REST OF E.AFRICA

The Primates of East Africa:

Hills Biodiversity Research Programme, Nanyuki, KenyaAbstract: Seventeen genera, 38 species, and 47 subspecies of primate occur in East Africa.

Tanzania holds the largest number of primate species (27), followed by Uganda (23), Kenya (19), Rwanda (15) and Burundi (13). Six percent of the genera, 24% of the species, and 47% of the subspecies are endemic to the region.

East Africa supports 68% of Africa’s primate genera and 41% of Africa’s primate species. In East Africa, Tanzania has the highest number and percentage of endemic genera (one, 7%) and endemic species (at least six, 22%). According to the IUCN Red List, 26% of the 38 species, and 17% of the 47 subspecies, are ‘threatened’ with extinction. No recent taxon of East African primate has become extinct and no recent taxon is known to have been extirpated from the region. Of the 18 threatened primate taxa (ten species, eight subspecies) in East Africa, all but four are present in at least one of the seven most ‘primate species-rich’ protected areas.

The most threatened primates in East Africa are Tana River red colobus Procolobus rufomitratus, Tana River mangabey Cercocebus galeritus, and kipunji Rungwecebus kipunji. The most threatened, small, yet critical, sites for primate conservation in East Africa are the Tana River Primate National Reserve in Kenya, and the Mount Rungwe Nature Reserve-Kitulo National Park block in Tanzania.

In order to further refine the present country-by-country primate lists for East Africa, as well as the priority actions for the conservation of primates in the region, research should focus on the prosimians of Burundi and Rwanda.Keywords: Burundi, East Africa, Kenya, primate, primate conservation, Rwanda, Tanzania, UgandaRésumé: L’Afrique de l’Est accueille 17 genres, 38 espèces et 47 sous-espèces de primates. La Tanzanie est la plus richement dotée avec 27 espèces, ensuite viennent l’Uganda (23), le Kenya (19), le Rwanda (15) et le Burundi (13).

Six pourcent des genres, 24% des espèces et 47% des sous-espèces sont endémiques dans la sous-région. L’Afrique de l’Est accueille ainsi 68% des genres de primates et 41% des espèces. De la sous-région la Tanzanie accueille le plus grand Nombre et le plus grand pourcentage de genres (une, 7%) et d’espèces endémiques (au moins six, 22%). Selon la Liste Rouge de l’UICN, 26% des 38 espèces et 17% des 47 sous-espèces sont « menacées » de disparition. Aucun taxon actuel de primate est-africain ne s’est éteint, ni a été éliminé dans la sous-région.

Des 18 taxons de primates menacés (dix espèces, huit sous-espèces) en Afrique de l’Est que quatre ne sont pas représentées dans au moins une des sept aires protégées les plus riches en espèces de primates. Les primates les plus en danger d’extinction en Afrique de l’Est sont le Colobe Bai du Tana Procolobus rufomitratus rufomitratus, le Mangabey du Tana Cercocebus galeritus et le Kipunji Rungwecebus kipunji.

Les sites de conservation des primates les plus menacés, les plus petits néanmoins essentiels sont la Réserve Nationale des Primates du Fleuve Tana au Kenya et l’ensemble de la Réserve Naturelle du Mont-Rungwe avec le Park National de Kitulo en Tanzanie. Afin d’améliorer les listes actuels des primates par pays en Afrique de l’Est, ainsi que pour l’élaboration des actions prioritaires pour la conservation des primates dans la sous-région, la recherche devrait se focaliser sur les prosimiens du Burundi et du Rwanda.

IntroductIonEast Africa is a large (1,705,880 km²), the tropical region comprised of Burundi, Kenya, Rwanda, Tanzania and Uganda (Figure 1, Table 1). The major geographic features of East Africa include the Albertine (Western) Rift Valley, Gregory (Eastern) Rift Valley, Africa’s three highest mountains (Mount Kilimanjaro, Mount Kenya, and Rwenzori Mountains; all >5100 m asl), Africa’s largest lake (Lake Victoria; 68,800 km²), and Africa’s second-largest and deepest lake (Lake Tanganyika; 32,900 km²; 1470 m deep; Figure 2).

The complex geography, together with wide ranges for rainfall and temperature, means that East Africa supports numerous habitat types as well as many barriers to the dispersal of species.The primary vegetation types of East Africa are grassland, bushland/ticket, and woodland, but much of the region’s species richness and endemism are associated with the more moist vegetation types, particularly forests (Britton, 1980; White, 1983). Although biological diversity is lower in East Africa than in the Guinea-Congolean Forest Block, species endemism is higher in East Africa (Jenkins, 1992).____________Correspondence to: Yvonne A.

de Jong, Eastern Africa Primate Diversity and Conservation Program & Lolldaiga Hills Biodiversity Research Programme, P.O. Box 149, Nanyuki 10400, Kenya. Email: yvonne@wildsolutions.nlAfrican Primates 7 (2): 135-155 (2012) Primates represent one of East Africa’s best-studied taxonomic groups of mammals. Nonetheless, the taxonomic status of many of the region’s primates is still under debate and the geographical distribution and conservation status of many of the primate taxa remain poorly understood.

The aim of this paper is to, for each East Africa country, provide an overview of primate diversity, highlight those primate taxa that are under greatest threat, list those sites most critical to primate conservation, and point out some of the more important biogeographical questions that remain.rESuLtS And discussion the taxonomy applied here is taken from Mammals of Africa (Butynski et al., in press)—which is based largely on Groves (2001, 2005, 2007) and Grubb et al. (2003). The vernacular (common) names used here also are taken from Mammals of Africa. A conservative approach has been taken in compiling the list of the primate taxa for East Africa (Tables 1 & 2). For a primate taxon that is likely present in a country, but for which a specimen, photograph or authoritative sighting is absent or unknown, a single question mark is shown after the abbreviation of the country name.

For a primate, a taxon reported to occur in a country, but for which a specimen, photograph or authoritative sighting is absent or unknown, and for which presence seems unlikely, two question marks are shown after the abbreviation of the country name.

Primate diversity in East African total of 17 genera (see Appendix), 38 species and 47 subspecies of primate occur in East Africa (Tables 1 & 2). Genera diversity is highest in Uganda (15) and Tanzania (14). The largest of the five countries, Tanzania, is the richest in primate species (27) and subspecies (28). Tanzania is particularly diverse in prosimians, with 11 species. Burundi and Rwanda, given their small size, are relatively rich in species of primates with 13 and 15, respectively.

There are 25 genera and 93 species of primate in Africa (Butynski et al., in press).

Thus, East Africa supports 68% of Africa’s primate genera and 41% of Africa’s primate species.136 / de Jong & ButynskiFigure 1. Map of Africa showing the location of the five countries of East Africa: Burundi, Kenya, Rwanda, Tanzania, and uganda.countryLand area (km²)GeneraSpeciesSubspeciesBurundi25,6501113 (4?¹, 1??²)6 (3?, 1??)Kenya569,6901219 (1?)24 (3?)Rwanda24,9501215 (3?, 1??)9 (3?, 1??)Tanzania886,0401427 (3?)28 (2?)Uganda199,5501523 (2??)19 (1?, 2??)Total1,705,8801738 47¹ Likely present but requires confirmation.²

Reported to be present but this seems unlikely. Requires confirmation.table 1 – The land area of each of the five countries of East Africa and the number of primate genera, species, and subspecies present in each country. The species and subspecies that require confirmation are not included in the totals. Genus/species/subspeciesVernacularcountry1Threat2GORILLA I. Geoffroy, 1852GorillasGorilla beringei Matschie, 1903Eastern gorilla, UENG.

b. beringei Matschie, 1903Mountain gorilla, UCRPAN oken, 1816chimpanzeesPan troglodytes (Blumenbach, 1775)Robust chimpanzees, R, T, UENP. t. schweinfurthii (Giglioli, 1872)Eastern chimpanzees, B,R, T, UENHOMO Linnaeus, 1758HumansHomo sapiens Linnaeus, 1758Modern human, K, R, T, ULCCOLOBUS Illiger, 1811Black colobus, black-and-white colobusColobus angolensis Sclater, 1860Angola colobus, R, K, T, ULCC. a. palliatus Peters, 1868*Peters’ Angola colobus, TLCC. a. Ruwenzori Thomas, 1901Rwenzori Angola colobus, R, T, UVUC. a. ssp. nov.**Mahale Mountains Angola colobusTNEColobus guereza Rüppel, 1835GuerezaK, T, ULCC. g. caudatus Thomas, 1885**?

Mount Kilimanjaro guereza K?3, TLCC. g. occidentalis (Rochebrune, 1887)Western guerezaULCC. g. matschiei Neumann, 1899*Mau Forest guereza K, T, ULCC. g. kikuyuensis Lönnberg, 1912**Mount Kenya guerezaKLCC. g. percivali Heller, 1913**Mount Uarges guereza KENPROCOLOBUS rochebrune, 1887olive colobus, red colobusProcolobus rufomitratus (Peters, 1879)Eastern red colobus K, T, ULCP.r. rufomitratus (Peters, 1879)**Tana River red colobus KENP.r. tephrosceles (Elliot, 1907)*Ashy red colobus T, UENP.r. ellioti (Dollman, 1909)Semliki red colobusUNEProcolobus gordonorum (Matschie, 1900)**Udzungwa red colobusTENProcolobus kirkii (Gray, 1868)**Zanzibar red colobus TENCERCOCEBUS É. Geoffroy, 1812drill-mangabeys-Cerco Cebus galeritus Peters, 1879**Tana River mangabeyKENCercocebus sanjei Mittermeier, 1986**Sanje mangabeyTENPrimates of East Africa / 137table 2 –

The primate taxa of East Africa, together with endemic status, countries of occurrence, and category of threat (Iucn, 2011). LOPHOCEBUS Palmer, 1903Baboon-mangabeysLophocebus johnstoni (Lydekker, 1900)Johnston’s grey-cheeked mangabey B, R, UNELophocebus Ugandan (Matschie, 1913)*Uganda grey-cheeked mangabey T, UNERUNGWECEBUS davenport, Stanley, Sargis, de Luca, Mpunga, Machaga & Olson, 2006**KipunjiRungwecebus kipunji (Ehardt, Butynski, Jones & Davenport, 2005)**KipunjiTCRPAPIO Erxleben, 1777BaboonsPapio Cynocephalus (Linnaeus, 1766)Yellow baboon K, TLCP. c. cynocephalus (Linnaeus, 1766)Central yellow baboon K?, TLCP. c. ibeanus Thomas, 1893Ibean yellow baboon K, T?LCP. c. kindae Lönnberg, 1919Kinda yellow baboonTLCPapio anubis (Lesson, 1827)Olive baboon B, K, R, T,

ULCERYTHROCEBUS Broussard, 1897Patas monkeyErythrocebus patas (Schreber, 1775)Patas monkey K, T, ULCE. p. pyrrhonotus (Hemprich & Ehrenberg, 1829)Eastern patas monkey K, UNEE. p. baumstarki Matschie, 1905**?Southern patas monkeyK?, TNECHLOROCEBUS Gray, 1870Savannah monkeysChlorocebus tantalus (Ogilby, 1841)Tantalus monkeyK, ULCC. t. budgetti (Pocock, 1907)Budgett’s tantalusK, ULCChlorocebus pygerythrus (F. Cuvier, 1821)Vervet monkeyB, K, R, T, ULCC. p. rufoviridis (I. Geoffroy, 1843)Reddish-green vervetB?, R?, T, ULCC. p. hilgerti (Neumann, 1902)Hilgert’s vervetK, T, ULCC. p. nesiotes (Schwarz, 1926)**Pemba vervet TNEC. p. excubitor (Schwarz, 1926)**

Manda vervet KNEALLOCHROCEBUS Elliot, 1913Mountain monkeysAllochrocebus lhoesti(Sclater, 1899)L’Hoest’s monkey B, R, UVUCERCOPITHECUS Linnaeus, 1758Arboreal guenon Cercopithecus neglectus Schlegel, 1876De Brazza’s monkey K, T?, ULCGenus/species/subspeciesVernacularcountry1Threat2table 2 continued:138 / de Jong & Butynski CERCOPITHECUS (continued): Cercopithecus denti Thomas, 1907Dent’s monkey B, R, ULCCercopithecus hamlyni Pocock, 1907Owl-faced monkey B?, R, U??4VUC. h. Hamlyn Pocock, 1907Nose-stripe owl-faced monkey B?, R, U??NECercopithecus mitis Wolf, 1822Gentle monkey B, K, R, T, ULCC. m. albogularis (Sykes, 1831)*Zanzibar Sykes’s monkey K, TLCC. m. monoides I. Geoffroy, 1841Tanzania Sykes’s monkeyTLCC. m. stuhlmanni Matschie, 1893Stuhlmann’s blue monkey K, ULCC. m. moloneyi Sclater, 1893 Moloney’s monkey TLCC. m. albotorquatus Pousargues, 1896**?Pousargues’s monkey KVUC. m. kolbi Neumann, 1902**

Kolb’s monkeyKLCC. m. kandti Matschie, 1905Golden monkey R, UENC. m. doggetti Pocock, 1907Doggett’s silver monkey B, R, T, ULCCercopithecus ascanius (Audebert, 1799)Red-tailed monkeyB, K, R, T, ULCC. a. schmidti Matschie, 1892Schmidt’s red-tailed monkey B, K, R, T, ULCPERODICTICUS Bennett, 1831PottoPerodicticus potto (Müller, 1766)PottoB, K, R, T?, ULCP. p. ibeanus Thomas, 1910Eastern pottoB, K, R, T?, ULCP. p. Stockley Butynski & De Jong, 2007**Mount Kenya pottoKDDOTOLEMUR coquerel, 1859Greater galagos Otolemur crassicaudatus (É. Geoffroy, 1812)Large-eared greater galago B, K, R, T, ULCO. c. monteiri (Bartlett in Gray, 1863)Miombo silver galago B, K, R, T, U?LCO. c. argentatus (Lönnberg, 1913)*Northern silver galago K, T, R?, ULCOtolemur garnettii (Ogilby, 1838)Small-eared greater galago K, TLCO. g. garnettii (Ogilby, 1838)**

Zanzibar small-eared galagoTLCO. g. lasiotis (Peters, 1876)White-tailed small-eared galago K, TLCO. g. pangenesis Matschie, 1905Pangani small-eared galago K, TLCO. g. kikuyuensis (Lönnberg, 1912)**Kikuyu small-eared galagoKLCGenus/species/subspeciesVernacularcountry1Threat2table 2 continued:

Primates of East Africa / 139 GALAGO E. Geoffroy, 1796Lesser galagosGalago senegalensis É. Geoffroy, 1796Northern lesser galago B?, K, R?, T, ULCG. s. senegalensis É. Geoffroy, 1796Senegal lesser galagoB?, K, R?, ULCG. s. braccatus Elliot, 1907*Kenya lesser galago K, TLCG. s. sotikae Hollister, 1920*Uganda lesser galagoK, T, ULCGalago moholi A. Smith, 1836Southern lesser galago B??, R??, T, U??LCG. m. moholi A. Smith, 1836Moholi lesser galago B??, R??, T, U??LCGalago gallarum Thomas, 1901Somali lesser galagoKLCGalago matschiei Lorenz, 1917Spectacled lesser galago B, R, T?, ULC GALAGOIDES, A. Smith, 1833dwarf galagosGalagoides zanzibaricus (Matschie, 1893)**

Zanzibar dwarf galago TLCG. z. zanzibaricus (Matschie, 1893)**Zanzibar dwarf galagoTLCG. z. udzungwensis Honess, 1996**Matundu dwarf galago TLC Galagoides rondoensis Honess, 1996**Rondo dwarf galagoTCRGalagoides orinus (Lawrence & Washburn, 1936)**?Mountain dwarf galago K?, TNTGalagoides granti (Thomas & Wroughton, 1907)Mozambique dwarf galagoTLCGalagoides cocos (Heller, 1912)*Kenya coast dwarf galago K, TLCGalagoides demidovii (G. Fischer, 1808) Demidoff ’s dwarf galago B?, R?, T, ULCGalagoides thomasi (Elliot, 1907)Thomas’s dwarf galago B?, R?, T, ULCGenus/species/subspeciesVernacularcountry1Threat2table 2 continued:140 / de Jong & Butynski* Endemic to East Africa.** Endemic to one East African county.**? Possibly endemic to one East African country. Requires confirmation.¹ B: Burundi; K: Kenya; R: Rwanda; T: Tanzania; U: Uganda.² CR: Critically Endangered; EN: Endangered; DD: Data Deficient; LC: Least Concern; NE:

Not Evaluated; NT: Near Threatened; V: Vulnerable.³ Likely present but requires confirmation.4 Reported to be present but this seems unlikely. Requires confirmation. Kenya endemic species Cercocebus Galerius (Tana River mangabey)Kenya endemic subspeciesColobus guereza kikuyuensis (Mount Kenya guereza) Colobus guereza Percival (Mount Uarges guereza)Procolobus rufomitratus rufomitratus (Tana River red colobus)Chlorocebus pygerythrus excubitor (Manda vervet)? Cercopithecus mitis albotorquatus (Pousargues’s monkey)Cercopithecus mitis kolbi (Kolb’s monkey)Perodicticus potto stockleyi (Mount Kenya potto)Otolemur garnettii kikuyuensis (Kikuyu small-eared galago)

Tanzania endemic generaRungwecebus (kipunji)Tanzania endemic speciesProcolobus gordonorum (Udzungwas red colobus)Procolobus kirkii (Zanzibar red colobus)Cercocebus sanjei (Sanje mangabey)Rungwecebus kipunji (kipunji)Galagoides zanzibaricus (Zanzibar dwarf galago)Galagoides rondoensis (Rondo dwarf galago)? Galagoides orinus (mountain dwarf galago)Tanzania endemic subspecies Colobus angolensis ssp. nov. (Mahale Mountains Angola colobus)? Colobus guereza caudatus (Mount Kilimanjaro guereza)? Erythrocebus patas baumstarki (southern patas monkey)Chlorocebus pygerythrus nesiotes (Pemba vervet)Otolemur garnettii garnettii (Zanzibar small-eared galago)Galagoides zanzibaricus zanzibaricus (Zanzibar dwarf galago)Galagoides zanzibaricus udzungwensis (Matundu dwarf galago)The following primate taxa are endemic to East Africa but present in more than one country:

SpeciesLophocebus Uganda (Uganda grey-cheeked mangabey)Galagoides cocos (Kenya coast dwarf galago)SubspeciesColobus angolensis palliatus (Peters’ Angola colobus)Colobus guereza matschiei (Mau Forest guereza)Procolobus rufomitratus tephrosceles (ashy red colobus) Cercopithecus mitis albogularis (Zanzibar Sykes’s monkey)Otolemur crassicaudatus argentatus (northern silver galago)Otolemur garnettii panganiensis (Pangani small-eared galago)Galago senegalensis braccatus (Kenya lesser galago)Galago senegalensis sotikae (Uganda lesser galago)One (6% of 17) genus, at least nine (24% of 38) species, and at least 22 (47% of 47) subspecies of primate are endemic to East Africa.

Tanzania has the highest number and percentage of endemic genera (one, 7%) and endemic species (at least six, 22%), whereas Kenya has the highest number and percentage of endemic subspecies (at least seven, 29%). Tanzania is particularly rich in endemic prosimians with at least two endemic species and three endemic subspecies. Although Colobus angolensis palliatus has been reported to occur in northern Malawi (e.g., Misuku Hills), this has never been confirmed (Ansell, 1974; Ansell & Dowsett, 1988). In this paper, therefore, this taxon is considered to be endemic to Kenya and Tanzania. Tim Davenport (pers. comm.) recently observed C. a. palliatus at Umalila, Tanzania, which is only ca. 15 km from Malawi. Umalila is the Tanzanian extension of Malawi’s Misuki Hills. The search for C. a. palliatus in northern Malawi should continue.

Threatened Primates of East Africa In East Africa, the genera with the highest number of ‘threatened species’ (i.e., ‘Vulnerable’, ‘Endangered’ or ‘Critically Endangered’; IUCN, 2011) are Procolobusand Cercocebus, each with two species (66% and 100% of total species, respectively; Tables 2 & 3). Ten (26%) of Primate endemism in East Africa There are no primate taxa endemic to Burundi, Rwanda or Uganda. The following primate taxa are endemic to Kenya, Tanzania, and East Africa: Primates of East Africa / 141 the 38 primate species, and eight (17%) of the 47 primate subspecies in East Africa are threatened (Tables 2 & 4). It should be noted, however, that two species and seven subspecies of primate in East Africa have not been evaluated to determine their category of threat.

Neither of the two ‘Not Evaluated’ species (Lophocebus Ugandan, Lophocebus johnstoni) is expected to be categorized as threatened. Five of the ‘Not Evaluated’ subspecies are, however, expected to be categorized as ‘threatened’ (Colobus angolensis ssp. nov., Erythrocebus patas baumstarki, Chlorocebus pygerythrus nesiotes, Chlorocebus pygerythrus excubitor, Cercopithecus Hamlyn Hamlyn).

As such, the number of threatened subspecies is likely to rise to about 13 (28%).Two (5%) of East Africa’s primate species are ‘Critically Endangered’ (Rungwecebus kipunji and Galagoides rondoensis; both endemic to Tanzania). Six (16%) of East Africa’s primate species are ‘Endangered’ (Gorilla beringei, Pan troglodytes, Procolobus gordonorum, Procolobus kirkii, Cercocebus galeritus, Cercocebus sanjei), while two species (5%) are ‘Vulnerable’ (Allochrocebus lhoesti, Cercopithecus Hamlyn; Tables 2 & 4).Of the East African countries,

Tanzania has the largest number of threatened primate species with six (22% of 27 species; Tables 2 & 5). Rwanda has the highest percentage of threatened primate species with four (27% of 15 species).142 / de Jong & ButynskiGenusnumber of species (%)number of subspecies (%)Gorilla1 (100)1 (100)Pan1 (100)1 (100)Homo0 0 Colobus02 (25)Procolobus2 (66)2 (66)Cercocebus2 (100)0Lophocebus00Rungwecebus1 (100)0Papio00Erythrocebus00Chlorocebus00Allochrocebus1 (100)0Cercopithecus1 (20)2 (20)Perodicticus00Otolemur00Galago00Galagoides1 (14)0to t a l10 (26)8 (17)table 3 – number and percentage of threatened species in each genus of primate in East Africa (Iucn 2011). The two ‘not Evaluated’ species and the seven ‘not Evaluated’ subspecies (Table 2) were included when calculating the percentages.

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