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Tetramorium boehmei sp. n. - a new ant (Hymenoptera: Formicidae) species from the Kakamega Forest, Western Kenya PDF

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Preview Tetramorium boehmei sp. n. - a new ant (Hymenoptera: Formicidae) species from the Kakamega Forest, Western Kenya

Bonn zoological Bulletin Volume 57 Issue 2 pp. 359-366 Bonn, November 2010 - Tetramorium boehmei sp. n. a new ant (Hymenoptera: Formicidae) species from the Kakamega Forest, Western Kenya & Francisco Hita Garcia, Georg Fischer, Patrick Kuck, Birthe Thormann Marcell K. Peters Zoological Research Museum Koenig, Adenaueralle 160, D-53113 Bonn, Germany. E-mails: [email protected]; [email protected]; [email protected]; [email protected]; [email protected] Abstract. Tetramorium boehmeiHita Garcia & Fischersp. n. -anew ant species from the Kakamega Forest inWestern Kenya is described. The new species can be placed in the Tetramorium camerunense species group and differs signifi- cantly from the other members ofthe group by its highly reduced sculpturation on head and mesosoma. With only two available specimens sampled in undisturbedprimary forest, Tetramorium boehmeisp. n. seemstobe arelatively rare en- demic species. Additionally, a first key to the Tetramorium species groups found in the Kakamega Forest is provided. Keywords.Ants,KakamegaForest, speciesgroupkey,taxonomy, Tetramorium, Tetramoriumcamerunensespeciesgroup. INTRODUCTION The ant genus Tetramorium Mayr, 1855 is almost global- species groups (Hita Garcia et al. 2009; FHG, unpub- ly distributed, andwith over430 described species one of lished). The Tetramorium camerunensespecies groupwas the most species-rich genera worldwide (Bolton 1995; B well represented with four species: Tetramorium lu- Bolton, Isle ofWight, pers. comm. 2010). The Afrotrop- cayanum Wheeler, W.M., 1905, Tetramorium cf. gegaimi ical zoogeographic regionholds the largest diversitywith Forel, 1916, and two undescribed species. over210 listed Tetramorium species (Bolton 1976, 1980, 1985, 1995; Hita Garcia et al. 2010). Recent taxonomic work was primarily focused on the Tetramorium weitzeckeri species group with the descrip- & The Kakamega Forest, one ofthe last indigenous forests tion ofTetramorium snellingi Hita Garcia, Fischer Pe- in Kenya, and its animal diversity have received consid- ters, 2010 and a species group revision for the whole erable scientific attention in the last decades (e.g. Claus- Afrotropicalregion(FHG, unpublished). However, around nitzer 1999, 2005; Copeland et al. 2005; Hita Garcia et 10 species or 25% of the Tetramorium fauna of the al. 2009; Kiihne 2008; Schick et al. 2005; Tattersfield et KakamegaForeststillremainundescribed. Withthiswork al. 2001; Wagner & Bohme 2007; Zimmermann 1972). we present a first preliminary key to the Tetramorium Generally, the forest is considered to be the eastern-most species groups present in the Kakamega Forest and de- relictofthe equatorial Guineo-Congolian lowlandrain for- scribe a new species belonging to the T. camerunense estbelt (Kokwaro 1988; Wagneretal. 2008; Zimmermann species group. 1972). The strong biogeographic affinities to West and Central African forests can be clearly seen in some fau- MATERIALAND METHODS nal elements like reptiles, dragonflies, and ants (Claus- nitzer 2005; Hita Garcia et al. 2009; Wagner et al. 2008). The ant fauna proved to be remarkably diverse with 288 The type material has been deposited in the following in- species from 52 genera and 11 subfamilies constitutingthe stitutions: second highest species richness reported fortheAfrotrop- ical zoogeographic region (Hita Garcia et al. 2009). By NMK: National Museums ofKenya, Nairobi, Kenya far the most species-rich genus in Kakamega Forest was ZFMK: ZoologicalResearch MuseumKoenig, Bonn, Ger- Tetramorium with more than 40 species belonging to 14 many ©ZFMK Bonn zoological Bulletin 57 (2): 359-366 360 Fancisco Hita Garcia et al. Both, holotype and paratype, were measured with an Postpetiole width (PPW): maximum width ofpostpetiole Olympus SZX 12 stereomicroscope equippedwith adual- measured in dorsal view. axis optical micrometeratamagnification of90x. The fol- HW lowing measurements and indices, in parts adapted from Ocular index (01): EL / * 100 Bolton (1980) and Glisten et al. (2006), were used: Cephalic index (CI): HW / HL * 100 HW Scape index (SI): SL / * 100 Head length (HL): maximum distance from the mid-point Propodeal spine index (PSLI): PSL / HL * 100 PTW PW ofthe anterior clypeal margin to the mid-point ofthe oc- Petiolar node index (PeNI): / * 100 cipital margin, measured in full-face view. Lateral petiole index (LPel): PTL / PTH * 100 Dorsal petiole index (DPel): PTW / PTL * 100 Head width (HW): width ofhead directly behind the eyes Postpetiolar node index (PpNI): PTW / PW * 100 measured in full-face view. Lateral postpetiole index (LPpI): PPL / PPH * 100 Dorsal postpetiole index (DPpI): PPW / PPL * 100 Scape length (SL): maximum scape length excludingbasal Postpetiole index (PPI): PPW / PTW * 100 condyle and neck. Measurements and indices arepresentedas minimum and Eye length (EL): maximum diameter of compound eye maximum values. Additionally, all measurements are ex- mm measured in oblique lateral view. pressed in and presented with three decimal places. Pronotal width (PW): maximum width ofpronotum meas- The digital colour images were produced with a Qlmag- ured in dorsal view. ing Micropublisher 5.0 RTV camera attached on a LEICA Z6 APO stereo-microscope and mounted with Syn- Weber's length (WL): diagonal length ofmesosoma in lat- croscopy Auto-Montage software (version 5.03). The eral view from the postero-ventral margin of propodeal mounted images were processed for publication with lobe to the anterior-most point ofpronotal slope, exclud- Adobe Photoshop CS2 and ImageJ. All images present- ing the neck. ed in this work are also online available atAntweb (Fish- er, 2002). Furthermore, holotype andparatype areunique- Propodeal spine length (PSL): in dorsocaudad view, the ly identified with specimen-level codes (e.g. tip ofthe measured spine, its base, and the centre ofthe CASENT0217239) affixed to each pin. propodeal concavity between the spines must all be in fo- DNA cus. Using a dual-axis micrometer the spine length is Total genomic was extracted fromtwo dissected sin- measured from the tip ofthe spine to a virtual point at its gle legs of the holotype, using the Qiagen base where the spine axis meets orthogonally with a line DNeasy-'Blood&Tissue Kit, followingthe manufacturers' DNA leading to the median point ofthe concavity. protocol. was eluted with 50 pi bufferAE; this step was repeated once to maximize yield. Petiole length (PTL): maximum length ofpetiolar node measured in dorsal view. A ca. 650 bp long fragment of the 5'-region of the cy- DNA tochrome c oxidase subunit I (COI), the standard Petiole height (PTH): maximum height ofpetiolar node barcode-marker for animals, was amplified using the measured in lateral view from the highest (median) point primers LCO 1490 and Nancy (5'-GGT CAA CAAATC ofthe node to the ventral outline. The measuring line is ATAAAG ATA TTG G-3' and 5 -CCC GGT AAAATT placed in an orthogonal angle to the ventral outline ofthe AAAATATAAACT TC -3 Folmer et al. 1994) and the node. Qiagen*Multiplex PCRKit.Amplificationreactionswere carried out in a 20 pi volume containing 10 pi QIAGEN Petiole width (PTW): maximum width of petiolar node Multiplex PCR Mastermix, 2 pi Q-Solution, 1.6 pi ofeach DNA measured in dorsal view. primer (both 10 pmol/pl), and 2.5 pi template, and filled up to 20 pi with sterile H 0. The PCR temperature 2 Postpetiole length (PPL): maximum length ofpostpetiole profile consisted ofan initial denaturation at 95° (15 min), measured in dorsal view. followed by 40 cycles at 94° (35 s, denaturation), 48.5° (90 s, annealing), 72° (90 s, extension), and a final exten- Postpetiole height (PPH): maximum height of the post- sion at 72° (10 min). PCR success was checked by elec- petiole measured in lateral view from the highest (medi- trophoresis on an 1.5% agarose gel containing ethidium an) point ofthe node to the ventral outline. The measur- bromide. The PCR product was purifiedusing 3 pi ofthe ing line is placed in an orthogonal angle to the ventral out- ExoSAP-IT* PCRpurification reagent followingthe man- line ofthe node. ufacturers' protocol. Bonn zoological Bulletin 57 (2): 359-366 ©ZFMK A new species ofTetramorium from the Kakamega Forest 361 The sample was bidirectionally sequenced by a commer- 8 Anterior clypeal margin with median impression. cial company (Macrogen Inc., Seoul, Republic ofKorea; T. dumezigroup (in parts) http://www.macrogen.com) using PCR primers. BLAST Anteriorclypeal margin entire 9 search confirmed belonging ofthe sequence to the genus Tetramorium. The sequence is deposited in GenBank(ac- 9 Tibiaewith short appressedpubescence 10 cession number HM753586). Tibiae with subdecumbent to erect pilosity orpubes- cence 11 KEY TO THE TETRAMORIUMSPECIES GROUPS 10 Hairs on dorsal mesosoma and gasterusually sparse, FOUND IN KAKAMEGA short, stout, andblunted T. simillimum group The following key to species groups is adapted from Hairs on dorsal mesosoma and gasterusually numer- Bolton (1976, 1980) and specific for the Kakamega For- ous, elongate and fine T. quadridentatum group est, though it also works for Western Kenya in general: 11 Frontal carinae long, usually reaching occiput. 1 Whole body covered with regularly branched hairs, T. dumezigroup (inparts) either bifid or trifid, giving the ant a woolly or furry Frontal carinae short and weakly developed, ending appearance 2 ateye level T. convexum group - Hairs generally simple, rarelybizarrelymodified, but never regularly branched bifid nor trifid as above . 3 12 Anteriorclypeal marginwithmedian impression. .. 13 Anterior clypeal margin entire, without amedian im- 2 Antennae 11-segmented; elongate simple hairs pres- pression 14 entalongthe antennal scapes andupperborders ofthe frontal carinae T. ericae group 13 Occipital region ofhead variablyrugose, rugulose or - Antennae 12-segmented; elongate simplehairs absent unsculptured, rarely with few anastomoses, without along the antennal scapes and upper borders of the arugo-reticulum T. camerunense group frontal carinae T. gabonensegroup - Occipital region of head distinctly rugo-reticulate. T. bicarinatum group 3 Antennae 11-segmented 4 - Antennae 12-segmented 5 14 Eyes larger, at least 9 ommatidia in the longest row. T. setigerum group 4 Petiolar node squamiform to high nodiform, never - Eyes smaller, at most 7 to 8 ommatidia in the longest blocky nodiform with sharply defined angles. row T.flabellum group T. weitzeckeri group - Petiolar node strongly blocky nodiform, generally with sharply defined angles. ... T. angulinode group Tetramorium camerunense species group 5 Lateral portion ofclypeusprominent, raisedto atooth Examination ofthenew species ledto the conclusion that or crest in full-face view; in dorsal view the lateral it can be easily placed in the T. camerunense species clypeal portions rise to a high peak in front ofthe an- group. Though the species group was well defined in tennal insertions andthen slope downtowardsthe me- Bolton (1980) it seems useful to reproduce it here: dianpartofthe clypeus T. sericeiventregroup - Lateral portion ofclypeus not modified as above. . 6 1. antennae 12-segmented 2. antennal scape relatively small to moderate (SI < 90) 6 Antennal scapes very long (always SI > 120); frontal 3. anterior clypeal margin generally with small median carinae weakly developed and short, at most reach- impression (absent in one species) ingtheposterioreyemargins T. aculeatum group 4. frontal carinae long andfine, generallyreachingpos- - Antennal scapes distinctly shorterthan above (always terioreyemargin, sometimesrunningto occipitalmar- SI < 110); frontal carinae variable 7 gin 5. antennal scrobe weakly developed 7 Propodeum armedwithapairofsmalltriangularteeth 6. propodeal spines of varying length, but always or denticles which at most are as large as the longer than propodeal lobes propodeallobes 8 7. mandibles generally smooth and shining, rarely fine- Propodeum armed with a pair of medium-sized to ly striate long spines which are noticeably larger than the 8. clypeus with three longitudinal rugae propodeallobes 12 9. cephalic dorsum usually finely longitudinally rugu- ©ZFMK Bonn zoological Bulletin 57 (2): 359-366 362 Fancisco Hita Garcia et al. Figs 1-2. Tetramoruim boehmei sp. n., holotype worker, CASENT0217238. 1 dorsum ofbody; 2 body in profde. ©ZFMK Bonn zoological Bulletin 57 (2): 359-366 A new species of Tetramorium from the Kakamega Forest 363 Figs 3—4. Tetramorium boehmei sp. n.. 3 holotype worker, CASENT0217238, full-face view of head; 4 paratype worker, CASENT0217239, full-face view ofhead. lose, without cross-meshes; occipital rugoreticulum Diagnosis. The highly reduced cephalic and mesosomal never developed sculpturation renders Tetramorium boehmei straightfor- 10. all dorsalbody surfaces withnumerous standinghairs wardly recognizable within the T. camerunense species 11. dorsal surfaces ofhind tibiae generally with decum- group. bent to appressed pubescence only, in two species suberect 12. sting appendage triangular, dentiform or pennant- DESCRIPTION shaped HL 0.700-0.772; HW 0.633-0.71 SL 0.533-0.578; EL 1; Priorto this study, the T. camerunense species group con- 0.122-0.139; PW 0.450-0.489; WL 0.822-0.900; PSL tained 12 species that were subdivided into two species 0.150-0.189; PTL 0.194-0.200; PTH 0.211-0.239; PTW complexes based on differences in sculpturation (Bolton 0.178-0.189; PPL 0.189-0.200; PPH 0.194-0.222; PPW 1980). The T. lucayanum complex, containing four 0.250-0.267; CI 90-92; SI 82-84; OI 19-20; PSLI21-24; species, canbe characterizedbythepresence ofsculptured PeNI 39^10; LPel 84-92; DPel 91-94; PpNI 55-56; LP- mandibles, petiole and postpetiole. One or both of the pl 90-97; DPpI 132-133; PPI 141 (2 measured). waist segments are generally strongly sculptured. The oth- er and larger complex, the T. camerunense complex with Head longerthanwide (CI 90-92). Anterior clypeal mar- eight species, possesses typicallyunsculptured waist seg- gin with small but distinct median notch. Frontal carinae ments and mandibles. fine andrelativelyweak, evenweakerbehind eye leveland significantly not reaching occipital margin. Antennal scrobe veryweakly developed, nearlyvestigial.Antennal & Tetramorium boehmei Hita Garcia Fischer sp. n. scape ofmoderate length, not reaching posterior margin (Figs 1-6) ofhead (SI 82-84). Eyes small to moderate (OI 19-20), with 8 to 9 ommatidia in longest row. Metanotal groove Holotype worker, KENYA, Western Province, Kakamega not impressed. Propodeal spines moderately sized (PSLI Forest, Colobus, 00° 21' 16" N, 34° 51' 36" E, 1650 m, 21-24), relatively thin, spinose and straight. Propodeal primary rain forest, hand collected, VII.2009, leg. G. Fis- lobes small, elongate-triangular and acute, always short- cher (NMK: CASENT0217238). Paratype worker, er than propodeal spines. Petiolar node nodiform, in pro- KENYA, Western Province, Kakamega Forest, Salazar, fileweakly higherthan long (LPel 84-92), in dorsal view 00° 19' 36" N, 34° 52' 14.6" E, 1650 m, Kakamega For- slightly longer than wide (DPel 91-94) and posteriorly est survey 2007, Transect 6, primary forest, Winkler leaf widerthan anteriorly. Postpetiole rounded, in dorsal view litter extraction, 21.VI.2007, leg. M. Peters (ZFMK: around 1.3 times wider than long (DPpI 132-133), and CASENT0217239). around 1.4 times wider than petiole (PPI 141); in lateral view weakly higher than long (LPpI 90-97). Sting ap- pendage triangular. Bonn zoological Bulletin 57 (2): 359-366 ©ZFMK 364 Fancisco Hita Garcia et al. Figs 5-6. Tetramorium boehmei sp. n., paratype worker, CASENT0217239. 5 dorsum ofbody; 6 body in profile. ©ZFMK Bonn zoological Bulletin 57 (2): 359-366 Anew species ofTetramoriwn from the Kakamega Forest 365 Mandibles either unsculptured, smooth and shining or group is the almost completely reduced sculpturation on finely striate. Clypeus with three longitudinal rugae, me- head and mesosoma. This reduction to a few weak rugu- dian ruga stronger developed than lateral rugae. Cephal- lae on the cephalic dorsum, and even less sculpturation ic sculpturation greatly reduced, laterally with only weak on the mesosomal dorsum, is unique inthe species group. partial rugulation, mostly smooth and shining; cephalic All otherspeciespossess a distinctly longitudinally rugose dorsum with 5-6 very weak and fine, widely spaced lon- or rugulose head and mesosoma, though variable from gitudinal nigulae between frontal carina, mostofthem bro- species to species, and sometimes irregularly shaped. ken along their length and never reaching occipital mar- gin, occipital region unsculptured. Cephalic ground Ithas to be mentionedthatthe holotype and paratype dif- sculpturation absent, generally smooth and shining. Lat- fer in some aspects that could be considered as sufficient eralmesosomaanteriorlymostlyunsculpmred, smooth and enough to divide them into two different species. First, the shiny, posteriorlywith weak irregularrugulation; dorsum mandibularsculpturation is completely smooth and shiny ofmesosoma unsculptured or with few weak rugulae, or in the holotype while it is longitudinally striate in the traces ofrugulae only, generally smooth and shining. Peti- paratype. This character is usually species-specific and ole eithercompletely unsculpmredorwithtraces ofsculp- could be considered as a good diagnostic tool to divide ture; postpetiole and gaster completely unsculpmred, them. Second, the paratype is larger and possesses more smooth and shiny. sculpture on head andmesosomathantheholotype which appears generally much more smooth and shining. Fur- All dorsal surfaces of head, mesosoma, both waist seg- thermore, the clypeal notch is distinct in both species but ments andgasterwith numerous long, simple, suberectto stronger developed in the paratype. However, at present, erect hairs. Fine pubescence on antennal scapes and tib- the observed variation is considered as intraspecific vari- ia appressed to subdecumbent. ation until more material becomes available. Apart from thenoteddifferencesthere is a strikingmorphological sim- Head, mesosoma, waist segments, and gaster very dark ilarity between both specimens and also the morphomet- brown to black, antennae, mandibles, and legs oflighter ric measurements of both are very close. Concluding, brownish colour. based on the analysis oftwo specimens, it would be pre- mature to describe them as different species. Queen and male unknown. Currently, the new species seems to be endemic to the Etymology. The new species is dedicated to Prof. Dr. Kakamega ForestinWestern Kenyawhere itwas sampled Wolfgang Bohme from Bonn, Germany, in honour ofhis in primary forest sites. Considering the high sampling ef- nearly four decades ofpassionate herpetological work at fort to assess the ant fauna ofthe Kakamega Forest (Hi- the Zoological Research Museum Koenig in Bonn. Fur- ta Garcia et al. 2009), and the only two available speci- thermore, with his encouraging, and always interesting, mens of T. boehmei, it seems to be a rather rare species. lectures, courses and excursions he had a significantpos- In addition, only little information is available on its bi- itive influence on the authors leading to their scientific ology. One specimenwas foundin aWinklerleaflitterex- dedication with zoological systematics and the Afrotrop- traction sample and one was hand collected from the ical zoogeographical region. ground. Consideringthis, thenew species couldberegard- ed as a rare terrestrial species, living either in the ground Notes. Generally, it is not recommendable for large and or the leaflitter. Though, it might also be possible that T. diverse genera as Tetramorium to describe single species boehmei lives in the lower vegetation or the canopy, and based only on few specimens outside a comprehensive the two specimenswere only accidentallycollectedbythe generic revision. Nevertheless, in the case of T. boehmei mentioned methods (the canopy ant fauna was consider- it seemsjustified for the following reasons. First, it does ablyless well sampledbythe authors thanthe groundliv- fit all group characters and can therefore easily be iden- ing ant fauna). Atpresent, taking into account ourknowl- tified as a I camerunense species group member, either edge ofthe leaflitter fauna ofthe Kakamega Forest, we byusing the species group keypresentedaboveorthe one considerthe leaflitterhypothesis as more likely. Theover- inBolton (1980). Withinthe T. camerunense species group all morphological appearancewith smallto moderate eyes it obviously belongs to the T. camerunense species com- and antennal scapes as well as the strongly reducedbody plex because ofthe unsculptured petiole and postpetiole. sculpturation are within the genus Tetramorium more of- Second, and more importantly, T. boehmei shows a re- tenfound inthe leaflitterthan inthe canopywhere species markable character combination that varies significantly tend to have largereyes andscapes. However, more spec- from the other members ofthe species group, and allows imens from more sampling events arenecessary to reveal an easy and clear identification. The single best diagnos- thepreferred stratum ofT. boehmei. Furthermore, it should tic character to separate T. boehmei from the rest ofthe benotedthat T. boehmeiwas sampled inthe two least dis- Bonn zoological Bulletin 57 (2): 359-366 ©ZFMK 366 Fancisco Hita Garcia et al. turbedprimary forest sites examined inthe Kakamega For- FisFhrearncBiLsc(o2,0U.0S2.)AA.n).twOenbli(nCealaitfohrtntipa:A/c/awwdwe.mayntowfebS.coierngcelsas,tSaacn- est (FHG, unpublished data). This might indicate that the cessed on July 20, 2010 new species prefers undisturbedprimary forest andreacts FormerO, BlackM, HoehW, LutzR,VrijenhoekR(1994)DNA negativelyto anthropogenic disturbance like selective log- primers for amplification ofmitochondrial cytochrome c ox- ging- idase subunit I from diverse metazoan invertebrates. Molec- ular Marine Biology and Biotechnology 3: 2947299 HitaGarciaF, FischerG, Peters MK, Wagele JW (2009)Apre- Acknowledgements. First, we want to thank Philipp Wagner, liminary checklist ofthe ants (Hymenoptera: Formicidae) of ZFMK, for making this more entomological and less herpeto- Kakamega Forest (Kenya). Journal of East African Natural logical publicationpossiblewithin the Festschrift. Then, we are History 98: 147-165 MK very grateful to Barry Bolton, Isle ofWight, U.K., who was so HitaGarciaF,FischerG, Peters (2010) Tetramoriumsnellin- kindtoreviewanearlydraftofthemanuscript, andprovidehelp- gi sp. n. -anew leaf-litterant species (Hymenoptera: Formi- ful comments. In addition, we thank Brian Fisher, California cidae) from a Western Kenyan rain forest. Myrmecological Academy ofSciences, San Francisco, U.S.A., for his coopera- News 13: 141-146 M tion with the images, Antweb, and also forreviewing the man- Glisten R, SchulzA, Sanetra (2006)RedescriptionofTeti-amo- uscript and helping to improve it. Furthermore, we would like riumforte Forel, 1904 (Hymenoptera: Formicidae), a west- tothankProf. Dr. JohannWolfgangWagele, ZFMK, forhisgen- ern Mediterranean ant species. Zootaxa 1310: 1-35 eral help. In addition, we arethankful forthe assistanceprovid- Kokwaro JO (1988) Conservation status ofthe Kakamega For- ed by the invertebrate lab ofthe National Museums ofKenya est in Kenya - the easternmost relict of the equatorial rain in Nairobi and the Kenya Wildlife Service in Kakamega. This forests ofAfrica. Monographs in Systematic Botany of the workwas fundedbythe German MinistryofEducationand Re- Missouri Botanical Garden 25: 471-489 search (BMBF) within the BIOLOG programme (BIOTA East Kiihne L (2008) Butterflies and moth diversity ofthe Kakame- ga Forest (Kenya). Brandenburgische Universitatsdruckerei, Africa subproject El6 [01LC0625A2]). Potsdam-Babelsberg MayrG (1855)Formicinaaustriaca. Beschreibungderbisherim oesterreichischen Kaiserstaate aufgefundenenAmeisennebst REFERENCES HinzufuegungjenerinDeutschland, inderSchweizundinIta- lienvorkommendenAmeisen.VerhandlungendesZoologisch- Bolton B (1976) The ant tribe Tetramoriini (Hymenoptera: Botanischen Vereins in Wien 5: 273-478 Formicidae). Constituentgenera,reviewofsmallergeneraand Schick S, VeithM, Lotters S (2005)Distributionpatternsofam- revision ofTriglyphothrix Forel. Bulletin ofthe British Mu- phibians from the Kakamega Forest, Kenya. African Journal seum (Natural History) Entomology 34: 281-379 ofHerpetology 54: 185-190 Bolton B (1980) The ant tribe Tetramoriini (Hymenoptera: TattersfieldP, Seddon MB, LangeCN(2001) Land-snail faunas Formicidae). The genus Tetramorium Mayr in the Ethiopian in indigenous rainforest and commercial forestry plantations zoogeographicalregion. Bulletinofthe British Museum (Nat- in Kakamega Forest, Western Kenya. Biodiversity and Con- ural History) Entomology 40: 193-384 servation 10: 1809-1829 W Bolton B (1985) The ant genus Triglyphotrix Forel a synonym Wagner P, Bohme (2007) Herpetofauna Kakamegensis-the ofTetramorium Mayr(Hymenoptera: Formicidae). Journal of amphibians andreptiles ofKakamegaForest, westernKenya. Natural History 19: 243-248 Bonner zoologische Beitrage 55: 123-150 W Bolton B (1995)Anewgeneralcatalogueoftheantsoftheworld. Wagner P, Kohler J, SchmitzA, Bohme (2008) The biogeo- Harvard University Press, Cambridge graphical assignment ofa west Kenyan rain forest remnant: ClausnitzerV(1999) Dragonfly(Odonata)recordsofKakamega further evidence from analysis ofits reptile fauna. Journal of Forest,Western Kenya, with notes onthe ecologyofrain for- Biogeography 35: 1349-1361 estspecies. Journal ofEastAfrican Natural History 88: 17-23 ZimmermanDA(1972)TheavifaunaofKakamegaForest,West- Clausnitzer V (2005) An updated checklist of the dragonflies ern Kenya, including a bird population study. Bulletin ofthe (Odonata) ofthe Kakamega Forest, Kenya. Journal of East American Museum ofNatural History 149: 257-337 African Natural History 94: 239-246 Copeland RS, Okeka W, Freidberg A, Merz B, White IM, De MeyerM, Luke Q (2005) Fruit flies (Diptera: Tephritidae) of Received: 05.VII.2010 KakamegaForest, Kenya. Journal ofEastAfricanNatural His- Accepted: 20.VII.2010 tory 94: 247-278 Bonn zoological Bulletin 57 (2): 359-366 ©ZFMK

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