Proc. Acad. Nat. Sci. Philadelphia, 7: 101. Phrynobatrachusaff. mababiensisFitzSimons, 1932 Specimens examined. ZFMK 77458, 81723-726, 81941- 1932 PhrynobatrachusmababiensisFitzSimons, Ann. Transvaal Mus., 942. 15: 40. Additional specimens. NMK A/1194, A/3850/1,3-10. Specimens examined. None. Remarks: ZFMK 81941-942 were collected within Remarks:This species has only been recorded from lit- Buyangu Village along a road under stones next to a erature (SCHICKet al. 2005) and is currently under inves- stream. NMK A/3850/5 was collected at the Isiukhu falls tigation by SCHICKet al. (2005) and other colleagues. in the Buyangu area. NMK A/1194 was from the Malava forest fragment. Phrynobatrachus natalensis (Smith, 1849) 1849 Stenorhynchus natalensis Smith, Ill. Zool. S. Afr., 3 (Appendix): Ranidae Rafinesque, 1814 24. Hoplobatrachus occipitalis(Günther, 1859) Specimens examined. ZFMK 81742-43. 1859 Rana occipitalisGünther, Arch. Naturgesch., 24: 320. Additional specimens. NMK A/3105/1, A/3863/1-4, Specimens examined. NMK A/3938. A/3931, A/3932. Key references:SCHICKet al. 2005. Remarks: NMK A/3105/1 was from the Kalunga glade within the Kakamega forest area. The series NMK A/3863 Remarks:The voucher was collected in a swamp in the and the ZFMK vouchers were from the Buyangu area. De- Buyangu area. Specimens, both adults and tadpoles, were tails on the other vouchers are unknown. found in an old swimming pool of the Serena Island Lodge in Kakamega town and were documented by a voucher specimen (NMK uncatalogued) and photographs. Phrynobatrachusaff. minutus (Boulenger, 1895) 1895 Arthroleptis minutusBoulenger, Proc. Zool. Soc. London, 1895: 539. Specimens examined. None Additional specimens. NMK A/3924/1-6, A/4310. Key references:LÖTTERSet al. 2006. Remarks: Details on the vouchers are unknown. NMK A/3924/1,2,5 are on permanent loan to the ZFMK. This taxon was not recorded by SCHICKet al. (2005). Afrana angolensis(Bocage, 1866) Fig. 2. Hoplobatrachus occipitalis from Kakamega town. Photo by Jörn KÖHLER. 1866 Rana angolensisBocage, J. Sci. Math. Phys. Nat., Lisboa, 1: 73. Specimens examined. NMK A/100/1-8, A/101, A/102/1- 2. Phrynobatrachus graueri(Nieden, 1911) Additional specimens. NMK A/1294/1, A/1314/1-8, 1911 Arthroleptis graueriNieden, Sitzungsber. Ges. Naturforsch. Freun- A/1649/1-3, A/3639, A/3937/1-2, A/4239. de Berlin, 1910: 441. Key references:LÖTTERSet al. 2006. Specimens examined. None. Remarks: NMK A/4239 was found along the Salazar Additional specimens. NMK A/198, A/3105/2. 07_Wagner&Böhme-2.qxd 12.08.2007 14:01 Uhr Seite 127 Bonner zoologische Beiträge 55 (2006) 127 road. The series NMK A/1314/1-8 and A/1649 were col- Additional specimens. NMK A/3955/1-2, A/4213. lected at the Ikuywa River. Details on the other specimens Key references:LÖTTERSet al. 2006. are unknown. Remarks:NMK A/4213 was found in a small swamp at the Buyangu area. Details on the other voucher are un- Amnirana cf. albolabris(Hallowell, 1856) known. This species was recorded for the first time for Kenya by LÖTTERS et al. (2006) and at present is only 1856 Hyla albolabrisHallowell, Proc. Acad. Nat. Sci. Philadelphia, 8: known to occur in Kakamega Forest within Kenya. 153. Specimens examined. None. Ptychadena aff. mascareniensis (Duméril & Bibron, Additional specimens. NMK A/196/1-2, A/1966. 1841) Key references:SCHICKet al. 2005; LÖTTERSet al. 2006. 1841 Rana mascariensis Duméril & Bibron, Erp. Gen., 8: 350. Remarks: The specimens NMK A/196/1-2 came from Specimens examined. ZFMK 81944. Kakamega town, near the Forest Department Pump House. Additional specimens. NMK A/3572, A/3840/4-5, A/3856, A/4214, A/4217/1-2, A/4221, A/4223, A/4227/1-10, Ptychadena anchietae(Bocage, 1867) A/4229/1-2, A/4232/1-4, A/4235/1-9. 1867 Rana anchietaeBocage, Proc. Zool. Soc. London, 1867: 843. Key references:VENCESet al. 2004; LÖTTERSet al. 2006. Specimens examined. NMK A/3845. Remarks:P. mascareniensiswas often found on the shore of the Isiukhu River in its small discharging streams and Additional specimens. A/4212, A/4216, A/4220/1-2, in the Buyangu area. NMK A/4227/1-3 was found near A/4224, A/4226/1-2, A/4234/1-3. the KEEPOffice at Isecheno, NMK A/4227/4-10 in a tem- Remarks:NMK A/4212 was collected in a small puddle porary swamp at Isecheno. This taxon was listed by LÖT- near the Buyangu view point; NMK A/4220/1-2 were col- TERSet al. (2006) as Ptychadena aff. mascareniensis. lected in amplexus on the 28.IV.2004 from the same pud- dle. NMK A/4216 was found in short grass habitat next to a small puddle within the Salazar secondary forest. Ptychadena oxyrhynchus(Smith, 1849) NMK A/4224 was collected on new Buyangu Campsite. 1849 Rana oxyrhynchusSmith, Ill. Zool. S. Afr., 3(Part 28): pl. 77, fig 2. NMK A/4234/1 was collected near the Keep office, A/4234/2-3 near the Isecheno Primary school. Details on Specimens examined. None. the other specimens are unknown. Additional specimens. NMK A/103/1-2, A/3846, A/4211, A/4215/1-4, A/4218, A/4219/1-3, A/4225/1-2, A/4228/1- 3, A/4230/1-2, A/4231/1-2, A/4233/1-4, A/4236/1-2. Ptychadena porosissima(Steindachner, 1867) Key references:LÖTTERSet al. 2006. 1867 Rana porosissimaSteindachner, Reise Österreichischen Fregatte Novara, Zool.: 18. Remarks:NMK A/4225/1-2, A/4228/1-3 and A/4236/1- 2 were collected in a pond on the new Buyangu Camp- Specimens examined. None. site, NMK A/4231/1-2 was found there in amplexus on Additional specimens. NMK A/3107/1-3, A/3574, A/4222. the 31. V. 2004 and the female laid eggs over night. NMK A/4233/1-4 were found at Isecheno in the southern part Key references:SCHICKet al. 2005; LÖTTERSet al. 2006. of the forest. Details on the other specimens are unknown. Remarks: NMK A/3107/1-3 were collected at Kalunga This taxon was listed by LÖTTERS et al. (2006) as Pty- glade. NMK A/4222 was found calling on wet mud near chadena aff. oxyrhynchus. a water edge in the Buyangu area. Details of the other specimen are unknown. Hyperoliidae Laurent, 1943 Afrixalus osorioi(Ferreira, 1906) Ptychadena taenioscelisLaurent, 1954 1906 Rappia osorioi Ferreira, J. Sci. Math. Phys. Nat., Lisboa, Ser. 2, 1954 Ptychadena taenioscelisLaurent, Ann. Mus. R. Congo Belge, Ter- 7: 162. vuren, Zool., 34: 25. Specimens examined. NMK A/3927/2. Specimens examined. None. 07_Wagner&Böhme-2.qxd 12.08.2007 14:01 Uhr Seite 128 128 PhilippWAGNER& WolfgangBÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya Hyperolius kivuensisAhl, 1931 1931 Hyperolius kivuensisAhl, Das Tierreich, 55: 280. Specimens examined. ZFMK 81745-746, 82183-184. Additional specimens. NMK A/3032/1-16, A/3103, A/3579, A/3709, A/3710, A/3748/1-4, A/3825/1-6, A/3864/4, A/3867/1-5, A/3953, A/4011/1-2, A/4026/1-2, A/4065. Key references:SCHICKet al. 2005; LÖTTERSet al. 2006. Remarks: NMK A/3103, A/3867, A/4011, A/4026 and ZFMK 81745-746 were from the Buyangu area. NMK A/4065 was collected at the BIOTACampsite. Details on the other vouchers are unknown. Fig. 3. Afrixalus osorioi from Kakamega Forest. Photo by Jörn KÖHLER. Additional specimens. A/3928, A/4017, A/4316/1-7. Key references:KÖHLERet al. 2005. Remarks: NMK A/3927/2 is on permanent loan to the ZFMK. NMK A/4017 and the series A/4316 were collect- ed at a pond within the Buyangu area. One specimen was additionally sighted in the South Nandi forest. This species was recorded for the first time for Kenya by KÖHLER et al. (2005) and at present only occurs in the Kakamega For- est complex within Kenya. Fig. 4. Hyperolius lateralisfrom Kakamega Forest. Photo by Jörn KÖHLER. Afrixalus quadrivittatus(Werger, 1908) 1908 ‘1907’Megalixalus leptosomus leptosomus Werger, Sitzungsber. Hyperolius lateralisLaurent, 1940 Akad. Wiss. Wien, Phys. Math. Naturwiss. Kl., 116: 1900. Specimens examined. NMK A/3933/2. 1940 Hyperolius lateralisLaurent, Rev. Zool. Bot. Afr., 34: 1. Additional specimens. NMK A/3933/1, A/4317/1-7. Specimens examined. ZFMK 81747-748. Key references:KÖHLERet al. 2005. Additional specimens. NMK A/2075/5-6, A/2075/8-10, A/3925/1-4, A/3936. Remarks: NMK A/3933/2 is on permanent loan to the ZFMK and was collected at a swamp in the Buyangu area. Remarks: NMK A/3925/1 is on permanent loan to The series NMK A/4317 was collected at the pond of the ZFMK. ZFMK 81747-748 were from the Buyangu area. Buyangu area. NMK A/3936 was from Rondo Retreat Centre in the southern part of the forest. Details on the remaining vouch- ers are unknown. Within Kenya, this taxon is currently on- Hyperolius acuticepsAhl, 1931 ly recorded from the Kakamega Forest. 1931 Hyperolius acuticepsAhl, Das Tierreich, 55: 282. Specimens examined. ZFMK 77616, 81749-750. Hyperolius viridiflavus(Duméril & Bibron, 1841) Additional specimens. NMK A/3922/2. 1841 Eucnemis viridi-flavus Duméril & Bibron, Erp. Gen., 8: 528. Key references:LÖTTERSet al. 2006. Specimens examined. ZFMK 77426, 81950. Remarks: ZFMK 81749-750 were from the Buyangu Additional specimens. NMK A/3056/1-3, A/3578, area. Details on the other vouchers are unknown. A/3866/5-11, A/668/1-4, A/1193/1-2, A/1444/1-5, A/3749, A/3809/1-5, A/3954/1-2, A/4013, A/4027/1-3, A/4066. 07_Wagner&Böhme-2.qxd 12.08.2007 14:01 Uhr Seite 129 Bonner zoologische Beiträge 55 (2006) 129 Remarks:Details on the vouchers are unknown, but most Kassina senegalensis(Duméril & Bibron, 1841) are from the Buyangu area. H. viridiflavus is the most 1841 Cystignathus senegalensisDuméril & Bibron Erp. Gen., 8: 418. common frog inside the forest. Specimens were observed at different ponds and also clearings for example, the BIO- Specimens examined. ZFMK 81741, 81946-949. TA field camp, where several specimens were sitting in- Additional specimens. NMK A/1158, A/1317, A/3711, side the lavatory. NMK A/1193/1-2 were from Malava for- A/3920. est. Remarks:All vouchers were from the Buyangu area and some were collected with a drift fence next to a pond. One specimen was collected on the Buyangu Hill outside the forest under stones. Fig. 5. Hyperolius cinnamomeoventrisfrom Kakamega Forest. Photo by Jörn KÖHLER. Hyperolius cinnamomeoventrisBocage, 1866 Fig. 7. Leptopelis mackayifrom Kakamega Forest. Photo by Arne SCHIØTZ. 1866 Hyperolius cinnamomeo-ventrisBocage, J. Sci. Math. Phys. Nat., Lisboa, 1: 75. Arthroleptidae Mivart, 1869 Specimens examined. ZFMK 77431-432, 81744. Leptopelis mackayi Köhler, Bwong, Schick, Veith & Additional specimens. NMK A/2095/1, A/3858/1-2, Lötters,2006 A/3918/1-2, A/4012. 2006 Leptopelis mackayiKöhler, Bwong, Schick, Veith & Lötters, Her- Key references:SCHICKet al. 2005; LÖTTERSet al. 2006. petological Journal 16: 183-189. Remarks:NMK A/3858, A/4012 and ZFMK 81744 are Specimens examined. ZFMK 83304-305 (paratypes), from the Buyangu area. This taxon was reported by LÖT- ZFMK 83306. TERSet al. (2006) as Hyperolius aff. cinnamomeoventris. Additional specimens. NMK A/3057/1 (holotype), NMK A/1407/1-3 (paratypes), NMK A/3072/1. Key references:KÖHLERet al. 2006. Remarks:This species was recently described by KÖH- LERet al. (2006) and is only known from Kakamega For- est and its vicinity and inhabits so far only forest habitats (also secondary and disturbed forest). It represents the sis- ter taxon of the West African L. modestusand was referred to this species in the past by SCHIØTZ(1975, 1999). Leptopelis aff.bocagii(Günther, 1865) 1865 “1864” Cystignathus bocagiiGünther, Proc. Zool. Soc. London, 1864: 481. Specimens examined. None. Fig. 6. Kassina senegalensis from Kakamega Forest. Photo by Jörn KÖHLER. Additional specimens. UZM R/074960-2. 07_Wagner&Böhme-2.qxd 12.08.2007 14:01 Uhr Seite 130 Remarks: The debate of the taxonomic status of this species is still ongoing. Many authors (e. g. BOULENGER 1896, KLAUSEWITZ1957, LOVERIDGE1957) have discussed differences or similarities between this taxon and Acan- thocercus cyanogaster RÜPPELL 1835. SPAWLS et al. (2002), LARGEN& SPAWLS(2006) and our own morphol- ogy studies of the two species support KLAUSEWITZ(1957) who regarded them as two distinct species. The reported distribution of both taxa is unclear because of the men- tioned taxonomic problems. Despite the works of KLAUSEWITZ(1954, 1957), a new re- view of the Acanthocercus- species complex is needed. The review will be a part of the PhD thesis of the senior Fig. 8. Leptopelisaff. bocagiiundescribed form from Kakame- author. ga Forest. Photo by Arne SCHIØTZ. The diagnoses of the subspecies of A. atricollis by KLAUSEWITZ(1957) are not adequate. Therefore, the ma- Key references:SCHIØTZ1975. terial is only preliminarily assigned to the subspecies ugandaensis because Kakamega Forest is geographical- Remarks: SCHIØTZ (1975) collected two males and one ly closer to the area of this subspecies than to minuta, ac- female (UZM R/074960-2). These vouchers were taken cording to KLAUSEWITZ’s (1957) map: A. a. minutainhab- from a savannah-like clearing near the Forest Station in its Ethiopia and eastern Kenya, while A. atricollis ugan- Kakamega Forest. One male was sitting in the bush when daensisoccurs within Uganda and western Kenya. calling. See SCHIØTZ(1975) for more details on the call. L. aff. bocagiiis an undescribed species, very close in mor- phology to L. bocagii. Only a few specimens are known, all from Kakamega (A. SCHIØTZ, pers. comm.). Reptilia Agamidae Spix, 1825 Acanthocercus atricollis (Smith, 1849) 1849 Agama atricollis Smith, llustrations of the Zoology of South Africa. 3 (Reptiles). Specimens examined. NMK L/2655, 2660/2-3; ZFMK 81952-963. Additional specimens. CAS 122731-122739. Key references:KLAUSEWITZ1957. Fig. 9. Acanthocercus atricollisfrom Kakamega Forest. Pho- Fig. 10. Agama kaimosaefrom Ngoromosi/ Nandi escarpment. to by Philipp WAGNER. Photo by Alexander BURMANN. 07_Wagner&Böhme-2.qxd 12.08.2007 14:01 Uhr Seite 131 Bonner zoologische Beiträge 55 (2006) 131 This agama subspecies was found in all areas surround- ing Kakamega Forest. It reaches the highest density in small villages and Ranger Stations and lives there on the clay huts, in syntopy with Trachylepis striata and Adol- fus jacksoni. It also occurs in hedges close to trees. It was not found inside the forest and also not in plantations and on forest edges. Agama kaimosaeLoveridge, 1935 1935 Agama agama kaimosaeLoveridge, Bull. Mus. Comp. zool., Har- vard 79:10. Specimens examined. ZFMK 83658-660; NMK L/2715/1,3,4; MCZ 40136- 40150. Key references: BURMANN2006, WAGNERet al. 2007. Remarks:After its synonymisation by LOVERIDGE(1936), this taxon was regarded as a synonym of Agama cau- dospinosa. But further investigations (BURMANN 2006, Wagner et al. 2007) have shown that Agama kaimosaeis a valid species. The taxon is a SWKenyan/N Tanzanian endemic (see map in WAGNER et al. 2007). The type lo- cality is a rocky highland three miles west of the Friends´ Africa Mission Station at Kaimosi. However no specimens were found there, neither by the senior author in 2003 nor by A. BURMANN in 2005. Our specimens were collected in Ngoromosi in the Nandi escarpment. Chamaeleonidae Gray, 1825 Fig. 11. Chamaeleo ellioti from Kakamega Forest. Photo by Philipp WAGNER. Chamaeleo gracilisHallowell, 1842 1842 Chamaeleo gracilisHallowell, J. Acad. Nat. Sci. Philadelphia 8: 324. Chamaeleo laevigatusGray, 1863 Specimens examined. NMK L/2203/1, NMK L/2653/1-2; ZFMK 82055-056. 1863 Chamaeleo laevigatusGray, Proc. Zool. Soc. London 1863: 95. Remarks:The species was not found in Kakamega For- Specimens examined. NMK L/1588. est directly but in forest edge areas of the South Nandi For- Remarks: The only specimen of this taxon was found est, located slightly east of Kakamega Forest. These forests 1981 by MADSENsix kilometres north of Kakamega town. were united with Kakamega until recently (MITCHELL Further data was not available. 2004), but in contrast to Kakamega the Nandi Escarpment has montane vegetation. The specimens were found in bushes next to the forest, on the western slopes of the Nan- Chamaeleo elliotiGünther, 1895 di escarpment. 1895 Chamaeleo elliotiGünther, Ann. Mag. Nat. Hist., London 15: 524, pl. 21, fig. A. Chamaeleo hoehneliiSteindachner, 1891 Specimens examined. NMK L/2480, L/1227, L/1273, 1891 Chamael(e)o höhnelii Steindachner, Anz. Akad. Wiss. Wien 28: L/2647, L/2653/1-2, L/2652/1-2, L/2658; ZFMK 54025, 141. 68494-97, 70835, 81974-80, 82057-58. Specimens examined. NMK L/252. Additional specimens. CAS 147912, 153247. Remarks:This voucher was collected in Kakamega town, Remarks: The species has a wide distribution in the so it is possible that the species also occurs in the sur- Kakamega area and specimens were found in several habi- rounding area of Kakamega Forest. tats in and around the forest. In the year of the fieldwork 07_Wagner&Böhme-2.qxd 12.08.2007 14:01 Uhr Seite 132 132 PhilippWAGNER& WolfgangBÖHME: The amphibians and reptiles of Kakamega Forest, western Kenya Shimba Hills and Athiplain. ZFMK 82022 was collected in a pitfall trap near the Buyangu Hill. This suggests that C. africanais not a strict tree dweller but also inhabits leaf litter during its search for food. Fig. 12. Rhampholeon boulengeri from Kakamega Forest. Pho- to by Philipp WAGNER. of the senior author, C. elliotiwas one of the species with the highest observed density. Only two years later it was very difficult to find them (P. HITA-GARCIApers. comm.). This was also corroborated by SPAWLS et al. (2002) and Fig. 13. Cnemaspis elgonensisfrom Kakamega Forest. Photo SCHMIDTet al. (2000) who also indicate that C. elliotihas by Philipp WAGNER. large population density dynamics. This chameleon was found in the surrounding areas of Kakamega and Nandi forests, and also inside the forest, on the shore of the Cnemaspis elgonensisLoveridge, 1935 Isiukhu River. Habitats were as follows: grassland, banana 1935 Cnemaspis elgonensisLoveridge, Proc. Zool. Soc. London 1935: and guava plantations, gardens, hedges, bushes, small 820. trees, secondary and riverside forest. Specimens examined. NMK L/1129, L/2263; ZFMK 82023. Rhampholeon boulengeriSteindachner, 1911 Additional specimens. USNM 158923. 1911 Rhampholeon boulengeriSteindachner, Anz. Akad. Wiss., Wien, Remarks:C. elgonensisis endemic to Uganda and Kenya 48: 178. and has a very restricted distribution area from the Mt. El- Specimens examined. NMK L/249, L/7270, L/2651, ; gon range to Kakamega in the south. Nothing is known ZFMK 54021-24, 77456, 81999. about its biology but similarities with other species of the genus can be anticipated. ZFMK 82023 was found at dawn Remarks:R. boulengeriis a typical dweller of the East on the lavatory of Udo´s Campsite in the northern part of African rain forests from the eastern parts of the DR Con- the forest. go in the west to Kenya in the east. Here it is, apart from Kakamega and Nandi forests, also found in the Cheran- gani Mts. (SPAWLSet al. 2002). Within Kakamega Forest Hemidactylus mabouia(Moreau de Jonnes, 1818) the pygmy chameleon was found in the northern part of the main fragment and in the Kisere fragment. 1818 Gekko mabouiaMoreau de Jonnes, Bull. Scient. Soc. Philomath. Paris, sér 3,5: 138. Specimens examined. NMK L/2648/1-2, L/2659/1, Gekkonidae Oppel, 1811 L/2659/3-4; ZFMK 70833-34, 81982-86. Cnemaspis africana(Werner, 1895) Remarks:This species inhabits nearly the entire sub-Sa- 1895 Gymnodactylus africanusWerner, Verh. zool.-bot. Ges. Wien 45: haran Africa. It has been transported by humans to South 190, pl. V, fig. 5. America (DIRKSEN 1995), Florida, USA (POWELL et al. 1998), Madagascar (GLAW & VENCES 1994) and to Specimens examined. NMK L/1987; ZFMK 82022. Madeira, Portugal (JESUSet al. 2002). In Kakamega it in- Remarks:This taxon is endemic to eastern Africa and is habits the houses of the villages around the forest and al- only distributed in Kenya and Tanzania. Next to so of Kakamega town. No specimens were found outside Kakamega Forest, the lizard is only known from Taita and of human habitations.