Primate Biogeography, Diversity, Taxonomy and Conservation of the Coastal Forests of Kenya

Yvonne A. de Jong Thomas M. Butynski

Eastern Africa Primate Diversity and Conservation Program

Report to the Critical Ecosystem Partnership Fund March, 2009

Y.A. de Jong & T.M. Butynski, 2009.

Primate Biogeography, Diversity, Taxonomy and Conservation of the Coastal Forests of Kenya.

Yvonne A. de Jong & Thomas M. Butynski Eastern Africa Primate Diversity and Conservation Program, Nanyuki, Kenya.

Report to the Critical Ecosystem Partnership Fund March, 2009.

Yvonne A. de Jong & Thomas M. Butynski Eastern Africa Primate Diversity and Conservation Program P.O. Box 149, 10400 Nanyuki, Kenya [email protected] & tbutynski&aol.com www.wildsolutions.nl

Cover photos Left: Adult male Chlorocebus pygerythrus hilgerti, west of Malindi, Kenya. Right: Galagoides cocos, Manda Island, Kenya.

Citation: De Jong, Y.A. & Butynski, T.M. 2009. Primate Biogeography, Diversity, Taxonomy and Conservation of the Coastal Forests of Kenya. Report to the Critical Ecosystem Partnership Fund. Eastern Africa Primate Diversity and Conservation Program, Nanyuki, Kenya.

Primates of the coastal forests of Kenya. 2 Y.A. de Jong & T.M. Butynski, 2009.

CONTENTS Executive summary 8 1. Introduction 11 2. Study area 13 3. Methods 15 3.1 Preparations 15 3.2 Field surveys 15 3.2.1 Diurnal primate surveys 15 3.2.2 Nocturnal primate surveys 18 3.2.3 Nocturnal listening survey 18 3.3 Data analysis 19 3.3.1 Galago vocalization analysis 19 3.3.2 Distribution mapping 19 3.4 Primate Photographic Maps 19 3.5 Definitions 21 4. Results 22 4.1 Survey site details, primate encounters and conservation 23 4.1.1 North Coast of Kenya 23 A. Lamu Archipelago 23 B. Kipini Conservancy and Witu Forest Reserve 25 C. Tana River Primate National Reserve 28 4.1.2 South Coast of Kenya 30 A. Kaya Rabai (Chijembeni), Mazeras 30 B. Diani 31 C. Mrima Hill Forest 32 D. Kaya Sega 33 4.1.3 Outside the coastal forests of Kenya 34 A. Mwea National Reserve 34 B. Kiboko Camp, Makindu 35 C. Tsavo West National Park 35 D. Tsavo East National Park 36 E. North Kilimangodo 37 4.2 Primates species of the coastal forests of Kenya 38 4.2.1 Otolemur garnettii lasiotis 38 4.2.2 Galago senegalensis braccatus 43 4.2.3 Galagoides cocos 46 4.2.4 Cercopithecus mitis albogularis 50 4.2.5 Cercopithecus mitis albotorquatus 57 4.2.6 Chlorocebus pygerythrus hilgerti 63 4.2.7 Chlorocebus pygerythrus excubitor 68 4.2.8 Papio cynocephalus ibeanus 71 4.2.9 Colobus angolensis palliates 77 4.2.10 Procolobus rufomitratus rufomitratus 80 4.2.11 Cercocebus galeritus 82 4.2.12 Hybrid, Chlorocebus pygerythrus hilgerti x 84 Cercopithecus mitis albogularis 5. Discussion 87 Acknowledgements 89 References 90

Primates of the coastal forests of Kenya. 3 Y.A. de Jong & T.M. Butynski, 2009.

Appendix 1. Distinguishing character tables (draft) of Papio anubis, Papio cynocephalus, Chlorocebus pygerythrus, Cercopithecus mitis, Colobus angolensis, Colobus guereza and their subspecies, in Kenya and Tanzania. 2. Primate groups/individuals encountered inside and outside the coastal forests of Kenya. 3. Diurnal road surveys conducted at the north coast of Kenya. 4. Diurnal road surveys conducted at the south coast of Kenya. 5. Diurnal road surveys, outside the coastal forests of Kenya. 6. Butynski, T.M., De Jong, Y.A., Perkin, A.W., Bearder, S.K. & Honess, P.E. 2006. Taxonomy, distribution, and conservation status of three species of dwarf galagos (Galagoides) in eastern Africa. Primate Conservation 21: 63-79. 7. Culverwell, J., Feely, J., Bell-Cross, S., De Jong, Y.A. & Butynski, T. M. 2008. A new pig for Tsavo. Swara 31: 50-52. 8. De Jong, Y.A. Culverwell, J. & Butynski, T.M. 2009. Desert warthog Phacochoerus aethiopicus found in Tsavo East National Park and Tsavo West National Park, southern Kenya. Suiform Soundings 8: 4-6.

Primates of the coastal forests of Kenya. 4 Y.A. de Jong & T.M. Butynski, 2009.

Tables 1 Primate species and subspecies occurring in the coastal forest of Kenya, 22 number of encounters, population trend, current IUCN Red List degree of threat category, and abundance in the coastal forest of Kenya. 2 Rates of encounter with Otolemur garnettii lasiotis from 2005 – 2008 in the 39 coastal forests of Kenya. 3 Rates of encounter with Galagoides cocos during 2005 – 2008 in the coastal 47 forests of Kenya. 4 Encounter rates of Cercopithecus mitis albogularis during 2005 – 2008 in 52 the coastal forests of Kenya and outside the Hotspot. 5 Field descriptions of C. m. albogularis Unguja Island, Tanzania, C. m. 53 albogularis Usa River, Tanzania, C. m. albogularis Mrima Hill, Kenya, Cercopithecus mitis albogularis Diani, Kenya and C. m. albogularis Gedi Ruins, Kenya. 6 Encounter rates of Cercopithecus mitis albotorquatus during 2005 – 2008 in 59 the Coastal Forests Kenya and while driving to the Hotspot. 7 Field descriptions of C. m. albogularis, Gedi Ruins, Kenya, C. m. 60 albotorquatus, Witu Forest Reserve, Kenya and C. m. albogularis, Unguja Island, Tanzania. 8 Encounter rates (groups/hour and groups/km) of Chlorocebus pygerythrus 65 hilgerti during 2005 – 2008 in the coastal forests of Kenya and travelling to the coastal area. 9 Field descriptions of Chlorocebus pygerythrus hilgerti obtained from the 66 extremes of the coastal forests of Kenya (i.e., Diani and Mpeketoni) and from Shaba National Reserve in central Kenya. 10 Encounter rates of Papio cynocephalus ibeanus during 2005 – 2008 in the 73 Kenya coastal forest Hotspot and while driving to the Hotspot. 11 Field descriptions of adult male P. c. ibeanus obtained from Manda Toto 76 Island, Lamu Archipelago (close to the type locality of P. c. ibeanus), Tana River Primate National Reserve, Diani Beach (S coast). 12 Encounter rates (groups/hour and groups/km) of Colobus angolensis 78 palliatus during 2005 – 2008 in the coastal forests of Kenya. 13 Phenotypic comparison of the Chlorocebus pygerythrus hilgerti x 85 Cercopithecus mitis albogularis hybrid, Chlorocebus pygerythrus hilgerti and Cercopithecus mitis albogularis using field descriptions and photographs obtained at Diani Beach, Kenya.

Primates of the coastal forests of Kenya. 5 Y.A. de Jong & T.M. Butynski, 2009.

Figures 1 Transects along which primate surveys were conducted from August 2005 through 16 July 2008 in the coastal forests of Kenya, and along roads leading to these forests. 2 Transects along which primate surveys were conducted from August 2005 through 17 July 2008 in Kipini Conservancy, Witu Forest Reserve, and the Lamu Archipelago, northeast Kenya, and along roads leading to these sites. 3 Transects along which primate surveys were conducted from August 2005 through 18 July 2008 in the coastal forests of southeast of the coastal forests of Kenya, and along roads leading to these sites. 4 Photo map for Chlorocebus pygerythrus. 20 5 Manda Toto Island. 25 6 Cercopithecus mitis albotorquatus in Witu Forest Reserve. 26 7 Otolemur garnettii lasiotis, Tana River Primate National Reserve, Kenya. 38 8 Approximate geographic distribution of Otolemur garnettii lasiotis in Kenya 38 9 Sonogram and oscillogram of the trailing advertising call of Otolemur garnettii 39 lasiotis from Diani Beach. 10 Colour of the tip of the tail of Otolemur garnettii lasiotis on the coast of Kenya. 41 11 Captive adult male Otolemur garnettii in Malindi, Kenya. 41 12 Galago senegalensis braccatus, Isiolo, central Kenya. 43 13 Approximate geographic distribution of Galago senegalensis braccatus in the 44 coastal area of Kenya. 14 Galagoides cocos, Takwa Ruins, Manda Island, Kenya. 46 15 Known distribution and encounters (2003-2008) with Galagoides cocos in Kenya. 47 16 Cercopithecus mitis albogularis, Diani, Kenya. 50 17 Approximate geographic distribution of Cercopithecus mitis albogularis in the 51 coastal area of Kenya. 18 Adult male C. m. albogularis at Jozani-Chwaka Bay National Park, Unguja Island, 55 Tanzania. 19 Adult male C. m. albogularis at Gedi Ruins, Kenya. 55 20 Cercopithecus mitis albotorquatus male from Witu Forest, Kenya. 57 21 Known distribution and encounters (2003-2008) with Cercopithecus mitis 58 albotorquatus. 22 Chlorocebus pygerythrus hilgerti male at Diani Beach, Kenya. 63 23 Approximate geographic distribution of Chlorocebus pygerythrus hilgerti in the 64 coastal area of Kenya. 24 Suspected geographic distribution and evidence for Chlorocebus pygerythrus 68 excubitor. 25 Adult male Papio cynocephalus ibeanus foraging on the beach of Manda Toto 71 Island, Lamu Archipelago, Kenya. 26 Approximate geographic distribution of Papio cynocephalus ibeanus over the coast 72 of Kenya. 27 Adult male Colobus angolensis palliatus at Mrima Hill, south coast of Kenya. 77 28 Known distribution and encounters (2003-2008) with Colobus angolensis palliatus 77 in the coastal forests of Kenya. 29 Cercocebus galeritus at Mchelelo Research Station, Tana River Primate National 82 Reserve, Kenya. 30 Adult male Chlorocebus pygerythrus hilgerti x Cercopithecus mitis albogularis 84 hybrid, Diani, Kenya. 31 Encounter site of C. p. hilgerti x C. m. albogularis hybrid. Diani, Kenya. 84

Primates of the coastal forests of Kenya. 6 Y.A. de Jong & T.M. Butynski, 2009.

32 Dzombo Hill Forest Reserve, surrounded by degraded bushland, agriculture and human settlements.

Primates of the coastal forests of Kenya. 7 Y.A. de Jong & T.M. Butynski, 2009.

EXECUTIVE SUMMARY Although primates represent one of the best-known taxonomic groups found in the “Eastern Arc Mountains and Coastal Forests of Tanzania and Kenya Biodiversity Hotspot” (EACF Hotspot), numerous important questions remain concerning taxonomy, distribution, abundance, conservation status, and priorities for conservation actions. The objectives of this study were to 1) determine the distribution, diversity, and taxonomy of the primate fauna in the coastal forests of Kenya, to 2) determine the conservation (Red List Degree of Threat) status of all taxa of primates in the coastal forests of Kenya, and to 3) determine the primary threats to all taxa of primates in the coastal forests of Kenya. Four survey areas were visited [1) Lamu Archipelago, 2) Kipini Conservancy and Witu Forest Reserve, 3) North Coast (Mombasa to Malindi), and 4) South Coast (Mombasa to Lunga Lunga)] in seven trips totalling 38 survey days. Additionally, regions to and from the survey areas, including sites outside the Hotspot, were surveyed. In total, 5439 km of diurnal and nocturnal surveys were conducted within and to/from the coastal forest of Kenya. Primates were encountered 239 times. Of these encounters, 178 were with groups and 61 were with solitary individuals. Of the 178 encounters with groups, 99 where in the coastal forests of Kenya, 79 were while travelling to or from these forests.

Nine genera, nine species, and 11 subspecies of primate occur in the coastal forest of Kenya; Otolemur garnettii lasiotis, Galago senegalensis braccatus, Galagoides cocos, Cercopithecus mitis albogularis, Cercopithecus mitis albotorquatus, Chlorocebus pygerythrus hilgerti, Chlorocebus pygerythrus excubitor, Papio cynocephalus ibeanus, Colobus angolensis palliatus, Procolobus rufomitratus rufomitratus, Cercocebus galeritus. Three out of the eleven subspecies are nocturnal.

Of the 11 subspecies, 55% (n=6) in are categorized by the 2008 IUCN Red List of Threatened Species as “Least Concern”, 9% (n=1) as “Vulnerable”, 18% (n=2) as “Endangered”, while 18% (n=2) were not assessed (Table x). The coastal forest of Kenya have been much reduced, fragmented and degraded, and what little forest remains is under increasing threat. The expansion of agriculture is the most critical threat, while the production of charcoal, taking of firewood and timber, and mining are additional serious causes of habitat loss and degradation (Obura, 2007).

Papio cynocephalus ibeanus was the most often encountered primate in the coastal forests of Kenya (32 groups), followed by Cercopithecus mitis albogularis (24 groups), and Cercopithecus mitis albotorquatus (16 groups). Otolemur garnettii lasiotis and Galagoides cocos are common while Galago senegalensis braccatus is present but rare in this part of Kenya.

This study found significant geographic range extensions for O. g. lasiotis, G. cocos and C. m. albotorquatus, and is the first to report a wild Cercopithecus mitis albogularis x Chlorocebus pygerythrus hilgerti hybrid.

With data collected during this study,we reviewed (Butynski et al., 2006) the complicated nomenclatural history for the Kenya coast galago, Galagoides cf. cocos, and examined whether ‘cocos’ is the valid species name for this recently resurrected taxon. We concluded that Galagoides cocos is the name that should be applied to the Kenya coast galago---not Galagoides zanzibaricus.

In Malindi, a phenotypically different looking captive O. g. lasiotis was observed.

We observed considerable phenotypic differences among Cercopithecus mitis in the type locality

Primates of the coastal forests of Kenya. 8 Y.A. de Jong & T.M. Butynski, 2009. for C. m. albogularis (Unguja Island, Tanzania), the western-most locality for ‘C. m. albogularis’ that we visited (Usa River, close to the type locality of kibonotensis), the northeastern-most locality that we visited (Gedi Ruins), and those in between these localities (e.g., Mrima Hill, Vanga, Diani, Lunga Lunga). These phenotypic differences appear to be in a cline.

C. m. albotorquatus is a poorly known subspecies. During this survey, C. m. albotorquatus were found to be common in the Witu Forest Reserve, Kipini Conservancy, and Tana River Primate National Reserve, but rare on Manda Island and on Lamu Island. The question of whether Cercopithecus mitis phylax is a valid subspecies remains unresolved.

Surprisingly little phenotypic variation was observed for C. p. hilgerti populations in the coastal forest of Kenya. We did, however, find variation in the intensity of coloration of the pelage. The question of whether C. p. excubitor is a valid subspecies remains unresolved. C. pygerythrus is present in the Lamu Archipelago but was not encountered during this study. If C. p. excubitor is a valid subspecies, than its geographical range is highly fragmented and it population size is critically low.

P. c. ibeanus is a widespread, common and opportunistic primate in the coastal forests of Kenya. Outside the Hotspot it is even more abundant, occurring in and outside protected areas. Phenotypic variation was observed throughout its range, both inside and outside the coastal zone. During this study, an extensive hybrid zone between P. anubis and P. c. ibeanus across Kenya was observed: 1) from the northeast and east Mt. Kenya to the Lower Tana River, and 2) along the Nairobi – Mombasa Highway, starting at least at Makindu (north of the Chyulu Hills National Park), through Tsavo, to the coast. P. c. ibeanus is present along the coast, but differs phenotypically from the north coast to the south coast.

Primate biodiversity is relatively high in the Tana River Primate National Reserve (seven species), Diani (six species), and Kipini Conservancy and Witu Forest Reserve (five species). Additional primate surveys in Boni and Dodori Forest Reserves, Patta Island, Kaya Gonja, Shimba Hills National Reserve, Buda Forest, and Mrima Hill (nocturnal surveys) are essential for compiling a list of ‘Primate Priority Conservation Sites in the coastal forest of Kenya. There can be no doubt, however, that the top priority site for primate conservation in the coastal forests of Kenya, indeed for all of Kenya, are the forests along the Lower Tana River. These forests not only hold the highest diversity of primate species in all of Kenya, they hold Kenya’s only two endemic species of primate. In addition, they are among the most threatened forests in all of Kenya and are of great importance for the conservation of many other taxa.

During surveys driving to/from the coastal forests of Kenya, the desert warthog Phacochoerus aethiopicus was found west of Garissa. This is the first record west of the Tana River and extends the known geographic range to the northwest ca. 265 km. We also obtained the first record for P. aethiopicus in Tsavo East and Tsavo West National Parks (range extension of ca. 390 km to the southwest). In addition, common warthog Phacochoerus africanus and P. aethiopicus were found to be sympatric in Tsavo West National Park. This is the first site at which P. aethiopicus and P. africanus are known to be sympatric.

To visually present the phenotypic diversity of the primates that occur in the coastal forests of Kenya, six photographic maps were developed (wildsolutions.nl; De Jong & Butynski, 2009). These comprise 288 of our photos taken of primates in Kenya and Tanzania, of which 72 are of Kenya coastal forest primates.

Primates of the coastal forests of Kenya. 9 Y.A. de Jong & T.M. Butynski, 2009.

Priority research questions for the primates of the coastal forests of Kenya include:

1) To what extent does phenotypic variation in O. g. lasiotis occur over its geographic range? 2) What is the southern boundary for O. g. lasiotis? 3) What is the geographical range of G. s. braccatus within and outside the coastal forests of Kenya? , 4) How far up the Tana River does the range of G. cocos extent? 5) Which subspecies of C. mitis occurs between the Tana River/Delta and the Galana River? 6) What subspecies of C. mitis occurs between the Galana River and Kilifi Creek? 7) Is C. m. phylax a valid subspecies and, if so, what is its geographic range? 8) What is the geographic range of C. p. hilgerti? 9) Is C. p. excubitor a valid subspecies and, if so, what is its geographic range? 10) What is the southern limit of the geographic range of C. m. albotorquatus? 11) Are there other reports of wild C. p. hilgerti x C. m. albogularis, and do they phenotypically resemble the ‘Diani hybrid’? 12) Does Galagoides zanzibaricus occur in Kenya? 13) What primate taxa occur in the forests along the extreme northern coast of Kenya (especially Boni-Dodori Forest)? 14) What primate taxa occur on Patta Island? 15) What is the taxonomic status of Papio in the forests along the Lower Tana River?

Primates of the coastal forests of Kenya. 10 Y.A. de Jong & T.M. Butynski, 2009.

1. INTRODUCTION Primates are of particular interest and importance because (1) they are essential components (often “keystone species”) of the ecosystems in which they occur, affecting the composition of the vegetation and accounting for a significant portion of the mammalian biomass; (2) they are vital to our understanding of human evolution and human diseases; (3) they are among the best indicators of the health of ecosystems, and (4) they are among the most important “flagship species” for those sites in which they are found. Indeed, the endemic and highly threatened primates have been highly instrumental in bringing the biological importance and plight of the Eastern Arc Mountains and the coastal forests of East Africa to the attention of the conservation community over the past 20 years. For conservation purposes it is important to identify species and subspecies. With natural history data, together with carefully assessed geographical species and subspecies ranges, adequate and effective conservation measures can be taken. Primates are an especially important taxonomic group for the focus of conservation actions in the “Eastern Arc Mountains and Coastal Forests of Tanzania and Kenya Biodiversity Hotspot” (EACF Hotspot). The EACF Hotspot supports no fewer than nine endemic species and five endemic (or near-endemic) subspecies of primates are reported to occur in the Hotspot. Several of these primate taxa are listed in the IUCN/SSC Red List as “Critically Endangered” or “Endangered”. Others are listed as “Data Deficient”. This means that, “…there is inadequate information to make a direct, or indirect, assessment of its risk of extinction based on its distribution and/or population status”. Although primates represent one of the best-known taxonomic groups found in the Hotspot, numerous important questions remain concerning taxonomy, distribution, abundance, conservation status, and priorities for conservation actions. In addition, field studies in which the phenotypic variation of primates species and subspecies is assessed are rare. “Undiscovered” species or subspecies of primates are still being found and the knowledge about how many species and subspecies of primates occur in the Hotspot, where they occur, and the level of extinction risk each taxon faces, is vital for primate conservation. In particular, there might well be additional unrecognized “cryptic” species in the genera Otolemur, Galago, and Galagoides. The main goal of this study is to prevent the loss of primate biodiversity in the coastal forests of Kenya. The objectives were to 1) determine the distribution, diversity, and taxonomy of the primate fauna in the coastal forests of Kenya, 2) determine the conservation (Red List Degree of Threat) status of all taxa of primates in the coastal forests of Kenya and 3) determine the primary threats to all taxa of primates in the coastal forests of Kenya. With support from CEPF we (1) undertook primate-focused surveys throughout the Kenya coastal forest part of this Hotspot (onwards refered to as coastal forests of Kenya), (2) conducted a search of the literature along with much detailed communications with colleagues, (3) undertook Red List assessments for the 2008 IUCN Red List of Threatened Species of 24 species and subspecies, and (4) published ‘primate photographic maps’,(5) established a website to communicate our data and results, and (6) published three articles based on research collected under this project. Primate surveys were conducted throughout the coastal forests of Kenya, as well as while driving to and from the Hotspot. The focus was on primate diversity, abundance, distribution, threats, and conservation status in the coastal forests of Kenya. The surveys served to provide information, for each taxon, on (1) phenotypic variation, (2) geographic range limits along the coast of Kenya, (3) a rough indication of abundance, (4) a baseline against which to monitor change, and (5) primary threats. The Co-Project Leaders took advantage of their time in the field to collect opportunistic data on the distribution and abundance on other mammal taxa within and outside the Hotspot, in particular the desert warthog Phacochoerus aethiopicus, common warthog Phacochoerus africanus, and dik dik

Primates of the coastal forests of Kenya. 11 Y.A. de Jong & T.M. Butynski, 2009.

Madoqua. Data on the taxonomy, abundance, distribution, and conservation status of some of Kenya’s ‘non-Hotspot’ primate taxa were also collected; particularly on the Somali galago Galago gallarum Thomas, 1901, Kenya lesser galago Galago senegalensis braccatus Elliot, 1907, and eastern patas monkey Erythrocebus patas pyrrhonotus (Hemprich & Ehrenberg, 1829). This study did not focus on the Tana red colobus Procolobus rufomitratus rufomitratus (Peters, 1879) or on the Tana mangabey Cercocebus galeritus Peters, 1879. Although both of these species are threatened, they are relatively well known, and have received considerable recent attention under various other projects.

Primates of the coastal forests of Kenya. 12 Y.A. de Jong & T.M. Butynski, 2009.

2. STUDY AREA This study was conducted in the Kenya part of the Eastern Arc Mountains and Coastal Forests of Tanzania and Kenya Hotspot (in this study referred to as ‘coastal forests of Kenya’). This Hotspot stretches from the border of Kenya with Somalia, to the border of Tanzania with Mozambique, and includes Unguja (Zanzibar) Island, Mafia Island, and Pemba Island. In this study, we focused on the Kenya coastal area, a ca 600 km long sea front strip from Ishakani (1º 41’S) in the north to Vanga (4 º 40’S) in the south. The intervening habitats between the coastal forest patches are included in the Hotspot (CI, 2003). The vegetation in the Hotspot falls within the “Zanzibar- Inhambane Regional Mosaic Zone” (White, 1983). The Hotspot corresponds for a large part (with the exception of Somalia) with the WWF’s “Northern Zanzibar-Inhambane Coastal Forest Mosaic Ecoregion”.

In 1990, 95 forest patches were recorded in the coastal forests of Kenya. These covered an area of 660 km² (Burgess et al., 2000a). The mean patch size was 6.7 km² in Kenya. The two largest coastal forests in Kenya are Arabuko-Sokoke (ca 370 km²) and the Shimba Hills National Reserve (ca 63 km2) (Younge et al., 2002). The coastal forests of Kenya support a high level of endemism, but are treatned due to clearance, fragmentation and degradation (by agriculture, settlement and tourism), illegal logging, charcoal burning, and other human activities.

Climate is an important natural factor and influence on the vegetation structure and growth (Clarke, 2000). The climate at the Kenya Coast is mainly influenced by the large-scale pressure systems of the Western Indian Ocean and monsoon winds. During December through March the winds blow from the northeast and during May through October they blow southeast. Inbetween there are 1-2 month transition periods with variable and lower winds (Kairo & Bosire, 2007). The mean annual temperature at Mombasa is 26.3°C, with a mean annual maximum of 30.3°C and a mean annual minimum of 22.4°C (Irebelo, 2006). The north coast receives less rain which increases towards the south coast, and becomes less seasonal in the south. The north coast has two rainy seasons while the south coast one (Hawthorne, 1993 in Clarke, 2000). The Lamu Archipelago receives a mean annual total rainfall of 889 mm, 80% of which falls from April to June, with 345 mm in May (Irebelo, 2006). Malindi receives 1022 mm/yr. Mombasa receives1040 mm/yr, with an average of 240 mm in May. Vanga, the extreme southeast corner of Kenya on the Tanzanian border, is wet from May to December (Irebelo, 2006).

The population of Kenya is estimated to be 32.0 million people, with ca 2.8 million (9.0%) residents in the coastal areas (Kairo & Bosire, 2007). The population growth rate at the coast is 3.1%, the national average is 2.9%. Population densities are highest in urban centers like Mombasa, Malindi, Lamu and Kilifi (Kairo & Bosire, 2007). Land use in the coastal region can be devided into four main categories, livestock ranches in the hinterland, agricultural settlement schemes, private land along the coastline, and un-alienated government land (Kamula & Ochiewo, 2007). Economic activities along the coast range from industries (e.g. salt industries and cement production), service (e.g. tourism), transport, fishing, agriculture and cottage industries (arts and crafts). The economic contribution of coastal activities to the national economy is ca 15% (of which 12.5% from tourism; Kairo & Bosire, 2007).

The two main rivers that flow through and drain the coast are the Tana River and the Sabaki River. The Tana River (850 km in lenght) originates at Mt Kenya and enters the Indian Ocean at Kipini. The Sabaki River originates as the Athi River near Nairobi (Central Highlands) which when joined with the Tsavo River (close to Tsavo town), becomes the Galana River. A few kilometers north of Malindi the Sabaki River enters the Indian Ocean. The entire Athi-Galana-Sabaki system extends for 390 km (Kairo & Bosire, 2007).

Primates of the coastal forests of Kenya. 13 Y.A. de Jong & T.M. Butynski, 2009.

National Parks and Reserves along the Kenya coast protect the Indian Ocean territorial waters and the areas that border the ocean (including the coral reefs, mangrove forests and beaches). On the north coast there are several parks and reserves, including Dodori Coastal Reserve, Boni National Reserve, Arabuko Sokoke Forest Reserve, Tana River Primate National Reserve, Kiunga Marine National Reserve, Malindi Marine National Park and Reserve, Watamu Marine National Park, and the Ras Tenewi Coastal Zone National Park (between the Tana River Delta and Lamu). On the south coast, the following protected areas are established, Shimba Hills National Reserve, Mombasa Marine National Park and Reserve, Diani-Chale Marine National Park and Reserve, Kisite Marine Park, and Mpunguti Reserve (Kenya Wildlife Service, 2009).

For this study we divided the coastal area of Kenya in to four survey areas: 1) Lamu Archipelago, 2) Kipini Conservancy and Witu Forest Reserve, 3) North Coast (Mombasa to Malindi), and 4) South Coast (Mombasa to Lunga Lunga). Additionally, regions to and from the survey areas, including sites outside the Hotspot, were surveyed.

Primates of the coastal forests of Kenya. 14 Y.A. de Jong & T.M. Butynski, 2009.

3. METHODS

3.1 Preparations The literature was searched and consulted in order to (1) prepare a list of questions related to the taxonomy and distribution of the primates of the coastal forests of Kenya and Tanzania, (2) obtain all information related to the taxonomy and distribution of Kenya’s primates, and (3) compile the distinguishing (i.e., diagnostic) characters for each taxon of primate in the coastal forests of Kenya and Tanzania as presented by various authors.

Distinguishing character (review and comparison) tables were drafted for Olive baboon Papio anubis (Lesson, 1827), yellow baboon Papio cynocephalus (Linnaeus, 1766), vervet monkey Chlorocebus pygerythrus (F. Cuvier, 1821), Sykes’s monkey Cercopithecus mitis Wolf, 1822, Angolan colobus Colobus angolensis Sclater, 1860, Colobus guereza Rüppel, 1835, and their subspecies, in Kenya and Tanzania (Appendix 1) for quick reference and comparison in the field. Additionally, these draft distinguishing character tables were distributed to several CEPF partners working in the Eastern Arc Mountains and Coastal Forests of Tanzania and Kenya Biodiversity Hotspot. The distinguishing character tables will, once finalized, be available on our website (www.wildsolutions.nl). A photo album was prepared and maintained for each primate taxon for quick reference and comparisons in the field.

3.2 Field surveys In order to confirm presence of diurnal and nocturnal primates, assess the relative abundance of primates, and meet the need to cover large areas in a limited time, rapid assessment survey methods were used. Differences in research conditions, constraints, and opportunities in the survey sites required a variety of methods and approaches. Field surveys were conducted from August 2005 through July 2008 by T. M. Butynski and Y. A. de Jong. A total of 302 survey hours were completed in 38 days, covering a distance of 5439 km.

3.2.1 Diurnal primate surveys A total of 218 h of diurnal surveys were completed during 38 surveys conducted during seven trips that covered a distance of 5265 km (Figures 1, 2 & 3). Surveys were conducted from a vehicle, boat, or on foot by two people. Surveys took place both in the Hotspot as well as during travels along roads to and from the Hotspot (referred to as ‘road surveys’ in this study). The number of primate groups encountered per kilometre and per hour were the indices used to assess relative abundance (Butynski & Koster, 1994; White & Edwards, 2000; Nekaris & Jayewardene, 2004).

Information collected during each survey included date, weather conditions, start time, finish time, places surveys (GPS), walking/driving speed (GPS), and distance covered (GPS). When primate groups were encountered during a survey, the following data were collected: date, time, GPS coordinates (Garmin GPSmap 60Cx), altitude (by GPS), primate species/subspecies, vegetation type, and tree density. The focus during every primate encounter was on obtaining a detailed description of as many individuals in a group as time and visibility allowed. Photos were taken with a Nikon D70 digital camera fitted with a 400 mm Nikon lens, and with a Canon EOS 40D digital camera fitted with a Canon 75-300 mm lens. Most of the photos were shot in ‘raw’. As many of the individuals as possible in each group were photographed. Each primate group was appointed a unique number (hereafter referred to as the ‘group number’). The track of the complete survey was saved in a GPS and downloaded in a Dell Latitude notebook using Garmin MapSource software (Figure 1, 2 & 3).

Primates of the coastal forests of Kenya. 15 Y.A. de Jong & T.M. Butynski, 2009.

Figure 1. Transects along which primate surveys were conducted from August 2005 through July 2008 in the coastal forests of Kenya, and along roads leading to these forests. Figure 2 and 3 show more details for northeast Kenya and the southeast of Kenya.

Primates of the coastal forests of Kenya. 16 Y.A. de Jong & T.M. Butynski, 2009.

Figure 2. Transects along which primate surveys were conducted from August 2005 through July 2008 in Kipini Conservancy, Witu Forest Reserve, and the Lamu Archipelago, northeast Kenya, and along roads leading to these sites.

Primates of the coastal forests of Kenya. 17 Y.A. de Jong & T.M. Butynski, 2009.

Figure 3. Transects along which primate surveys were conducted from August 2005 through July 2008 in the coastal forests of southeast of the coastal forests of Kenya, and along roads leading to these sites.

3.2.2 Nocturnal primate surveys The presence of galagos was recorded during nocturnal surveys. Nocturnal surveys were conducted for a total of 20 h, covering 175 km. These surveys were conducted from a vehicle and/or by foot at all study sites. These surveys were conducted between 18.45 - 23.00 h and 4.00 - 06.30 h. Reflection from the eyes of galagos can be observed at >100 m in suitably open habitats. Torches (Maglights and Petzl Tikka XP headlamps) were used to scan the vegetation. Walks and drives were conducted slowly (ca 0.5-1 km/h on foot and ca 5 -10 km/h by vehicle) with pauses to scan the vegetation, observe primates, and/or record vocalisations. The following were recorded: date, weather conditions, moon phase, start time, finish time, localities surveyed (GPS), walking/driving speed (GPS), and distance covered (GPS). When galagos were encountered, binoculars (Zeiss Victory 10x42 and Zeiss Dialyt 7x42B) were used. The following data were collected when primates were encountered: date, time, moon phase, GPS coordinates, altitude, primate species/subspecies, vegetation type, tree density, number of individuals, and height of individuals in vegetation. Additionally, phenotypic descriptions were obtained and photos were taken using a Canon EOS 40D digital camera with a 75-300 mm Canon lens combined with a Canon Speedlite 420EX flash.

3.2.3 Nocturnal listening survey A total of 64.0 listening surveys were completed during 38 survey days. The advertisement call of galagos provides species specific information that can be used for species identification (Zimmermann, 1994; Bearder et al, 1995). Listening from a fixed point can reveal the presence of galago species. Audio recordings of galago vocalizations (and other nocturnal mammals and birds), preferably the loud advertisement call, were made during surveys, or opportunistically, using two

Primates of the coastal forests of Kenya. 18 Y.A. de Jong & T.M. Butynski, 2009.

Marantz Digital PMD660 recorders with Sennheiser Shot-Gun ME-66 microphones. The time and date of every recording is automatically saved within the audio file and notes were kept in our notebooks. Nocturnal listening surveys were mainly conducted from camp or from a higher point at dusk, dawn, and before and after nocturnal road or walking surveys.

3.3 Data analysis

3.3.1 Galago vocalization analysis Audio files were transferred from the Marantz digital audio recorder to a laptop. The best quality recordings were used to identify the species vocalizing. When confirmation was needed, vocalizations were send to S. K. Bearder and/or A. Perkin, Nocturnal Primate Research Group, Oxford Brookes University, Oxford, UK. Sonograms and spectrograms, as well as the numerical acoustic parameters, were produced from each vocalization using Avisoft-SASLab Pro software (R. Spect, Berlin; version 4.51).

3.3.2 Distribution mapping Locality records [date, time, locality name, latitude (decimal), longitude (decimal), altitude (m asl), primate species/subspecies, group code, vegetation type, notes] for all primates encountered during surveys were stored in a Microsoft Access database (XP). All records were plotted on a map using Garmin MapSource (6.10.2) and MapInfo Professional 8.0.

3.4 Primate Photographic Maps To present the phenotypic diversity of primates occurring in the coastal forests of Kenya, six photographic maps (‘photo maps’) were developed (Figure 4). These comprise 288 photos of out the thousands of photos taken of primates in Kenya and Tanzania. Of these, 72 are of primates in the coastal forests of Kenya. The photos were sorted into six taxonomic groups (Galagonidae, Papio, Cercopithecus mitis, Colobinae, Chlorocebus pygerythrus, and Chlorocebus pygerythrus hilgerti x Cercopithecus mitis albogularis hybrid). All photos were ‘geotaged’. The associated coordinates were either obtained automatically with a phototracker GPS (Gisteq PhotoTrackr) or a handheld GPS (Garmin, GPSmap 60Cx) using Picasa (version 2.7) software, and Google Earth (version 4.3.7284.3916 Beta) software to geotag the individual photos. The photos were uploaded to a Picasa Web Album which shows them automatically on a photo map (Figure 4).

Viewers of the photo maps can select a road, a terrain, or a satellite map and then zoom in or out on specific areas. Photos can be enlarged and viewed separately on a detailed map. Additionally, viewers can change from “Map View” to “Album View” which gives an overview of all photos on the photo map. Each photo is labeled with the taxonomic name (genus, species, subspecies), name of the locality were the photo was taken, vegetation type, altitude, and various comments. Viewers are invited to leave their feedback and comments with each photo. The photo maps are available for download in Google Earth KML format.

The first draft of the photo map for each of the six primate taxonomic groups is now available online. The ‘official version’ will be launched in March 2009. The web addresses will be distributed to a large number of primatologists. These primatologists will be invited to leave their comments with the photos and to discuss primate biogeography, diversity, taxonomy, and conservation. These six photo maps will serve as the beginning of a ‘living’ photo collection of eastern Africa’s primates. These six maps will be updated soon after new photos are available. Photo maps for other primate taxa in eastern Africa will gradually be added to this collection. This

Primates of the coastal forests of Kenya. 19 Y.A. de Jong & T.M. Butynski, 2009. resource can be used by anyone interested in the biogeography, diversity, taxonomy, and conservation of eastern Africa’s primates.

Figure 4. Screenshot of the ‘photo map’ for Chlorocebus pygerythrus.

Overview of the six photo maps:

Galagonidae: Total of 28 photos uploaded of which seven were taken in the coastal forests of Kenya. Taxa photographed and mapped include: Otolemur garnettii lasiotis, Otolemur garnettii panganiensis, Otolemur garnettii garnettii, Otolemur monteiri monteiri, Galago senegalensis braccatus, Galago gallarum, Galagoides cocos, Galagoides granti. Web address: http://picasaweb.google.com/wildsolutions/Galagonidae?feat=email#

Papio: Total of 103 photos uploaded of which 17 were taken in the coastal forests of Kenya. Taxa photographed and mapped include: Papio cynocephalus ibeanus, Papio cynocephalus cynocephalus, Papio anubis, hybrids between P. cynocephalus and P. anubis. Web address: http://picasaweb.google.com/wildsolutions/PapioAnubisPapioCynocephalus?feat=email#

Cercopithecus mitis: Total of 46 photos uploaded, of which 20 were taken in the coastal forests of Kenya. Taxa photographed and mapped include: Cercopithecus mitis albogularis, Cercopithecus mitis albotorquatus, Cercopithecus mitis kolbi. Web address: http://picasaweb.google.com/wildsolutions/CercopithecusMitis?feat=email#

Colobinae: Total of 17 photos uploaded, of which three were taken in the coastal forests of Kenya. Taxa photographed and mapped include: Colobus angolensis palliatus, Colobus

Primates of the coastal forests of Kenya. 20 Y.A. de Jong & T.M. Butynski, 2009.

guereza kikuyuensis, Colobus guereza caudatus, Procolobus kirkii, Procolobus gordonorum. Web address: http://picasaweb.google.com/wildsolutions/Colobinae?feat=email#

Chlorocebus pygerythrus: Total of 89 photos uploaded, of which 20 were taken in the coastal forests of Kenya. The only taxon photographed and mapped is Chlorocebus pygerythrus hilgerti. Web address: http://picasaweb.google.com/wildsolutions/ChlorocebusPygerythrus?feat=email

Chlorocebus pygerythrus hilgerti x Cercopithecus mitis albogularis hybrid: Total of five photos uploaded, all from Diani, Kenya. Web address: http://picasaweb.google.com/wildsolutions/ChlorocebusPygerythrusHilgertiXCercopithecusMit isAlbogularisHybrid?authkey=Gv1sRgCMqKmI2R_d6zXA&feat=email#

All photo maps will become accessible in March 2009 through our website: www.wildsolutions.nl

3.5 Definitions In this study we apply the following definitions;

Genus: “A genus is a monophyletic group of species (or a single species), which separated from other such groups earlier than the Miocene-Pliocene boundary.” (Groves, 2006).

Species: “A species is a population (or group of populations), distinguished by the possession of one more consistent (fixed, absolute) heritable differences from other such populations.” (Groves, 2006).

Subspecies: “A subspecies is a geographic segment of a species, distinguished by the possession at high frequencies, but not a much as 100%, of one or more heritable differences from other such segments.” (Groves, 2006).

Extent of occurrence: “..the area contained within the shortest continuous imaginary boundary which can be drawn to encompass all the known, inferred or projected sites of present occurrence of a taxon, excluding cases of vagrancy” (IUCN, 2001).

Area of occupancy: “..the area within its ‘extent of occurrence’ which is occupied by a taxon, excluding cases of vagrancy.” (IUCN, 2001).

Kenya coastal forests: The Kenya coastal forest part of the Eastern Arc Mountains and Coastal Forests of Tanzania and Kenya Biodiversity Hotspot. The intervening habitats between the coastal forest patches are included.

Primates of the coastal forests of Kenya. 21 Y.A. de Jong & T.M. Butynski, 2009.

4. RESULTS During this project, primates were encountered 239 times during seven field trips that totalled 38 survey days. Of these encounters, 178 were with groups and 61 were with solitary individuals. Of the 178 encounters with groups, 99 where in the coastal forests of Kenya, 79 were while travelling to or from the coastal forests of Kenya (Table 1).

Nine genera, nine species and 11 subspecies of primates occur in the coastal forest of Kenya. Eight of the subspecies are diurnal and three are nocturnal. Of the 11 subspecies, 45% (n=5) have more or less stable populations, 36% (n=4) have declining populations, and 18% (n=2) are data deficient (IUCN, 2008). Fifty-five percent (n=6) are categorized (IUCN, 2008) as “Least Concern”, 9% (n=1) as “Vulnerable”, and 18% (n=2) as “Endangered”, and 18% (n=2) were not assessed for the 2008 IUCN Red List of Threatened Species (Table 1).

Table 1. Primate species and subspecies occurring in the coastal forest of Kenya, number of encounters, population trend, current IUCN Red List degree of threat category, and abundance in the coastal forest of Kenya. No. Subspecies Number Number of Population Degree of Abundance in encounters encounters trend (IUCN, threat the coastal in the coastal outside the 2008) category forest of forests of coastal forests (IUCN, 2008) Kenya Kenya of Kenya Nocturnal primates

1 Otolemur garnettii >40 Present, not Stable Least Concern Very common lasiotis individuals counted 2 Galago senegalensis 0 Present, not Stable Least Concern Uncommon braccatus counted 3 Galagoides cocos >20 0 Stable Least Concern Common individuals

Diurnal primates

4 Cercopithecus mitis 24 groups 3 groups Decreasing Least Concern Common albogularis 5 Cercopithecus mitis 16 groups 0 Decreasing Vulnerable Common in albotorquatus along the Lower Tana River and Witu area. 6 Chlorocebus 11 groups 33 groups Stable Not assessed Common pygerythrus hilgerti outside of the coastal forests. 7 Chlorocebus 0 0 ? Not assessed Rare pygerythrus excubitor 8 Papio cynocephalus 32 groups 39 groups Stable Least Concern Very common ibeanus 9 Colobus angolensis 11 groups 0 ? Least Concern Common in palliatus the coastal forest south of Mombasa 10 Procolobus 2 groups 0 Decreasing Endangered Endemic to rufomitratus the Lower rufomitratus Tana River

Primates of the coastal forests of Kenya. 22 Y.A. de Jong & T.M. Butynski, 2009.

where it is common. 11 Cercocebus galeritus 3 groups 0 Decreasing Endangered Endemic to the Lower Tana River where it is common.

Other Chlorocebus 1 individual 0 - - Rare pygerythrus hilgerti x Cercopithecus mitis albogularis hybrid Cercopithecus mitis 4 groups Total 99 groups, 61 79 groups, solitary solitary individuals individuals not counted

Appendix 2 presents all primate groups/individuals encountered inside and outside the coastal forests of Kenya, including the date of encounter and the vegetation type they were found in.

The rate of encounter with primates was highest on in Mrima Hill Forest (6.6 groups/h; 0.9 groups/km; n=19), followed by Kipini Conservancy (1.3 groups/h; 0.3 groups/km; n=32).

4.1 Survey site details, primate encounters and conservation This chapter is divided into three sections: (1) North Coast of Kenya, (2) South Coast of Kenya, and (3) areas in Kenya but outside of the coastal forests. Road survey details are presented in Appendix 3, 4 & 5. Only road surveys in which primates were encountered are included.

4.1.1 North Coast of Kenya

A. Lamu Archipelago

Latitude/longitude: Lamu Town (Lamu Island): S2.27036 E40.90200; Manda Island (Airport): S2.25713 E40.91091; Manda Toto (manager’s building): S2.22151 E40.97880.

Altitude: 0 – 10 m asl

Vegetation: Mangrove forest, Acacia bushland, beach, dense and diverse coastal shrub, including some taller trees (Acacia, Commiphora) and wild fruit trees on coral rag.

Survey dates: 1 - 8 July 2008.

Survey details: Total of 135.0 km, 65.2 h (47.0 h diurnal, 26.7 h nocturnal) over 8 days. A local boat (dhow) was hired to travel from Manda Island (airport) to east Lamu Island, north Lamu Island, Manda Toto Island, west Manda Island, and east Manda Island. Diurnal surveys: 128.1 km, 47.0 h 54.4 km, 35.4 h by foot

Primates of the coastal forests of Kenya. 23 Y.A. de Jong & T.M. Butynski, 2009.

73.7 km, 11.6 h by boat (speed 4.9 – 6.3 km/h). Nocturnal surveys: 6.9 km, 3.7 h by foot Nocturnal listening survey: 14.7 h

Primate species encountered: O. g. lasiotis, G. cocos, C. m. albotorquatus, P. c. ibeanus. Strong evidence was obtained for the presence of C. p. excubitor but this subspecies was never encountered.

Encounter rates: Lamu Island: No diurnal or nocturnal primates seen. C. m. phylax: vocalizations heard from mangrove forest. Evidence was collected for the presence of C. p. excubitor. Residents stated that this taxon is present in very low numbers and occasionally hunted and kept/sold as pets.

Manda Toto Island: Twice a group (3 individuals) of P. c. ibeanus was encountered on this small island. No nocturnal primates were heard or observed.

Manda Island: The primate density on Manda Island is very low. C. mitis vocalizations (‘pyows’) heard once from the mangrove forest during a boat survey in the NW. In the SE one group of C. mitis was encountered (and ‘pyows’ heard) of which one individual was seen once during a ground survey. This group was in dense and diverse coastal shrub (including Acacia, Commiphora, baobab, and wild fruit trees) on coral rag on the edge of a mangrove forest. People report C. mitis to occur mainly in the mangrove forests.

W Manda Island: G. cocos heard.

Manda Island, at Takwa Ruins: G. cocos [4.1 individuals/h (n=6), 1.6 individuals/km (n=6)] and O. g. lasiotis [2.0 individuals/h (n=3), 0.8 individuals/km (n=3)].

Airport, Manda Island: several people said that C. p. excubitor visits the fresh water sources at the airport on a regular basis to drink.

Audio recordings: Advertisement calls recorded of G. cocos and O. g. lasiotis

Conservation The primate density in the Lamu Archipelago is very low. C. m. albotorquatus seems to be depended on the extensive mangrove forests surrounding the Islands. It is uncertain how much the other primate taxa depend on this ecosystem. Mangrove forests and other coastal wetlands occur for the most part on the north coast of Kenya, on the Lamu Archipelago, and at the tidal mouths of the Tana and Sabaki Rivers. Smaller wetlands occur in the mouths of semi perennial and seasonal coastal rivers on the south coast (Obura, 2007). The total area of mangroves forests in Kenya is estimated at 530 - 610 km². Sixty-sevel percent of Kenya’s mangroves occur in Lamu District, 10% in Kilifi District, and 10% in Kwale District. A total of 103 km² of mangrove forest has been lost [Abuodha & Kairo (2001) in Obura (2007)].

Mangrove forests have been extensively utilised for firewood, building poles (Rhizophora nzucronata is the preferred species), and masts (Heritiera littoralis) for Arab dhows (Irebelo, 2006). H.littoralis has been almost completely cut out from many sites. Mangrove swamps have been cleared, degraded and fragmented for human settlements and activities in several places (Irebelo, 2006; Obura, 2007).

Primates of the coastal forests of Kenya. 24 Y.A. de Jong & T.M. Butynski, 2009.

Remarks In the past, the islands of the ‘Lamu Archipelago’ (i.e., Lamu, Manda, Manda Toto, and Patta Islands), have also been referred to as the ‘Witu Islands’ (e.g., Groves, 2001). The channels that separate the islands of the Lamu Archipelago Islands from the mainland are shallow and narrow (Patta Island ca 3 km; Manda Island ca 20 m; Lamu Island ca 150-200 m). As such, Figure 5. Manda Toto Island on the left, Manda Island on the right. these islands are not particularly isolated. During low tide the two islands are almost connected. The extensive mangrove around all islands are probably a much greater barrier to primate movement than is the ocean (i.e., the channels).

There are two endemic subspecies of primates described for the Lamu Archipelago, but the validity of both taxa is in question (Groves 2001, Grubb et al. 2003). Manda Island is the type locality for C. p. excubitor. The geographical range of C. p. excubitor is fragmented among the islands and its numbers appears to be extremely low (see Chapter 4.2.7). The type locality of C. mitis phylax is Patta Island. It is unclear if C. m. phylax is endemic to Patta Island (see Chapter 4.2.5).

The density of diurnal primates on Lamu Island, Manda Toto Island, and Manda Island is very low. The diurnal primates that we observed were shy and the groups appeared to be small. C. mitis is present (apparently at low density) in the extensive mangrove forests of N Lamu Island (ca 150-200 m away from the mainland), and on NW and SW Manda Island. These are the first reported observation for C. mitis on these two islands. C. mitis were not detected on Manda Toto Island and are almost certainly absent from that tiny island (1 km²). It remains unknown as to which subspecies the C. mitis of Lamu Island and Manda Island belongs. The one C. mitis that we briefly observed on Manda Island could have been either C. m. albotorquatus or C. m. phylax.

During this survey we collected the first record of G. cocos for the Lamu Archipelago. This record extended the range of this species 70 km to the east (Butynski et al., 2006; Chapter 4.2.3; Appendix 6; Figure 15).

Further research: - Nocturnal and diurnal primate surveys on Patta Island to obtain presence/absence data, abundance data, and detailed field descriptions of all primate taxa.

- The validity of C. m. phylax and of C. p. excubitor need to be assessed. This will not be easy given the low abundance and shyness of both species in the Lamu Archipelago.

B. Kipini Conservancy and Witu Forest Reserve

Latitude/longitude: S2.49910; E40.60710 (Kipini Conservancy headquarters)

Altitude: 0-20 m asl

Vegetation: Coastal thickets, open grassland with doum palm and bushes, patches of medium to dense moist and diverse coastal forest. Sandy soils.

Primates of the coastal forests of Kenya. 25 Y.A. de Jong & T.M. Butynski, 2009.

Survey dates: 21-25 June 2006

Survey details: Total of 132.5 km, 48.4 h (25.4 diurnal, 23.0 nocturnal) in 5 days.

Diurnal surveys: 121.9 km; 113.9 km, 19.1 h by vehicle, 8.0 km, 6.3 h by foot Nocturnal surveys: 11.6 km; 10.6 km, 2.0 h by vehicle, 1 km, 0.9 h by foot Nocturnal listening hours: 20.1 h

Primate species encountered: P. c. ibeanus, C. m. albotorquatus, C. p. hilgerti, G. cocos, and O. g. lasiotis.

Encounter rates: Total of 32 diurnal primate groups encountered; 1.3 groups/h, 0.3 groups/km (n=32) P. c. ibeanus: 0.8 groups/h, 0.2 groups/km (n=20). C. m. albotorquatus: 0.4 groups/h, 0.1 groups/km (n=10). C. p. hilgerti: 0.1 groups/h, 0.02 groups/km (n=2). O. g. lasiotis: 0.3 individual/h, 0.3 individual/km (n=3). unidentified nocturnal primate: 0.04 individual/h, 0.1 individual/km (n=1)

Audio recordings: Audio recordings of the advertisement calls of G. cocos and O. g. lasiotis were obtained.

Conservation Kipini Conservancy was created in 2006. That same year, hundreds of illegal settlers arrived in the area, clearing a large portion of the natural vegetation. Other threats to the conservation of the area are illegal logging, tapping for palm wine, and poaching (Dowsett-Lemaire & Dowsett, 2007).

In Witu Forest Reserve, a high level of illegal logging was observed. Some parts of the forest have been cleared of large trees. The biodiversity of Witu Forest Reserve and the water catchment of the forest are seriously threatened (Nielsen & Sick, 2008). Figure 6. Cercopithecus mitis albotorquatus in Witu Forest Reserve

Both G. cocos and O. garnettii are at low densities in Kipini Conservancy. P. c. ibeanus and C. p. hilgerti are common and often found in the vicinity of human settlements (in towns/villages, gardens). C. m. albotorquatus, a poorly-known subspecies, is common. Some authorities suggest state that C. m. albotorquatus is confined to the forests of the Lower Tana River (Chapter 4.2.5). That this is not the case as this subspecies is abundant in the Witu and Kipini forests. This is one

Primates of the coastal forests of Kenya. 26 Y.A. de Jong & T.M. Butynski, 2009. of the more significant finding of this project and one that bodes well for the long-term conservation of C. m. albotorquatus.

In Kipini Forest we obtained the first solid evidence (good recordings of the advertisement call) for G. cocos north of the Tana River---extending the range ~60 km to the NE. This species is otherwise only known from the coastal forests of extreme northern Tanzania, and southern and central Kenya (Chapter 4.2.3, Appendix 6). The presence of G. cocos north of the Tana River/Tana Delta is of considerable conservation significance and suggests that this species may occur in extreme southern Somalia.

Remarks Over the past 5 years, we have received reports of Procolobus in the Witu Forest Reserve and Kipini Forest. Kipini is ca. 90 km southeast of the known range of P. r. rufomitratus. Chapter 4.2.10 describes in more detail our search for Procolobus in Kipini Conservancy. If Procolobus are present at Witu and/or Kipini, they must be in very low numbers. This, coupled with the fact that the area of Kipini Forest in which Procolobus are said to occur has recently been given over by the Kenya Wildlife Service (KWS) and Kipini Conservancy (with funding from World Bank) for resettlement of people from the Lower Tana River, means that time is a critical factor as the small area of remaining (relatively species-rich) forest is likely to be destroyed in the near future. If Procolobus are present, then immediate actions will be necessary to prevent resettlement and to find an alternative solution for the people being resettled. During February - March, 2007, members of the Danish Zoological Society (with CEPF funding) conducted biodiversity surveys in Witu Forest Reserve (Nielsen & Sick, 2008). Twenty mammal species, including four primates (Cercopithecus mitis, Papio cynocephalus, “Otolemur crassicaudatus” and ”Galago gallarum or senegalensis bracattus”, and eighty-five species of birds were recorded. Other recorded mammals include Lion Panthera leo, Bush Elephant Loxodonta africana, African Buffalo Syncerus caffer, Harveys Duiker Cephalophus harveyi, Suni Neotragus moschatus, and the Lesser Elephant Shrew Elephantulas rufescens. Duiker densities were low compared to other locations in East Africa. This was likely due to differences in habitat type and quality (Nielsen & Sick, 2008).

Nielsen & Sick (2008) mention the possible presence of Galago gallarum in Witu Forest Reserve. Chapter 4.2.2 mentions that this is higly unlikely, given the habitat preference of G. gallarum (dry Acacia-Commiphora woodland/bushland and thorn scrub; Butynski & De Jong, 2004). It is more likely that the galago species they encountered during their surveys was either G. cocos or G. s. braccatus (Chapter 4.2.2), with G. cocos being the more likely. Nielsen & Sick (2008) report the presence of Otolemur crassicaudatus in Witu Forest Reserve. We encountered O. g. lasiotis in the area and strongly doubt the presence of O. crassicaudatus in the Reserve since this species is not known to occur in Kenya (Groves, 2001; T. M. Butynski & Y.A. de Jong, unpubl. data). The photos of Otolemur as presented in the report appear to be of O. g. lasiotis.

Dowsett-Lemaire & Dowsett (2007) list the mammals they encountered during their vegetation, birds and mammals surveys in Kipini Conservancy during October-December, 2006. Five primate species are listed; Galagoides zanzibaricus, Otolemur garnettii, Papio cynocephalus, Cercopithecus mitis albogularis, and Cercopithecus aethiops. If we apply the taxonomy we use in this study to all five primate species, and add our knowlegde about their subspecies, the list becomes: G. cocos, O. g. lasiotis, P. c. ibeanus, C. m. albotorquatus and C. p. hilgerti. According to Dowsett-Lemaire & Dowsett (2007), all five primate species are common in the Conservancy.

Primates of the coastal forests of Kenya. 27 Y.A. de Jong & T.M. Butynski, 2009.

G. senegalensis was not recorded in Kipini Conservancy by Dowsett-Lemaire & Dowsett (2007). They observed G. senegalensis “in the bush country” of the Tana River Primate National Reserve (Andrews et al., 1975) and concluded that this species is probably not in Kipini Conservancy (in the absence of thornbush).

Neither Nielsen & Sick (2008) nor Dowsett-Lemaire & Dowsett (2007) found Procolobus in the Witu - Kipini area, nor did we find evidence for Procolobus in this region. It now seems highly unlikely that Procolobus occur in this region.

Further research There are no priorities for further primate research in the forests of the Witu - Kipini area.

C. Tana River Primate National Reserve

Latitude/longitude: S1.87587; E40.13804

Altitude: 40 – 50 m asl

Vegetation: Riverine evergreen gallery forest surrounded by grassland, wooded grassland, and arid medium-dense bushland on sandy soil (floodplain). Characteristic trees include Ficus spp, Phoenix reclinata, Acacia robusta, Populus ilicifolia, Blighia unijugata, Sorindeia madagascariensis, Diospyros mespiliformis, and Mimusops obtusifolia (Roberts & Luke, 1993 in Bennun & Njoroge, 1999).

Survey dates: 6-7 August 2005 and 20-21 June 2006

Survey details: In total 30.3h [15.0 h (11.0 diurnal, 4.0 nocturnal) and 15.3 h (9.3 diurnal, 6.0 nocturnal)].

Diurnal surveys: ca 2 km, vehicle, along the edge of the forest Nocturnal surveys: ca 1 km, foot Nocturnal listening survey: 8 h.

Recordings, photos, and phenotypic descriptions made of primates in the vicinity of Mchelelo Research Camp.

Primate species encountered: P. c. ibeanus, C. m. albotorquatus, P. r. rufomitratus, C. galeritus, G. cocos, and O. g. lasiotis.

Chlorocebus pygerythrus hilgerti and Galago senegalensis braccatus are present (Andrews et al., 1975; Butynski & Mwangi, 1994; Bearder et al., 2008), but were not encountered during this survey.

Encounter rates 6-7 August 2005: Total of six diurnal primate groups encountered; 0.6 groups/h (n=6). P. c. ibeanus: 0.1 groups/h (n=1). C. m. albotorquatus: 0.3 groups/h (n=3). P. r. rufomitratus: 0.1 groups/h (n=1). C. galeritus: 0.1 groups/h (n=1).

Primates of the coastal forests of Kenya. 28 Y.A. de Jong & T.M. Butynski, 2009.

Encounter rates 20-21 June 2006: Total of seven diurnal primate groups encountered: 0.8 groups/h and 25 nocturnal primates; 4.2 individuals/h. P. c. ibeanus encountered; 0.1 groups/h (n=1). C. m. albotorquatus encountered; 0.3 groups/h (n=3). P. r. rufomitratus encountered; 0.1 groups/h (n=1). C. galeritus encountered; 0.2 groups/h (n=2). O. g. lasiotis encountered; 15 individuals/h (n=15; estimate of 15 animals/ha). G. cocos encountered; 10 individuals/h (n=10; estimate of 15 animals/ha).

Audio recordings The adult male loud call (‘whoop-gobble’) of C. galeritus was recorded and send to Carolyn Ehardt, Project Leader of the CEPF-funded project ’Taxonomy and Conservation Genetics of the Threatened Mangabey Taxa of the Eastern Arc Mountains and Coastal Forests of Tanzania and Kenya Biodiversity Hotspot’ (a joint project with T. Butynski). The recordings were also sent to Dr. Jean-Pierre Gautier, the authority on the vocalizations of Cercocebus. Other species whose calls were recorded in the Tana River Primate National Reserve are O. g. lasiotis and G. cocos.

Conservation Primate densities in Tana River Primate National Reserve are high. Six primate taxa were encountered and two more are known to occur in the Reserve. Two out of the eight primate taxa are listed as “Endangered” on the 2008 IUCN Red List of Threatened Species (C. galeritus and P. r. rufomitratus) and one is listed as “Vulnerable” (C. m. albotorquatus). The other taxa are listed as “Least Concern” (IUCN, 2008).

Habitat loss due to agricultural clearing and extraction of forest products by local communities is the main threat to all primate taxa in the Tana River Primate National Reserve (Butynski & Mwangi, 1994, 1995; Mbora, 2003; Oates et al., 2008; Struhsaker & Grubb, in press). Additionally, there are major changes in the flow volume and flood cycles of the Tana River as a result of the construction of five hydroelectric power dams upriver (Butynski 1995). This is thought to have major impacts on the establishment and survival of the forests of the Lower Tana River. Recent agricultural developments in the area (e.g., two proposed large sugar cane plantations and one proposed large oil palm plantations) will result in a large influx of people to the region with a resultant increased demand for forest products and forest land. The proposed Tana Integrated Sugar Project in Tana River and Lamu Districts alone threatens more than 200 km² of natural habitat.

Remarks During this study, we obtained photos and field descriptions for several primate taxa, including C. m. albotorquatus and P. c. ibeanus. Intra group phenotypic variation for P. c. ibeanus here is exceptionally great. Additionally, the phenotype of P. c. ibeanus here varies from that observed at Manda Toto Island, which is, apparently, close to the type locality of the subspecies (which is said to be Lamu Island). Chapter 4.2.8 describes the morphology of P. c. ibeanus in the Kenya coastal forests in more detail.

Further research More individual P. c. ibeanus along the Lower Tana River need to be described, photographed, and compared with P. c. ibeanus in the coastal forests to the north and south of the Tana Delta and on the Lamu Archipelago.

Primates of the coastal forests of Kenya. 29 Y.A. de Jong & T.M. Butynski, 2009.

Presence-absence data of G. s. braccatus should be collected in or in the vicinity of the Reserve

Details about the diurnal road surveys conducted at the north coast of Kenya are presented in Appendix 3

4.1.2 South Coast of Kenya

A. Kaya Rabai (Chijembeni), Mazeras

Latitude/longitude: S3.94465 E39.58189

Altitude: 210 m asl

Vegetation: Dry coastal forest, coconut palms Cocos nucifera, cashew nut Anacardium occi and mango Mangifera indica common on the forest edge. Agricultural land, human settlements and bush lands surround Kaya Rabai (Chijembeni).

Survey date: 20-21 February 2006

Survey details: Nocturnal listening hours and audio recording: 10.3 h

Primate species encountered: G. cocos, O. g. lasiotis. C. m. albogularis, C. p. hilgerti, and P. c. ibeanus are reported by residents to be present in the area but were not encountered during this survey. There are reports of C. a. palliates from Kaya Rabai (last report 1945; Anderson, 2007).

Encounter rate: individuals G. cocos were heard very often and occasionally seen.

Audio recordings: Digital and tape recordings were acquired of advertising calls and other vocalizations of G. cocos and O. g. lasiotis. Recordings were sent for analysis to the Nocturnal Research Group in Oxford. The sonogram and oscillogram of the vocalizations recorded were used to confirm that G. cocos is the name that should be applied to the Kenya coast galago---not Galagoides zanzibaricus (Butynski et al., 2006).

Conservation: Kaya Rabai consists of a number of coastal forest fragments (totalling 8 km²) and is part of the Sacred Mijikenda Kaya Forests. In 2008 these forests received UNESCO World Heritage Site status [criteria (iii)(v)(vi); Unesco.org, 2009].

The forests in and around Mazeras are highly fragmented, degraded, and surrounded by agriculture. Some of the forests are sacred and at least somewhat protected. There are five primate taxa in and around these forests. The three forest-depended species (G. cocos, O. g. lasiotis, and C. m. albogularis) are likely declining as a result of habitat degradation and loss.

Remarks: Data collected during this survey, together with earlier data from this area, have led to the following publication:

Butynski, T.M., De Jong, Y.A., Perkin, A.W., Bearder, S.K. & Honess, P.E. 2006. Taxonomy, distribution, and conservation status of three species of dwarf galagos (Galagoides) in Eastern Africa. Primate Conservation 21: 63-79.

Primates of the coastal forests of Kenya. 30 Y.A. de Jong & T.M. Butynski, 2009.

Abstract: This paper reviews the complicated nomenclatural history for the Kenya coast galago, Galagoides cf. cocos, and examines whether ‘cocos’ is the valid species name for this recently resurrected taxon. This paper also reviews the phenotypic and vocal differences among G. cocos; the Zanzibar galago (Galagoides zanzibaricus zanzibaricus); the Udzungwa galago (Galagoides zanzibaricus udzungwensis); and the Mozambique galago (Galagoides granti), as well as their geographic ranges and conservation status. The following are among the findings: (1) Galagoides cocos’ is the name that should be applied to the Kenya coast galago; (2) in the field, the loud calls of these three species are diagnostic and remain the best means for identification; (3) there is a suite of phenotypic characters that, when taken together, can be used to distinguish among these three species when in the hand or viewed in the field in good light at close range; (4) G. z. zanzibaricus is phenotypically distinct from G. z. udzungwensis; (5) the three species are parapatric or, perhaps, narrowly sympatric; (6) the three species are endemic to the coastal forests of eastern Africa with G. cocos in the north (Kenya and northeastern Tanzania), G. zanzibaricus in Tanzania, and G. granti from southern Tanzania to southern Mozambique; and (7) none of the three species is threatened at this time, although G. z. zanzibaricus meets the IUCN Red List criteria for an Endangered subspecies.

The complete article is attached in Appendix 6.

Further research: The subspecies status of the C. mitis that occurs in Kaya Rabai needs to be determined.

B. Diani

Latitude/longitude: S4.33461; E39.56251 and S4.2848; E39.5913

Altitude: 0 to ca 20 m asl

Vegetation: Gardens with tall exotic and indigenous trees, edge with patches of deciduous coral rag forest.

Survey date: 23 April 2008 and 14 December 2008

Survey details: 23 April 2008 Nocturnal surveys: 0.8 km, 3.25 h on foot. Nocturnal listening survey: 2.72 h. Due to bad weather conditions further nocturnal surveys were cancelled

14 December 2008 Opportunistic observations: ca. 4 h.

Primate species encountered: P. c. ibeanus, C. m. albogularis, C. a. palliates, C. p. hilgerti, C. p. hilgerti x C. m. albogularis hybrid, O. g. lasiotis, G. cocos.

Encounter rates: 23 April 2008 O. g. lasiotis: 6.0 individuals/h, 3.6 individuals/km (n=3) G. cocos: 7.6 individuals/h, 4.8 individuals/km (n=4)

Primates of the coastal forests of Kenya. 31 Y.A. de Jong & T.M. Butynski, 2009.

Audio recordings: The advertisement call and other vocalizations of O. g. lasiotis and G. cocos recorded.

Conservation: Originally, Diani was one of the most diverse areas of forest along the Kenya coast with a rich coral rag flora (Rovertson & Luke, 1993 in Bennun & Njoroge, 1999). Six primate species occur in and around Diani and primate densities are high. Diani is a popular tourist destination with many large hotels and houses on large green compounds. All six primate species encountered in Diani were observed on hotel compounds, foraging on flowers, and sleeping in indigenous and exotic tall trees. Most primate species, but in particular P. c. ibeanus, C. m. albogularis, and, C. p. hilgerti, feed on hand-outs and raid tourist facilities and garbage bins for human food items. The destructive effects of tourist development on the natural vegetation is obvious in Diani. The Colobus Trust, an NGO concerned with primate (particular C. a. palliates) conservation along the south coast of Kenya, has established several practical conservation activities to protect the primates and the forests of the area.

Coastal forest within the Diani-Chale Marine Reserve are being cleared, fragmented and degraded by big tourist companies and hotels, despite the protected status of the Reserve (colobus.wildlifedirect.org, 2008).

Remarks: One adult male C. p. hilgerti x C. m. albogularis hybrid was encountered once (14 December 2008 at 12:45 h) during 4 days of opportunistic surveys. The hybrid was with two adult C. m. albogularis of which at least one was a male. Chapter 4.2.12 presents the details and photos of this hybrid.

During a survey of C. a. palliates in 2001, Anderson et al. (2007) found 50 groups comprised of 332 individuals in Diani. This is the second biggest population in Kenya after Shimba Hills National Reserve. The biggest threat to their survival is at Diani is loss of habitat due to tourism activities. Other threats are illegal logging, charcoal production, firewood collection and hunting (Anderson et al., 2007).

Further research: More surveys should be conducted in and around Diani in search of additional C. p. hilgerti x C. m. albogularis hybrids. The density, phenotypic characters, ecology and behaviour of these hybrids should be studied.

C. Mrima Hill Forest

Latitude/longitude: S4.48714; E39.26224

Altitude: 100-300 m asl

Vegetation: Degraded and secondary coastal forest. Road surveys on edge of forest and small agricultural settlements with trees such as mango, cashew, baobab and coconut palm).

Survey dates: 7 February 2006, 17:00 - 18:50 h 20 February 2006, 11:00 - 12:02 h

Primates of the coastal forests of Kenya. 32 Y.A. de Jong & T.M. Butynski, 2009.

Survey details: Diurnal surveys: 21.2 km, 2.9 h by vehicle, 10.0 km/h.

Primate species encountered: C. m. albogularis and C. a. palliatus

Encounter rates: Total of 19 primate groups encountered; 6.6 groups/h, 0.9 groups/km (n=19). C. m. albogularis: 3.4 groups/h, 0.5 groups/km (n=10). C. a. palliates: 3.1 groups/h, 0.4 groups/km (n=9).

Audio recordings: None

Conservation: Mrima Hill Forest is a Forest Reserve, Nature Reserve and National Monument. Despite its protected status, Mrima Hill Forest has been severely degraded due to selective logging and pole cutting and is surrounded by intensively cultivated farmland and a high human population density (e.g. Bennun & Njoroge, 1999). Mrima Hill Forest holds high densities of C. m. albogularis and C. a. palliatus. In 2001, Anderson et al. (2007) counted 13 groups (82 individuals) of C. a. palliatus in Mrima Hill Forest. Illegal logging, charcoal burning and firewood collection are reported to occur only on a low scale by Anderson et al. (2007) and are the only threats to C. a. palliatus in Mrima Hill Forest. This contradicts the information on conservation threats as given by Bennun & Njoroge (1999). Further information on the threats to the Mrima Hill Forest needs to be obtained.

Remarks: The area needs to be revisited to conduct nocturnal primate surveys, surveys on foot within the forest (opposed to vehicle surveys from the edge of the forest), and to describe and survey the surrounding area to assess the conservation threats.

Further research: - Conduct nocturnal surveys in Mrima Hill Forest - Assess conservation issues that threaten Mrima Hill Forest - The subspecies status of the C. mitis that occurs in Mrima Hill Forest needs to be determined.

D. Kaya Sega

Latitude/longitude: S4.54612; E39.09309

Altitude: 76 m asl

Vegetation: Dense to medium-dense Commiphora bushland surrounded by patches of dry coastal forest. Seasonally dry river bed with tall trees.

Survey date: 22 April 2008

Survey details: In total 3.8 h nocturnal surveys

Nocturnal survey: 1.0 h, 1.6 km on foot Nocturnal listening hours and audio recording: 2.8 h

Primates of the coastal forests of Kenya. 33 Y.A. de Jong & T.M. Butynski, 2009.

Primate species encountered: O. g. lasiotis

Encounter rate: O. g. lasiotis: 1.9 individuals/h, 1.3 individuals/km (n=2) Several individuals were heard, observed, and recorded from camp.

Audio recordings: Advertisement call and other vocalizations of O. g. lasiotis recorded from camp.

Further research: Diurnal surveys should be conducted to obtain presence/absence and relative abundance figures of the diurnal primates.

Details about the diurnal road surveys conducted at the south coast of Kenya are presented in Appendix 4

4.1.3 Outside the coastal forests of Kenya

A. Mwea National Reserve

Latitude/longitude: S0.80487; E37.59122

Altitude: 1015-1120 m asl

Vegetation: Mostly Acacia-Commiphora bush-woodland but some dense, tall riverine vegetation.

Survey dates: 3-4 August 2005, 28 April 08

Survey details: 3-4 August 2005 A total of 11.33 h, >26.4 km were surveyed in 1,5 days

Diurnal surveys: >15.4 km; 5.63 h by vehicle Nocturnal surveys: 11 km; 1.18 h by vehicle Nocturnal listening hours: 4.52 h

28 April 2008 Diurnal surveys: 40.21 km; 2.95 h by vehicle

Primate species encountered: C. mitis, Papio sp., and C. p. hilgerti

Encounter rates: 3-4 August 2005 A total of 11 diurnal primate groups were encountered: 1.0 groups/h (n=11)

C. mitis: 0.7 groups/h (n=4) C. p. hilgerti: 0.7 groups/h (n=3) Papio: 0.7 groups/h (n=4)

Primates of the coastal forests of Kenya. 34 Y.A. de Jong & T.M. Butynski, 2009.

28 April 2008 Papio: 0.7 groups/h, 0.05 groups/km (n=2)

Remark No galagos were encountered or heard in the 5.7 nocturnal hours we surveyed despite what appeared to be very suitable habitat for O. g. lasiotis or G. s. braccatus.

Further research Photos and field descriptions of Papio taken in Mwea National Reserve need to be analyzed and compared with photos and field descriptions of P. c. ibeanus and P. anubis to determine if Mwea National Reserve is part of the hybrid zone for Papio.

Photos and field descriptions of C. mitis taken in Mwea National Reserve need to be analyzed and compared with photos and field descriptions of C. m. albotorquatus and C. m. kolbi to determine if C. mitis occurring in Mwea National Reserve are intermediates (as or preliminary analysis suggests).

Additional nocturnal surveys need to be conducted to determine which nocturnal primates occur in Mwea National Reserve---if any.

B. Kiboko Camp, Makindu

Latitude/longitude: S2.20346; E37.71383

Altitude: 956 m asl

Vegetation: Acacia xanthophloea woodland, medium dense Acacia bushland.

Survey dates: 6 February 2006

Survey details: A total of 5.79 h of surveys were completed in one night

Nocturnal survey: 2.23 h Nocturnal listening survey: 1.33 h

Primate species encountered: O. g. lasiotis and G. s. braccatus. Other primate species observed in this area are Papio and C. p. hilgerti (Y.A. de Jong & T. M. Butynski, pers. obs.).

Encounter rates: A total of two nocturnal primates were encountered: 2.22 individuals/h (n=2)

O. g. lasiotis: 1.11 individuals/h (n=1) G. s. braccatus: 1.11 individuals/h (n=1)

C. Tsavo West National Park

Latitude/longitude: S2.68341; E38.17749 (Mtito Andei, head quarters of the National Park)

Primates of the coastal forests of Kenya. 35 Y.A. de Jong & T.M. Butynski, 2009.

Altitude: 700 – 1300 m asl

Vegetation: Dense to open Acacia – Commiphora bushland, ground water forests, plains.

Survey dates: 19-21 April 08

Survey details: Total of 150.7 km, 9.51 h (7.0 diurnal, 2.5 nocturnal) in 2 days.

Diurnal survey: 150.7 km; 7.01 h by vehicle Nocturnal survey: 0.5 km; 0.50 h by foot Nocturnal listening hours: 2.0 h

Primate species encountered: C. m. albogularis, C. p. hilgerti, Papio, and G. s. braccatus

Encounter rates: A total of 15 diurnal primate groups were encountered: 1.8 groups/h, 0.1 groups/km (n=15).

C. m. albogularis: 0.4 groups/h, 0.02 groups/km (n=3) C. p. hilgerti: 0.7 groups/h, 0.03 groups/km (n=5) Papio: 1.0 groups/h, 0.05 groups/km (n=7)

G. s. braccatus: 4 individuals/h (n=2)

Remarks: Desert warthog P. aethiopicus and common warthogs P. africanus were found sympatric in Tsavo West National Park. Data from this trip were published in Swara (Culverwell et al., 2008. A new pig for Tsavo Swara 31: 50-52) and in Suiform Soundings, the the newsletter of the IUCN/SSC Pigs, Peccaries, and Hippos Specialist Group (De Jong et al., 2009. Desert warthog Phacochoerus aethiopicus found in Tsavo East National Park and Tsavo West Pational Park, southern Kenya, Suiform Soundings 8: 4-6). Both articles are attached in Appendix 7 and 8.

D. Tsavo East National Park

Latitude/longitude: S3.36731; E38.56476 (Voi, head quarters of the National Park)

Altitude: 150 – 1200 m asl

Vegetation: Dense to open Acacia – Commiphora savannah bushland, open plains.

Survey dates: 26-27 April 08

Survey details: Total survey hours: 10.7 h Diurnal surveys: 135 km; 7.7 h by vehicle Nocturnal listening hours: 3.0 h

Primate species encountered: C. p. hilgerti, Papio and O. g. lasiotis

Primates of the coastal forests of Kenya. 36 Y.A. de Jong & T.M. Butynski, 2009.

Encounter rates: Two groups of diurnal primates were encountered, 0.65 groups/h, 0.02 groups/km (n=2) C. p. hilgerti: 0.3 groups/h, 0.01 groups/km (n=1) Papio: 0.3 groups/h, 0.01 groups/km (n=1)

Audio recordings: none

E. North Kilimangodo

Latitude/longitude: S4.28659; E39.05576

Altitude: 223 m asl

Vegetation: Degraded Acacia bushland surrounded by agriculture and human settlements

Survey dates: 21 April 08

Survey details: Nocturnal listening survey: 4 h

Primate species encountered: Vocalizations heard of O. g. lasiotis and G. s. braccatus, no primates encountered.

Audio recordings: Advertisement call recorded of G. s. braccatus.

Details about the diurnal road surveys conducted outside the coastal forests of Kenya are presented in Appendix 5

Primates of the coastal forests of Kenya. 37 Y.A. de Jong & T.M. Butynski, 2009.

4.2 Primates species of the coastal forests of Kenya

4.2.1 Otolemur garnettii lasiotis (Peters, 1876)

Vernacular name: White-tailed greater galago Subspecies type locality: Mombasa, Kenya Synonym: hindei Elliot, 1907. Kitui, Kenya

Field characters: Large galago with ears of intermediate size (relative to the other subspecies of O. garnettii). Muzzle often darker than rest of face. Inside of ears hairless, pinkish. Dorsum, sides and outer limbs vary from dark brown, brown, brownish-grey, to silvery grey. Ventrum paler than dorsum, varying from brown to grey or off- white. Tail bushy and similar to dorsum in colour as dorsum. Tail tip varies from black to off Figure 7. Otolemur garnettii lastiotis present in good white. densities (>15 individuals/ha) in riverine forest in the Tana River Primate National Reserve, Kenya.

Distribution: Lower Jubba River, Somalia, south to Kenya. Along the Kenya coast inland to Kibwezi and the Taita Hills. South to Tanga, NE Tanzania (Figure 8; Schwarz, 1931; Olson, 1979; Groves, 2001). O. g. lasiotis is the most north-eastern subspecies of O. garnettii.

Extent of occurrence: ca. 80,100 km². The area of occupancy is but a very small portion of the extent of occurrence as the habitat suitable for O. g. lasiotis is limited within its geographic range.

Figure 8: Approximate geographic distribution of Otolemur garnettii lasiotis in Kenya with sites of encounters (2003-2008) also depicted. Based on Y. de Jong & T. Butynski (unpubl. data) and Butynski et al. (2008a).

Primates of the coastal forests of Kenya. 38 Y.A. de Jong & T.M. Butynski, 2009.

Photographic map: Photo map for Galagonidae, contains 28 photographs, of which four are of O. g. lasiotis (T. M. Butynski & Y. A. de Jong). http://picasaweb.google.com/wildsolutions/Galagonidae?feat=email# or via: www.wildsolutions.nl

Encounter sites: During this study, O. g. lasiotis was encountered in all six areas we conducted nocturnal surveys in the Coastal Forest of Kenya.

Encounter sites in the coastal forests of Kenya and to the Hotspot were; Chyulu Hills National Park; Kiboko Camp, Makindu; Ndololo Camp, Tsavo East National Park; Mazeras; Kaya Sega; north Kilimangodo; Diani; Gedi Ruins; Watamu; Arabuko Sokoke Forest; Malindi; Tana River Primate National Reserve; Kipini Conservancy; NE of Witu Forest; Manda Island.

Encounter rate: A total of at least 40 O. g. lasiotis were observed in the coastal forests of Kenya. Twenty-five individuals were observed during survey hours. Hundreds of O. g. lasiotis advertisement calls were heard (Table 2; Figure 8).

Table 2. Rates of encounter with Otolemur garnettii lasiotis from 2005 – 2008 in the coastal forests of Kenya. Area Individuals/hour Individuals/km Vehicle/Foot Kaya Sega 1.9 (n=2) 1.3 (n=2) Foot Diani 6.0 (n=3) 3.6 (n=3) Foot Tana River PNR 15 (n=15) - Foot Kipini Conservancy 0.8 (n=1) 0.1 (n=1) Vehicle NE of Witu Forest 4.3 (n=2) 4.0 (n=2) Foot Manda Island 2.0 (n=3) 0.8 (n=3) Foot

Vegetation: O. garnettii is a forest-depended species. During this study, O. g. lasiotis was encountered in dry coastal forest, evergreen forest, semi-evergreen riverine forest, secondary forest, fragmented forest, degraded coastal forest, gardens with tall trees, and on the forest edge.

Altitude range: Sea level to ca 1300 m asl (Chyulu Hills National Park).

Vocalizations: The advertisement call of O. g. lasiotis is a ‘trailing call’; a patterned sequence of units arranged into a phase with a loud beginning and which trails away over the last few units (Bearder et al., 1995; Honess, 1996). Calls are given singly both by males and females and are often replied to by other individuals (also Honess, 1996). Each trailing call comprises six to 10 loud cries reaching a crescendo and then trails off for the last three or four units (also Nash et al., 1989). The frequency of calling varies and depends on the population density (Bearder et al., 2003). The oscillogram and sonogram of the advertisement call of O. g. lasiotis are presented in Figure 9.

Figure 9. Sonogram and oscillogram of the trailing advertising call of Otolemur garnettii lasiotis from Diani Beach, 34 km south of Mombasa, SE Kenya, the type locality of this subspecies. Call recorded by Y. de Jong and T. Butynski. A number of advertisement calls were heard in the beginning of

Primates of the coastal forests of Kenya. 39 Y.A. de Jong & T.M. Butynski, 2009. the evening before heavy weather made it impossible to record. The call follows the typical series of loud cries ending with a trailing off (last four units). For the above recording: call length = 3.9 seconds; frequency range = 0.38–4.41 kHz; fundamental frequency = 0.57 kHz; number of units = 9; unit duration: x= 0.27 sec (range: 0.209-0.360, n=9, sd=0.06).

Conservation status: O. g. lasiotis occurs locally in high densities and is able to use secondary, fragmented and/or degraded vegetation. This subspecies occurs in a number of protected areas in Kenya, including the Tana River Primate National Reserve, Tsavo West National Park, Tsavo East National Park, Arabuko Sokoke Forest, Kipini Conservancy, Kaya Rabai, and other Mijikenda Kaya Forests (UNESCO World Heritage site since 2008; Unesco, 2009). No major treats for the species are reported (also IUCN, 2008), however, its fragmented distribution makes it vulnerable to local extinction due to vegetation clearing for agriculture, tourism, etc. (also Harcourt & Perkin, in press).

IUCN Red List: Least Concern (Butynski et al., 2008a; www.iucnredlist.org/details/136900). Assessors: T.M. Butynski, S. Bearder, & Y.A. de Jong.

Discussion: Otolemur garnettii is a common species in the coastal forests of Kenya. The taxonomy of Otolemur has been debated for many years. This study recognised O. g. lasiotis as the subspecies present in the coastal forests of Kenya. This is consistent with Hill & Meester (1974), Jenkins (1987), Groves (2001, 2005), Grubb et al. (2003), and Harcourt & Perkin (in press). Based on analysis of the geographic variation (phenotypic variation and skull measurements) found in O. garnettii, Olson (1979) considers lasiotis to be a synonym of Otolemur garnettii garnettii [Ogilby, 1838, type specimen from Unguja Island (Zanzibar), Tanzania]. According to Olson (1979), O. g. garnettii occupies the forests of the lower reaches of both the Juba and Webi Shabelle Rivers in Somalia, the lower Tana and Sabaki Rivers in Kenya, and isolated forests in the Chyulu and Taita Hills. In Tanzania, O. g. garnettii inhabits the entire length of the coastal region but is highly discontinuous (Olson, 1979).

Jenkins (1987), Groves (2001), Harcourt & Perkin (in press) consider O. g. garnettii to be restricted to Unguja, Pemba, and Mafia Islands. The dorsum of this subspecies is rich reddish-brown and the ventrum is yellow with a slightly greenish tone. Tail brown or greyish-brown with a dark brown or blackish tail tip (Jenkins, 1987; Groves, 2001; Harcourt & Perkin, in press). Olson (1979) accepts that there are phenotypic differences between O. garnettii on Unguja and Pemba with those of the coastal mainland, but considered the differences to be no greater than those occurring between O. garnettii on the two islands.

Several authors considered the tail tip in O. garnettii to be one of the diagnostic characters of the subspecies. A pale tail tip (in contrast to the colour of the dorsum and proximal part of the tail) is said to be typical for O. garnettii on the coast of the mainland. However, white tail tips are also found in the island populations (Olson, 1979). On both Pemba and Unguja Islands we only encountered O. g. garnettii with dark tail tips (n=10) during a brief survey of four nights (Butynski & De Jong, unpubl. data). Olson (1979) reports a high percentage of white tail tips in O. garnettii between the Pangani River, Tanzania, and Mombasa, Kenya. During our field studies, however, we observed seven O. garnettii in this region, six of which had a dark brown or blackish tail tip (Kaya Gonja, Kenya, and just a few kilometres south of the Kenya-Tanzania border) and one with an all brown tail (Diani, Kenya). Of all the 40 O. g. lasiotis that we described on the coast of Kenya, two had a whitish or greyish tail tip, 28 had a dark brown-blackish tail tip, and one had an all brown tail (Figure 10). In conclusion, in the coastal forests of Kenya, the colour of the tip of the tail O. g. lasiotis varies from whitish to greyish to blackish. Throughout the geographical range of O.

Primates of the coastal forests of Kenya. 40 Y.A. de Jong & T.M. Butynski, 2009. garnettii in East Africa, outside the Kenya coastal area, we encountered whitish, greyish and blackish tail tips, sometimes at the same site (Butynski & De Jong, unpubl. data). We conclude that the colour of the tip of the tail cannot be used as a character for distinguishing among subspecies of O. garnettii.

Figure 10: Colour of the tip of the tail of Otolemur garnettii lasiotis on the coast of Kenya (Y. de Jong & T. Butynski, unpubl. data; Butynski et al., 2008).

In April, 2008, we encountered a captive adult male Otolemur garnettii in Malindi. The individual looked phenotypically different (Figure 11) from any O. g. lasiotis we had observed along the coast of Kenya. The caretaker of this animal informed us that it had been brought in when a juvenile (2.5 years ago). Despite the unsuitable and unhygienic enclosure, the animal seemed healthy.

Figure 11. Captive adult male Otolemur garnettii in Malindi, Kenya. Field characters: Face grizzled silvery grey with a dusky silver wash on the cheeks and around the muzzle. Eye rings absent. Area around the eyes pinkish. Rhinarium dark grey. Area around rhinarium pinkish and sparsely haired. Inside and outside ears hairless and pink. Crown and dorsum silvery grey with mouse grey under fur. Lower back, outer hind legs, and tail dusky silver. Ventrum and inner limbs silvery white. Hands hairless and pinkish. Finger tips brownish. Ear length ca. 41 mm.

Primates of the coastal forests of Kenya. 41 Y.A. de Jong & T.M. Butynski, 2009.

The mean ear length of O. g. lasiotis is 47 mm (range: 43-54 mm for males, n=3; Taita Hills, Kenya; A. Perkin, pers. obs. in Harcourt & Perkin, in press). The mean ear length of O. garnettii is 45 mm (range: 34-55 mm for both sexes combined, n= 356; Olson & Nash, 2002). The ear length of the Malindi Otolemur falls well within this range and is close to the measurements taken by Perkin in the Taita Hills.

Olson (1979, p. 315) notes the considerable variation in the pelage colour of O. g. lasiotis and contrasts this to the consistency in colour of the highland subspecies (O. g. kikuyuensis). The National Museums of Kenya holds eight brown specimens collected in the Gedi area (Olson, 1979). Olson (1979) describes O. g. lasiotis as ”typically brown” but found five specimen (n = 29) with a greyish pelage, all collected along the Kenya coast in the vicinity of Gede (= Gedi, Polhill, 1988; ca. 15 km south of Malindi town). Three of those specimen are ”predominantly brown but possess a light greyish wash to the pelage on the sides of the body”. The other two specimen are ”unmistakably grey in colour with only the slightest hind of a brownish wash to the dorsal hairs”. According to Olson (1979), these specimens resemble Galago crassicaudatus monteiri (silver greater galago Otolemur monteiri in current taxonomy; Groves, 2001; Grubb et al., 2003) but the external and cranial dimensions, as well as the texture of the pelage, indicate that they are to O. garnettii. In 1963 (J. Buettner-Janusch & V. Buettner-Janusch, pers. comm. in Olson, 1979) reported high variation in pelage colour in the Sokoke Forest (Gedi area) population of O. garnettii. Olson (1979) suggests that further studies be conducted into the ”nature and source of this exceptionally wide range of variation” in O. garnettii for this part of Kenya.

Several nocturnal primate specialists (including Simon Bearder, Leanne Nash, Caroline Harcourt and Andrew Perkin) were contacted during this study in order to identify the Malindi Otolemur. L. Nash (pers. comm.) commented that six out of the 12 O. garnettii that she described from Gedi were ”silvery”, ”grey with brown”, or ”grey-brown”. She analysed the photographs of the Malindi Otolemur and suggested that, on the basis of the face/ear relative proportions, that this is O. garnettii. S. Bearder (pers. comm.) suggested that this individual is an O. garnettii based on “the small ears, the shape of the face and the lack of black on the inside of the ears (which is a characteristic of Otolemur crassicaudatus).” He commented that colour variation is often found in Otolemur and describes this as an ”interesting variation”.

On the basis of (1) the considerable colour variation observed for O. g. lasiotis, (2) ear size, (3) communication with nocturnal primate specialists, and (4) its geographical origin, we conclude that the captive Otolemur male observed in Malindi is an O. g. lasiotis.

Nielsen and Sick (2008) report the presence of Otolemur crassicaudatus in Witu Forest Reserve. We encountered O. g. lasiotis in the area and strongly doubt the presence of O. crassicaudatus in the Reserve since this species is not known to occur in Kenya (Groves, 2001; T. M. Butynski & Y.A. de Jong, pers. comm.). Photos of Otolemur taken in the Reserve published in the report (Nielsen and Sick, 2008) resemble O. g. lasiotis.

Future research priorities:

• Is the Pangani River, Tanzania, the southern boundary of O. g. lasiotis? • To what extent does phenotypic variation in O. g. lasiotis occurs throughout its geographical range?

Primates of the coastal forests of Kenya. 42 Y.A. de Jong & T.M. Butynski, 2009.

4.2.2 Galago senegalensis braccatus Elliot, 1907

Vernacular name: Kenya lesser galago Subspecies type locality: Tsavo River, near Mt Kilimanjaro, Kenya Synonyms: None

Field characters: Medium-sized galago with bony, short-haired tail (Figure 12). Long, thick, and woolly pelage. Back of ears brown or greyish. Front of ears hairless, pinkish. Muzzle Figure 12. Galago senegalensis braccatus in Euphorbia pinkish or pale grey. Interocular stripe white. tirucalli hedge around human settlement, Isiolo, central Eye rings dark grey. Sides of the face and lower Kenya. cheeks off-white or pale grey. Crown, neck and shoulders grey or brownish-grey. Outer arms grizzled rufus-grey. Outer legs rufus with a sharp delineation between the grey upper thigh and dorsum. Ventrum, throat and chest, off-white with yellow wash. Tail grey to brown.

Distribution: Highlands of Kenya and Tanzania, east of Eastern (Albertine) Rift Valley. In NE and NC Tanzania, south to Pangani River (T. M. Butynski & Y. A. de Jong, pers. obs.; Nash & Zimmermann, in press). In Tsavo region (Groves, 2001) and coastal area of Kenya (Jenkins, 1987). North to South Horr, Kenya (I. Mwenja, pers. comm.), were it is sympatric with Galago gallarum. The northern limit is unknown (Figure 13).

Extent of occurrence: unknown

Photographic map: Photo map for Galagonidae, contains 28 photographs, of which seven are of G. s. braccatus (T. M. Butynski & Y. A. de Jong). http://picasaweb.google.com/wildsolutions/Galagonidae?feat=email# or via: www.wildsolutions.nl

Encounter sites: During this study, G. s. braccatus was not encountered in the coastal forests of Kenya between 2005 – 2008. Outside the hotspot it was encountered in the Chyulu Hills National Park; Kiboko Camp, Makindu; Komboyo Camp, Tsavo West National Park; north Kilimangodo; Mbuvu.

Encounter rate: G. s. braccatus was not encountered in the coastal forests of Kenya.

Vegetation: Outside the Hotspot, G. s. braccatus was encountered in dense to open Acacia or Acacia-Commiphora wood/bushland, thorn scrub, and savannah.

Altitude range: About 100 – 1900 m asl (Laikipia Plateau, central Kenya; Y. A. de Jong & T. M. Butynski, pers. obs.)

Vocalizations: The advertisement call of G. s. braccatus is a medium loud repetitive call comprised of single, low pitched units, repeated at regular intervals. The number of units in a call varies greatly and can last for up to 1 hr (also Harcourt, 1984; Nash et al., 1989; Bearder et al., 1995). The advertisement call is referred to as a ‘woo’ by Zimmerman (1990), a ‘bark’ by Harcourt (1984), but called a ‘honk’ in this study. Calls are given by males and females and made throughout the night with a peak right after dusk and right before dawn.

Primates of the coastal forests of Kenya. 43 Y.A. de Jong & T.M. Butynski, 2009.

Figure 13: Approximate geographic distribution of Galago senegalensis braccatus in the coastal area of Kenya with sites of encounters (2003-2008) depicted (Bearder et al., 2008; Y. de Jong & T. Butynski, unpubl. data). There is not enough data available to produce a more accurate map of the distribution of G. s. braccatus.

Conservation status: G. s. braccatus is most likely patchily distributed in the coastal forests of Kenya. There seem to be no serious threats to the survival of G. s. braccatus in the coastal area of Kenya, where this subspecies is present in low densities. Forest clearing for agriculture, degradation and fragmentation could result in an expansion of dryer bush/woodland and thorn bush which might support more G. s. braccatus in the coastal strip.

IUCN Red List: Least Concern (Bearder et al., 2008; www.iucnredlist.org/details/136863). Assessors: S. K. Bearder, T.M. Butynski, & Y.A. de Jong

Discussion: The limits of the geographical range of G. s. braccatus are not known. Jenkins (1987) described its range as from S Ethiopia to N and coastal Kenya. Groves (2001) only mentions the Tsavo Region as its range. Nash and Zimmermann (in press) mentions the highlands of Kenya and Tanzania east of the Eastern Rift Valley, south to Pangani River (Tsavo region) as the geographical range of G. s. braccatus. She notes that the northern limit is undetermined.

G. s. braccatus is an uncommon species in the Coastal Forest of Kenya. During this study we did not encounter G. s. braccatus in the area. The main reason for the lack of encounters in SE Kenya is most likely due to the few nocturnal surveys we conducted in savannah, thorn bush, and open woodland in SE Kenya. G. s. braccatus does inhabit forests in certain areas but only in forest were other small- to medium-sized galago species are absent (Bearder et al., 2008). Five kilometers across the Kenya – Tanzania border, in NE Tanzania, we encounter eight G. s. braccatus in dense Acacia shrub (Figure 13) within a distance of ca 6 km (rates of encounter: 10.7 ind/h, 1.3 ind/km; travelling by vehicle; T. M. Butynski & Y. A. de Jong, unpubl. data). We

Primates of the coastal forests of Kenya. 44 Y.A. de Jong & T.M. Butynski, 2009. suspect that G. s. braccatus is present in low densities in the coastal region of Kenya, on the edge with coastal forest and in the semi-arid strip farther inland.

G. senegalensis is a widespread species in Kenya. Its range is fragmented but it is locally common and occurs in high densities at many sites. G. s. braccatus was encountered at rates of: 0.1 ind/km (1.0 ind/h) by vehicle in the Ol Pejeta Conservancy, Laikipia Plateau (1,800-1,900 m asl), 1.1 ind/h by foot in Makindu (960 m asl), 1.2 ind/km (1.4 ind/h) by foot in Kajiado, S Kenya (1,500 m asl), and 1.1 ind/km (3.4 ind/h) by vehicle in Kora National Park (500 m asl). At Mpala Ranch, Laikipia Plateau, Off et al. (2008) estimated 0.4 and 2.4 animals/ha. The highest densities were in riverine vegetation, the lowest in dry bush areas. On the Laikipia Plateau, G. s. braccatus densities can locally be as high as 300 to 500 individuals/km² (Y. A. de Jong & T. M. Butynski, pers. comm.).

In Tanzania, the encounter rates with G. s. braccatus were 0.1 ind/km (2.4 ind/h) by vehicle in Tanga (10 m asl), 0.9 ind/h by foot in Meia Meia (1,330 m asl), 4.1 individuals/km (3.5 individuals/h) by foot near Mikumi National Park (470 m asl), and 4.0 individuals/h by foot at Mto wa Mbu, Lake Manyara National Park (970 m asl; T. Butynski & Y. de Jong, pers. comm.).

In Witu Forest Reserve, the Danish Zoological Society encountered either G. s. braccatus or Galago gallarum (Nielsen & Sick, 2008). During this study, we did not encounter any of the two lesser galagos when we visited Witu Forest Reserve but did find low densities of Galagoides cocos, a species not mentioned in their report. Given the fact that their surveys were conducted in a forest habitat, it is most likely that the species they encountered was either G. cocos (known to inhabit coastal forest) or G. s. braccatus (inhabiting mainly savannah, thorn bush, riverine vegetation, open woodland but occasionally found in forest were no other small- to medium-sized galagos are present; Bearder et al., 2008; Off et al., 2008; Nash, in press). G. gallarum inhabits dry Acacia-Commiphora wood/bushland and thorn bush (Butynski & De Jong, 2004). Dowsett- Lemaire and Dowsett (2007) did not encounter G. s. braccatus on Kipini Conservancy during their surveys in 2006.

G. s. braccatus was not encountered in the Tana River Primate National Reserve during this study. This subspecies does, however, occur in this Reserve (Kingdon, 1971; Andrews et al., 1975; Homewood, 1976; Marsh, 1978; Wolfheim, 1983; Decker, 1994; Butynski & Mwangi, 1994; Bearder et al., 2008; T. M. Butynski, pers. obs.). We think that G. s. braccatus is present only in very low densities within this Reserve.

Harcourt (1984) reports G. senegalensis in the coastal forest of Kenya but that this species is absent in the coastal forest of Diani were she studied Otolemur garnettii lasiotis and G. cocos (then referred to as Galago zanzibaricus) for 18 months.

Future research priorities; • What is the extent of occurrence or area of occupancy of G. s. braccatus? • Where in the coastal forests of Kenya does G. s. braccatus occur? • How will forest degradation and clearance affect G. s. braccatus?

Primates of the coastal forests of Kenya. 45 Y.A. de Jong & T.M. Butynski, 2009.

4.2.3 Galagoides cocos (Heller, 1912)

Vernacular name: Kenya coast galago Species type locality: Mazeras, Kenya. Synonyms: none

Field characters: Small, reddish-brown or buffy- brown galago with mouse grey under fur (Figure 14). Ears relatively large (compared to Galago senegalensis and Galago gallarum), dark brown/grey inside. Ear tips slightly bent backwards. Muzzle pointed and turned slightly upwards at tip. Patch on either side of muzzle dark brown and prominent. Chin pinkish-grey. Nose stripe off white, Figure 14. Galagoides cocos in Acacia – baobab extending to slightly above eyes. Eye rings grey. woodland at Takwa Ruins, Manda Island, Kenya. Throat, cheeks and ventrum off white to pale grey. Limbs, hands and feet buffy-brown. Dorsum buffy-brown. Proximal ca. 25% of tail same color as dorsum (i.e., buffy-brown) becoming gradually dark brown towards the tip of the tail (Butynski et al., 2006).

Distribution: G. cocos is restricted to the coastal forests of Kenya and N Tanzania. Occurs along the coastal strip of Kenya, as far north as the Tana River Primate National Reserve, Kipini Conservancy (on the coast just north of the Tana Delta) and Manda Island southward to at least the Mgambo Forest Reserve in N Tanzania at the north end of the East Usambara Mountains (between 1º 43' and 4 º 49' S; Butynski et al., 2006; Figure 15). Reported to occur in S Somalia along the Webi Shabeelle River (Nash et al., 1989) but this requires confirmation.

Extent of occurrence: ca. 13 900 km². Area of occupancy is very much less than this.

Photographic map: Photo map for Galagonidae, contains 28 photographs, of which three are of G. cocos (T. M. Butynski & Y. A. de Jong). http://picasaweb.google.com/wildsolutions/Galagonidae?feat=email# or via: www.wildsolutions.nl

Encounter sites: During this study, G. cocos was encountered in five out of the six areas we conducted nocturnal surveys in the coastal forests of Kenya; Kaya Rabai, Mazeras; Tana River Primate National Reserve; Kipini Conservancy; Manda Island; Diani.

Primates of the coastal forests of Kenya. 46 Y.A. de Jong & T.M. Butynski, 2009.

Figure 15: Known distribution and encounters (2003-2008) with Galagoides cocos in Kenya. The other source used for this map is Butynski et al. (2006).

Encounter rate: During this study, at least 20 G. cocos were encountered (excluding Kaya Rabai, Mazeras). All encounters were within the coastal forests of Kenya (Table 3; Figure 15).

Table 3. Rates of encounter with Galagoides cocos during 2005 – 2008 in the coastal forests of Kenya. Area Individuals/hour Individuals/km Vehicle/Foot Note Kaya Rabai, - - Foot No counts Mazeras conducted, many individuals heard and seen. Tana River PNR 10 (n=10) Foot About 15 individuals/ha Kipini Conser- 0 0 Vocalisations heard vancy only Manda Island 4.1 (n=6) 1.6 (n=6) Foot Diani 7.6 (n=4) 4.8 (n=4) Foot

Vegetation: G. cocos was encountered in evergreen and semi-evergreen riverine forest, dry coastal forest, fragmented, disturbed and degraded coastal vegetation, secondary forest, gardens and mosaics of mixed agriculture were some natural forest remains.

Altitude range: Sea level to at least 350 m asl in the foothills of the East Usambara Mountains, Tanzania (Butynski et al., 2006).

Primates of the coastal forests of Kenya. 47 Y.A. de Jong & T.M. Butynski, 2009.

Vocalizations: The advertisement call of G. cocos is an incremental call which is described as “A patterned phrase in which the units are arranged into distinct groups or ‘unit sequences’ which tend to get longer as the call progresses. Calls are given singly.” (Bearder et al., 1995). We describe the advertisement call of G. cocos in Butynski et al., (2006) as, “starts with a series of high-pitched, rapidly uttered, ’chirrups’ followed by units arranged in phrases that are high in frequency and amplitude, and that gradually become lower in amplitude. The number of units within each phrase increases incrementally until the end of the call. Often, as in this case, phrases with same number of units are repeated. The number of units per phrase rarely decreases. Units are often frequency modulated”. Butynski et al. (2006) presents the sonogram and oscillogram of G. cocos recorded in Diani, Kenya.

Conservation status: G. cocos is the most abundant and widespread galago in the coastal forests of Kenya, with densities of approximately 170 –180 individuals/km² both at Gedi and Diani (Harcourt & Nash, 1986). G. cocos is able to persist in fragmented, disturbed and degraded coastal vegetation, secondary forest and mosaics of mixed agriculture were some natural vegetation remains. There is a need to better determine the edges of the geographical range of G. cocos along the coast of N Kenya, S Somalia and in the vicinity of Nairobi (Butynski et al., 2006).

IUCN Red List: Least Concern (Butynski et al., 2008b; www.iucnredlist.org/details/136212). Assessors: T.M. Butynski, A. Perkin, S. Bearder, Y.A. de Jong, & P. Honess. G. cocos was only recently resurrected to a full species (Grubb et al., 2003; Butynski et al., 2006). Previously, ttaxonomists considered G. cocos as a subspecies or synonym of Galagoides zanzibaricus or Galago zanzibaricus (Butynski et al., 2006; Appendix 6).

Discussion: Data on G. cocos, partly obtained during this survey, were published in Butynski et al. (2006). The abstract of this article is given below and the full article is presented in Appendix 6). This publication includes a map and presents a thorough review of the geographical range of G. cocos. After publication of the map, this study found two more localities for the species; Kipini Conservancy and Manda Island. The Manda Island locality extended the range 70 km to the east of the distribution shown in Butynski et al. (2006; Figure 15).

Primate Conservation (2006) 21: 63 –79 Taxonomy, Distribution, and Conservation Status of Three Species of Dwarf Galagos (Galagoides) in Eastern Africa. Thomas M. Butynski, Yvonne A. de Jong, Andrew W. Perkin, Simon K. Bearder, and Paul E. Honess

Abstract: This paper reviews the complicated nomenclatural history for the Kenya coast galago, Galagoides cf. cocos, and examines whether ‘cocos’ is the valid species name for this recently resurrected taxon. This paper also reviews the phenotypic and vocal differences among G. cocos; the Zanzibar galago (Galagoides zanzibaricus zanzibaricus); the Udzungwa galago (Galagoides zanzibaricus udzungwensis); and the Mozambique galago (Galagoides granti), as well as their geographic ranges and conservation status. The following are among the findings: (1) ‘Galagoides cocos’ is the name that should be applied to the Kenya coast galago; (2) in the field, the loud calls of these three species are diagnostic and remain the best means for identification; (3) there is a suite of phenotypic characters that, when taken together, can be used to distinguish among these three species when in the hand or viewed in the field in good light at close range; (4) G. z. zanzibaricus is phenotypically distinct from G. z. udzungwensis; (5) the three species are parapatric or, perhaps, narrowly sympatric; (6) the three species are endemic to the coastal forests of eastern Africa with

Primates of the coastal forests of Kenya. 48 Y.A. de Jong & T.M. Butynski, 2009.

G. cocos in the north (Kenya and northeastern Tanzania), G. zanzibaricus in Tanzania, and G. granti from southern Tanzania to southern Mozambique; and (7) none of the three species is threatened at this time, although G. z. zanzibaricus meets the IUCN Red List criteria for an Endangered subspecies.

The complete article can be found in Appendix 6.

Future research priorities: • Does G. cocos occur in Ololua Forest, Nairobi? • How for north, north west does G. cocos along the Tana River?

Primates of the coastal forests of Kenya. 49 Y.A. de Jong & T.M. Butynski, 2009.

4.2.4 Cercopithecus mitis albogularis (Sykes, 1831)

Vernacular name: Zanzibar Sykes’s monkey Species type locality: Zanzibar Island Synonyms: kibonotensis Lönnberg, 1908; maritime Heller, 1913; kima, Heller 1913

Field characters: Medium-size monkey. Face dark grizzled grey with hairless face mask (Figure 16). Eyes reddish-brown. Ear Figure 16. Cercopithecus mitis albogularis adult male at fringe narrow, pale grey. Brow band grizzled Diani, Kenya. grey or grizzled grey-olive, directed forward and upward. Crown grizzled olive-grey with pale rufus wash. Cheeks grizzled pale grey, grizzled grey or grizzled olive-grey. Collar varying from pale grey to bright white, 55 - 80% complete. Collar line in Kenya populations well defined and extends from upper jaw, across the lower cheeks to ca 4-5 cm below the ears then down onto the sides of the neck, across the top of the shoulders to the nipples. Lower shoulders and upper outer arms dark grey. Lower outer arms black. Inner arms varying from grey to black. Outer legs dark grey, inner legs varying from pale grey (Kenya coast) to dark grey (Tanzania). Back of the upper legs varying from grizzled grey (Tanzania) to pale grey with rusty wash. Rusty wash most intense towards base of the tail in Kenya coast population. Dorsum grizzled grey between shoulders. Behind shoulders grizzled rusty red extending onto the upper tail. Ventrum varying from blackish or dark grey (Tanzania) to pale grey to off white (Kenya coast). Tail blackish or black. Base of the tail rusty all around.

Distribution: From Gedi Ruins (central coast of Kenya), southwards along the coast to N Tanzania (including Unguja and Mafia Islands), west to Mount Kilimanjaro, Mount Meru, Tsavo West National Park, Chyulu Hills National Park, Taita Hills, and Mount Kasigau. Along the coast of Tanzania as far south as NE Mozambique (Figure 17).

Extent of occurrence: ca. 209,000 km². The area of occupancy is but a very small portion of the extent of occurrence as the habitat suitable for C. m. albogularis is limited within its geographic range.

Photographic map: Photo map of Cercopithecus mitis, contains 46 photographs, of which 15 in the coastal forests of Kenya (T. M. Butynski & Y. A. de Jong). http://picasaweb.google.com/wildsolutions/CercopithecusMitis?feat=email# or via: www.wildsolutions.nl

Primates of the coastal forests of Kenya. 50 Y.A. de Jong & T.M. Butynski, 2009.

Figure 17. Approximate geographic distribution of Cercopithecus mitis albogularis in the coastal area of Kenya with sites of encounters (2003-2008) also depicted. Based on Y. de Jong & T. Butynski (unpubl. data) and Kingdon et al. (2008c).

Encounter sites: - Gedi Ruins (354, 355; Appendix 2) - Diani (346, 348, 350, 352; Appendix 2) - Gazi (185; Appendix 2) - Buda Forest (345; Appendix 2) - Mrima Hill (187, 190, 191, 194, 195, 197, 198, 248; Appendix 2) - Kaya Sega (343, Appendix 2) - Lunga Lunga (341, Appendix 2) - Vanga (342; Appendix 2) - Kaya Rabai (no number, Appendix 2) - Tsavo West National Park (330, 333, 338; Appendix 2)

Encounter rate: Twenty-two groups of C. m. albogularis were encountered in the coastal forests of Kenya. Four groups were encountered in Tsavo West National Park and Mazeras while travelling to the Hotspot (Table 4; Figure 17).

Primates of the coastal forests of Kenya. 51 Y.A. de Jong & T.M. Butynski, 2009.

Table 4. Encounter rates (groups/hour and groups/km) of Cercopithecus mitis albogularis during 2005 – 2008 in the coastal forests of Kenya and outside the Hotspot. This table only includes those surveys during which C. m. albogularis was encountered (n=number of groups). Area Groups/hour Groups/km Vehicle/Foot Note Gedi Ruins One group seen Foot twice at the ruins Mrima Hill 3.4 (n=10) 0.5 (n=10) Vehicle Using road around forest edge. Kaya Rabai, Mazeras 0 0 Foot Vocalisations heard Tsavo West NP 0.4 (n=3) 0.02 (n=3) Vehicle Road Surveys Kiboko to Lunga 0 0 Vehicle Highway Lunga (excluding Mrima Hill) Lunga Lunga to 0 0 Vehicle Main road and Mazeras (including bush tracks Shimba Hills NR, excluding Mrima Hill) Mtito Andei to 0 0 Vehicle Bush track Kilimangodo North Kilimangodo North 0.6 (n=3) 0.03 (n=3) Vehicle Using main roads to Kaya Sega (Vanga, Lunga Lunga, Kaya Gonja, Godo) Kaya Sega to Diani 0.5 (n=4) 0.02 (n=4) Vehicle Using tracks and (Vanga, Kaya main roads. Three Dzombo, Kaya encounters in Gonja, Buda Forest) Diani town.

Diani, Tiwi to Nyali 0.4 (n=1) 0.02 (n=1) Vehicle Using main roads Nyali to Watamu 0.3 (n=1) 0.01 (n=1) Vehicle Using main roads Watamu to Mamburi 0.2 (n=1) 0.01 (n=1) Vehicle Using main roads (including Malindi) Watamu to Tsavo 0 0 Vehicle East NP (including Malindi, Arabuko Sokoke Forest, Sala Gate Tsavo East NP)

Vegetation: C. m. albogularis is primarily a forest species and was encountered in coastal forest that varied from dry to moist coastal forest. This taxon is also encountered in miombo woodland, submontane, riverine, groundwater, secondary and plantation forests. Within populated coastal areas present on forest edged, in gardens, and in small patches of degraded coastal forest.

Altitude range: 0 – 2,900 m asl (Mount Kilimanjaro, Tanzania; T. M. Butynski & J. M. Butynski, pers. comm.).

Conservation status: C. m. albogularis is widespread and common in suitable habitat. Major threats to the survival of this taxa are habitat loss, degradation and fragmentation throughout those parts of its range that are outside protected areas.

Primates of the coastal forests of Kenya. 52 Y.A. de Jong & T.M. Butynski, 2009.

IUCN Red List: Cercopithecus mitis albogularis: Least Concern (Kingdon et al., 2008a; www.iucnredlist.org/details/40645) Assessors: Kingdon, J., Butynski, T.M. & De Jong, Y.A.

Discussion: The taxonomy of Cercopithecus mitis (or Cercopithecus albogularis) has been debated for many years and is still not agreed on. In this study we consider Cercopithecus mitis albogularis to be the subspecies that occurs in the coastal forests of Kenya (Booth, 1963; Grubb, et al., 2003, Grubb, 2006; Kingdon et al., 2008a, and Lawes et al., in press). We consider C. m. kibonotensis to be a synonym of C. m. albogularis (contra Hill, 1966; Dandelot, 1968, 1974; Rahm, 1970;, Napier, 1981; Lernould, 1988; Kingdon, 1997; Groves, 2001) who accept C. m. kibonotensis as the name of the mainland subspecies (distribution: southern Kenya coast south of Kilifi Creek, and northern Tanzania inland to Mt Kilimanjaro and Mt Meru) and C. m. albogularis to be the island (Unguja & Mafia) equivalent, with the main diagnostics characters being the larger body size of C. m. kibonotensis and the banding of the hairs (Groves, 2001, 2006). Grubb (2006) analysed the body sizes of both subspecies and did not find significant differences---but the sample size was small. Booth (1968, p.46) considers all subspecies of C. m. albogularis east of the rift valley as one subspecies because there are no distinguishing differences between the forms/subspecies; there is a cline going from the Kenya highlands to Zanzibar. For this reason, she claims that kolbi, kibonotensis and monoides are synonyms of C. m. albogularis. Booth (1963) makes no comment on the status of C. m. albotorquatus.

Table 5 present field descriptions obtained at five localities. The main phenotypic differences within C. m. albogularis is the colour of the ventrum and inner arms and legs which ranges from pale grey in the NE of the geographic range to blackish or dark grey in the Tanzania animals. The cheeks of the Kenya animals are more slender than for the Tanzania animals. The white collar of the Kenya animals is 60 - 80% complete, compared to 55-60% complete in the Tanzania population. Compared to the Tanzanian animals, the collar of the Kenya animals is more sharpely demarcated, runs through the cheeks, and closer to the ear. Unlike the Tanzanian animals, those in Kenya have red wash on the back of the upper hind legs.

Table 5. Field descriptions of C. m. albogularis Unguja Island, Tanzania, C. m. albogularis Usa River, Tanzania, C. m. albogularis Mrima Hill, Kenya, Cercopithecus mitis albogularis Diani, Kenya and C. m. albogularis Gedi Ruins, Kenya. Variable C. m. albogularis C. m. albogularis C. m. albogularis Cercopithecus C. m. Unguja Island, Usa River, Mrima Hill, mitis albogularis albogularis Gedi Tanzania Tanzania Kenya Diani, Kenya Ruins, Kenya Altitude 20 1195 80 16 20 (m asl) Photo

Ears Narrow, pale grey, Pale grey tufts Narrow, pale grey, Narrow, pale grey, Fringe absent or fringe fringe fringe (some narrow, pale whitish) grey. Face Grizzled pale grey. Grizzled pale grey. Grizzled grey. Bare Grizzled grey. Bare Grizzled grey.

Primates of the coastal forests of Kenya. 53 Y.A. de Jong & T.M. Butynski, 2009.

Bare black skin Bare black skin black skin around black skin around Bare black skin around eyes. around eyes eyes. eyes. around eyes. Cheeks Whiskers full, Whiskers full, Whiskers not full, Whiskers not full, Whiskers not grizzled pale grey. grizzled olive-grey. grey. Collar line grey. Collar line full, grey. Collar No collar line No collar line through the cheeks. through the cheeks. line through the crossing the lower crossing the lower cheeks. cheeks. cheeks. Brow band Projects forward Projects forward Projects forward Projects forward Projects forward and upwards. and upwards. and upwards. and upwards. and upwards. Grizzled pale grey. Grizzled olive-pale Grizzled grey. Grizzled grey. Grizzled grey. grey. Crown Grizzled olive-grey Grizzled olive-grey Grizzled olive-grey Grizzled olive-grey Grizzled olive- with rufus wash. with rufus wash. with pale rufus with pale rufus grey. wash. wash.

Collar Whitish-pale grey, Absent. White White, ca 75-80% Not sharply White, ca 75% ca 55-60% throat patch complete. Collar defined. Around complete. Collar complete. present. line sharp, running lower ca. 60% of line sharp, from upper jaw, neck. Throat off- running from across cheeks ca 4 white. Sides of upper jaw, across cm below the ears, neck light grey. cheeks to ca 4 onto neck, down cm below the the shoulders to ears onto neck, include the throat down shoulders, and upper chest. then grading into pale grey ventrum. Throat Whitish White White Off white White Shoulders Grizzled dark grey Black Dark grey Grizzled dark grey Dark grey Outer Black Black Black Black Black arms Inner arms Dark grey Black Dark grey Blackish Dark grey Outer legs Grizzled grey, feet Dark grey Dark grey Grizzled grey Dark grey blackish Inner legs Grizzled grey Dark grey Upper part white, Pale grey Pale grey lower part dark grey Back of Grizzled grey Grizzled grey Pale grey with Off white with Pale grey with hind legs rufus wash rufus wash rufus wash Dorsum Grizzled grey Grizzled olive-grey Grizzled grey Grizzled grey Grizzled grey between shoulders. between shoulders. between shoulders. between shoulders. between Distal half rich red Behind shoulders Behind shoulders Behind shoulders shoulders. extending onto rich red extending grizzled rusty red grizzled rusty red Behind shoulders upper tail. onto upper tail. extending onto extending onto grizzled rusty red upper tail. upper tail. extending onto upper tail. Ventrum Grizzled dark grey. Blackish Grey Grey. Center of the Pale grey ventrum pale grey. Sides Grizzled reddish- Grizzled reddish- Grizzled grey Grizzled grey Grizzled grey grey grey Tail Blackish except Blackish except Blackish except Black except bright Blackish, except pale red around rich red around bright rufus around rufus around base. bright rufus base. base. base. around base. Diagnostic Hairless face mask, Collar absent, Grey ventrum, Grey to pale grey Off white to pale

Primates of the coastal forests of Kenya. 54 Y.A. de Jong & T.M. Butynski, 2009. characters dark grey ventrum, blackish ventrum, white upper inner ventrum grey ventrum. rich red dorsum, rich red dorsum, legs. full cheek full cheek whiskers. whiskers.

Heller (1913) provides a description of a specimen of C. m. albogularis from Mazeras (then referred to as Lasiopyga albogularis maritima) which notes the lighter coloration overall and the absence of a black lining or black tips to the hairs of the ventrum. Although we did not observe C. m. albogularis well enough in Mazeras to make a complete field description, the C. m. albogularis populations we found on the Kenya coast certainly have a pale grey or off-white ventrum.

Groves (2001) notes in his description of C. a. albogularis and C. a. kibonotensis (which he accepts as a full subspecies of C. albogularis), that C. a. kibonotensis is very similar to C. a. albogularis (which he considers to be endemic to Unguja Island) except that C. a. albogularis has an almost complete collar, and the collar of C. a. kibonotensis is less extensive. Napier (1981) also describes C. m. kibonotensis with a less extensive collar compared to C. m. albogularis. However, during this study (which is based on field descriptions and photographs as opposed to the museum descriptions of Napier (1981) and Groves (2001), we found the opposite; the animals on Unguja Island have a whitish (males) or pale grey (juveniles and females) collar that is ca 55-60% complete; Figure 18) where as the animals on the Kenya coast have a collar with is ca 60-75% complete (white in males and off white or pale grey in juveniles and some females). In addition, the Kenya coast animals have a collar which forms a well demarcated line from the upper jaw, across the cheeks, under the ears onto the neck, and crosses the top of the shoulders down onto the chest (Figure 19).

Figure 18. Adult male C. m. albogularis at Jozani-Chwaka Bay National Park, Unguja Island, Tanzania. Note the grizzled grey cheeks and the whitish to pale grey collar which is ca 60% complete.

Figure 19. Adult male C. m. albogularis at Gedi Ruins, Kenya. Note that the white of the collar crosses the cheek and that the collar is >70% complete.

Primates of the coastal forests of Kenya. 55 Y.A. de Jong & T.M. Butynski, 2009.

In conclusion, we observed considerable phenotypic differences among the animals found in the type locality for C. m albogularis (Unguja Island, Tanzania), the western-most locality for ‘C. m. albogularis’ that we visited (Usa River, close to the type locality of kibonotensis), the northeastern-most locality that we visited (Gedi Ruins), and those in between these localities (e.g., Mrima Hill, Vanga, Diani, Lunga Lunga). These phenotypic differences appear to be in a cline. The phenotypic differences between the extremes of the geographic range of C. mitis from Gedi south to Unguja Island differ in degrees and seem to be part of the natural occurring variability within C. m. albogularis.

Future research priorities: • Which subspecies of C. mitis occurs between the Tana River and the Galana River?

Primates of the coastal forests of Kenya. 56 Y.A. de Jong & T.M. Butynski, 2009.

4.2.5 Cercopithecus mitis albotorquatus Pousargues, 1896

Vernacular name: Pousargues’s white-collared monkey Subspecies type locality: not known (zoo animal) Synonyms: phylax Schwarz, 1927, Patta Island, Lamu Archipelago rufotinctus Pocock, 1907, “British East Africa (? Mombasa).”

Field characters: Medium-size monkey (Figure 20). Appears to be slender as compared to the other subspecies of C. mitis. Face dark grizzled grey, slender, with short hair. Eyes reddish-brown. Ears visible with narrow grey fringe. Browband grizzled greyish-olive, directed far forward. Upper jaw dark grey. Collar bright white and almost (ca. 85%) complete Collar line very well defined and extends Figure 20. Cercopithecus mitis albotorquatus male from upper jaw, across the cheeks to ca 2 cm below the from Witu Forest, Kenya, where this subspecies is ears then down onto the sides of the neck, across the top common. of the shoulders to the nipples. Crown grizzled greyish-olive with rufus teint. Lower shoulders and upper outer arms grizzled dark grey. Lower outer arms dark grey to blackish. Outer legs grey. Ventrum and inner limbs pale grey to off-white. Back of upper legs with long rusty hairs, most intense red towards the base of the tail. Dorsum grizzled grey between the shoulders becoming increasingly grizzled olive-russet towards the tail. Tail blackish. Base of the tail rusty all around.

Distribution: Restricted to the forest along the north coast of Kenya and inland along the lower Tana River into Meru National Park. Perhaps along the coast in extreme Southern Somali but this needs confirmation (de Beaux, 1937; Hill, 1966; Gartland & Brain, 1968; Rahm, 1970; Gippoliti, 2002). Present on the Lamu Archipelago and patchily distributed throughout the Tana River forests as northwest as Meru National Park (Figure 21; De Jong & Butynski, 2004; Butynski & De Jong, 2008). Not known to occur south of the Tana River or Tana Delta where it appears to be replaced by C. m. albogularis. Along the Juba River and Shebelli River of S. Somalia it is replaced by C. m. zammaranoi.

Extent of occurrence: ca. 11 500 km². Area of occupancy is very much less than this due to the severely fragmented geographic range and limited extent of forest in this semi-arid region.

Primates of the coastal forests of Kenya. 57 Y.A. de Jong & T.M. Butynski, 2009.

Figure 21. Known distribution and encounters (2003-2008) with Cercopithecus mitis albotorquatus. The other source used for this map is Butynski & De Jong (2008).

Photographic map: Photo map for Cercopithecus mitis, contains 46 photographs, of which five are of C. m. albotorquatus (T. M. Butynski & Y. A. de Jong). http://picasaweb.google.com/wildsolutions/CercopithecusMitis?feat=email# or via: www.wildsolutions.nl

Encounter sites: During this study, C. m. albotorquatus was encountered in three out of the four areas we conducted diurnal surveys in the coastal forests of Kenya;

- Tana River Primate National Reserve (17, 17A, 18A, 254, 254A, 255A; Appendix 2) - Witu Forest Reserve (269, 271, 271A; Appendix 2) - Kipini Conservancy (277A, 273, 278A to 278E, Appendix 2) - Takwa Ruins, Manda Island (367, Appendix 2) - Lamu Island (heard but not seen)

Encounter rate: During this study, 16 groups of C. m. albotorquatus were encountered all in the coastal forests of Kenya (north coast; Table 6; Figure 17).

Primates of the coastal forests of Kenya. 58 Y.A. de Jong & T.M. Butynski, 2009.

Table 6. Encounter rates (groups/h and groups/km) of Cercopithecus mitis albotorquatus during 2005 – 2008 in the Coastal Forests Kenya and while driving to the Hotspot. Only includes the surveys in which C. m. albotorquatus was encountered. Area Groups/hour Groups/km Vehicle/Foot Note Tana River PNR 0.30, n=6 Foot Witu Forest 7.14, n=3 0.6 n=3 Vehicle Using existing roads Reserve Kipini 1.60, n=5 1.2, n=5 Foot Vehicle surveys not Conservancy included Manda Island 0.16, n=1 0.1, n=1 Foot Calls heard from at least one other group Lamu Island 0 0 Foot Calls heard from at least one group.

Vegetation: Evergreen and semi-evergreen riverine forests of the Tana River, surrounded by semi- arid medium-dense bush land on sandy soil (Tana River floodplain). Forest-grassland mosaic, coastal forest and dry coastal shrub. Mangrove forest.

Altitude range: 25 – 400 m asl (De Jong & Butynski, 2004)

Conservation status: C. m. albotorquatus is a poorly known subspecies. Threats to their conservation are habitat loss, degradation and fragmentation due to forest clearance and agriculture. During this survey, C. m. albotorquatus were found to be common in the Witu Forest Reserve, Kipini Conservancy, and Tana River Primate National Reserve, but rare on Manda Island and on Lamu Island. Said to be present on Patta Island but we did not visit Patta Island. During primate surveys in Meru National Park and Kora National Park in 2003, C. m. albotorquatus were found to be common in the riverine forests of the Tana River (De Jong & Butynski, 2004). In short, during our study we found C. m. albotorquatus to be much more widespread than previously known and also that there are some sites where this subspecies is at high densities. These significant findings bode well for the long-term conservation and survival of C. m. albotorquatus.

IUCN Red List: Cercopithecus mitis albotorquatus: Vulnerable B1ab (i,ii,iii,v; Butynski & De Jong, 2008; www.iucnredlist.org/details/39990). Assessors: T. M. Butynski, & Y.A. de Jong Previous assessment: Data Deficient (DD; ver 2.3, 1994), year assessed, 2000 (Butynski & De Jong, 2008)

Discussion: C. m. albotorquatus is a relatively poorly-known but distinctive subspecies that is , recognised as a subspecies by most primate taxonomist (Pocock, 1907; Hill, 1966; Rahm, 1970; Kingdon, 1971; Dandelot, 1974; Napier, 1981; Lernould, 1988; Kingdon, 1997; Groves, 2001, Grubb, 2001, Grubb et al., 2003). Several authors suggested that this subspecies was confined to the forests along the lower Tana River forests (Gartlan & Brain, 1968; Rahm, 1970; Kingdon, 1971; Andrews et al., 1975; Napier, 1981; Lernould, 1988; Groves, 2001). During a primate survey in 2003 by T. Butynski & Y. de Jong, this subspecies was found to be present and common as far inland as Meru National Park and Kora National Park. This is much farther inland (to the northwest) of the previous known range (De Jong & Butynski, 2004). This finding extended the known range ca 100 km to the west (Saka being the closest record; Napier, 1981).

Primates of the coastal forests of Kenya. 59 Y.A. de Jong & T.M. Butynski, 2009.

The southern limit of the geographical range of C. m. albotorquatus is uncertain. Figure 21 shows a gap of 80 km between the ranges of C. m. albotorquatus and C. m. albogularis, based on the lack of reports from this area. The closest record for C. m. albotorquatus (Witu Forest Reserve) to C. m albogularis is for the Gedi Ruins (108 km SW of Witu Forest Reserve). We find, however, that the ‘C. m. albogularis’ at Gedi Ruins are phenotypically different from C. m. albotorquatus and from the description of the holotype for C. m. albogularis (which was collected on Zanzibar). Table 7 presents the field descriptions we obtained of C. m. albogularis from Gedi Ruins, Kenya, C. m. albotorquatus from Witu Forest Reserve, Kenya and C. m. albogularis from Unguja Island, Tanzania (type locality of C. m. albogularis).

Table 7. Field descriptions of C. m. albogularis, Gedi Ruins, Kenya, C. m. albotorquatus, Witu Forest Reserve, Kenya and C. m. albogularis, Unguja Island, Tanzania. Description C. m. albogularis, Unguja C. m. albogularis, Gedi Ruins C. m. albotorquatus, Witu Island, Tanzania Kenya Forest Reserve, Kenya Altitude 20 m asl 20 m asl Foto

Ears Narrow pale grey fringe Absent or narrow pale Narrow fringe, grey. grey fringe. Face Grizzled pale grey. Bare black Grizzled grey. Bare black skin Triangular shape with skin around eyes. around eyes. short hair. Grizzled grey. Bare black skin around eyes. Cheeks Full, grizzled pale grey. Grizzled grey, slender Grizzled grey, slender appearance. Collar line appearance. Collar line crossing the lower cheeks crossing the lower cheeks Brow band Projecting forward and upwards. Projecting forward and Projects forward and Grizzled pale grey upwards. Grizzled grey. upwards. Grizzled greyish- olive Crown Grizzled olive-grey with rufus Grizzled olive-grey. Grizzled olive-grey with teint. rufus teint.. Collar Whitish-pale grey, ca 55-60% Collar white, ca 75% complete Bright white, almost (ca. complete. Collar line 85%) complete. Collar sharp, running from upper jaw, line very well defined, across the cheeks to extends from upper jaw, ca 4 cm below the ears across the cheeks to ca 2 cm into the neck, down crossing below the ears then down the top of the shoulders grading onto the sides of the neck, into the pale ventrum. across the top of the shoulders to the nipples. Throat Whitish White Bright white Shoulders Grizzled dark grey Dark grey Partly white from collar, remainder grizzled grey

Primates of the coastal forests of Kenya. 60 Y.A. de Jong & T.M. Butynski, 2009.

Outer arms Black Black Outer arms dark grey to blackish. Inner arms Dark grey Dark grey Pale grey Outer legs Grizzled grey, feet Dark grey. Grey. blackish Inner legs Grizzled grey Pale grey Pale grey to off white. Back of hind Grizzled grey. Pale grey with rufus wash Long, off white, hairs with limbs distinguishing rusty wash. Rust darkest around base of tail. Dorsum Grizzled grey between Grizzled grey between Grizzled grey between shoulders. Half way the back shoulders. Below shoulders. Behind shoulders onto the upper tail rich red. shoulders grizzled rusty grizzled olive-grey-rusty red extending onto the upper becoming increasingly rusty tail. towards the tail. Ventrum Grizzled dark grey. Pale grey Pale grey to off white. Sides Grizzled reddish-grey Grizzled grey Grizzled grey Tail Pale red around base of Bright rufus around the base of Intense red around base of tail, rest of tail blackish. the tail, rest of blackish tail (especially ventrally). Terminal part blackish. Overall The hairless face mask The off white-pale grey The short hair, triangular noticable, dark grey ventrum, ventrum is distinguishable for face and the bright and wide rich red dorsum and full cheeks this population collar that is almost complete are distinguishable for this is distinguish-able for this population subspecies.

C. mitis in the Lamu Archipelago The type locality of C. m. phylax is Patta Island. The distribution of C. m. phylax is the Lamu Archipelago (“Patta and Witu Islands.” (=Lamu Archipelago; Groves, 2001), however, Hill (1966) notes that this taxa is only known from the type locality (Patta Island). We found no published locality records of C. mitis for Manda and Lamu Islands to clarify this inconsistency.

During a survey on Lamu Island, the density of primates was found to be very low. The observed primate groups were small and the animals were shy. We collected evidence that C. mitis is present in very low densities in the extensive mangrove forests on N Lamu Island (ca 150-200 m away from the mainland). Calls were heard from at least one group in the mangrove forest but we failed see these monkeys during our 2 day visit (mainly due to bad weather). The local people on this part of the Island are familiar with C. mitis and, we were told, they occasionally hunt and keep them as pets.

During a survey on Manda Island, the density of C. mitis was found to be very low. As on Lamu, this is likely due to the lack of year round fresh water. On one occasion we heard C. mitis vocalizations (including ‘pyows’) from the mangrove forest during a boat survey on NW Manda Island. On SE Manda Island we encountered one group of C. mitis (and heard ‘pyows’). One individual was seen during our ground survey. The animal was in dense and diverse coastal shrub (including Acacia, Commiphora, and baobab) on coral rag on the edge of a mangrove forest. People here report that C. mitis to occur mainly in the mangrove forests.

If Hill (1966) is correct in that phylax is restricted to Patta Island than this survey is the first to confirm that C. mitis is present both on Lamu Island and Manda Island.

Primates of the coastal forests of Kenya. 61 Y.A. de Jong & T.M. Butynski, 2009.

Hill (1966), Rahm (1970), Kingdon (1971), Napier (1981), Lernould (1988), Kingdon (1997) and Groves (2001) consider phylax to be valid subspecies. Dandelot (1968) considers C. m. phylax as a synonym of C. m. albotorquatus on the basis of the ‘similarity of characters and the proximity of the Island to Somalia’. Grubb (2001) agrees with this on the basis of skull measurements and the examination of pelage coloration. Like Dandelot (1968 in Groves, 2001, Grubb et al. (2003), Grubb (2006), and Lawes et al. (in press), we suspect that phylax is a synonym of C. m. albotorquatus. We suspect this for two reasons:

1) The channel that separates the Lamu Archipelago Islands from the mainland is shallow and narrow (Patta Island ca 3 km; Manda Island ca 20 m; Lamu Island ca 150-200 m). Therefore, these Islands are not particularly isolated. The extensive mangrove around all islands might be more of a barrier to primate movement than is the water. If C. mitis on the Lamu Archipelago is an endemic subspecies, then any differences from the mainland forms may be due more to the need to live in a very 'fresh water restricted' habitat and in mangrove forest, than to any isolation related to distance from the mainland. C. m. albotorquatus is known to occur on the mainland ca 45 km away (Witu Forest Reserve) from Lamu Island and ca 55 km away from Manda Island. It seems likely that in the past this taxon experienced gene flow with C. m. albotorquatus on the mainland on more than one occasion.

2) C. m. phylax is described as “having a dark olive head, upper parts dark reddish brown, russet tuft under base of tail, black arms, mouse grey legs, under parts white, and a white part-collar round neck” (Grubb, 2001). According to Groves (2001) phylax is “small in size”. Grubb (2001) compared the skull measurements of the female holotype to C. m. albotorquatus (female from Tana River) and found that the phylax holotype is slightly larger than the mainland specimen. He also notes that the pelage falls within the range of the Tana River specimens. Grubb (2001) states that “Some of these measurements may have been taken differently, but they are quite insufficient to suggest that insular and mainland populations are subspecifically different.” The animal we encountered on Manda Island looked similar to C. m. albotorquatus we encountered on the mainland. Unfortunately, we only (briefly) saw one C. mitis on Manda Island and we failed to obtain a photo. The group we encountered was shy and only the crown, face and under parts were seen. The crown of the individual we encountered on appeared no darker than the crown of C. m. albotorquatus. During this study we did not visit Patta Island. Future surveys should include a visit to Patta Island to further describe the C. mitis there and to collect preliminary ecological data on this taxon.

Future research priorities: • What is the southern limit of the geographical range of C. m. albotorquatus? • What is the geographical range of C. m. phylax? • How is C. m. phylax phenotypically different from C. m. albotorquatus (in Patta and Manda Islands) and is it a valid subspecies? • Which subspecies of C. mitis occurs between the Tana River (west Meru National Park, Ngaia Forest, Meru Forest and Mwea National Park) and the confirmed range of Cercopithecus mitis kolbi (Mt Kenya Forests).

Primates of the coastal forests of Kenya. 62 Y.A. de Jong & T.M. Butynski, 2009.

4.2.6 Chlorocebus pygerythrus hilgerti (Neumann, 1902)

Vernacular name: Eastern vervet Type locality: Gobeli River, Webbi Shibeli, Harar, Ethiopia Synonyms: ellenbecki (Neumann, 1902), Lake Zwai, Suksuki and Maki Rivers, Ethiopia. johnstoni (Pocock, 1907), Moshi, S Mt Kilimanjaro, Tanzania. 1500 m asl callida (Heller, 1912), S Lake Naivasha, Kenya arenaria (Heller, 1913), Merille water holes, Marsabit Road, N Kenya Figure 22. Chlorocebus pygerythrus hilgerti male contigua (Heller, 1920), Changamwe, 10 W at Diani Beach, Kenya. Mombasa, Kenya luteus (Elliot, 1910), Wambugu, SW Mt Kenya, Kenya rubellus (Elliot, 1909), Fort Hall, Kenya. tumbili (Heller, 1913), Ndi, Taita District, S Kenya.

Field characters: Medium-size, slender monkey (Figure 22). Face and ears blackish to jet black. Brow band, cheeks and throat off-white to white, forming a complete ring around the black face. Cheek whiskers medium to long sweeping upwards and backwards, covering part of the ears. Crown and dorsum grizzled olive-grey, intense olive-grey, or pale sandy brown. Ventrum and inner limbs off-white to white. Sides in some populations show a ‘white stripe’ between the olive-grey dorsum and the white ventrum. Upper outer arms and legs grizzled olive-grey. Lower outer arms and legs grizzled grey. Hands and feet blackish to jet black in males and grey to blackish in females and juveniles. Top of tail grizzled mouse grey, distal ca. 10% darker grey to jet black. Red patch under base of the tail of males in some populations. Underside of tail pale grey.

Distribution: Throughout Kenya (excluding the Lamu Archipelago). Into E Uganda, S Ethiopia, S Somalia, and N Tanzania (Figure 23).

Extent of occurrence: Not known.

Photographic map: Photo map of Chlorocebus pygerythrus, contains 86 photographs, of which 20 in the coastal forests of Kenya (T.M. Butynski & Y.A. de Jong). http://picasaweb.google.com/wildsolutions/ChlorocebusPygerythrus?feat=email or via: www.wildsolutions.nl

Primates of the coastal forests of Kenya. 63 Y.A. de Jong & T.M. Butynski, 2009.

Figure 23. Approximate geographic distribution of Chlorocebus pygerythrus hilgerti in the coastal area of Kenya with sites of encounters (2003-2008) depicted. Based on Y. de Jong & T. Butynski (unpubl. data) and Kingdon et al. (2008c). There is not enough data available to produce a more accurate distribution map of C. p. hilgerti.

Encounter sites: - Wenje (15, 251A; Appendix 2) - Tana River Primate National Reserve north (256; Appendix 2) - Tana River Primate National Reserve north (252A; Appendix 2) - Kipini Conservancy (259; Appendix 2) - Mpeketoni (265; Appendix 2) - Tiwi (353; Appendix 2) - Nyali (353A; Appendix 2) - Malindi (356; Appendix 2) - Malindi West (358; Appendix 2) - Diani (373; Appendix 2) - Lamu Island (none seen but evidence collected for their presence (Appendix 2) - Manda Island (none seen but evidence collected for their presence (Appendix 2)

Encounters while travelling to the Hotspot: - Tsavo West National Park (326, 328, 331, 334, 337; Appendix 2) - Tsavo East National Park (362; Appendix 2) - Voi south (249; Appendix 2) - Kora National Park (20, 21, 22, 23, 24, 25, 27A, 304; Appendix 2) - Meru National Park (27, 28, 307, 308A; Appendix 2) - Mwingi (12; Appendix 2) - Mwea National Reserve (1, 3, 5; Appendix 2)

Primates of the coastal forests of Kenya. 64 Y.A. de Jong & T.M. Butynski, 2009.

Encounter rate: A total of 44 groups of C. p. hilgerti were encountered of which 11 groups were in the coastal forests of Kenya. Thirty-five groups were encountered while travelling to and from the Hotspot (Table 8; Figure 23).

Table 8. Encounter rates (groups/hour and groups/km) of Chlorocebus pygerythrus hilgerti during 2005 – 2008 in the coastal forests of Kenya and travelling to the coastal area. Only included the surveys C. p. hilgerti was encountered. Area Groups/hour Groups/km Vehicle/Foot Note Nyali, Haller’s 1 (n=1) 0.78 (n=1) Foot Park. Kipini 0.1 (n=2) 0.02 (n=2) Vehicle Conservancy /Mpeketoni/ Witu Forest Reserve Tsavo West NP 0.7 (n=5) 0.03 (n=5) Vehicle 2 days survey Tsavo East NP 0.3 (n=1) 0.01 (n=1) Vehicle Kora NP 0.1 (n=1) 0.01 (n=1) Vehicle 2 days survey North Kitui 1. 7 (n=5) 0.08 (n=5) Vehicle NR/Kora NP Meru NP (2005) 0.8 (n=2) 0.03 (n=2) Vehicle One additional group seen from Meru NP on Kora NP side of Tana River Meru NP (2007) 0.4 (n=2) 0.02 (n=2) Vehicle Mwea NR 0.7 (n=3) Vehicle

Road Surveys Diani – Nyali, 0.4 (n=1) 0.02 (n=1) Vehicle, 25.2 km/h Travelling slowly south coast on highway Tana River PNR – 0.2 (n=1) 0.01 (n=1) Vehicle, 27.5 km/h Kipini Conservancy Garissa – Tana 0.4 (n=2) 0.01 (n=2) Vehicle, 38.3 km/h River PNR Watamu – N. 0.2 (n=1) 0.01 (n=1) Vehicle, 23.5 km/h Malindi – W. Malindi - Watamu Tanga (Tz) – Kaya 0.4 (n=3) 0.01 (n=3) Vehicle Through Mrima Rabai, Mazeras Hill and Shimba Hills NR Kaya Rabai, 0.1 (n=1) 0.002 (n=1) Vehicle Highway Mazeras - Nairobi Mwea NR – 0.2 (n=1) 0.01 (n=1) Vehicle Mwingi East Mwingi East – 0.2 (n=2) 0.10 (n=2) Vehicle Tana River PNR

Vegetation: Savannah, open to medium dense Acacia and Acacia-Commiphora woodland, close to permanent water sources containing tall Acacia trees (A. xanthophloea and A. tortilis). Found in miombo woodland, riverine vegetation and forest-grassland mosaic. Able to persist in degraded and fragmented vegetation. Throughout its range is able to live close to human

Primates of the coastal forests of Kenya. 65 Y.A. de Jong & T.M. Butynski, 2009. settlements and hotel compounds, attracted by water, agricultural crops, human food, and security from predators. Requires a year round source of water and tall trees in which to sleep and escape from predators.

Altitude range: 0 – 2000 m asl (Naivasha, Kenya; Y. A. de Jong & T. M. Butynski, unpubl data)

Conservation status: C. p. hilgerti is a very patchily distributed subspecies that is, however, often common in the open and medium dense Acacia woodlands of the coastal strip of Kenya. There are no threats to their conservation. Hunted in many places because of the damage it causes to crops. C. p. hilgerti can be especially common and abundant in the vicinity of tourist lodges and hotels on the coastal strip in Kenya.

IUCN Red List: Chlorocebus pygerythrus: Least Concern (Kingdon et al., 2008b; www.iucnredlist.org/details/136271). Assessors: Kingdon, J., Gippoliti, S., Butynski, T.M. & De Jong, Y.A. C. p. hilgerti is recognised but not assessed by the 2008 IUCN Red List of Threatened Species.

Discussion: Chlorocebus pygerythrus is a patchily distributed but locally common species in the coastal forests of Kenya. The taxonomy of this taxon has been debated for many years. This study recognised C. p. hilgerti as the subspecies present in the coastal strip of Kenya. This is consistent with Groves (2001, 2005, 2006), and Kingdon et al., 2008b but at odds with the taxonomy of Hill (1966), Kingdon (1971, 1997), Napier (1981), and Grubb et al. (2003). We fully agree with Grubb et al. (2003) and Groves (2006) that more study and a full revision of the genus is required.

During this study we found surprisingly little phenotypic variation between C. p. hilgerti populations in the extremes of their distribution in coastal forests of Kenya. We did find variation in the intensity of their colours in the region (and throughout Kenya and Tanzania; T. M. Butynski & Y. A. de Jong, unpubl data). Table 9 presents field description of C. p. hilgerti obtained from Mpeketoni (NE Kenya coast; number 265, Appendix 2), Diani (S Kenya coast; number 373, Appendix 2) and, as a comparison, Shaba National Reserve (central Kenya, outside the coastal area; number 370, Appendix 2).

Table 9. Field descriptions of Chlorocebus pygerythrus hilgerti obtained from the extremes of the coastal forests of Kenya (i.e., Diani and Mpeketoni) and from Shaba National Reserve in central Kenya. Body Part Diani Mpeketoni Shaba NR Photo

Face Jet black, hairless Blackish, pale skin around Black

Primates of the coastal forests of Kenya. 66 Y.A. de Jong & T.M. Butynski, 2009.

eyes Ears Black Blackish Blackish Whiskers White, medium length, White, long, sweep upwards White, medium long, sweep upwards and and backwards covering the sweep upwards and backwards covering part ears backwards covering part of the ear. of the ears. Brow band White, forming a White, forming a complete White, forming a complete white ring white ring around face with complete white ring around face with cheeks cheeks and throat around face with cheeks and throat and throat Crown Grizzled olive-grey Grizzled olive-grey Grizzled olive-grey Throat Off white White White Dorsum Grizzled olive-grey. Grizzled olive-grey. Grizzled olive-grey. Ventrum Off white White White Sides Grizzled olive-grey. Intense olive in contrast with Intense olive in contrast Narrow off-white lateral white ventrum. Narrow white with white ventrum. stripe. lateral stripe. Narrow white lateral stripe. Hands and feet Black in males, grey to Blackish in males, pale grey Blackish in males, pale dark grey in juveniles to dark grey in juveniles and grey to dark grey in and females. females. juveniles and females. Outer arms Upper arms grizzled Upper arms grizzled olive- Upper arms grizzled olive-grey, lower arms grey, lower arms grizzled olive-grey, lower arms grizzled grey grey grizzled grey Inner arms Off-white White White Outer legs Grizzled mouse grey Upper legs grizzled olive- Upper legs grizzled grey. Lower legs grizzled olive- grey. Lower legs grey. grizzled grey. Inner legs Off-white White White Tail Grizzled mouse grey, Grizzled grey, terminal end Grizzled grey, tail tip distal ca. 10% darker black. Red patch under base black. Under tail slightly grey. Red patch under of the tail in males. Under tail lighter grey. base of the tail in males. slightly lighter grey. Under tail slightly lighter grey. Overall Phenotypically like most Relatively strong contrast Phenotypically like most other populations of C. p. between dorsum and ventrum other populations of C. hilgerti. due to more intense colours. p. hilgerti. Long whiskers stand out.

The main phenotypic differences we found in the extremes of the geographical range of C. p. hilgerti in Kenya are the ‘white lateral stripes’ in some populations, the length of the whiskers, the intensity of fur colour (including the black and white body parts).

Future research priorities:

• What is the extent of occurrence or area of occupancy of C. p. hilgerti?

Primates of the coastal forests of Kenya. 67 Y.A. de Jong & T.M. Butynski, 2009.

4.2.7 Chlorocebus pygerythrus excubitor (Schwarz, 1926)

Vernacular name: Manda vervet monkey Subspecies type locality: Manda Island, Witu Islands (=Lamu Archipelago), off coast of N Kenya. Synonyms: Chlorocebus voeltzkowi, Matschie, 1923 (nomen nudum).

Field characters: No data

Distribution: Lamu Archipelago, including Lamu Island, Manda Island, Patta Island, and probably Manda Toto Island (Figure 24).

Extent of occurrence: < 200 km², based on the size of Lamu, Manda, and Patta Islands, excluding the extensive mangrove forests around the Islands which is unsuitable habitat for C. pygerythrus. The area of occupancy is highly fragmented and expected to be much smaller than this.

Figure 24. Suspected geographic distribution and evidence for Chlorocebus pygerythrus excubitor. The actual area of occupancy is probably very much smaller than shown.

Photographic map: We have no photos of C. p. excubitor.

Encounter sites: C. p. excubitor was not observed during this study but evidence was collected that this taxon is present in the following areas: - Matondoni and the surrounding area, NE Lamu Island - Lamu Town and vicinity - Manda Airstrip and vicinity, E Manda Island

Primates of the coastal forests of Kenya. 68 Y.A. de Jong & T.M. Butynski, 2009.

- SE Manda Island - Takwa Ruins, SE Manda Island - Manda Toto Island occasionally (reported by local fisherman, n=3). Said to sometimes cross the narrow channel between Manda Island and Manda Toto Island during low tide.

During this study we were not in the position to survey Patta Island.

Encounter rate: C. p. excubitor was not encountered during this study. On all Islands C. p. excubitor occurs at very low densities

Vegetation: No data. Evidence was collected, however, that C. p. excubitor occupies savannah wood/bushland and is attracted to farms and human settlements.

Altitude range: sea level to ca 10 m asl.

Conservation status: C. p. excubitor occurs in very low numbers in the Lamu Archipelago. One of the causes for this is likely the scarcity of year round fresh water sources on the Islands. Residents stated that C. p. excubitor is hunted and kept or sold as pets. It is unknown if the pet trade is a serious threat to this subspecies but this seems unlikely. The geographical range of C. p. excubitor is highly fragmented (between the Islands) and its total numbers must be very low.

IUCN Red List: Chlorocebus pygerythrus: Least Concern (Kingdon et al., 2008b; www.iucnredlist.org/details/136271). Assessors: Kingdon, J., Gippoliti, S., Butynski, T.M. & De Jong, Y.A. C. p. excubitor is recognised but not assessed by the 2008 IUCN Red List of Threatened Species.

The maximum possible area of occupancy for C. p. excubitor is < 200 km². Given this small and fragmented distribution, C. p. excubitor qualifies as Endangered (EN, B1) on the basis of the present criteria (IUCN, 2008).

Discussion: Chlorocebus pygerythrus excubitor (Schwartz, 1926) is accepted as a valid subspecies by Hill (1966), Kingdon (1971, 1997), Napier (1981), Groves (2001, 2005), Grubb et al. (2003), and Kingdon et al. (2008b). The holotype was collected on Manda Island. Hill (1966) describes this subspecies as “A small insular race of dull reddish pelage, very similar to Chlorocebus pygerythrus nesiotis but redder and, in the darker phase, heavily speckled with black.” During this study, we visited the Lamu Archipelago (Lamu, Manda, and Manda Toto Islands) for 8 days, collected evidence for the presence of C. pygerythrus, but did not see or hear this animal.

Due to the absence of observations of C. pygerythrus in the Lamu Archipelago during this study, additional visits and surveys need to be conducted to either support or reject C. p. excubitor as a valid subspecies. If C. p. excubitor is indeed a valid subspecies, than its geographical range is highly fragmented and it population size is critically low.

In this study we threat C. p. excubitor as a valid subspecies due to the lack of evidence to the contrary. However, having surveyed a part of the geographical range of C. p. excubitor, we have the following concerns about its current subspecies status:

Primates of the coastal forests of Kenya. 69 Y.A. de Jong & T.M. Butynski, 2009.

- C. p. hilgerti occurs on the mainland only ca 30 km away (Mpeketoni) from Manda Island. It is likely that the geographical range extents even farther towards Lamu and Manda Islands. - Manda Island is separated from the mainland by a very narrow and shallow channel that, in some places, is <20 m wide. During low tide, Manda Island is connected to the mainland. Lamu Island is ca. 15-200 m from the mainland. Therefore, both Islands are not particularly isolated. The extensive mangrove around all Islands might, however, be much more of a barrier to primate movement than is the open water.

Future research priorities: • What is the geographical range of C. p. excubitor? • Obtain field observations, photos, and detailed descriptions of specimens of C. pygerythrus from Lamu, Manda and Patta Islands and C. pygerythrus from the adjacent mainland to confirm the validity of C. p. excubitor.

Primates of the coastal forests of Kenya. 70 Y.A. de Jong & T.M. Butynski, 2009.

4.2.8 Papio cynocephalus ibeanus Thomas, 1893

Vernacular name: Ibean yellow baboon Subspecies type locality: Lamu Island, Lamu Archipelago, Kenya Synonyms: Papio ruhei Zukowsky, 1942. Banks of Webbi Shabeli, 40 km NW of Mogadishu, Somalia; Papio thoth ibeanus, Thomas 1893, Kenya: Lamu

Field characters: Large, slender, mostly terrestrial primate Figure 25. Adult male Papio cynocephalus ibeanus (Figure 25). Males have a trace of mane, are foraging on the beach of Manda Toto Island, Lamu larger and more robust than females. Muzzle Archipelago, Kenya. almost hairless and pointed forward and slightly upwards with dark grey to blackish skin. In most populations, pale grey ‘muzzle-patches’ are present on either side of the muzzle just below the eyes. Cheeks and chest yellowish to off-white. Ears sparsely haired, dark grey to blackish, pointed in some populations. Crown grizzled yellowish-brown. Neck, dorsum, sides, and outer limbs grizzled yellowish-brown, yellowish-grey, or olive-brown. Ventrum and inner limbs pale yellowish-grey or off-white. Back of the upper legs pale yellow to golden-yellow. Hands and feet grizzled yellowish-brown to blackish. Tail grizzled yellowish-brown, base upwards, followed by a kink or a bend, than dropping down. Tail shape and length variable within groups. Callosities hairless, dark grey. Hair relatively wavy and long, especially on the shoulders, sides and upper legs.

Distribution: S Somalia, SE and coastal Kenya (Figure 26; Jolly, 1993; Kingdon et al., 2008; Altmann et al., in press).

Extent of occurrence: no data

Photographic map: Photo map of Papio cynocephalus ibeanus, contains 103 photographs, of which 17 coastal forests of Kenya (T.M. Butynski & Y.A. de Jong). http://picasaweb.google.com/wildsolutions/PapioAnubisPapioCynocephalus?feat=email# or via: www.wildsolutions.nl

Primates of the coastal forests of Kenya. 71 Y.A. de Jong & T.M. Butynski, 2009.

Figure 26: Approximate geographic distribution of Papio cynocephalus ibeanus over the coast of Kenya with sites of encounters (2003-2008) also depicted. Based on Y. de Jong & T. Butynski unpubl. data) and Kingdon et al. (2008e).

Encounter sites:

Encounters with P. c. ibeanus along the coast of Kenya. - Gazi (186; Appendix 2) - Diani (349, 351; Appendix 2) - Malindi (357; Appendix 2) - Tana River Primate National Reserve (18, 282; Appendix 2) - Mwenje (16; Appendix 2) - Tana River Primate National Reserve S (281; Appendix 2) - Garsen (257, 280; Appendix 2) - Kipini Conservancy (258, 260, 261, 262, 263, 264, 274, 275, 276, 277, 277A, 278, 278F; Appendix 2) - Witu Town (272A, 279, Appendix 2) - Witu Forest Reserve (270, 272; Appendix 2) - Witu Forest Reserve E (266, 267, 268; Appendix 2) - Manda Toto Island (365, 366; Appendix 2)

Encounters with Papio while travelling to the Hotspot - Garissa (283; Appendix 2) - Garissa SW (14, 19B, 284, 286; Appendix 2) - Garissa S (19; Appendix 2) - Kibwezi (252; Appendix 2) - Makindu (174, 176, 178, 179; Appendix 2) - E Sala Gate, Tsavo East National Park (359; Appendix 2)

Primates of the coastal forests of Kenya. 72 Y.A. de Jong & T.M. Butynski, 2009.

- Meru NP P. anubis (26, 84, 301, 308, 309; Appendix 2) - Mtito Andei (180; Appendix 2) - Mwardimu (340; Appendix 2) - Mwea National Reserve (2, 6, 9, 11, 363, 364; Appendix 2) - Tsavo West National Park (182, 325, 327, 329, 332, 335, 336, 339; Appendix 2) - Tsavo East National Park (360, 361; Appendix 2) - Voi N (250, 251; Appendix 2) - Voi S (183, 184; Appendix 2)

Encounter rate: A total of 71 Papio groups were encountered of which 32 were P. c. ibeanus groups within the Kenya coastal forests. The remaining 39 groups were encountered while travelling to and from the Hotspot (Table 10; Figure 26).

Table 10. Encounter rates (groups/hour and groups/km) of Papio cynocephalus ibeanus during 2005 – 2008 in the Kenya coastal forest Hotspot and while driving to the Hotspot. Only includes the surveys in which C. c. ibeanus was encountered. Area Groups/hour Groups/km Vehicle/Foot Tana River PNR 2005 0.1, n=1 Vehicle Tana River PNR 2006 0.1, n=1 Vehicle Kipini Conservancy & 0.8, n=20 0.16, n=20 Vehicle Witu Forest Reserve Manda Toto Island 0.2, n=2 0.13, n=2 Foot Mwea NR 2005 0.7, n=4 Vehicle Mwea NR 2008 0.7, n=2 0.05, n=2 Vehicle, 13.6 km/h Tsavo West NP 1.0, n=7 0.05, n=7 Vehicle, 20.1 km/h Tsavo East NP 0.3, n=1 Vehicle, 23.3 km/h Road Surveys

Mwea NR – Tana 0.1, n=2 0.004 Vehicle, ca 45 km/h River PNR Tana River PNR – 0.2, n=1 0.01, n=1 Vehicle, ca 45 km/h Garissa Tana River Primate 0.3, n=1 0.01, n=1 Vehicle, 30 km/h Reserve –Garsen – Witu Witu – Garsen – Tana 1.1, n=4 0.03, n=3 Vehicle, 40.2 km/h River PNR Nairobi – Makindu – 0.7, n=5 0.13, n=1 Vehicle, ca 65 km/h Mrima Hill Mazeras – Nairobi 0.4, n=3 0.01, n=1 Vehicle, 43.8 km/h Tana River PNR – 0.3, n=3 0.01, n=3 Vehicle, ca 50 km/h Garissa – Nanyuki Garissa – Kora NP 0.1, n=1 0.003, n=1 Vehicle Kora NP – Meru NP 0.4, n=1 0.02, n=1 Vehicle Mtito Andei – 0.2, n=1 0.004, n=1 Vehicle, 41.5 km/h Kilimangodo Kaya Sega – Diani 0.3, n=2 0.01, n=2 Vehicle, 27.9 km/h Mazeras – Nairobi 0.4, n=3 0.01, n=3 Vehicle Malindi – Tsavo East 0.3, n=2 0.01, n=2 Vehicle, 31.8 km/h NP

Vegetation: Savannah, open woodland, Brachystegia woodland (Miombo), dry bushland, and forest-grassland mosaic. Avoids forest but occupies forest edges. Encountered on beaches (Manda

Primates of the coastal forests of Kenya. 73 Y.A. de Jong & T.M. Butynski, 2009.

Toto Island, Kenya, and on beaches in Saadani National Park, coastal Tanzania; T. M. Butynski & Y. A. de Jong, pers. obs.) and uses mangrove forests (also Kingdon et al., 2008e). P. c. ibeanus is able to persist in secondary and highly fragmented vegetation, including cultivated areas. Often encountered close to human settlements, attracted by water, human food, crops, and security from predators.

Altitude range: Sea level to at least 1140 m asl

Conservation status: P. c. ibeanus is a widespread, common and opportunistic primate in the coastal forests of Kenya. Outside the Hotspot it is even more abundant, occurring in and outside protected areas. No major threats to the survival of this subspecies are known. P. cynocephalus is listed as a vermin under the African Convention due to their tendency to raid crops and forage in and around human settlements.

IUCN Red List: Papio cynocephalus ibeanus: Least Concern (Kingdon, 2008; http://www.iucnredlist.org/details/136862). Assessors: J. Kingdon

Discussion: P. c. ibeanus is locally common in the coastal forests of Kenya where it is often found near human settlements to which it is attracted by water human food, crops and security from predators. This species appears to be limited mainly by year around supplies of water and suitable sites for sleeping (i.e., large, steep, cliffs and/or tall trees).

The taxonomy of species with large geographical ranges and which occupy many different habitats are often a topic of debate. Often these widespread species show considerable phenotypic variation but are genetically linked due to intermediate populations (Sarmiento, 1998). During this study we found phenotypic variation among groups of P. c. ibeanus throughout its range inside and outside coastal forests of Kenya (see Table 11.). Groves (2001, 2005), Grubb et al. (2003), Kingdon et al. (2008e), and Altmann et al. (in press) recognize three subspecies for Papio cynocephalus: P. c. cynocephalus, P. c. ibeanus, and P. c. kindae.

Table 11. shows some of the variation between P. cynocephalus groups at Diani [S coast; which Hill (1967) referred to P. c. cynocephalus] and groups on Manda Toto Island [N coast; which Hill (1967) referred to P. c. ibeanus]. The variation observed between Diani and Manda Toto Island (ca 300 km apart) is, however, less than the variation found within groups at Tana River Primate National Reserve (ca 100 km from Manda Toto Island), which indicates that P. c. ibeanus is present in Diani. Hill (1967) indicates that the Galana River is the boundary between P. c. ibeanus and P. c. cynocephalus (with P. c. cynocephalus occurring in the south part of the Kenya coastal strip and into Tanzania). He, thus, considers the Diani population to be P. c. cynocephalus, contradictive to our findings.

P. c. ibeanus differs from P. c. cynocephalus in that its pelage is wavy---not straight (Jolly, 1993). Additionally, Jolly (1993) reports a trace of mane in P. c. ibeanus which is absent in P. c. cynocephalus. Zinner et al. (2008) also indicate that P. c. cynocephalus is present at Diani. Results of the molecular work on Papio in eastern Africa by Zinner et al. (2008) shows that Diani P. cynocephalus is genetically distinct from P. cynocephalus in Amboseli.

Primates of the coastal forests of Kenya. 74 Y.A. de Jong & T.M. Butynski, 2009.

P. c. ibeanus from S Somalia (once referred to as P. c. ruhei but now considered a synonym of P. c. ibeanus; Groves, 2001, Grubb et al., 2003, Zinner et al., 2008), is genetically closely related to P. anubis from Kenya and N Tanzania, and P. c. ibeanus from Amboseli. Zinner et al. (2008) speculate that P. c. ibeanus is a hybrid form between P. anubis and P. c. cynocephalus and that their P. cynocephalus sample from Diani represents P. c. ibeanus. Further research is needed along the south coast of Kenya and the north coast of Tanzania in order to determine where P. c. ibeanus gives way to P. c. cynocephalus, or whether there is a cline from one form into the other. Acknowledging that P. c. cynocephalus and P. c. ibeanus are hybridizing, we strongly suspect that no sharp boundary between these two subspecies exists and that what is present is a cline of intermediate forms. As such, the validity of P. c. ibeanus and of P. c. cynocephalus would be called into question as, indeed, would the validity of P. anubis.

The morphological features displayed by each ‘subspecies’ of Papio includes pelage, body size, and dentition (Hill, 1967; Jolly & Brett, 1973, Hayes et al., 1990 in Kamilar, 2006). According to Kamilar (2006), there is no clear differentiation among the ecologies of savannah baboon subspecies. This is in support of the hypothesis by Jolly (1993) that non-hamadryas baboon taxa lack ecological separation. The morphological variation that is found in the subspecies of Papio are due to a latitudinal cline and that all savannah baboons are of one species (Jolly, 1993; Kamilar, 2006).

We agree with Jolly (1993), Kamilar (2006) and Zinner (2008) on several points. However, we think it is useful in this study to classify the primates in the Kenya portion of this Hotspot as subspecies for the present time---pending further study. Therefore, we accept the taxonomy of Groves (2001, 2005), Grubb et al. (2003), Kingdon et al. (2008e), and Altmann et al. (in press), and recognise P. c. ibeanus to be the subspecies of P. cynocephalus present in the Kenya part of the Hotspot.

Natural hybridization between P. anubis and P. c. ibeanus in the extensive contact zone running across Kenya is often reported (Maples & McKern, 1967; Kingdon, 1971, 1997; Maples, 1972; Napier, 1981; Samuels & Altmann, 1992; Jolly, 1993, 1998; Sarmiento, 1998; Alberts & Altmann, 2001; Groves, 2001; Grubb et al., 2003; Newman et al., 2003; Kamilar, 2006; Kingdon et al., 2008e; Zinner et al., 2008; Altmann et al., in press; Palombit, in press; Butynski & de Jong, pers. obs.). The coastal forests of Kenya is occupied by P. c. ibeanus, no P. c. ibeanus x P. anubis hybrids were encountered within to the Hotspot. Hybrids were however encountered over an extensive area while driving to the Hotspot from the Kenya Highlands where P. anubis occurs.

Research has been conducted on hybrid groups in Amboseli National Park, Kenya. P. c. ibeanus is known to occur in Amboseli but P. anubis sometimes move into Amboseli from either the south (Mt Kilimanjaro), southeast (Namanga), or north (hybrid zone occurring around Shimba and Kiboko, along the Nairobi – Mombasa Highway; Maples & McKern, 1967). More research is needed to more clearly delineate the P. c. ibeanus x P. anubis hybrid zone. However, this study confirms at least sections of the hybrid zones:

1. NE & E of Mt. Kenya towards the Tana Delta. Papio across this >200 km wide area are readily recognized as phenotypically intermediate between P. anubis and P. c ibeanus. They also exhibit much intragroup variation in coat colour and body shape. From the interior of Kenya towards the coast, baboons become phenotypically more like the coastal P. c. ibeanus.

2. Hybrids occur along the Nairobi – Mombasa Highway, starting at least 35 km E of Makindu (Mapes & Mc Kern, 1967), N of the Chyulu Hills National Park , through Tsavo. Along this hybrid cline, baboons gradually become phenotypically more like P. c. ibeanus towards the coast.

Primates of the coastal forests of Kenya. 75 Y.A. de Jong & T.M. Butynski, 2009.

Table 11. Field descriptions of adult male P. c. ibeanus obtained from Manda Toto Island, Lamu Archipelago (close to the type locality of P. c. ibeanus), Tana River Primate National Reserve, Diani Beach (S coast). Field P. c. ibeanus P. c. ibeanus P. c. ibeanus description Manda Toto Island Tana River PNR Diani Beach Foto

Cheeks Yellowish to off-white Yellowish-white to pale Pale yellowish-grey grey Nostrils Directed upwards Directed upwards Directed upwards Pale grey Present Present but vague Present muzzle patch Head Pointed Pointed Pointed shape Mane Trace Absent Trace Dorsum Grizzled yellowish- grey Grizzled yellowish- grey. Grizzled yellowish- grey Some with brighter yellow than others Ventrum Greyish to off-white with Pale yellowish-white to pale Pale yellowish-grey and inner yellow wash grey limbs Callosite Medium U-shape Medium U-shape Medium U-shape shape Tail Kinked Bent (arched) Kinked Body Robust compared to Tana Slender (gracile) Robust compared to Tana shape River PNR Papio River PNR Papio Overall Phenotypically similar to Very different from other Phenotypically similar to population at Diani, SE populations of Papio population at Manda Toto Kenya. observed in East Africa. Island (close to type locality) Also, more phenotypic variation seen within groups than observed for other populations of Papio in East Africa.

Further research priorities: • Where does P. c. ibeanus give way to P. c. cynocephalus, or is there a cline from one form into the other? • What is the exact location and size is the P. c. ibeanus x P. anubis hybrid zone? • Are all savannah baboons part of one species?

Primates of the coastal forests of Kenya. 76

Y.A. de Jong & T.M. Butynski, 2009.

4.2.9 Colobus angolensis palliatus Peters, 1868

Vernacular name: Peter’s Angola Colobus Subspecies type locality: East Africa, opposite Zanzibar (=Unguja Island). Synonyms: sharpei, Thomas, 1902. Fort Hill, Malawi. langheldi, Matschie, 1914. Ujiji, E Lake Tanganyika.

Field characters: Medium-sized, arboreal, wholly black-and-white monkey with white cheeks, shoulders, and tail tip (Figure 27).

Distribution: Discontinuously distributed in SE Kenya. Restricted to coastal forests of Kwale District. Northern boundary is the the border between Kwale District and Mombasa town. Southern Highlands, Eastern Arc Mountains, coastal forests and gallery forests of eastern Tanzania (Figure 28; Napier, 1985; Kingdon, 1997; Groves, 2001; Anderson et al., 2007; Kingdon et al., 2008d). Figure 27. Adult male Colobus angolensis palliatus at Mrima Hill, south coast of Kenya, where this Extent of occurrence: ca 272,000 km². Area of subspecies is common. occupancy in Kenya ca. 249 km² (Anderson et al., 2007).

Figure 28. Known distribution and encounters (2003-2008) with Colobus angolensis palliatus in the coastal forests of Kenya. The other sources used for this map are Kingdon et al., (2008d) and Anderson et al., (2007).

Primates of the coastal forests of Kenya. 77

Y.A. de Jong & T.M. Butynski, 2009.

Photographic map: The photo map for African Colobinae contains 17 photos, of which three are of C. a. palliatus (T. M. Butynski & Y. A. de Jong). http://picasaweb.google.com/wildsolutions/Colobinae?feat=email# or via: www.wildsolutions.nl

Encounter sites: - Mrima Hill (188, 189, 192, 193, 196, 199, 246, 247, 344; Appendix 2) - Diani (347, 375; Appendix 2)

Encounter rate: During this study, 11 groups of C. a. palliatus were encountered, all in the coastal forests of Kenya (Table 12; Figure 28).

Table 12. Encounter rates (groups/hour and groups/km) of Colobus angolensis palliatus during 2005 – 2008 in the coastal forests of Kenya. This table only includes the surveys in which C. a. palliatus was encountered. Area Groups/hour Groups/km Vehicle/Foot Mrima Hill 3.1, n=9 0.42, n=9 Vehicle Kaya Sega - 0.3, n=2 0.01 (n=2) Vehicle, 27.9 km/h Diani, road survey Diani Opportunistically, 1 Opportunistically, 1 Foot group group

Vegetation: C. a. palliatus was encountered in coastal forest, on the edge of forest and agricultural land, and in gardens of hotels and residencies. This taxon is able to survive in small patches of degraded and disturbed forest.

Altitude range: sea level – at least 550 m asl (Amani Nature Reserve, W. Newmark, pers. comm.)

Conservation: The major threat for the survival of C. a. palliates in Kenya is habitat loss and degradation due to clearance for agriculture, local settlements, tourism and coral block mining (Anderson et al., 2007; Bocian & Anderson, in press). The geographical range of C. a. palliates is highly fragmented. The largest forest, Shimba Hills National Reserve, is ca 150 km2 (Anderson et al., 2007). C. a. palliates is a flagship species for SE Kenya (Kahumbu, 1997; Cunneyworth & Rhys- Hurn, 2004, in Anderson et al., 2007). C. a. palliates is present in several protected areas (e.g., Shimba Hills National Reserve) and in some of the Mijikenda Kaya Forests (UNESCO World Heritage site since 2008; Unesco, 2009). Anderson et al. (2007) reports that C. a. palliates does not raid crops.

IUCN Red List: Colobus angolensis palliates: Least Concern (Kingdon et al., 2008d; http://www.iucnredlist.org/details/5148). Assessors: J. Kingdon, T.T. Struhsaker, J. Oates, J.F. Hart, T.M. Butynski, Y.A. de Jong, & C.P. Groves. C. a. palliates was ‘Data Deficient’ in the 2000 Red List Assessment.

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Discussion: C. a. palliates is a locally common subspecies in the coastal forests of south Kenya where it sometimes occurs in very small patches of forest. In 2001, C. a. palliates was found in 55 populations in forest fragments in SE Kenya (Anderson et al., 2007). The total population in Kenya was estimated to be 3100-5000 individuals. Shimba Hills National Reserve supports the largest C. a. palliates population (17% of the animals) and is the largest protected forest in the geographical range (density estimate: 2.9±0.52 groups/km2 and 15.3±2.88 individuals/km² in the Reserve; Anderson et al., 2007).

Density of C. a. palliates varies widely among sites and is significantly affected by forest area, forest loss over 12 years, and the availability of 14 major food tree species. Density in the Diani Forest is ca. 31 individuals/km2; in the Mwache F. R., ca. 6 individuals/km2. Are absent from some forest fragments (Anderson et al., 2007a). Historically, was present north of Mombasa to Arabuko Sokoke Forest (Anderson et al., 2007). Within Kenya, currently restricted to Kwale District, south of Mombasa.

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4.2.10 Procolobus rufomitratus rufomitratus (Peters, 1879)

Vernacular name: Tana River Red Colobus Subspecies type locality: Muniuni, Tana River, Kenya Synonyms: Piliocolobus rufomitratus (Peters, 1879)

Field characters: Medium-sized, arboreal monkey with russet crown and greyish-brown back and tail.

Distribution: From Kipende in the north to Mitipani 60 km to the south, where the Lamu-Garsen road enters the Tana River floodplain, Lower Tana River, Kenya (Butynski et al., 2008c)

Extent of occurrence: Sixty kilometres along the lower Tana River in riverine forest. Area of occupancy is considerably smaller as the forests here are highly fragmented and small (Butynski et al., 2008c).

Map: Map of the geographical range of P. r. rufomitratus by Butynski et al. (2008c) can be found on http://www.iucnredlist.org/details/39993/rangemap

Photographic map: No photos taken of P. r. rufomitratus

Encounter sites: - Mchelelo Research Station, (17B, 255; Appendix 2)

Encounter rate: No data.

Vegetation: Dependent upon riverine / gallery forest (Struhsaker & Grubb, in press).

Altitude range: 20-40 m asl

Conservation status: Habitat loss due to agricultural clearing and extraction of forest products by local communities is the main threat to P. r. rufomitratus (Butynski & Mwangi, 1994, 1995; Mbora, 2003; Mbora & Butynski, 2007; Butynski et al., 2008c; Struhsaker & Grubb, in press). Additionally, the alteration of river flow volume and cycles by five hydroelectric power dams upriver is likely to negatively affect the establishment, longevity and productivity of the forests along the lower Tana River (Butynski, 1995). Recently proposed agricultural ‘development’ projects (oil palm and sugar cane plantations) in the area are an indirect threat as they will greatly increase the number of people in the region. The proposed Tana Integrated Sugar Project in Tana River and Lamu Districts threatens more than 200 km² of semi-natural habitat in the area just to the south of the area in which the P. r. rufomitratus lives. The insecurity of the area and the lack of protection to the forests on which the P. r. rufomitratus depends could move this species into the Critically Endangered category in the near future (Butynski et al., 2008c; Wieczkowski & Butynski, in press).

IUCN Red List: Endangered B1ab(iii,v) ver 3.1 (Butynski et al., 2008c; www.iucnredlist.org/details/39993) Assessors: T. M. Butynski, T. T. Struhsaker & Y.A. de Jong.

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Discussion: Since 2002, we have received reports of Procolobus in the Witu Forest Reserve and Kipini Conservancy on the north coast of Kenya. Kipini is ca. 90 km southeast of the known range of P. rufomitratus. Brief visits to the nearby Witu Forest Reserve by T. Butynski (pers. obs.) in 1994 and 2004 failed to reveal any Procolobus, but Procolobus was not the focus of his and no time was given to questioning local people about the presence of Procolobus. During this study, in June 2006, T. Butynski and Y. de Jong visited Kipini Conservancy for 1.5 days. Four residents of the Conservancy claimed to have seen Procolobus in Kipini. One of these people, Bakari Kawa Buya, was once a field assistant on a P. rufomitratus research project along the Tana River. As such, he is very familiar with P. rufomitratus. The informants independently claimed that there were at least two small groups of Procolobus in that part of the Conservancy that we visited, and that, unlike P. rufomitratus, these animals had reddish backs and tails. That is, their pelage color is noticeably different from P. rufomitratus. We found only three species of monkeys in Kipini Conservancy: yellow baboon Papio cynocephalus, Sykes’s monkey Cercopithecus mitis, and vervet Cercopithecus aethiops. All three are readily distinguished from Procolobus, and all three are well known to the local people---yet they claim that a fourth species, a Procolobus, is also present. These comments by the local people, together with the proximity of Kipini to the forests of the lower Tana River, strongly suggests that there is a fourth species of monkey in Kipini Forest. We searched for Procolobus for 2.5 h on 24 June and for 3 h on 25 June. Although no Procolobus were encountered, we did find several sites with numerous green, compound, leaves on the ground, the petioles of which had been chewed off in a fashion characteristic of Procolobus feeding. Although we had considerable doubt as to the presence of Procolobus before we surveyed Kipini Conservancy, we realized that there was some chance that Procolobus are present---and that this was either a second population of P. rufomitratus or, more likely, a new taxon. If Procolobus are present, they must be in very low numbers. This, coupled with the fact that the area of Kipini Forest in which Procolobus are said to occur has recently been given over by the Kenya Wildlife Service (KWS) and Kipini Conservancy for resettlement of people from the lower Tana River area, means that time is a critical factor as the small area of remaining forest is likely to be destroyed in the near future. As a result of our preliminary survey on Kipini Conservancy, we applied for a grant from the Margot Marsh Biodiversity Foundation which allowed us to follow up on the reports of Procolobus in Kipini Conservancy and Witu Forest Reserve. Additionally we will survey two other areas in East Africa for the presence of Procolobus. The primary objectives of the project are to (1) confirm or refute reports that populations of Procolobus occur in the Kipini Forest, Ngezi Forest, and Rufiji River Forests, and, if confirmed, (2) assess both the taxonomic and conservation status of these populations, and (3) make recommendations for conservation action.

During February and March, 2007, the Danish Zoological Society (with funds from CEPF) conducted surveys in Witu Forest Reserve (Nielsen & Sick, 2008). Aware of the P. rufomitratus reports they found no evidence of their presence and consider it unlikely that additional surveys will record P. r. rufomitratus in Witu forest Reserve due to the condition and isolated nature of Witu relative to other forest containing P. r. rufomitratus (Nielsen & Sick, 2008). . During this study no data was collected at the Tana River Primate National Reserve on P. r. rufomitratus.

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4.2.11 Cercocebus galeritus Peters, 1879

Vernacular name: Tana River Mangabey Subspecies type locality: Mitole, Tana River, Kenya Synonyms: Cercocebus galeritus ssp. galeritus, Peters, 1879

Field characters: A semi-terrestrial, medium-sized, greyish monkey with a long, lax, ‘crest’ of hair on the crown.

Distribution: Endemic to a 60 km stretch of floodplain forests (in 27 patches) along the lower Figure 29. Cercocebus galeritus at Mchelelo Tana River, SE Kenya, from Kanjonja in the north Research Station, Tana River Primate to Garsen in the south (01.40S-02.15S, 40.10E, National Reserve, Kenya. 20-40 m asl; Butynski & Mwangi 1994, 1995; Butynski et al. 2008d).

Extent of occurrence: Sixty square kilometres, but area of occupancy is considerably smaller (Butynski & Mwangi, 1994).

Map: map of the geographical range of C. galeritus by Butynski et al. (2008d) can be found on http://www.iucnredlist.org/details/4200/rangemap

Photographic map: none

Encounter sites: - Mchelelo Research Station, (16A, 19A, 253; Appendix 2)

Encounter rate: No data. Found to be common within its small range (also Butynski et al., 2008d)

Vegetation: restricted to riverine gallery forests and adjacent patches of bush (Butynski et al., 2008d)

Altitude range: 20-40 m asl (Butynski et al., 2008d)

Conservation status: C. galeritus only contains of one population of ca. 1,200 individuals. C. galeritus is severely threatened by deforestation for agricultural land, timber, burning of adjacent grasslands preventing forest regeneration, overgrazing of forest understory by livestock, and changes to the flow of the Tana River and water table by hydropower dams and irrigation projects. Recently proposed agricultural ‘development’ projects (oil palm and sugar cane plantations) in the area are an indirect threat as they will greatly increase the number of people in the region. The proposed Tana Integrated Sugar Project in Tana River and Lamu Districts threatens more than 200 km² of semi-natural habitat in the area just to the south of the area in which the mangabey lives. The insecurity of the area and the lack of protection to the forests on which the mangabey depends could move this species into the Critically Endangered category in the near future (Butynski et al., 2008d; Wieczkowski & Butynski, in press).

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IUCN Red List: Endangered C2a(ii) ver 3.1 (Butynski et al., 2008d; www.iucnredlist.org/details/4200) Assessors: T.M. Butynski, T.T. Struhsaker, J. Kingdon & Y.A., de Jong

Discussion: During this study there were no data collected for C. galeritus. During a primate survey in June, 2006, at Mchelelo Research Station, were adult male loud calls (‘whoop-gobble’) of C. galeritus were recorded and sent to Prof. Carolyn Ehardt, Co-Project Leader of the CEPF-funded project ’Taxonomy and Conservation Genetics of the Threatened Mangabey Taxa of the Eastern Arc Mountains and Coastal Forests of Tanzania and Kenya Biodiversity Hotspot’ (a joint project with T. M. Butynski). The recordings were also sent to Dr. Jean-Pierre Gautier, the authority on the vocalizations of Cercocebus.

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4.2.12 Hybrid Chlorocebus pygerythrus hilgerti x Cercopithecus mitis albogularis

Field description: Slender, medium-size, greyish monkey (Figure 30). Bare, dark grey, skin around eyes and along top of muzzle. Cheeks and brow band pale grizzled grey. Ears dark grey with narrow pale grey. Throat patch and ventrum off white. Shoulders and outer limbs grizzled mouse grey.

Lower outer front limbs slightly darker grey. Inner limbs Figure 30. Adult male Chlorocebus pygerythrus hilgerti light grey. Hands and feet dark grey. Crown, dorsum, sides x Cercopithecus mitis albogularis hybrid on a large and proximal ca. 10% upper part of tail grizzled greyish-olive beach hotel compound in Diani, Kenya. with rufus wash. Remainder of tail grading into dark grey or blackish. Pelage around the ischial callosities pale grey with rusty wash. Scrotum dark grey to dark purple, partly hidden under the pelage.

Distribution: Leopard Beach Hotel, Diani, Kenya (Figure 31).

Figure 31. Encounter site of C. p. hilgerti x C. m. albogularis hybrid. Diani, Kenya (14 December, 2008).

Encounter sites: - Diani, S4.2848; E39.5913, 16 m asl

Encounter rate: One adult male C. p. hilgerti x C. m. albogularis hybrid was encountered once (14th December 2008 at 12.45 h) during a four days period. The hybrid was with two adult C. m. albogularis of which at least one was a male.

Photographic map: Photo map contains five photographs (T. M. Butynski & Y. A. de Jong). http://picasaweb.google.com/wildsolutions/ChlorocebusPygerythrusHilgertiXCercopithecusMitis AlbogularisHybrid?authkey=Gv1sRgCMqKmI2R_d6zXA&feat=email# or via: www.wildsolutions.nl

Vegetation: The hybrid was encountered in the gardens of the (large) Leopard Beach Hotel, about 100 m from the Indian Ocean. The vegetation includes many exotic and indigenous plants (including Terminalia sp, Cocos nucifera, Plumeria sp.). The hybrid and the two C. m. albogularis were feeding on flowers and human foods around the tourist rooms.

Altitude: 16 m asl

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Discussion: Only one C. p. hilgerti x C. m. albogularis hybrid was encountered during this study. Although C. p. hilgerti and C. m. albogularis are sympatric at many sites, there are no reports of hybrids between these two species or between these two genera in the wild (Lernould, 1988; Erhart et al., 2004; Lawes et al., in press). C. pygerythrus and C. mitis are ecologically, morphologically, behaviourally and genetically very different from each other (e.g., Dutrillaux et al., 1980; Erhart et al., 2004). Table 13. presents a comparison of the phenotypic characters of C. p. hilgerti x C. m. albogularis, C. p. hilgerti, and C. m. albogularis.

Bernstein (1971, cited in Gray, 1972) reported a dead full-term Cercopithecus aethiops x Cercopithecus mitis foetus at the Yerkes Regional Primate Research Center Field Station, Lawrenceville, Georgia, USA. Erhart et al. (2004) report two male Chlorocebus pygerythrus x Cercopithecus albogularis (in this study referred to as Cercopithecus mitis) hybrids, born in 1981 and 1986, which were the subject of their behavioural study at the University of Texas in Austin, USA. The parents originated in Uganda. One of the findings of their study is that the developmental trajectories of the hybrid males were more similar to C. albogularis than to C. pygerythrus (the mothers of the hybrids were C. pygerythrus). No behavioural observations were conducted on the Diani hybrid during our study but this particular hybrid was encountered with two C. m. albogularis.

Table 13. Phenotypic comparison of the Chlorocebus pygerythrus hilgerti x Cercopithecus mitis albogularis hybrid, Chlorocebus pygerythrus hilgerti and Cercopithecus mitis albogularis using field descriptions and photographs obtained at Diani Beach, Kenya. Phenotypic pygerythrus x mitis Chlorocebus pygerythrus Cercopithecus mitis charac- hilgerti albogularis teristics Photo

Ear fringe Narrow pale grey None Narrow, grey, some whitish Cheeks Medium long hair, White, medium long Medium long hair, grizzled pale grey. whiskers that sweep grizzled pale grey. upwards and backwards covering part of the ear. Face Triangular-shaped area of Jet black, hairless Grizzled grey. Bare black bare grey skin around eyes skin around eyes. and nose. Grizzled pale grey pelage around the mask. Eyes Mahogoney brown. Mahogoney brown. Light brown Brow band Projecting forward and White, narrow and short Projecting forward and narrow. Grizzled pale upwards. Grizzled pale grey. grey Crown Grizzled olive-grey with Grizzled olive-grey. Grizzled greyish-olive pale rufus wash. with pale rufus wash

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Collar Vague and not sharply Absent Present but not distinct or defined. Around lower ca. sharply defined. Around 50% of the neck. Throat lower ca. 50% of the neck. off white. Sides of neck Throat off-white. Sides of mouse grey. neck light grey. Throat Off white throat-patch Off white Off white throat-patch Shoulders Grizzled mouse grey Grizzled mouse grey Grizzled grey, becoming blackish on the arms. Outer Grizzled mouse grey. Grizzled mouse grey Black. arms Lower arms slightly darker grey. Inner arms Inner limbs light grey. Off-white Blackish. Outer legs Grizzled mouse grey. Grizzled mouse grey grizzled grey Inner legs Inner limbs light grey. Off-white pale grey. Hands and Dark grey. Black in males. Dark grey Black feet in females and young. Back of Hair slightly longer. Off Off white to pale grey. Hair longer. Off white legs white to pale grey. Rusty Red patch under base of with rufus wash. wash around ischial tail in males. callosities. Dorsum Grizzled olive-grey with Grizzled olive-grey. Grizzled grey between rufus wash extending unto shoulders. Below the upper tail. shoulders grizzled rusty extending onto the upper tail. Ventrum Off white Off white Grey. Center of the ventrum pale grey. Sides Grizzled olive-grey with Grizzled olive-grey. Mouse grey rufus red wash. Narrow off-white stripe between side and ventrum Tail Grizzled olive-grey with Grizzled mouse grey, Proximally rusty red from rufus wash over the distal 10% darker grey. dorsum. Distal part black. proximal 10% of upper tail Red patch under base of Intense red around base of (10%) grading into dark the tail in males tail. grey then blackish distally. Scrotum Purple to dark grey. Partly Bright turquoise to blue. Dark grey, largely hidden hidden behind fur. under fur. Overall Very grey animal. Body Phenotypically like other Phenotypically like other shape most resembles C. p. populations of C. p. populations of C. m. hilgerti. Generally hilgerti. albogularis. intermediate in colour between C. p. hilgerti and C. m. albogularis.

Future research priorities:

• Are there other reports of wild hybrids between C. p. hilgerti and C. m. albogularis and do they phenotypically resemble the ‘Diani hybrid’?

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5. DISCUSSION During this study we encountered 99 groups of diurnal primates and more than 60 nocturnal primates in the coastal forest of Kenya. We recognise nine species and 11 primate subspecies in the coastal forests of Kenya. All 11 subspecies are described and discussed in separate sections in Chapter 4.2.

Papio cynocephalus ibeanus was the most often encountered diurnal primate in the coastal forests of Kenya (32 groups), followed by Cercopithecus mitis albogularis (24 groups) and Cercopithecus mitis albotorquatus (16 groups). Three nocturnal primates species occur in the coastal forest of Kenya, Otolemur garnettii lasiotis and Galagoides cocos are common, Galago senegalensis braccatus is present but rare in this part of Kenya. This study is the first to report a wild Cercopithecus mitis albogularis x Chlorocebus pygerythrus hilgerti hybrid.

This study resulted in additional insights into the geographical range of all 11 subspecies. Considerable geographical range extensions for O. g. lasiotis, G. cocos and C. m. albotorquatus were found. Field descriptions were obtained of almost all subspecies and these have been compared with the descriptions of individuals encountered in parts of the geographic ranges outside of the coastal forests of Kenya. Data collected during this study, combined with museum and literature research, will be used to write additional articles and notes on the distribution, diversity, taxonomy, and conservation status of six primate species (Papio anubis, Papio cynocephalus, Cercopithecus mitis, Chlorocebus pygerythrus, Otolemur garnetttii, and Galago senegalensis) in Kenya and Tanzania, and in the ‘Kenya Primate Atlas’ that we are now compiling.

Primate encounters during this project formed the basis of our distribution maps, and species databases. In addition, locality data for all primate taxa occurring in Kenya have been obtained from a thorough search of the literature and museum specimens, as well as from communications with other field workers, and entered into these databases. Each Microsoft Access database holds one primate species and its subspecies. The databases will be used to compile and maintain distribution maps for each primate taxon. These databases will be updated soon after new information is received. As such, they will serve as ‘living’ databases to support primate research and conservation initiatives, such as the periodic IUCN Red List degree of threat assessments that are undertaken by the IUCN/SSC Primate Specialist Group. These databases will be made accessible on our website (wildsolutions.nl).

Priority research questions for the primates of the coastal forests of Kenya include:

1) To what extent does phenotypic variation in O. g. lasiotis occur over its geographic range? 2) What is the southern boundary of O. g. lasiotis? 3) What is the geographical range of G. s. braccatus within and outside the coastal forests of Kenya? , 4) How far up the Tana River does the range of G. cocos extent? 5) Which subspecies of C. mitis occurs between the Tana River/Delta and the Galana River? 6) What subspecies of C. mitis occurs between the Galana River and Kilifi Creek? 7) What is the geographical range of C. m. phylax? 8) Is C. m. phylax a valid subspecies? 9) What is the geographic range of C. p. hilgerti? 10) Is C. p. excubitor a valid subspecies? 11) What is the geographic range of C. p. excubitor? 12) What is the southern limit of the geographic range of C. m. albotorquatus?

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13) Are there other reports of wild hybrids between C. p. hilgerti and C. m. albogularis and do they phenotypically resemble the ‘Diani hybrid’? 14) Does Galagoides zanzibaricus occur in Kenya? 15) What primate taxa occur in the forests along the extreme northern coast of Kenya (especially Boni-Dodori Forest)? 16) What primate taxa occur on Patta Island? 17) What is the taxonomic status of Papio in the forests along the Lower Tana River?

We established a website – www.wildsolutions.nl; De Jong & Butynski (2009), ‘Primate Biogeography, Diversity, Taxonomy and Conservation in Eastern Africa’ to communicate our photos, audio recordings, species databases, publications, and other data/information to colleagues and institutions. This website will be updated on a regular basis, and will be used by T. M. Butynski and Y.A. de Jong as a tool and resource for their research and conservation projects in eastern Africa.

The combination of gradual climatic desiccation and increasing human activities accounts for much of the forest loss along the coast of East African (Clarke & Karoma, 2000). The majority of the coastal forests in Kenya are gazetted as Forest Reserves (Rodgers & Burgess, 2000a), however, the problems that occur in forest management and protection often go hand-in-hand with conflicts of interests among a) the local people who utilised the forest resourches in the past, b) commercial companies who exploit the forest, and c) the Forest Department which is responsible for forest protection and management (Burgess & Mbwana, 2000). As a result, the coastal forest of Kenya are under continuous and increasing threat. These forests are being degraded, fragmented, shrinking and disappearing. The expansion of agriculture is recognised as the most critical threat, while the production of charcoal, taking of firewood and timber, and the mining of limestone, lead, marble and several minerals is causing additional serious habitat loss (Obura, 2007).

The diurnal, forest–dependent, primates are the most threatened primate taxa in the coastal forests of Kenya. Of the 11 primate subspecies present, 55% (n=6) are categorized as “Least Concern”, 9% (n=1; C. m. albotorquatus) is “Vulnerable”, 18% (n=2; P. r. rufomitratus and C. galeritus) are “Endangered”, and 18% (n=2) were not assessed for the 2008 IUCN Red List of Threatened Species. The conservation threats to all primate taxa and survey sites in the coastal forests of Kenya are described in Chapter 4.1 and 4.2. Conservation activities should focus on the areas with the highest biodiversity or with endemic taxa (e.g. Rodgers & Burgess, 2000b). Of the sites visited during this study, primate densities were very low in the Lamu Archipelago, but these Islands hold five species of primate and up to two endemic subspecies. Primate biodiversity is relatively high in the Tana River Primate National Reserve (seven species), Diani (six species), and Kipini Conservancy and Witu Forest Reserve (five species). Additional primate surveys in Boni and Dodori Forest Reserves, Patta Island, Kaya Gonja, Shimba Hills National Reserve, Buda Forest and Mrima Hill (nocturnal surveys) are essential for compiling a list of ‘Primate Priority Conservation Sites in the coastal forest of Kenya. There can be no doubt, however, that the top priority site for primate conservation in the coastal forests of Kenya, indeed for all of Kenya, are the forests along the Lower Tana River. These forests not only hold the highest diversity of primate species in all of Kenya, they hold Kenya only two endemic species of primate. In addition, they are among the most threatened forests in all of Kenya.

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ACKNOWLEDGEMENTS We are grateful to Conservation International and CEPF for their financial support. We thank the Kenyan Ministry of Education, Science & Technology for permission to conduct field work in Kenya, and John Salehe, Ian Gorden, Lorna Depew, John Watkin, Russ Mittermeier, Stephen Nash, Andrew Perkin, Nike Doggart, Kristina Razon, Farouk Sherman, Quentin Luke, Trish Luke, Jean-Pierre Dekker, Juliet King, Bernard Agwanda, Jake Butynski, Tine van de Velde, George Eshiamwata, Simon Bearder, Todd Olson, Carolyn Harcourt, Carolyn Ehardt, Arnoud de Jong, Leanne Nash, David Mbora, Julie Wieczkowski, Noel Rowe, Jonathan Kingdon, Dennis Milewa, Paul Honess, James Culverwell, Jim Feely, Sheila Bell-Cross, Iregi Mwenja, Helen Dufresne and Tom Struhsaker for their support to this Project.

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Anderson, J., Cowlishaw, G. & Rowcliffe, J.M. 2007. Effects of forest fragmentation on the abundance of Colobus angolensis palliatus in Kenya’s coastal forests. International Journal of Primatology 28: 637-655.

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Bearder, S.K., Butynski, T.M. & De Jong, Y.A. 2008. Galago senegalensis. In: 2008 IUCN Red List of Threatened Species. . Downloaded on 08 February 2009.

Bearder, S.K., Honess, P.E. & Ambrose, L. 1995. Species diversity among galagos with special reference to mate recognition. In Creatures of the Dark: The Nocturnal Prosimians (eds L. Alterman, G.A. Doyle & M.K. Izard, M.K.), Plenum, New York. Pp. 331-52.

Bearder, S.K., Ambrose, L., Harcourt, C., Honess, P., Perkin, A.W., Pimley, E., Pullen, S. & Svoboda, N. 2003. Species-typical patterns of infant contact, sleeping site use and social cohesion among nocturnal primates in Africa. Folia Primatologica 74: 337-54.

Bennun, L. & Njoroge, P. 1999. Important Bird Areas in Kenya. Nature Kenya (East Africa Natural History Society), Nairobi.

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Appendix 1. Distinguishing character tables (draft) of Papio anubis, Papio cynocephalus, Chlorocebus pygerythrus, Cercopithecus mitis, Colobus angolensis, Colobus guereza and their subspecies, in Kenya and Tanzania.

Distinguishing characters in Papio anubis - DRAFT

P. anubis P.a. furax P.a. neumanni Face Purple-black (1), naked, dark grey to black (3), ruffs of hair at cheeks (3), Eastern Uganda and western Kenya Below the Chanler Falls on the Ewaso - Colour cheek dark grey (4), black (5), hair on cheeks prominent (5) (above the Chanler Falls on the Northern Ngiro, then southward through Kenya - Colour naked Ewaso Ngiro). and Tanzania. skin Mane Short to medium (1), male large manes (4), Upper parts Olive-brown (1), dark grey to olive-brown sometimes grading into olive or Darker than neumanni (6) Pattern and colour generally similar (6) light brown (khaki or grey) (3), Under parts Olive-brown (1), dark grey to olive-brown sometimes grading into olive or light brown (khaki or grey) (3), dark (5) Tail Broken (1), hairy, sometimes bent at acute angle midway appearing as if broken (3), directed up for proximal one-fourth then falls away as if broken (4), blackish (5) Limbs Hands and feet black in some populations (3), feet olive-brown (1), forelimbs blackish (5) Pelage overall Olive-brown (1), dark grey to olive-brown sometimes grading into olive or light brown (khaki or grey) (3), olive brown (4), furax darker than neumanni (?), dark greyish brown, fading into black (5) Male - Female Canines smaller in females (3) Nostrils Protruding beyond plane of upper lip (6) Other Shorter limbed, more robust and muscular (5) Shorter muzzle than neumanni (6)

Literature 1. Sarmiento, E. 1997. Current problems with Papio taxonomies. African Primates 3: 48-52 2. MOA ms, Papio cynocephalus draft 2002 3. MOA ms, Papio anubis draft 2003 4. Groves, C. 2001. Primate Taxonomy p.238-240. 5. Maples, W.R. 1994. Systematic reconsideration and a revision of the nomenclature of Kenya Baboons. Am. J. Phys. Anthrop. 36: 9-20 6. Dandelot 1971. An identification manual Primates In: Meester, J, & Setzer, H.W. The mammals of Africa. p. Distinguishing characters in Papio cynocephalus - DRAFT

P. cynocephalus P.c. typical P.c. ibeanus P.c. kinda Face Purple-black (1), muzzle greyish to black (2), cheeks almost white and hair sparse (2), Mikumi (T), Ruaha (T) Amboseli (K), Tana River (K) Angola, Zambia (west of - Colour cheek cheeks white (4), cheeks very pale approaching a yellow-white(5), hair short on top of head Typical and Ibean) - Colour naked and lays flat (5) skin Mane None, nape and neck yellow-fringed (1), no manes (4) Upper parts Brownish-yellow (1), light brown to yellow brown (2), yellow to yellow-brown (4). Juvenile reddish-brown (2) with whitish coat Under parts Almost white and hair sparse (2), white (4) Tail Variably arched (1), light brown to yellow brown (2), long tail tapered and sharp bend or Curved (4) hook, very variable (2), varying from simply curved in P. c. Kindae to more usually broken but held low (4) Limbs Longer limbs than anubis (2), outer limbs light brown to yellow brown (2), inner limbs almost white and hair sparse (2)(4). Pelage overall Brownish-yellow (1), long (2), straight and soft, silky wavier and coarser pelage straight and soft, silky pelage (2) (2) pelage (2) Male - Female Females 50% of male body size and 80% of limb length (2), Other Armpit and crotch colour ranges from pinkish to almost white (2). More variable in colour than anubis, from fairly bright yellow with black bands on the hairs to a rufus brown (5), fur long but less dense than anubis.

Literature 1. Sarmiento, E. 1997. Current problems with Papio taxonomies. African Primates 3: 48-52 2. MOA ms, Papio cynocephalus draft 2002 3. MOA ms, Papio anubis draft 2003 4. Groves, C. 2001. Primate Taxonomy p.238-240. 5. Maples, W.R. 1994. Systematic reconsideration and a revision of the nomenclature of Kenya Baboons. Am. J. Phys. Anthrop. 36: 9-20

Distinguishing characters for Chlorocebus pygerythrus – DRAFT.

Species C. tantalus C.p. excubitor C. aethiops C.p. johnstoni

Distribution Volta River to Sudan, Uganda and L. Turkana Witu Islands, Kenya Sudan & Ethiopia NE Tanzania & Kenya district. Limbs Grey (1). Lateral sides mouse grey Arms: forearm to elbow and on legs Laterally more grey than body (5) tights

from hindlimb to knee purely mouse yellowish wash (5). Greyer than body (6) grey. Upperarm speckled grey with Legs sometimes washed with yellow (6) black and the tight white (5) Hand/Feet Dorsum of h/f darker Black (5) Wrists and h/f black. Feet less black than hands (5). Black, also the wrists, sharply

defined (6) Feet less black than hands (6) Under parts White (1) Cream (5) Pale olive-yellow (4) Dirty white (5)(6) Upper parts Grizzled, gold to greenish (1). Golden green (5) Resembling hilgerti (1) Grey-green, frequently with yellowish Washed out tawny grey (5)(6)

tinge. Lighter bands of dorsal hairs between dark olive-buff and old gold (5) Tail Creamy white tip, white basal tuft (1). Long shiny Above dark grey with basal part End of tail whitish (4) Grey above, pure black at tip, rusty red at hairs (5). Whitish/yellow upper side and lightly showing extension of back colour, base (5). Grey above, scarcely washed speckled with black tips underside. Terminal end terminal end black, beneath yellowish with yellow black at tip (6) Large rusty

creamy white (5) white tuft at root (5) white with red patch at root (5) rusty-red basal patch Face Black (1) Black line from corner eye running back Whitish moustache Jet black (5)(6) along temples (1) Brow band & frontal White (1). Frontal band distinct, narrow in the Distinct but narrow (5) Frontal band greyish white to tawny white band centre and white (5) separate from the whiskers (5). Greyish-white or tawny-white

(5). Long black brow hairs border it in front (5) browband (6) Cheek Long, stiff and yellowish with short black tips (1). Whiskers long, falciform, pure white (4) Long whiskers passing gradually above whiskers White with yellow at tips, directed upwards and into colour of crown (5). Greyish-white or backwards conceiling the ears (5) Ends of longer tawny-white extending from browband (6) hairs speckled black & yellow, tips black (5) Long cheek hairs overlaying ears (6)

Crown Grizzled, gold to greenish (1). Golden green (5) Washed out tawny grey (5) Neck

Ears Body size Smaller than hilgerti (2)(4)

Scrotum Sky blue (1) Turquoise blue

Overall Sides golden green (5) rusty patch on pubic and Differs from hillgerti in that there is no perineal area and around scrotum (5) rufous in the pelage (5) 1. Groves, 2001. Primate Taxonomy, p. 202-203, 2. MOA ms Cercopithecus (a) pygerythrus, draft 03, 3. Lonnberg, 1913 Mammals collected by the Swedish zoological expedition to East Africa. K. Svenska Vetensk Aixad, Handl. 48(5): 37-39, 4. Dandelot 1971. An identification manual Primates In: Meester, J, & Setzer, H.W. The mammals of Africa. P. 27-29., 5. Hill, O.W.C. 1953. Primates. Comparative Anatomy and Taxonomy. 6. Pocock, R.I. 1907. A revision of the monkeys of the genus Cercopithecus, Proc. Zool. Soc London

Distinguishing characters in Chlorocebus pygerythrus - DRAFT

Species C.p. arenarius C.p. callidus C.p. centralis Distribution N. Kenya, from Tana to L. Rudolf, Somalia, central Tanzania, Kenya between L. Victoria and Rift NW Tanzania, SW Uganda, Rwanda, Burundi, DRC. Ethiopia Valley to Mt Elgon and Mt Debasien; Elgeyo and Mau esc. Limbs Deep mouse grey, upper arms tinge of dorsal Laterally iron grey (5) Light iron grey laterally, sharp demarcation with dorsal (5). colour (5) Forearm below elbow and hindleg below knee ashy grey speckled (6) Hand/Feet Deep brownish-black (5) Black (4). Almost completely black (5) Wrist, H/F black mixed with grey (5). Wrist and hand black, ankle grey, foot blackish (6) Under parts Well haired dirty white (5) Brownish fawn (4) Upper parts Pale olive green (5) Paler than johnstoni (5) Greenish (5). Greenish speckled with black (6)

Tail Brownish-red at root tail (5) Bicolour, grey above Grey speckled above. Greenish of back continues Grey with yellow speckling near root dorsally. Terminal end black

with greenish tinge; below white or yellowish white. on tail (5). Terminal end black (5) (5). Few rusty hairs at root (5)(6) Top black (5) Face Black (5) Chin black (5) Nose & muzzle with black hairs

Brow band & frontal band Black brow hairs (5)

Cheek White (5) Not white, individual variation (5) Short, not concealing ears (5)(6)

- whiskers

Crown Pale olive green (5) Greenish (5)

Neck Ears Body size Scrotum Surrounded with white hairs (5)

Overall Long pelage esp. on shoulders (5). Pubic area tinged with red

(6)

1. Groves, 2001. Primate Taxonomy, p. 202-203, 2. MOA ms Cercopithecus (a) pygerythrus, draft 03, 3. Lonnberg, 1913 Mammals collected by the Swedish zoological expedition to East Africa. K. Svenska Vetensk Aixad, Handl. 48(5): 37-39, 4. Dandelot 1971. An identification manual Primates In: Meester, J, & Setzer, H.W. The mammals of Africa. P. 27-29., 5. Hill, O.W.C. 1953. Primates. Comparative Anatomy and Taxonomy. 6. Pocock, R.I. 1907. A revision of the monkeys of the genus Cercopithecus, Proc. Zool. Soc London Distinquishing characters in Cercopithecus mitis - DRAFT Species albotorquatus kibonotensis stuhlmanni elgonis L. l. mauae C. l. phylax C. kolbi hindei Distribution Tana River (Kenya) S. Kenya & N. Tanz to Kilifi, W. Kenya & Uganda W. Kenya & Highlands & U Mau (Kenya) Patta & Witu Island (K) Tutha, Kenya district ? Taitas, Mazeras Under base of tail Red patch extending to thighs Russet tuft (2) sometimes present (2)(3) Under parts Dark ashy grey, buffy, olive grey, ashy grey or Black (2)(3) White (2) grey with buffy median line, grey with black speckled, grey speckled tufts of buff (2), grey (4), ashy grey or with white, black (2), grey white (6). cream (7). Anterior part of chest snow Blackish chest (10) white (9) Upper parts Orange suffusion rump (3), speckled Darker than under parts (3), frosty Pale (2) Olive (2) dark reddish-brown (2) grey with a wide band of russet (4), pale grey (6), steel blue grey sometimes yellow (5), dark olive (7). Speckled faintly greenish (7) yellow (9) Crown Very yellow or not yellow (2), dark olive Black (3)(7) Black, distinctive with Dark (3) Reddish-yellow (9) (7). Head speckled yellowish-grey with brow band(10). Back of the head black (9) black (10) Brow-band Dark olive (7) Grey – black, varies a lot(3), grey (7), grey white (2) Speckled as cheeks (10) Ear White (2), ear tufts yellowish, pale- No ear tufts (2)(3) Yellowish and banded (9) - tufts yellowish-white, no tufts, short grey (2) usually white (3) Almost bare ears, little hair (10) Cheeks Whitish whiskers, face black (6). Face dark olive (2) Speckled and sides of neck as brow band(10) Collar broad & white (almost complete (2)(3), small white (2), not complete (3) No collar (10) Black (3) white & partly (2) Less less white and less distinctive ¾ (4) not always present (7). Snow than kolbi (9) white half collar (9) Throat White (4)(7). Snow white contrasting Less white on throat than Kolbi , it Grey white (6), white (7) Greyish- black patch (2) with grey of cheeks (9) does not reach down to neck white patch (10) (good indicator)(8) Shoulders Dove-grey (5), dark olive (7). Speckled wholly black from crown to nape, speckled (2) yellowish-grey with black (9). Nape speckled, unspeckled (2) thick mass speckled yellowish-grey with black (9) of speckled hair on shoulders from cheeks (10) Limbs sometimes dark grey (2), dark grey (4), Hindlegs are much blacker than sometimes speckled down to feet, dark hind limbs (2)(3) Legs mouse grey (2), arms dove-grey (5). Inside fore leg ashy grey kolbi (8) black, dark grey (2), partly black (3) black (2) becoming sooty grey towards wrists. Outer limbs speck smokey grey (9). Inner limbs pale grey to white(9) Tail Dark grey (4). Base speckled yellow for Terminal half black (3) 0-60% of distal black (2), frosty grey less black & speckled (2) 2 inch, gradually rufouslaterally and (6) inferiorly, rest dark speckled. Term end black (9) Rusty red at root above and outside (9) Other Inner thighs may be reddish-brown (7). Very like albogularis (3), neck black Smaller in size (3) Cheeks reddish-yellow (9). Co Lumbar region dark brown, red brown, (7). No brown color in fur but grey deep ochre, reddisch ochre, olive black (10) Eyes dul muddy brown speckled (2). Cheeks speckled (10) yellowish-grey with black (little greyer than head) (9)

Distinquishing characters in Cercopithecus mitis - DRAFT Species neumani moloneyi monoides kolbi C.a. albogolaris doggetti Distribution Nyando Valley, 30 miles east of Mtn region of S. Tanz., Zambia, Coastal forest T (SE) Kenya highlands east of rift Zanzibar (T) NE Tanzania, DRC, SW Uganda, S Kisumu Malawi Burundi, Rwanda Under base of tail Reddish hair (3) Reddish hair (3) No red-brownish patch in adults, Russet tuft (3) Reddish (3) younger animals do show this patch (8) Under parts speckled grey (2) Dark (1), dark slate grey (7). Dark (7) Not black (1) Smokey grey (9) Upper parts bright russet (1), dark red saddle Reddish-brown (3)(7). speckled Redder than albogularis (3)(7), dark red Reddish-brown (3) Light-silvery brown (3), grizzled grey (7). Ruddy chestnut brown (9) and rich reddish orange-yellow (9) (5), russet (7) (5), light grey brown (7) Crown Dark grey (3). Blackish speckled Yellowish grey (3), yellowish-olive Darker than body (3) Black (3)(5)(7) with yellow (9) (7). Head thickly speckled with black (9) Brow-band narrowly banded, dusky grey Light speckled grey (3). White Yellowish gray (3) Brow fringe (3) Pale & grizzled (5) (2) above brow (9) Ear Grey (2) Tufts present (8) Tufts Long & bright - Tufts (5)(7), white (8)(9)

Cheeks Whiskers light speckled grey (3). Cheek whiskers yellowish grey (3), Greyish (2), cheek whiskers (3) Long grizzled whiskers (5) Cheeks speckled greyish yellow cheeks yellowish-olive (7) (9) Collar No collar (10) Creamy (3) No whitish half colar (9) Nearly complete (3), broad snow-white Nearly complete (3) (5)(7) Less sharp definitions between dark and white compared to albotorquatus (9) Throat Patch creamy (3)(7) White (1), little patch (3), varies in White (2) (3) size (7) Shoulders Speckled with greyish yellow (9). Yellowish-olive (7). Behind Black (5) Greyish (2), speckled yellowish grey In front of shoulders black (9) shoulders speckled and rich (3) reddish orange-yellow (9) Limbs slightly speckled (2) Thighs speckled grey (9). Black Thighs dark grey speckled (9) Hind legs speckled, looking dark grey (8) Dark grey, shiny black (5), hindlimbs upperarms (9). Grey speckled (9) Reddish tinge on hindlegs blackish grey (7) H/F black speckled (9) Tail 66% of distal half black (3), ventral 40% of terminal end black (3) No red on on anal region and base tail 60% distal part black 30% terminal end black surface reddish (7) Black, (9) speckled towards base, reddish above root (9) Other Speckled gray with dark reddish Neck yellowish-olive (7). Neck Longer fur than albotorquatus (9) Nape black (3)(7). Buff speckled saddle (3), nape dark grey (3), thickly speckled with black (9) frontal band (3). Possibly the closest sides and thighs light grey (7)(1). Closely resembling C. a. resemblance with ancestral type (5) Sides speckled grey (9) albogularis (9)

Literature 1. Dandelot 1971 In: Meester, J, & Setzer, H.W. The Mammals of Africa. p. 18-21., 2. Grubb, P. 2001. Synonyms reduce the number of subspecies in the guenon Cercopithecus mitis. African Primates 5: 24-33., 3. Groves, C. 2001. Primate Taxonomy p.206-211., 4. Harcourt, C.S. 1974. Ecology & social organisation of the Sykes monkey in the Tana River Gallery Forest (MS). 5. Kingdon, J. 1984. East African Mammals. Pp. 234-247., 6. Rudran, R. 1978. Socioecology of the Blue Monkeys (C.m. stuhlmanni) of the Kibale Forest, Uganda. 7. Lawes, Cords & Lehn (ms) Cercopithecus mitis, Mammals of Africa., 8. Lonnberg, 1913 Mammals collected by the Swedish zoological expedition to East Africa. K. Svenska Vetensk Aixad, Handl. 48(5): 37-39. 9. Pocock, R.I. 1907. A revision of the monkeys of the genus Cercopithecus, Proc. Zool. Soc London 10. Booth, 1963 Nat. Geo soc. Research report.

Distinguishing characters in Colobus angolensis – DRAFT

Species C.a. palliatus C.a ruwenzorii (adolfifriederici) C.a sharpei (syn. palliates) Distribution SE Tanzania into SE Kenya DRC, Ruanda, E shores Lake Tanganyika, Tanzania S Tanzania Tail Tail tip white , bushy, occupying about 33% of tail Distal 5-10cm of tail greyish (1), tail almost entirely Distal 25% of tail grizzled (2), tail brush rather full, - tip length (1), no distinct brush at end of tail, the distal black, distal 5% grizzled (2), tail almost entirely black white and 33% of tail length but the speckling - base third being white but the area up to the middle (3)(4) extends farther up to about the middle of the tail (4) - brush being distinctly speckled (4) Cap Cap hairs lying flat toward back with nuchal parting Hairs of the occiput broadly and evenly lengthened (3), hairs of occiput lengthened into a distinct tuft (4) (4) Brow band White brow band fairly broad, broadly connected Narrow brow band (2)(4) Distinct white brow band (2) via full cheek whiskers with epaulettes (1) Cheek Epaulettes large (1)(3), full cheek whiskers Whiskers and epaulettes forming a broad, continuous Whiskers, throat and chest fringe very long and full, - epaulettes connected with forehead band and epaulettes white band, sometimes overlain with long black hairs broadly connected with the shoulder mane (4), long - whiskers (1), whiskers not very long, epaulettes poor, (1), large epaulettes (3). epaulettes (2) broadly connected with shoulder mane (4) Frontal band White frontal band (3) White fontal band moderately broad (4) White frontal band moderately broad (4) Shoulders Whiskers, throat and chest fringe very long and full, broadly connected with the shoulder mane (4) Pubic region White pubic band, broad and oval in males, narrow a 6 to 10 cm wide band of white or greyish in pubic White broad pubic patch (2) in females. White perineal fringe (3), perineal region (1), pubic patch small and grizzled (2)(3) patch small (4) Pelage Coat long, thick and soft (4) Very long pelage (3), Long pelage (2), coat very long, thick and soft but rather short (4) Literature 1. Groves, 2001. Primate Taxonomy, p. 250-251 2. O’Leary, R. 2003. An Annotated Catalog of the African Primate, Genera Colobus and Procolobus in the Collections of the American Museum of Natural History. 3. Dandelot 1971. In: Meester, J, & Setzer, H.W. The mammals of Africa. P. 32. 4. Schwarz, 1929. On the Local Races and Distribution of the Black and White Colobus Monkeys. Proceedings of the Zoological Society of London. Distinguishing characters in Colobus guereza - DRAFT Species C.g. caudatus C.g. matschiei C.g. kikuyensis C.g. percivali C. abyssinicus roosevelti Distribution Mt Kilimanjaro, Mt Meru, Momela Kenya, west of Rift and in the Rift, From Ngong Escarpment to Mt Mount Gargues, Mathews Range, Mau Forest (8) Lakes and Kahe. west to Mt Elgon, south to Kenya and Aberdares, Kenya Kenya Ngorongoro Crater and the Grumeti River Tanzania Tail White tail tuft making up over White tail tuft short (45% of tail White tail tuft very bushy, extending White tail tuft extends over 63% of More extensively black tail than - Length 80% of tail length, only extreme length), rest black (1)(6), tail over 72% of tail length (1), tail tail (1), tail as long as head plus C. g. matschiei (8). White - Tuft proximal part of tail black (1)(6). longer than head and body (1), proximally gray (1), tail short, about body (1), tuft full (3)(6) and bushy, terminal end of tail 25%, other - Base Base of tail black, slightly shorter tuft copious, 50% of the length of equal to head plus body (1), Base of occupies 66% of the whole tail (3). 75% black short hair (8) than in kikuyensis (3). Tail tuft tail (3)(4). Tail base black, tail tuft tail grey, slightly longer than in Tail tuft smaller than caudatus, longer longer than kikuyensis (3), tail full covering 33% of total length caudatus (4). Tail completely white tail, larger body than caudatus (8) completely white, often yellow in (5). with tuft almost from base (4). Base colour (4). 3 to 4 inches of base of tail black, white tuft voluminous black, the rest white tuft (20-21 forming about 66% of the length of inch)(7) tail (5)(6). Veil Flank veil longer than kikuyensis Flank veil much yellowed, not Flank veil very long extending well up Flank veil very long, creamy yellow, - length (1)(3). Veil covers black part of extending up onto back but on back and beyond base of tail tuft only just reaching tail base (1)(3). tail completely covering base of tail (1), posterior (1)(3), veil covering 30% of the tail portion of veil about as long as the (4). Veil covering up to a quarter of tail tuft, extending to the basal the tail (5) portion of the tail (3)(4)(5), occasionally as far down as the beginning of the tuft (3). Shoulders White hairs sprinkled over No white sprinkling on shoulders shoulders, nearly linking veil to below veil (1) white of throat (1) Throat White (1) Under parts Fur less woolly than kikuyensis (1)(3) Other Longer and distinct thigh-band Hair short (1) Hair long; thigh stripe short (1), Hair very long (1). Temporal ridges Resembling C. g. matschiei but (compared to kikuyensis) (3) Thigh-band short and pale (3), fur into sagittal crest (8) smaller. Sagittal cres on the soft and woolly (3) skull when aged (8)

Literature 1. Groves, 2001. Primate Taxonomy, p. 250-251 2. Oates, J.F. & Trocco, T.F. 1983. Taxonomy and Phylogeny of Black-and-White Colobus Monkeys. Folia Primatologica: 40:83-113. 3. Schwarz, 1929. On the Local Races and Distribution of the Black and White Colobus Monkeys. Proceedings of the Zoological Society of London. 4. Dandelot 1971. An identification manual Primates In: Meester, J, & Setzer, H.W. The mammals of Africa. P. 31. 5. O’Leary, R. 2003. An Annotated Catalog of the African Primate, Genera Colobus and Procolobus in the Collections of the American Museum of Natural History. 6. MOA ms Colobus guereza, draft 04 7. Thomson 1885, Report on the mammals obtained and observed by Mr H.H. Johnston and Mt Kilimanjaro. Proc. Zool. Soc. Lond. p. 219 8. Heller, 1913. Smiths. Misc. Coll. lxi, No. 17, p11.

Appendix 2. Encounters with Primate groups within and outside the coastal forest of Kenya (2005-2008).

Encounters with Cercopithecus mitis albogularis and Cercopithecus mitis albotorquatus within and outside the coastal forests of Kenya (2005-2008).

Number Species Locality name Latitude Longitude Altitude m Vegetation Date Time 185 C. m. albogularis Gazi -4.40361 39.50647 9 Along road 07-02-06 16.37 187 C. m. albogularis Mrima Hill -4.49444 39.26333 81 Secondary forest edge, coastal and shamba mango cashew 07-02-06 17.16 baobab cocos palm 190 C. m. albogularis Mrima Hill -4.48444 39.27057 110 Secondary forest edge, coastal and shamba mango cashew 07-02-06 17.27 baobab cocos palm 191 C. m. albogularis Mrima Hill -4.48361 39.27133 82 Secondary forest edge, coastal and shamba mango cashew 07-02-06 17.29 baobab cocos palm 194 C. m. albogularis Mrima Hill -4.49069 39.25439 101 Secondary forest edge, coastal and shamba mango cashew 07-02-06 17.55 baobab cocos palm 195 C. m. albogularis Mrima Hill -4.48881 39.25431 124 Secondary forest edge, coastal and shamba mango cashew 07-02-06 18.07 baobab cocos palm 197 C. m. albogularis Mrima Hill -4.48656 39.25418 130 Secondary forest edge, coastal and shamba mango cashew 07-02-06 18.19 baobab cocos palm 198 C. m. albogularis Mrima Hill -4.48428 39.25394 146 Secondary forest edge, coastal and shamba mango cashew 07-02-06 18.20 baobab cocos palm 248 C. m. albogularis Mrima Hill -4.48694 39.27056 79 Secondary forest edge, coastal and shamba mango cashew 20-02-06 11.13 baobab cocos palm - C. m. albogularis Kaya Rabai, Mazeras -3.94494 39.58164 212 Coastal forest edge including Coconut palms cashew nut (Anacardium occidentale), mango (Mangifera indica) 330 C. m. albogularis Tsavo West NP, -2.93328 37.90464 798 Ground water forest 20-4-08 12.10 Finch Hatton camp 333 C. m. albogularis Tsavo West NP, -2.93171 37.90537 806 Ground water forest 20-4-08 13.40 Finch Hatton camp 338 C. m. albogularis Tsavo West NP, -2.98343 38.02153 804 Ground water forest including Raphia farinifera and 20-4-08 17.25 Mzima Springs Phoenix reclinata palms. 341 C. m. albogularis Lunga lunga -4.55784 39.12630 48 In tall fig tree in riverine forest, close to village along the 22-4-08 10.30 highway 342 C. m. albogularis Vanga -4.66056 39.21556 0 Mangrove forest 22-4-08 12.15 343 C. m. albogularis Vanga W -4.54278 39.06028 123 Dense dry bushland 22-4-08 15.37 345 C. m. albogularis Buda Forest -4.46444 39.41497 77 Forest edge with highway 23-4-08 15.15 348 C. m. albogularis Diani -4.30525 39.58041 22 In small patch of trees/gardens along a fairly quiet beach 23-4-08 16.32 road in town 350 C. m. albogularis Diani -4.33461 39.56251 8 On hotel compound/garden 23-4-08 18.15 352 C. m. albogularis Diani -4.33584 39.56615 8 along the highway/beach road 24-4-08 8.12 354 C. m. albogularis Gedi Ruins -3.30912 40.01770 20 Coastal forest 24-4-08 15.32 355 C. m. albogularis Gedi Ruins -3.30912 40.01770 20 Coastal forest 25-4-08 8.39 346 C. m. albogularis Diani -4.31021 39.57641 11 Along main beach road 23-4-08 16.22 374 C. m. albogularis Diani -4.2848 39.5913 16 On hotel compound 13-12-08 12.50

17 C. m. albotorquatus Tana River Primate NR -1.87700 40.13878 40 Riverine gallery forest 06-08-05 07.00 17A C. m. albotorquatus Tana River Primate NR -1.87700 40.13878 40 Riverine gallery forest 06-08-05 08.50 18A C. m. albotorquatus Tana River Primate NR -1.87700 40.13878 40 Riverine gallery forest 06-08-05 17.22 254 C. m. albotorquatus Tana River Primate NR -1.87642 40.13756 40 Riverine gallery forest 20-06-06 15.00 254A C. m. albotorquatus Tana River Primate NR -1.87642 40.13756 40 Riverine gallery forest 20-06-06 17.00 255A C. m. albotorquatus Tana River Primate NR -1.87642 40.13756 40 Riverine gallery forest 20-06-06 16.00 269 C. m. albotorquatus Witu forest -2.38850 40.51738 25 Dense dry costal forest edge 23-06-06 08.28 271 C. m. albotorquatus Witu forest -2.38702 40.49663 25 Dense dry costal forest edge 23-06-06 08.50 271A C. m. albotorquatus Witu forest -2.38702 40.49663 25 Dense dry costal forest edge 23-06-06 08.58 273 C. m. albotorquatus Kipini Conservancy -2.43615 40.46513 14 On road in small dense forest patch in populated area 23-06-06 10.12 278A C. m. albotorquatus Kipini Conservancy Ca -2.50 Ca 40.56 14 Forest-grassland mosaic 24-7-06 09.47 278B C. m. albotorquatus Kipini Conservancy Ca -2.50 Ca 40.56 14 Forest-grassland mosaic 24-7-06 Ca 10.00 278C C. m. albotorquatus Kipini Conservancy Ca -2.50 Ca 40.56 14 Forest-grassland mosaic 24-7-06 Ca 10.00 278D C. m. albotorquatus Kipini Conservancy Ca -2.50 Ca 40.56 14 Forest-grassland mosaic 24-7-06 Ca 10.05 278E C. m. albotorquatus Kipini Conservancy Ca -2.50 Ca 40.56 14 Forest-grassland mosaic 24-7-06 Ca 10.15 367 C. m. albotorquatus Manda Island SE -2.29886 40.95814 3 Dense and diverse coastal shrub, including some taller trees 7-7-08 13.15 including Acacia, Commiphora, wild fruit trees on coral rag

4 C. mitis ? Mwea NR -0.79703 37.63078 1012 Very dense riverine vegetation 03-08-05 16.12 7 C. mitis ? Mwea NR -0.84478 37.64056 1017 Mixed dense Acacia woodland 04-08-05 08.47 8 C. mitis ? Mwea NR -0.84372 37.64008 109 Mixed dense Acacia woodland 04-08-05 09.05 10 C. mitis ? Mwea NR -0.83631 37.63294 108 Riverine, dense Acacia woodland 04-08-05 09.50 Encounters with Chlorocebus pygerythrus hilgerti within and outside the coastal forests of Kenya (2005-2008).

Number Species Locality name Latitude Longitude Altitude m Vegetation Date Time 1 C. p. hilgerti Mwea NR -0.82114 37.64000 1032 03-08-05 13.10 3 C. p. hilgerti Mwea NR -0.81444 37.62541 Dense Acacia- Commiphora bushland 03-08-05 15.57 5 C. p. hilgerti Mwea NR -0.79703 37.63078 Dense Acacia- Commiphora bushland 03-08-05 16.12 12 C. p. hilgerti Mwingi E -0.91331 38.12856 04-08-05 13.45 13 C. p. hilgerti Garissa SW -0.66986 39.12436 Acacia- Commiphora woodland 05-08-05 08.43 15 C. p. hilgerti Wenje (Mwiti) -1.78528 40.09735 45 Riverine vegetation 05-08-05 15.40 20 C. p. hilgerti Kora NP -0.24972 38.58889 Acacia- Commiphora bushland/woodland 08-08-05 16.08 21 C. p. hilgerti Kora NP -0.20778 38.63444 08-08-05 16.41 22 C. p. hilgerti Kora NP -0.07250 38.61083 08-08-05 17.22 23 C. p. hilgerti Kora NP -0.03750 38.61667 Bushland close to river 08-08-05 17.39 24 C. p. hilgerti Kora NP -0.03306 38.59194 Bushland close to river 08-08-05 17.49 25 C. p. hilgerti Kora NP -0.02553 38.57444 Bushland close to river 08-08-05 17.59 27A C. p. hilgerti Kora NP -0.07161 38.42039 Riverine vegetation 09-08-05 09.12 27 C. p. hilgerti Meru NP -0.07161 38.42039 875 Acacia- Commiphora bushland/woodland 09-08-05 08.58 28 C. p. hilgerti Meru NP 0.93360 37.21028 538 Riverine vegetation 09-08-05 11.00 Campsite, riverine forest and dry 85 C. p. hilgerti Meru NP 0.16597 38.20756 600 bushland 13-10-05 Makindu, Hunters Woodland, including Acacia xanthophloea, ficus. Short 175 C. p. hilgerti lodge -2.20999 37.71606 grass undergrowth. 06-02-06 15.13 Makindu, Kiboko Woodland, including A. xanthophloea, ficus. Short 177 C. p. hilgerti camp -2.19331 37.67158 927 grass undergrowth. 06-02-06 17.06 Makindu, Kiboko Woodland, including A. xanthophloea, ficus. Short 178A C. p. hilgerti camp -2.19331 37.67158 927 grass undergrowth. 07-02-06 06.40 181 C. p. hilgerti Tsavo -2.99344 38.45839 462 Along highway 07-02-06 10.31 249 C. p. hilgerti Voi S -3.57500 38.77694 249 Along highway, Commiphora-Acacia bush-woodland 21-02-06 08.22 251A C. p. hilgerti Wenje (Mwiti) -1.78122 40.10539 25 Acacia bushland, sandy soil, close to village 20-06-06 10.08 Tana River Primate 252A C. p. hilgerti NR N -1.84272 40.06139 42 Acacia bushland, sandy soil, close to village 20-06-06 10.47 Tana River Primate 256 C. p. hilgerti NR S -1.93767 40.08487 46 Acacia bushland sandy soil 21-06-06 10.48 Grassland with doumpalm, greenbush whith some 259 C. p. hilgerti Kipini Conservancy -2.49178 40.53170 14 taller trees 22-06-06 10.07 265 C. p. hilgerti Mpeketoni -2.39103 40.69322 14 Village 22-06-06 17.02 285 C. p. hilgerti Garissa SW -0.67555 39.10205 339 26-06-06 8.03 287 C. p. hilgerti Garissa SW -0.78003 38.73337 26-06-06 10.00 288 C. p. hilgerti Garissa SW -0.79605 38.60290 564 26-06-06 10.26 308A C. p. hilgerti Meru NP 0.16598 38.20754 599 Camp site, riverine forest, Acacia- Commiphora bushland/woodland 17-01-07 11.48 321 C. p. hilgerti Meru NP 0.13338 38.25423 13-2-07 7.40 322 C. p. hilgerti Meru NP 0.16298 38.21313 590 13-2-07 8.07 Makindu, Hunters 324 C. p. hilgerti lodge -2.21298 37.71455 Garden of a lodge, around pond 19-4-08 12.45 Tsavo West NP, 326 C. p. hilgerti Severin -3.00220 37.98203 721 Acacia bushland, medium to open. Close to river 20-4-08 10.45 328 C. p. hilgerti Tsavo West NP -2.98758 37.94847 751 Riverine Forest 20-4-08 11.24 Tsavo West NP, 331 C. p. hilgerti Finch Hatton camp -2.93328 37.90464 798 Ground water forest 20-4-08 12.10 334 C. p. hilgerti Tsavo West NP -2.94401 37.90562 797 Acacia- Commiphora bushland/woodland 20-4-08 14.25 Tsavo West NP, Ground water forest including Raphia farinifera and 337 C. p. hilgerti Mzima Springs -2.98343 38.02153 804 Phoenix reclinata palms. 20-4-08 17.25 On the edge of the beach, medium dense bushland 353 C. p. hilgerti Tiwi Beach -4.23965 39.60154 10 and a campsite/lodge 24-4-08 9.30 353A C. p. hilgerti Nyali -4.00000 39.70000 30 Planted diverse forest 24-4-08 11.43 356 C. p. hilgerti Malindi -3.20695 40.11279 Captivity 25-4-08 13.13 358 C. p. hilgerti Malindi W -3.21743 40.07061 13 Mango plantation/cattle pasture 25-4-08 15.00 Tsavo East NP, Campground with tall trees, surrounded by medium-open 362 C. p. hilgerti Ndololo camp -3.36131 38.64594 510 Acacia bushland 27-4-08 06.30 373 C. p. hilgerti Diani -4.2848 39.5913 16 In forest patch next to green hotel compound 13-12-08 12.45

Encounters with Papio within and outside the coastal forests of Kenya (2005-2008)

Number Species Locality name Latitude Longitude Altitude m Vegetation Date Time 26 P. anubis Meru NP -0.07333 38.41361 09-08-05 08.36 301 P. anubis Meru NP 0.26747 38.12800 757 Meru NP entrance 14-01-07 13.22 308 P. anubis Meru NP -0.13125 38.25040 540 Grass, edge with riverine forest 17-01-07 11.27 309 P. anubis Meru NP 0.13923 38.17225 Meru HQ 18-01-07 8.00 84 P. anubis Meru NP 0.16597 38.20756 599 On and around campsite, riverine forest and dry bushland 13-10-05 (including Acacia, commiphora)

19B P. c. ibeanus Garissa SW -0.73556 38.97806 Medium dense Acacia bushland, 08-08-05 08.21 14 P. c. ibeanus Garissa SW -0.64844 39.27019 Medium dense Acacia - Commiphora woodland 05-08-05 09.19 16 P. c. ibeanus Wenje (Mwiti) N -1.73361 40.08889 05-08-05 17.38 18 P. c. ibeanus Tana River Primate -1.87700 40.13878 35 Dry bushland, near riverine forest 06-08-05 11.29 NR 19 P. c. ibeanus Garissa S -0.78694 39.66833 Degraded open Acacia bushland 07-08-05 15.25 186 P. c. ibeanus Gazi -4.40361 39.50647 9 Along road 07-02-06 16.37 257 P. c. ibeanus Garsen -2.27600 40.22712 43 Dense Acacia coastal shrub, Ficus sycamorous 21-06-06 12.54 woodland 258 P. c. ibeanus Kipini Conservancy -2.54205 40.60223 13 Forest edge with agricultural land and small village 21-06-06 16.42 260 P. c. ibeanus Kipini Conservancy -2.48272 40.55528 Grassland with doumpalm, greenbush with 22-06-06 10.45 some taller trees 261 P. c. ibeanus Kipini Conservancy -2.47582 40.55717 13 Grassland with doumpalm, greenbush with some 22-06-06 10.55 taller trees, on edge with agricultural land 262 P. c. ibeanus Kipini Conservancy -2.47265 40.55675 Grassland, open bush/woodland with doumpalm 22-06-06 10.58 bordering agricultural land 263 P. c. ibeanus Kipini Conservancy -2.42517 40.61950 Grassland, open bush/woodland with doumpalm 22-06-06 14.35 bordering agricultural land 264 P. c. ibeanus Kipini Conservancy -2.42258 40.62205 16 Grassland, open bush/woodland with doumpalm 22-06-06 14.39 bordering agricultural land 266 P. c. ibeanus Witu forest E -2.34858 40.59468 16 Coastal forest clearing, moving into agricultural land and 23-06-06 07.48 tall grass 267 P. c. ibeanus Witu forest E -2.37585 40.53937 Open woodland with doumpalm 23-06-06 08.08 268 P. c. ibeanus Witu forest E -2.37767 40.53607 23-06-06 08.13 270 P. c. ibeanus Witu forest -2.38817 40.50380 25 Dense dry costal forest edge 23-06-06 08.45 272 P. c. ibeanus Witu forest -2.38475 40.49023 25 Dense dry costal forest edge 23-06-06 09.13 274 P. c. ibeanus Kipini Conservancy -2.48328 40.55850 Forest-grassland mosaic 23-06-06 11.11 275 P. c. ibeanus Kipini Conservancy -2.48420 40.56627 Forest-grassland mosaic 23-06-06 11.20 276 P. c. ibeanus Kipini Conservancy -2.48767 40.58597 Forest-grassland mosaic 23-06-06 11.29 277 P. c. ibeanus Kipini Conservancy -2.49470 40.60373 Forest-grassland mosaic 23-06-06 11.41 278 P. c. ibeanus Kipini Conservancy -2.48517 40.57365 Forest-grassland mosaic 24-06-06 11.00 279 P. c. ibeanus Witu town -2.38812 40.42112 25 Forest-grassland mosaic, on edge with town 25-06-06 11.57 280 P. c. ibeanus Garsen E -2.29057 40.25813 25-06-06 12.26 281 P. c. ibeanus Tana River Primate -1.98548 40.09865 Town 25-06-06 13.40 NR S 282 P. c. ibeanus Tana River Primate -1.87642 40.13892 Riverine gallery forest edge with dry bushland 25-06-06 14.35 NR 283 P. c. ibeanus Garissa -0.46300 39.62180 Close to human settlement 26-06-06 6.48 284 P. c. ibeanus Garissa SW -0.58888 39.34577 Open woodland 26-06-06 7.27 286 P. c. ibeanus Garissa SW -0.76335 38.80568 471 Medium dense bushland, red soil with little 26-06-06 9.18 to no undergrowth 340 P. c. ibeanus Mwardimu -4.17304 39.01811 251 Close to agricultural, outside medium dense Acacia 21-4-08 17.06 bushland 349 P. c. ibeanus Diani -4.33785 39.56533 Along beachroad 23-4-08 17.01 351 P. c. ibeanus Diani -4.33461 39.56251 8 Hotel compound/garden 23-4-08 18.15 357 P. c. ibeanus Malindi -3.20695 40.11279 Captivity 25-4-08 13.13 365 P. c. ibeanus Manda toto Island -2.22719 40.98672 9 Beach, mangrove forest 4-7-08 10.40 366 P. c. ibeanus Manda toto Island -2.22719 40.98672 9 Beach, mangrove forest 5-7-08 8.35 272A P. c. ibeanus Witu town S -2.49906 40.52739 25 Mainroad 25-06-06 11.57 277A P. c. ibeanus Kipini Conservancy -2.45803 40.63002 9 23-06-06 11.41 278F P. c. ibeanus Kipini Conservancy -2.49180 40.56206 15 Open woodland with tall grass, dense forest 23-06-06 11.41

2 Papio Mwea NR -0.80556 37.62556 Dense Acacia – Commiphora bushland 03-08-05 15.30 6 Papio Mwea NR -0.79508 37.61667 Open Acacia tortolis woodland, lots of undergrowth 03-08-05 16.48 9 Papio Mwea NR -0.83939 37.63725 Dense Acacia woodland 04-08-05 09.19 11 Papio Mwea NR -0.84075 37.62903 Riverine, medium Acacia woodland 04-08-05 10.03 174 Papio Makindu, -2.19331 37.67158 927 Highway 06-02-06 13.10 Kiboko camp 176 Papio Makindu, -2.20999 37.71606 Lodge compound with large yellow fever 06-02-06 15.54 Hunters lodge 178 Papio Makindu, -2.19331 37.67158 927 A. xanthophloea, ficus woodland, tall trees, short 06-02-06 17.45 Kiboko camp grass undergrowth. 179 Papio Makindu, -2.19331 37.67158 927 A. xanthophloea, ficus woodland, tall trees, short 07-02-06 08.25 Kiboko camp grass undergrowth. 180 Papio Mtito Andei -2.74000 38.23000 710 Highway 07-02-06 09.50 182 Papio Tsavo West NP -3.00180 38.46244 471 Highway, close to garbage dump 07-02-06 10.55 183 Papio Voi S -3.46458 38.65333 537 Highway 07-02-06 11.44 184 Papio Voi S -3.61031 38.83939 444 Highway, Acacia - Commiphora bushland 07-02-06 12.23 250 Papio Voi N -3.18639 38.49639 590 Highway 21-02-06 09.00 251 Papio Voi N -3.00444 38.46889 490 Highway 21-02-06 09.18 252 Papio Kibwezi -2.41139 37.94861 937 Highway 21-02-06 10.24 325 Papio Tsavo West NP, -2.90508 38.03984 787 Acacia - Commiphora bushland with dense tall grass 19-4-08 16.16 Chyulu Gate 327 Papio Tsavo West NP, -3.00218 37.97029 725 Open and low bush with short grass and bare 20-4-08 11.13 Severin ground. Edge with riverine vegetation. 329 Papio Tsavo West NP -2.96189 37.91249 787 20-4-08 11.52 332 Papio Tsavo West NP, -2.92463 37.91076 808 Garden of lodge. Ground water forest and water 20-4-08 12.39 Finch Hatton camp around 335 Papio Tsavo West NP -2.98965 37.95239 751 Open Acacia - Commiphora bushland 20-4-08 15.10 336 Papio Tsavo West NP -3.00074 37.98199 721 20-4-08 15.45 339 Papio Tsavo West NP, -2.98343 38.02153 804 Ground water forest including Raphia farinifera 20-4-08 17.25 Mzima Springs and Phoenix reclinata palms. 359 Papio Tsavo East NP, -3.12033 39.36525 145 Acacia bushland, medium 26-4-08 11.08 Sala Gate E 360 Papio Tsavo East NP -3.05814 38.86868 268 Open grass and bushland 26-4-08 15.45 361 Papio Tsavo East NP, -3.36131 38.64594 510 Tall trees, campground, medium/open Acacia 27-4-08 06.30 Ndololo camp bushland 363 Papio Mwea NR -0.78226 37.63596 1018 Riverine Forest, edge with dense Acacia 28-4-08 10.45 bushland and agricultural land 364 Papio Mwea NR -0.81176 37.60661 1075 Dense Acacia woodland with tall grass 28-4-08 13.50

Encounters with Colobus angolensis palliates within and outside the coastal forests of Kenya (2005-2008)

Number Species Locality name Latitude Longitude Altitude m Vegetation Date Time 188 C. a. palliatus Mrima Hill -4.49231 39.26725 80 Secondary forest edge, coastal and shamba mango cashew baobab cocos palm 07-02-06 17.23 189 C. a. palliatus Mrima Hill -4.48667 39.27094 86 Secondary forest edge, coastal and shamba mango cashew baobab cocos palm 07-02-06 17.26 192 C. a. palliatus Mrima Hill -4.48361 39.27133 82 Secondary forest edge, coastal and shamba mango cashew baobab cocos palm 07-02-06 17.29 193 C. a. palliatus Mrima Hill -4.49378 39.25539 81 Secondary forest edge, coastal and shamba mango cashew baobab cocos palm 07-02-06 17.50 196 C. a. palliatus Mrima Hill -4.48881 39.25431 124 Secondary forest edge, coastal and shamba mango cashew baobab cocos palm 07-02-06 18.07 199 C. a. palliatus Mrima Hill -4.48428 39.25394 146 Secondary forest edge, coastal and shamba mango cashew baobab cocos palm 07-02-06 18.20 246 C. a. palliatus Mrima Hill -4.49222 39.26722 72 Secondary forest edge, coastal and shamba mango cashew baobab cocos palm 20-02-06 11.10 247 C. a. palliatus Mrima Hill -4.48944 39.26917 80 Secondary forest edge, coastal and shamba mango cashew baobab cocos palm 20-02-06 11.12 344 C. a. palliatus Mrima Hill -4.49423 39.25727 83 Secondary forest edge, coastal and shamba mango cashew baobab cocos palm 23-4-08 11.24 347 C. a. palliatus Diani -4.30525 39.58041 22 In small patch of trees/gardens along a fairly quiet beach road in town 23-4-08 16.32 375 C. a. palliatus Diani -4.2848 39.5913 16 In forest patch next to green hotel compound 13-12-08 12.45

Encounters with Procolobus rufomitratus rufomitratus within the coastal forests of Kenya (2005-2008)

Number Species Locality name Latitude Longitude Altitude m Vegetation Date Time Tana River Primate 17B P. r. rufomitratus NR -1.87641 40.13891 46 Riverine gallery forest Tana River Primate 255 P. r. rufomitratus NR -1.87641 40.13891 46 Riverine gallery forest 20-06-06 15.30

Encounters with Cercocebus galeritus within the coastal forests of Kenya (2005-2008)

Number Species Locality name Latitude Longitude Altitude m Vegetation Date Time 16A C. galeritus Tana River Primate NR -1.87642 40.13756 46 Riverine gallery forest 6-8-05 05.00 19A C. galeritus Tana River Primate NR -1.87642 40.13756 46 Riverine gallery forest 7-8-05 07.30 253 C. galleritus Tana River Primate NR -1.87642 40.13756 46 Riverine gallery forest 20-6-06 14.10

Appendix 3. Diurnal road surveys conducted at the north coast of Kenya.

Only road surveys in which primates were encountered are included.

A. Tana River Primate National Reserve to Garsen to Witu, Kenya.

Survey date: 21 June 2006 Survey hours: 3.7 h Trip distance: 129 km Road: Partly tarmac road (B8, C112), 30.0 km/h

Primate encounter rates: Total of two diurnal primate groups encountered; 0.5 groups/h, 0.02 groups/km (n=2) C. p. hilgerti: 0.3 groups/h, 0.01 groups/km (n=1). P. c. ibeanus: 0.3 groups/h, 0.01 groups/km (n=1).

B. Witu to Garsen to Tana River Primate National Reserve, Kenya.

Survey date: 25 June 2006 Survey hours: 3,7 h Trip distance: 130 km Road: Partly tarmac road (B8, C112), 40.0 km/h

Primate encounter rates: P. c. ibeanus: 1.1 groups/h, 0.03 groups/km (n=4).

C. Nyali to Watamu (including Gedi Ruins)

Survey date: 24 April 2008 Survey hours: 4.4 h Trip distance: 129.4 km Road: Main highway (B8), 42 km/h.

Primate encounter rate: C. m. albogularis: 0.3 groups/h (n=1), 0.01 groups/km (n=1)

D. Watamu to north Malindi to Watamu

Survey date: 25 April 2008 Survey hours: 6.6 h Trip distance: 163.6 km Road: Main highway and several side roads (B8 & C103), 24 km/h.

Primate encounter rate: Two groups of diurnal primates encountered, 0.3 groups/h (n=2), 0.01 groups/km (n=2)

C. m. albogularis: 0.2 groups/h, 0.01groups/km (n=1) C. p. hilgerti: 0.2 groups/h, 0.01groups/km (n=1)

P. c. ibeanus, C. p. hilgerti and O. g. lasiotis found in captivity in Malindi. All three individuals said to come from the vicinity of Malindi. Photos and descriptions obtained.

The Otolemur garnettii found in captivity was an adult male. The individual looked phenotypically different from any O. g. lasiotis we had observed along the coast of Kenya (Chapter 4.2.1).

E. Watamu to Tsavo East National Park (Ndololo Camp).

Survey date: 26 April 2008 Survey hours: 8.1 Trip distance: 260 Road: secondary road (C103), 32 km/h.

Primate encounter rate: Two groups of diurnal primates encountered, 0.3 groups/h (n=2), 0.01 groups/km (n=2) Papio: 0.3 groups/h, 0.01 groups/km (n=2)

Appendix 4. Diurnal road surveys conducted at the south coast of Kenya

A. Nairobi to Makindu to Mrima Hill Forest

Survey date: 6-7 February 2006 Survey hours: 7.6 h Trip distance: 535 km Road: Highway (A109 & A14), ca 65 km/h

Primate encounter rate: Total of seven diurnal primate groups encountered; 0.9 groups/h, 0.01 groups/km (n=7). C. p. hilgerti: 0,1 groups/h, 0.002 groups/km (n=1). Papio: 0.7 groups/h, 0.01 groups/km (n=5). C. m. albogularis: 0.1 groups/h, 0.002 groups/km (n=1).

Remarks: Phenotypic characters of Papio change from ‘good’ Papio anubis in Nairobi to Papio cynocephalus ibeanus on the Kenya coast, with animals with intermediate characteristics in between Nairobi and the coast. The extent of the P. anubis x P cynocephalus hybrid zone needs to be determined (Chapter 4.2.8)

B. Mrima Hill Forest to Mazeras (including Shimba Hills National Reserve)

Survey date: 20 February 2006 Survey hours: 8.2 h Trip distance: 311.0 km Road: Secondary roads and highways (A14, C106, C107, A109), 44 km/h

Primate species encountered: None

Remark During this road survey we drove through and partly around Shimba Hills National Reserve. No primates were encountered during this survey. In 2001 the Shimba Hills National Reserve is reported to hold the largest population of C. a. palliates in Kenya (458 groups, 2436 individuals; Anderson et al., 2007).

C. North Kilimangodo to Kaya Sega

Survey date: 22 April 2008 Survey hours: 5.3 h Trip distance: 107.7 km Road: Secondary roads (including C106), 19 km/h.

Primate encounter rate: C. m. albogularis: 0.6 groups/h, 0.03 groups/km (n=3).

Remarks During this road survey we drove through Kaya Gonja, a 4.2 km² Forest Reserve close to Kaya Sega. In 2001, Kaya Gonja held seven groups (33 individuals) of C. a. palliates (Anderson et al., 2007). Main threats to this population are illegal logging and hunting. Other threats are charcoal making, firewood collection and encroachment (Anderson et al., 2007).

D. Kaya Sega to Diani

Survey date: 23 April 2008 Survey hours: 7.5 h Trip distance: 221.1 km Road: Secondary roads and partly highway (including C106, C108, A14), 28 km/h.

Primate encounter rate: In total eight diurnal primate groups encountered, 1.1 groups/h, 0.04 group/km (n=8) C. m. albogularis: 0.5 groups/h, 0.02 groups/km (n=4) C. a. palliates: 0.3 groups/h, 0.01 groups/km (n=2) P. cynocephalus: 0.3 groups/h, 0.01 groups/km (n=2)

Remarks During this road survey we briefly visited Dzombo Hill Forest, a 5 km² Forest Reserve and National Monument which reaches an altitude of 470 m asl. The Hill is covered with coastal mixed forest but is surrounded by degraded bushland, agriculture and human settlements (Figure 32). Despite its protected status, the forest covering Dzombo Hill is under threat from timber and pole extraction, bark stripping and agricultural encroachment (Bennun & Njoroge, 1999; Anderson et al., 2007). Anderson et al. (2007) encountered four groups Figure 32. Dzombo Hill Forest Reserve, surrounded by (23 individuals) of C. a. palliates in the forest. degraded bushland, agriculture and human settlements. Hunting is a serious threat to this small population (Anderson et al., 2007).

During this road survey we passed Buda Forest Reserve (6.8 km²) and encountered C. m. albogularis on the edge of the forest. In 2001, Anderson et al. (2007) counted eight groups (52 individuals) of C. a. palliates in Buda Forest. The main threat to Buda Forest is illegal logging. Other threats are charcoal making, firewood collection, and hunting (Anderson et al., 2007).

Further research Nocturnal and diurnal surveys should be conducted in Dzombo Hill Forest and Buda Forest to assess primate presence/absence and densities.

E. Diani to Tiwi to Nyali

Survey date: 24 April 2008 Survey hours: 2.2 h Trip distance: 64.9 km Road: Highway (A14) and some secondary side roads, 28 km/h.

Primate encounter rate: In total, two groups of diurnal primates encountered, 0.8 groups/h, 0.03 groups/km (n=2) C. m. albogularis: 0.4 groups/h, 0.02 groups/km (n=1) C. p. hilgerti: 0.4 groups/h, 0.02 groups/km (n=1)

Appendix 5. Diurnal road surveys, outside the coastal forests of Kenya

A. Mwea National Reserve to Tana River Primate National Reserve.

Survey date: 4-5 August 2005 Survey hours: 15.3 h (13.8 diurnal, 1.5 nocturnal) Trip distance: 547 km (541 diurnal, 5.8 nocturnal) Road: highways and dirt roads (B7, A3, B8), ca 45 km/h

Primate encounter rates: A total of five diurnal primate groups encountered; 0.36 groups/h, 0.01 groups/km (n=5). A total of two nocturnal primates were encountered; 1.33 individuals/h, 0.34 individuals/km (n=2) Papio: 0.14 groups/h, 0.004 groups/km (n=2). C. p. hilgerti: 0.22 groups/h, 0.006 groups/km (n=3) G. s. braccatus: 0.67 individuals/h, 0.17 individuals/km (n=1). Galago sp: 0.67 individuals/h, 0.17 individuals/km (n=1).

Remarks: Papio changes from ‘good’ P. anubis to the east of Mwea National Reserve to ‘good’ P. c. ibeanus at Kipini on the Kenya coast with intermediate (hybrid) individuals in between. The size of the Papio hybrid zone needs to be determined. Chapter 4.2.8 presents more details on the Papio hybrid zone.

Two solitary desert warthogs P. aethiopicus were encountered 15 km and 80 km west of Garissa town in medium-dense Acacia bushland. These are the first records west of the Tana River and extend the geographic range to ca. 265 km northwest of Mkokoni, the nearest locality (d’Huart & Grubb, 2001). These encounteres are published in two publications (Culverwell et al., 2008; Appendix 6, and De Jong et al., 2009; Appendix 7).

B. Tana River Primate National Reserve to Garissa.

Survey date: 8 August 2005 Survey hours: 5.13 h Trip distance: 202 km Road: highways and dirt roads (B8, A3, B7), ca 45 km/h

Primate encounter rates: Papio encountered; 0.19 groups/h, 0.01 groups/km (n=1)

C. Mazeras to Nairobi

Survey date: 21 February 2006 Survey hours: 7.0 h Trip distance: 440 km Road: Highway (A109)

Primate encounter rates: Four diurnal primate groups encountered; 0.57 groups/h, 0,01 groups/km (n=4). C. p. hilgerti: 0.14 groups/h, 0.002 groups/km (n=1). Papio: 0.43 groups/h, 0.01 groups/km (n=3).

Remarks: Papio changes from ‘good’ Papio anubis in Nairobi to P. c. ibeanus on the Kenya coast with intermediate (hybrid) individuals in between. The size of the Papio hybrid zone needs to be determined. Chapter 4.2.8 presents more detail on the Papio hybrid zone.

D. Nanyuki to Garissa to Tana River Primate National Reserve

Survey date: 19-20 June 2006 Survey hours: 15.0 h (12.1 diurnal, 2.9 nocturnal) Trip distance: 614 km Road: Highways and dirt roads (A2, C74, B7, A3, B8); 57.7 km/h and 38.3 km/h

Primate encounter rates: C. p. hilgerti: 0.16 groups/h, 0.003 groups/km (n=2).

E. Tana River Primate National Reserve to Garissa to Nanyuki

Survey date: 25-26 June 2006 Survey hours: 11.1 h Trip distance: 650 km Road: Highways and dirt roads (B8, A3, B7, C74, A2); 45.0 km/h & 57.0 km/h

Primate encounter rates: Total of six diurnal primate groups encountered; 0.54 groups/h, 0.01 groups/km (n=6) C. p. hilgerti: 0.27 groups/h, 0.01 groups/km (n=3). Papio: 0.27 groups/h, 0.01 groups/km (n=3).

Remark: Papio changes from ‘good’ P. anubis in Nanyuki to P. c. ibeanus at Kipini on the Kenya coast. About 100 km west of Garissa we encountered Papio hybrids with a striking golden yellow ventrum, cheeks, sides, inner limbs and back of their hind legs. The hybrid zone between the two species is obviously very wide with much phenotypic variation within and among groups between Garissa to at least Mwingi. The size of the Papio hybrid zone needs to be determined. Chapter 4.2.8 presents more detail on the Papio hybrid zone.

F. Nanyuki to Mtito Andei

Survey date: 19 April 2008 Survey hours: 7.4 h Trip distance: 451.5 km Road: Highways (A2, A109); 59 km/h

Primate encounter rates: C. p. hilgerti: 0.14 groups/h, 0.002 groups/km (n=1)

G. Mtito Andei to north Kilimangodo

Survey date: 21 April 2008 Survey hours: 6.5 h Trip distance: 285.8 km Road: Secondary roads and highway (A109,), 42 km/h

Primate encounter rates: P. cynocephalus: 0.15 group/h, 0.004 (n=1). Primate Conservation 2006 (21): 63 –79

Taxonomy, Distribution, and Conservation Status of Three Species of Dwarf Galagos (Galagoides) in Eastern Africa

Thomas M. Butynski1, Yvonne A. de Jong2, Andrew W. Perkin3, Simon K. Bearder3, and Paul E. Honess4

1Eastern Africa Regional Program, Conservation International, Nairobi, Kenya 2Kenya Institute of Primate Research, Ol Kalou, Kenya 3Nocturnal Primate Research Group, Anthropology Department, Oxford Brookes University, Oxford, UK 4Department of Veterinary Services, University of Oxford, Oxford, UK

Abstract: This paper reviews the complicated nomenclatural history for the Kenya coast galago, Galagoides cf. cocos, and exam- ines whether ‘cocos’ is the valid species name for this recently resurrected taxon. This paper also reviews the phenotypic and vocal differences among G. cocos; the Zanzibar galago (Galagoides zanzibaricus zanzibaricus); the Udzungwa galago (Galagoides zanzibaricus udzungwensis); and the Mozambique galago (Galagoides granti), as well as their geographic ranges and conserva- tion status. The following are among the findings: (1) Galagoides‘ cocos’ is the name that should be applied to the Kenya coast galago; (2) in the field, the loud calls of these three species are diagnostic and remain the best means for identification; (3) there is a suite of phenotypic characters that, when taken together, can be used to distinguish among these three species when in the hand or viewed in the field in good light at close range; (4) G. z. zanzibaricus is phenotypically distinct from G. z. udzungwensis; (5) the three species are parapatric or, perhaps, narrowly sympatric; (6) the three species are endemic to the coastal forests of eastern Africa with G. cocos in the north (Kenya and northeastern Tanzania), G. zanzibaricus in Tanzania, and G. granti from southern Tanzania to southern Mozambique; and (7) none of the three species is threatened at this time, although G. z. zanzibaricus meets the IUCN Red List criteria for an Endangered subspecies. Key Words: Dwarf galagos, Galagoides, cocos, granti, zanzibaricus, udzungwensis, taxonomy, conservation

Introduction Nomenclatural History for the Kenya Coast Galago

Many of the species and subspecies of the family Galagi- On 16 December 1911, Edmund Heller (1912) col- dae (galagos or bushbabies) have been subjected to repeated lected an adult male dwarf galago (one of 10 specimens) taxonomic revisions and name changes over the past century at Mazeras, Kenya, which he named Galago moholi cocos. (for example, Elliot 1913; Allen 1939; Hill 1953; Groves 1977, This taxon was raised to species status (Galago cocos) by 1993, 2001, 2005; Grubb et al. 2003). The Kenya coast galago Elliot (1913), but later placed as a subspecies of the Somali (or Diani small galago), Galagoides cocos, is no exception galago (Galago gallarum cocos), as a subspecies of the (Figs. 1 and 2). Galagoides cocos is a recently revived, highly Zanzibar galago (Galago zanzibaricus cocos), or simply not cryptic species of the coastal forest of eastern Africa. recognized as a valid taxon and placed as a junior synonym In this paper we (1) review the nomenclatural history for of the Zanzibar galago (Galago senegalensis zanzibaricus the Kenya coast galago, (2) provide new information that con- or Galago zanzibaricus zanzibaricus or Galago zanzibari- firms that cocos‘ ’ is the valid name for this recently revived cus or Galagoides zanzibaricus) (Table 1). Most recently, species, (3) summarize the phenotypic and the main qualitative this taxon has been provisionally referred to as ‘Galagoi- vocal differences among G. cocos, the Zanzibar galago (Gala- des cf. cocos’ (Bearder et al. 2003; Grubb et al. 2003). This goides zanzibaricus), and the Mozambique or Grant’s galago binomial is ‘provisional’ because the validity of the use of (Galagoides granti), (4) review their geographic ranges, and the name ‘cocos’ requires confirmation. (5) examine their conservation status (Figs. 1–7).

63 Butynski et al.

Figure 1. Adult (sex not known) Kenya coast galago (Galagoides cocos) from Arabuko-Sokoke Forest Reserve, southeastern Kenya (near Gedi). Note the muzzle patches and buffy-brown dorsum. Photograph by Harald Schuetz.

Figure 2. Adult female Kenya coast galago (Galagoides cocos) from Diani, southeastern Kenya. Note the muzzle patches. Photograph by Andrew Perkin.

Figure 4. Adult (sex not known) Udzungwa (or Matundu) galago (Galagoides zanzibaricus udzungwensis) from Matundu Forest Reserve, Udzungwa Moun- Figure 3. Adult male Zanzibar galago (Galagoides zanzibaricus udzungw- tains, south-central Tanzania (near Ifakara). Note that the hairs of the tail are of ensis) from Pande Game Reserve, Tanzania (near Dar es Salaam). Note the even length, sparse, and wiry, that the bone of the tail is visible, and that the tip absence of muzzle patches. Photograph by Nike Doggart. of the tail is dusky. Photograph from Honess (1996).

64 Galagoides in Eastern Africa

Many of the more recent taxonomic studies on Galagi- 1990) or of Galagoides zanzibaricus (for example, Honess dae make no mention of cocos, but presumably they consider 1996; Anderson 1999, 2000; DelPero et al. 2000; Masters and cocos to be a synonym of Galago zanzibaricus (for example, Bragg 2000; Masters and Brothers 2002). Groves 1977; Nash et al. 1989; Masters 1998; Zimmermann The Need for Confirmation of the Name ‘cocos’ for the Kenya Coast Galago

Until recently, G. zanzibaricus was considered a poly- typic species of the coastal forests from southern Somalia through Kenya and Tanzania (including Unguja Island, Zanzi- bar, Tanzania) to southern Mozambique, and inland to central

Table 1. Summary of the nomenclature changes for the Kenya coast galago (Galagoides cocos).

Authority Latin name

Heller (1912) Galago moholi cocos Elliot (1913), Hollister (1924) Galago cocos Allen and Loveridge (1927) Galago gallarum cocos Schwarz (1931), Hill (1953), Allen cocos a synonym of Galago (1939), Hill and Meester (1977) senegalensis zanzibaricus cocos a synonym of Galago Jenkins (1987) zanzibaricus zanzibaricus cocos a synonym of Galago Kingdon (1971, 1997), Groves (2005) zanzibaricus cocos a synonym of Galagoides Groves (1993) zanzibaricus Groves (2001) Galago zanzibaricus cocos Bearder et al. (2003), Grubb et al. Galagoides cocos (2003)

Figure 5. Adult (sex not known) Mozambique galago (Galagoides granti) from Rondo Forest Reserve, southeastern Tanzania (near Lindi). This individual is emerging from a tree hole and, thus, its tail is not visible. Note the relatively large, blackish ears and buffy-brown dorsum. Photograph by Simon Bearder.

Figure 7. Two adult Zanzibar galagos (Galagoides zanzibaricus zanzibaricus) Figure 6. Typical adults of three Galagoides spp. at the British Museum of (left) from Unguja Island, Zanzibar, Tanzania, and two adult Udzungwa galagos Natural History, London. Bottom to top: Kenya coast galago (Galagoides (Galagoides zanzibaricus udzungwensis) (right) from Kissarawe, Tanzania. All cocos) from Gande, Kenya, Udzungwa galago (Galagoides zanzibaricus ud- four specimens are at the British Museum of Natural History (BMNH), Lon- zungwensis) from Kissarawe, Tanzania, and Mozambique galago (Galagoides don. The two specimens of G. z. zanzibaricus represent the extremes in pelage granti) from Coguno, Mozambique. Note the great similarity in the color of the coloration among the 10 adult specimens at the BMNH. Note that the dorsum, dorsum, and that G. granti is the largest of the three species and has a wider, tail, and outer front limbs are medium to bright cinnamon in G. z. zanzibaricus fuller, tail. Photograph by Tom Butynski. and buffy-brown in G. z. udzungwensis. Photograph by Tom Butynski.

65 Butynski et al.

Tanzania, Malawi, and extreme eastern Zimbabwe (Hill 1953; tion and taxonomic tool (Zimmermann et al. 1988; Courtenay Groves 1977; Hill and Meester 1977; Smithers and Wilson and Bearder 1989; Harcourt and Bearder 1989; Nash et al. 1979; Jenkins 1987; Courtenay and Bearder 1989; Skinner 1989; Zimmermann 1990; Masters 1991; Bearder et al. 1995, and Smithers 1990; Groves 2001, 2005; Bearder et al. 2003; 2003; Honess 1996; Honess and Bearder 1996; Butynski et Grubb et al. 2003). The ecology, behavior, and vocal repertoire al. 1998; Ambrose 1999, 2003; Bearder 1999; Groves 2001; of the mainland subspecies, G. z. cocos, is well known, hav- Perkin et al. 2002), it was judged that the level of difference ing been the focus of detailed field studies at Diani and Gedi between the advertising calls of G. z. cocos and G. z. zanzi- Forests, Kenya (Harcourt 1986; Harcourt and Nash 1986a, baricus/G. z. udzungwensis is far greater than can be accom- 1986b). Far less well known is the nominotypical subspecies, modated at the subspecies level. In fact, the advertising call of G. z. zanzibaricus, an endemic of Unguja Island, Zanzibar. G. z. cocos is far more similar to the advertising ‘incremental’ It was not until A. Perkin visited Unguja Island in 1998 and call of G. granti (formerly G. zanzibaricus granti) (Fig. 10) recorded the species-specific advertising call of topotypical than it is to the ‘single unit rolling’ call of G. z. zanzibaricus G. zanzibaricus (Fig. 8) that it became clear that these two (see Bearder et al. 1995). As such, G. z. cocos was reassigned forms were different: species status, G. cf. cocos (Bearder et al. 2003; Grubb et al. 2003). Interestingly, the geographic range of G. zanzibaricus (1) The species-specific advertising call of G. z. zanzibari- is located between the geographic ranges of G. cocos and cus is very different from that of G. z. cocos. Galagoi- G. granti (see below). des z. zanzibaricus has a ‘single unit rolling call’ (Fig. 8), Grubb and co-authors give a succinct overview of this and G. z. cocos has an ‘incremental call’ (Fig. 9) (Hon- complex situation. ess 1996; Honess and Bearder 1996; Perkin et al. 2002; “Galagos at Diani, which were thought to be Grubb et al. 2003). Galagoides zanzibaricus (Harcourt and Nash, 1986a, b) are vocally distinct from true or topotypical G. zan- (2) The species-specific advertising call ofG. z. zanzibaricus zibaricus of Zanzibar. They are provisionally identi- is identical, or nearly so, to the species-specific advertis- fied as Galagoides cf. cocos, and we assign them to ing call of the recently named Udzungwa (or Matundu) the G. granti group. Galagos from the Udzungwa galago, Galagoides udzungwensis (see A. Perkin unpubl. Mtns and other localities in Tanzania have been data, cited in Bearder 1999). This led to the realization named Galagoides udzungwensis (Honess, 1996) that G. udzungwensis may not be a new species, but rather but, on the basis of their vocalization, do not differ synonymous with, or a subspecies of, G. zanzibaricus from those of topotypical G. zanzibaricus of Zanzibar (see Perkin et al. 2002; Bearder et al. 2003; Grubb et al. (A. Perkin unpubl. data, cited in Bearder, 1999). The 2003). Here we treat the Udzungwa galago as a distinct form udzungwensis may prove to be a valid taxon at mainland subspecies (G. z. udzungwensis), but emphasize the subspecific level but until the systematics is clari- that the taxonomic status of the Udzungwa galago is far fied, we relegate it to the synonymy of Galagoides from resolved (see below). zanzibaricus.” (Grubb et al. 2003, pp.1315–1316).

Based on a considerable body of knowledge concerning And below the species-specific advertising calls of the Galagidae, and “The galago recorded from Diani is vocally dis- their wide use and acceptance as a robust species recogni- tinct from Galagoides zanzibaricus and has been

Figure 8. Sonogram and oscillogram of the ’single unit rolling’ advertising call of the Zanzibar galago (Galagoides zanzibaricus zanzibaricus) from Unguja Island, Zanzibar, Tanzania, the type locality for this species. Call recorded by Andrew Perkin. This call is comprised of a series of ‘rolling’ trill units that, after a few units, increase in frequency and amplitude before reaching a mild crescendo and then trailing off with trill units of lower amplitude and frequency. The lowering of the frequency is achieved by eliminating the higher frequency elements. The lowest frequency elements remain constant. Each trill unit is made up of a very rapid series of trill subunits. The number of units per ‘single unit rolling’ call varies considerably (Honess 1996; A. Perkin, pers. obs.). For the above recording: Call length = 10.8 seconds. Frequency range = 0.62–11.12 kHz. Fundamental frequency = 0.75 kHz. Range of unit frequency modulation = 0.81–3.57 kHz. Number of phrases = 0. Number of units = 19. For the Udzungwa galago (Galagoides zanzibaricus udzungwensis) population in the Matundu Forest Reserve, south-central Tanzania, the type locality for this subspecies: Mean number of units per single unit rolling call = 14 (SE = 0.17, range = 1– 46, n = 2,122). Mean unit interval = 0.28 seconds (n = 181). Mean unit length = 0.22 seconds (n = 196). Range of fundamental frequency = 0.95 to 1.00 kHz (Honess 1996, Honess and Bearder 1996, A. Perkin unpubl. data). Oscillograms of the single unit rolling call of G. z. udzungwensis are presented in Bearder et al. (1995), Honess (1996), Honess and Bearder (1996), and King- don (1997).

66 Galagoides in Eastern Africa

Figure 9. Sonogram and oscillogram of the ’incremental’ advertising call of the Kenya coast galago (Galagoides cocos) from Diani Beach, southeastern Kenya. Call recorded by Simon Bearder. This call often, but not always, starts with a series of high-pitched, rapidly uttered, “chirrups” followed by units arranged in phrases that are high in frequency and amplitude, and that gradually become lower in amplitude. The number of units within each phrase increases incrementally until the end of the call. Often, as in this case, phrases with same number of units are repeated. The number of units per phrase rarely decreases. Units are often frequency modulated. For the above recording: Call length = 4.7 seconds. Frequency range = 0.65–11.15 kHz. Fundamental frequency = 0.98 kHz. Range of unit frequency modulation = 0.68–10.37 kHz. Number of phrases = 6 (with three introductory “chirrup” units and one incipient unit at the end). Mean number of units per phrase = 2.8. For the G. cocos population at Diani: Mean call length = 4.3 seconds (range = 1.7–8.6 seconds, n = 12). Frequency range = 0.8–9.3 kHz. Fundamental frequency = 0.8–1.2 kHz. Mean number of phrases = 6 (range = 3 –11, n = 13). Mean number of units per phrase = 2.5 (range 1–11, n = 60). Mean unit interval = 0.35 seconds (range 0.20 –0.59 seconds, n = 27). Mean unit length = 0.41 seconds (range 0.15–0.57 seconds, n = 33) (Courtenay and Bearder 1989). See also the acoustic measurements presented in Zimmermann (1990). Additional sonograms and oscillograms of the incremental advertising call, and other calls of G. cocos, are presented in Courtenay and Bearder (1989), Harcourt and Bearder (1989), Zimmermann (1990), Bearder et al. (1995), and Kingdon (1997).

Figure 10. Sonogram and oscillogram of the incremental advertising call of the Mozambique galago (Galagoides granti) from Rondo Forest Reserve, Rondo Plateau, southeastern Tanzania (near Lindi). Call recorded by Paul Honess. This sonogram is an example of a full incremental call in which the numbers of units in each phase gradually increase incrementally. Compared with the incremental call of G. cocos, the incremental call of G. granti maintains relatively consistent amplitude, has more units per phrase, and is more staccato. In the above example, the amplitude increases slightly during the middle phrases and decreases slightly during the last phrase. For the above recording: Call length = 4.9 seconds. Frequency range = 0.56 –11.18 kHz. Fundamental frequency = 0.75 kHz. Range of unit frequency modulation = 1.03 –5.08 kHz. Number of phrases = 6. Mean number of units per phrase = 3.6 (range 3 –5). In the G. granti population of the Rondo Forest Reserve: Mean number of phrases per incremental call = 5.8 (SE = 0.2, range = 1–17, n = 211). Mean unit interval = 0.55 seconds (n = 41). Mean unit length = 0.41 seconds (n = 53) (Honess 1996, Honess and Bearder 1996). Additional oscillograms of the incremental advertising call, and other calls, of G. granti are presented in Honess (1996), Honess and Bearder (1996), and Kingdon (1997).

67 Butynski et al.

Figure 11. Sonogram and oscillogram of the incremental advertising call of the Kenya coast galago (Galagoides cocos) from Kaya Chijembeni (Rabai), 4 km north- east of Mazeras, southeastern Kenya, the type locality for this species. Call recorded by Yvonne de Jong and Tom Butynski. Although numerous full incremental calls were heard and recorded at Kaya Chijembeni during two nights, there was much background noise and wind, and none of the recordings of the full incremental call were suitable for the production of a clear sonogram. The sonogram presented here is of an incomplete or incipient incremental call. Nonetheless, the first five phrases of incremental units are present and identifiable as the species-specific advertising call ofG. cocos. In this case, there are no “chirrup” phrases and the number of units does not increase incrementally but the call still follows the typical high-pitched, rapid or staccato pattern of the G. cocos incremental call. For the above recording: Call length = 2.8 seconds. Frequency range = 0.98–12.4 kHz. Fundamental frequency = 0.77 kHz. Number of phrases = 5. Number of units per phrase = 2. Frequency modulation is not detectable, probably due to the low amplitude of the call.

recorded elsewhere in Kenya and in Tanzania, where Kaya Fungo respectively) and some others might be A. Perkin (in litt.) reported it from the northern tip visited (Butynski unpubl., including information from of the East Usambara Mtns, seemingly close to G. Q. Luke). Museum specimens of Galagoides cocos zanzibaricus (Table IV). The form Galago moholi should be compared with the Diani galago further to cocos Heller, 1912, was described from Mazeras ensure that we are dealing with a single taxon. We pro- (Manzeras), relatively close to Diani (Table IV). visionally consider the Diani galago to be conspecific Groves (2001) recognized cocos as a mainland form with Galagoides cocos, under the vernacular name of Galagoides zanzibaricus. Vocalizations typical of Kenya coast galago.” (Grubb et al. 2003, p.1317). G. zanzibaricus have not been recorded in Kenya. Therefore, it seems highly likely that the Diani galago In other words, in considering the Kenya coast galago is a separate species, Galagoides cf. cocos. Never- once again a valid species, one important action remains: the theless it is important to confirm this. Vocalizations confirmation of cocos’‘ as the species name. This is necessary recorded from as near to the type locality of Gala- because the tape recordings used to describe the loud call for goides cocos as possible should be compared with the Kenya coast galago were made at Diani, Kenya (04°18′S, the voice of the Diani galago. The forest at the type 39°35′E) (Zimmermann 1990; Bearder et al. 1995), c.40 km locality (Kaya Mazeras) has been destroyed, but for- south of the type locality for G. cocos (i.e., Mazeras). There ests 5 km and 20 km distant (Kaya Mtswakana and are three rivers between these two sites that are potential barri-

68 Galagoides in Eastern Africa ers to dispersal (Maji ya Chumvi, Mambome, and Pemba/Cha T. Butynski and Y. de Jong heard about 100 G. cocos Shimba). A visit to Mazeras was, therefore, required to deter- incremental calls during one night at Kaya Chijembeni. As at mine whether the species-specific loud call of G. cocos at the other sites where C. cocos is similarly common, there was a type locality is the same as that of the dwarf galago at Diani. distinct ‘dusk chorus’ (c. 19:05–19:15h) of incremental calls, a Note that, in the above quotation, Grubb et al. (2003) much lower rate of incremental calls throughout the night, and assume that the holotype of G. cocos was collected in the for- a slight ‘dawn chorus’ (c. 05:45–05:55h) of incremental calls. est of ‘Kaya Mazeras’, which has since been destroyed. What The only other species of galago heard at Kaya Chi- Heller (1912) actually says about the type locality is: jembeni was O. garnettii. Thus, only two species of galagos “The Mazeras specimens were all obtained on were seen or heard in the Mazeras/Kaya Chijembeni/Rabai the brushy borders of the cocoa-palm groves. These area — G. cocos and O. garnettii. Y. de Jong and T. Butynski groves are the dominant feature in the landscape of returned to Kaya Chijembeni on 20 February 2006 to obtain the coast belt, and extend almost unbroken from the better (digital) recordings of the dwarf galago (Fig. 11). Again, sea beaches inland a distance of ten or fifteen miles. the only galagos heard were G. cocos and O. garnettii. They mark the tropical littoral zone more precisely Incremental calls identical to those of topotypical G. cocos than any other plant growth.” (Heller 1912, p.2). at Mazeras have been recorded (by A. Perkin, T. Butynski, Y. de Jong, S. Bearder, N. Cordiero, N. Svoboda, A. Kempson In short, Heller obtained the holotype (and nine other and S. Gregory) at several localities along the Kenya coast specimens) of G. cocos in an area of coconut palms (Cocos both north and south of Mazeras, as well as in the northern nucifera) and bushland, and not in forest. lowland coastal forests of the East Usambara Mountains in northeastern Tanzania. These calls were analyzed by A. Perkin Confirmation of the Name ‘cocos’ for the Kenya Coast and S. Bearder. Galago Visual examination of the type G. cocos and eight other adult specimens of G. cocos from Mazeras (housed at the T. Butynski and Y. de Jong visited the Mazeras area on United States National Museum, Washington, DC) reveal 10 –12 February 2004. The first night was spent searching that they are not phenotypically different from the three spec- for G. cocos in the Mazeras Botanical Garden (03°57′58′′S, imens of G. cocos at the National Museums of Kenya that 39°33′05′′E, 134 m a.s.l.) in Mazeras town. No dwarf galagos were collected along the Tana River (NMK 992), and on the were heard or seen, although the small-eared greater galago, Kenya coast at Gedi (NMK 5351) and Kipendi (no specimen (Otolemur garnettii), was common (about eight individuals number). One of the specimens at the National Museums of seen or heard). Kenya (MR14) was collected at Mrima Hill, Kenya, very near The second night was spent searching for G. cocos the border with Tanzania. This specimen has an intact penis, on the edge of Kaya Chijembeni (Rabai) (03°56′42′′S, the morphology of which helps to identify this population as 39°34′54′′E, 210 m a.s.l.), a relatively large coastal forest G. cocos (see below). located about 4 km northeast of Mazeras town, 17 km from We conclude that there is now no doubt that the type the Indian Ocean, and 40 km north of Diani. Coconut palms of G. cocos that Heller (1912) described from Mazeras is are an extremely common species there on the forest edge conspecific with the dwarf galago found along much of the and, with cashew nut (Anacardium occidentale) and mango coast of Kenya and into northern Tanzania. In short, the name (Mangifera indica), are scattered throughout the farmlands ‘cocos’ can correctly be used as the species name of the Kenya and bush lands that surround Kaya Chijembeni. Many dwarf coast galago. galagos were heard and seen at this site (both on the forest This clarification of the correct name for the Kenya coast edge and inside the forest), and tape recordings were acquired galago has at least three important implications for previ- of ‘incremental’ advertising calls and other vocalizations. To ous research conducted on G. cocos, G. zanzibaricus, and our ears, the advertising call recorded in Kaya Chijembeni G. granti. First, most of the field research that has been con- matched that recorded for the dwarf galago at Diani, as well ducted on the distribution, abundance, behavior, and ecology as the advertising call that T. Butynski has heard many hun- of ‘G. zanzibaricus’ was, in fact, conducted on G. cocos (for dreds of times (and recorded) at others sites on the coast of example, Harcourt 1984, 1986; Harcourt and Nash 1986a, Kenya (for example, Kilifi, Watamu, Gedi), and in forests 1986b; Harcourt and Bearder 1989). Second, the distinc- along the lower Tana River (02°30′S, 40°30′E), c.150 km to tive rolling calls of galagos on mainland Tanzania that led the north of Mazeras. Honess (1996) and Honess and Bearder (1996) to name this That the incremental advertising call recorded from form G. udzungwensis, belong to G. zanzibaricus (i.e., G. z. G. cocos at Kaya Chijembeni is the same as that recorded udzungwensis). Third, those researchers who obtained data from the dwarf galago at Diani is confirmed through compari- from specimens initially assigned to ‘G. zanzibaricus’ have sons of the sonograms and oscillograms of these calls (Figs. sometimes, unknowingly, combined data from two species 9 and 11). The advertising calls from these two sites have a (G. cocos and G. zanzibaricus), and, on occasion, from a third very similar incremental structure pattern, frequency range, species (G. granti). This is especially the case for specimens and call duration. collected from coastal Kenya, from coastal Tanzania south of

69 Butynski et al.

the East Usambara Mountains, and from Unguja Island, Zan- ies lead us to conclude that there is (1) too much intraspecific zibar. This means that (1) the results and conclusions of some variation and, especially, (2) too much interspecific overlap for previous studies of ‘G. zanzibaricus’ may need to be reviewed these characters to serve as diagnostic features. and reevaluated, and (2) that the providence of each and every The full range of phenotypic variation present in G. cocos, specimen labeled ‘G. zanzibaricus’ must be known in order G. z. zanzibaricus, G. z. udzungwensis, and G. granti remains to help ensure that the specimen is not, in fact, G. cocos or unknown. Although they need to be examined quantitatively, G. granti. and with sample sizes far larger than those currently available, the following phenotypic characters hold promise for distin- Morphological Differences Among G. cocos, G. zanzibari- guishing among G. cocos, G. z. zanzibaricus, G. z. udzungwen- cus and G. granti sis, and G. granti, especially when taken together:

Galagoides cocos, G. zanzibaricus, and G. granti are Muzzle patches among the most cryptic of primate species. That they are simi- G. cocos – patch on either side of muzzle dark, blackish, and prominent (Figs. 1 and 2). lar phenotypically and morphologically is demonstrated by the G. z. zanzibaricus – patch on either side of muzzle less dark, grayish, and less inability of some of the foremost primate taxonomists of their prominent. time to differentiate among them. For example, Schwarz (1931), G. z. udzungwensis – patch on either side of muzzle less dark, grayish, and in reference to G. senegalensis zanzibaricus, states (p. 56): less prominent (Figs. 3 and 4). G. granti – patch on either side of muzzle less dark, grayish, and less prominent “There can be no doubt that Heller’s cocos is (Fig. 5). identical with this race. The size, coloration, and the Ears large upper M³ are found both in the series at Berlin G. cocos – seldom longer than 38 mm, dusky behind. and the one of cocos studied by Heller and Hollister. G. z. zanzibaricus – seldom longer than 35 mm, dusky behind. G. z. udzungwensis – seldom longer than 33 mm, dusky behind. By the identification of the two the known range of G. granti – seldom shorter than 37 mm, blackish behind. Ears not only long zanzibaricus is considerably extended. There is no but also relatively broad (Fig. 5). difference between the island and coast specimens.” Dorsum G. cocos – hairs c.10 mm long, tipped buffy-brown (Figs. 1 and 6). G. z. zanzibaricus – hairs c.8 mm long, tipped cinnamon or rufous-cinnamon What is needed next is a detailed comparison of large (Fig. 7). numbers of G. cocos and G. zanzibaricus specimens to deter- G. z. udzungwensis – hairs c.9 mm long, tipped buffy-brown (Figs. 6 and 7). mine their morphological differences, coupled with ecologi- G. granti – hairs c.12 mm long, tipped buffy-brown with slight pinkish tint cal, behavioral, acoustic, and molecular studies. Other than (Figs. 5 and 6). the highly distinctive species-specific advertising call, one of Tail G. cocos – even length hairs over tail; hairs dense, c.14 mm long, soft. the differences noted thus far is that G. cocos is slightly larger Proximal c.25% of tail same color as dorsum (i.e., buffy-brown); distal than G. z. zanzibaricus. For example, the mean length of the c.33% dark buffy-brown in some (Mazeras) specimens, but same color as head+body for G. cocos from southeast Kenya and northeast dorsum in other specimens (Fig. 6). G. z. zanzibaricus – even length hairs over tail; hairs sparse, c.13 mm long, Tanzania is 158 mm (n = 46, range = 142–183 mm) (Appendix wiry, rufous-cinnamon, cinnamon, or dusky-cinnamon (highly variable). Tail I, Tables A and B), while the mean length of the head+body for darker/brighter cinnamon than dorsum, and either evenly colored or with G. z. zanzibaricus from Unguja Island, Zanzibar, is 143 mm gradual darkening to reddish or dusky toward tip (Fig. 7). G. z. udzungwensis – even length hairs over tail; hairs sparse, c.11 mm, wiry. (n = 11, range 125–150) (Appendix I, Table C). This size dif- Proximal c.75% of tail same color as dorsum (i.e., buffy-brown); distal c.25% ference extends to body weight; G. cocos has a mean body slightly darker brown or dusky. Some with tail tipped white (Figs. 4, 6, and weight of 144 g (n = 78, range = 117–172), whereas G. z. 7). zanzibaricus has a mean body weight of 127 g (n = 10, range G. granti – bushy, wider over distal c.80%; hairs dense, c.15 mm long, soft. Tail darker than dorsum with distal c.10 – 60% blackish-brown. Some with = 104–172). tail tipped white (Fig. 6). Of the two subspecies of G. zanzibaricus, it appears that the island form, G. z. zanzibaricus, is smaller than the main- land form, G. z. udzungwensis (Appendix I, Tables C and Of these four taxa, G. z. zanzibaricus and G. granti are D). Mean length of the head+body for G. z. udzungwensis is phenotypically the most distinctive. The dorsum of G. z. zan- 162 mm (n = 17, range = 139–180) and mean body weight is zibaricus is cinnamon and the tail has at least some rufous, 145 g (n = 6, range = 118–105). The data available suggest whereas the dorsum of the other three taxa is buffy-brown and that G. granti is larger than G. z. zanzibaricus, and very simi- all lack rufous in the tail (Fig. 7). The color of the dorsum of G. lar in size to G. z. udzungwensis and G. cocos (Appendix I, cocos, G. z. udzungwensis, and G. granti is virtually identical, Tables E, F and G). although there is a slight pinkish tint to the dorsum of G. granti Color of the nose stripe, chin, throat, cheeks and ven- when seen in good light (Fig. 6). trum, length of the nose stripe, and length of the tail relative When observed at close range in the field, G. granti is to length of the head+body, have all been proposed as useful distinguished from G. cocos, G. z. zanzibaricus, and G. z. for distinguishing among G. cocos, G. z. zanzibaricus, G. z. udzungwensis by its relatively large, broad, round, and black- udzungwensis, and G. granti (for example, Elliot 1913; Nash ish (behind) ears, and by the very full, bottlebrush-shaped tail, et al. 1989; Honess 1996; Groves 2001). However, our stud- which is blackish-brown over the distal part (Figs. 5 and 6).

70 Galagoides in Eastern Africa

The particularly large ears of G. granti have been noted previ- ess (1996), Honess and Bearder (1996), Anderson (2000), and ously (for example, Honess 1996; Masters and Bragg 2000). Perkin (in press). See sketches in Kingdon (1997). As a species, G. zanzibaricus is probably best distin- Vision plays an important role in the life histories of all guished phenotypically from G. cocos and G. granti by the galagos, perhaps especially for species recognition. Detailed relatively short, wiry, stiff hairs over the tail. This hair type study of the light and dark facial markings of these four taxa makes it relatively easy to see the skin of the tail through the may reveal that they are species-specific and, therefore, a pelage (Fig. 4). Galagoides cocos and G. granti have rela- useful diagnostic tool. This is not only a priority topic for tively long, soft, lax hairs over the tail. research related to the search for species-typical differences Of the four taxa considered here, G. cocos and G. z. among G. cocos, G. z. zanzibaricus, G. z. udzungwensis, and udzungwensis are, phenotypically, the most difficult to distin- G. granti, but also among the many other cryptic taxa within guish from one another (Fig. 6). The presence in G. cocos of the Galagidae (Bearder 1999; Bearder et al. 2006). a prominent dark, blackish patch on either side of the muzzle The differences noted here among G. cocos, G. zanzibari- is probably the best phenotypic character available for distin- cus, and G. granti in their species-specific advertising calls, guishing C. cocos and G. z. udzungwensis in the field (Figs. body measurements, and phenotypic characters are consistent 1–4) (A. Perkin pers. obs.). As already stated, however, all of with the species-level differences observed for other species the characters listed above are in need of detailed quantitative in the Galagidae (Honess 1996; Honess and Bearder 1996; study in order to determine their reliability, both in the field Masters and Bragg 2000; Masters and Brothers 2002). and in the museum. In contrast to the great similarity among G. cocos, G. zan- The penile morphology of G. cocos, G. zanzibaricus, zibaricus, and G. granti, these three species are readily distin- and G. granti is diagnostic (Fig. 12). For details, see Hon- guished from the other seven species of galagos with which one or all are sympatric or parapatric. These are O. garnet- tii, the thick-tailed (or large-eared) greater galago (Otolemur crassicaudatus), Somali lesser galago (Galago gallarum), northern lesser galago (Galago senegalensis), southern lesser galago (Galago moholi), mountain dwarf galago (Galagoides orinus), and Rondo dwarf galago (Galagoides rondoensis). The main morphological characters for distinguishing among G. cocos, G. gallarum, and G. senegalensis are summarized in Butynski and De Jong (2004).

Geographic Ranges of G. cocos, G. zanzibaricus and G. granti

Galagoides cocos occurs in evergreen forest all along the coastal strip (plain) of Kenya, south of the Tana River (Nash et al. 1989; Bearder et al. 2003; Grubb et al. 2003) southward to at least the Mgambo Forest Reserve in northern Tanzania at the north end of the East Usambara Mountains (A. Perkin unpubl. data) (Fig. 13). Galagoides cocos is reported to occur as far north as the Webi Shabeelle River in southern Somalia (Nash et al. 1989), but this needs confirmation. Thomas Butynski recorded the advertising call of a galago in the Ololua Forest, Nairobi, that S. Bearder identi- fied as that of the G. cocos. Ololua Forest is c.390 km inland from the coast of Kenya and, at 1,850 m a.s.l., well above the known altitudinal range for G. cocos elsewhere (0 –350 m). This record for Ololua Forest requires confirmation. Galagoides cocos and G. z. udzungwensis are parapatric or, perhaps, sympatric at a few sites c.2–8 km to the north of the East Usambara Mountains in the coastal strip of north- eastern Tanzania (Figs. 13 and 14) (A. Perkin in litt. in Grubb et al. 2003). Although there is a complex mosaic of habitat Figure 12. Schematic drawings showing the penile morphology of: 1 – Ga- types in this region, preliminary observations indicate that lagoides cocos; 2 – Galagoides granti; 3 – Galagoides zanzibaricus udzun- G. cocos is present in the dry mixed coastal forests and mixed gwensis. A – ventral view; B – dorsal view; C – lateral view (dorsum right). Scale is indicated on the right. See Perkin (in press) for details. Adapted from woodland of the northernmost forests of Tanzania’s coastal Perkin (in press). strip (for example, Bombo East I and Bombo East II For-

71 Butynski et al.

T a " n a

R

i Mwingi v e - 1° r

r e " iv R Nairobi i th Kitui Hola A G a Machakos la n a R iv e r

- 2°

Kiboko KENYA Garsen

- 3° Sabaki River Malindi

Arusha Gedi Voi

Mazeras - 4° Mombasa N Same A E TANZANIA C Diani O N IA D IN

LEGEND - 5° 0 G.50 cocos distribution100 Tanga Amani kilometers Pemba G. cocos locality Pa ng an i R iv " G. cocos unconfirmed er

0 50 100 Zanzibar kilometers (Unguja)

37° 38° 39° 40°

Figure 13. Approximate geographic distribution of the Kenya coast galago (Galagoides cocos). The shaded area extends out 5 km from the center of each locality point. The distribution of this species remains poorly known. Names of the sites plotted on this map, and the sources of these data, are available from Yvonne de Jong (e-mail: ).

72 Galagoides in Eastern Africa

KENYA - 3° Mombasa Same

Babati

P an g an i R i ve r

Kondoa Pemba - 4° Amani Tanga

Farkwa Handeni

Zanzibar (Unguja) - 5° Dodoma TANZANIA Wami River "

r e iv R u v Dar es Salaam Morogoro u R Kilosa INDIAN OCEAN INDIAN - 6°

G Mikumi Kisiju reat R uah a R i ver R ufiji Rive Iringa r Mafia Mang'ula - 7° Utete r e Ifakara iv R ji fi u R

Kihansi LEGEND Mahenge G. zanzibaricus distributionKilwa Kivinje G. zanzibaricus locality - 8° " G. zanzibaricus unconfirmed 0 50 100 kilometers

36° 37° 38° 39° 40°

Figure 14. Approximate geographic distributions of the Zanzibar galago (Galagoides zanzibaricus) The shaded area extends out 5 km from the center of each locality point. The distribution of this species remains poorly known. Names of the sites plotted on this map, and the sources of these data, are available from Yvonne de Jong (e-mail: ).

73 Butynski et al.

est Reserves), as well as in the more moist (but relatively The known southern limit for G. granti is the Limpopo tree-species and bird-species poor) groundwater forests and River in southern Mozambique. The western limit in the south- adjacent woodlands on the lower northern slopes of the East ern part of the range appears to be extreme eastern Zimbabwe Usambara Mountains (for example, Mgambo Forest Reserve) (Smithers and Lobão Tello 1976; Smithers and Wilson 1979; (A. Perkin and N. Cordeiro unpubl. data). In this region, G. z. Skinner and Smithers 1990). udzungwensis appears to be confined to the more moist (and In summary, present information indicates that G. cocos, relatively tree-species and bird-species rich) forest on the G. zanzibaricus, and G. granti are parapatric, or narrowly sym- eastern slopes of the East Usambara Mountains (for example, patric, species of the evergreen forests of the coastal strip of Kambai, Segoma, Manga, and Marimba forest reserves) (A. eastern Africa from northern Kenya (perhaps southern Somalia) Perkin unpubl. data). to extreme southern Mozambique and extreme eastern Zimba- Galagoides z. udzungwensis and G. granti appear to be bwe. Galagoides cocos is the northern species, G. zanzibaricus parapatric at the Kilombero-Rufiji River with G. z. udzun- is the central species, and G. granti is the southern species. gwensis reaching its southern limit on the north (left) bank (Fig. 14) and G. granti reaching its northern limit on the Conservation Status of G. cocos, G. zanzibaricus, and south (right) bank (Fig. 15) (Honess 1996; A. Perkin in litt. G. granti in Grubb et al. 2003). In addition, the geographic ranges of G. z. udzungwensis and G. granti either approach one another Galagoides cocos, G. zanzibaricus, and G. granti now or meet at the southern end of the Udzungwa Mountains in survive in highly fragmented, probably declining, populations south-central Tanzania. Here, G. granti is present at 1,500 to as a result of the extensive (>65%) loss of eastern Africa’s 1,800 m a.s.l. in the Lulanda Forest Reserve at the headwa- original coastal forest cover. More than 90% of the original ters of the Kilombero River (A. Perkin in litt. in Grubb et al. coastal forest of Kenya and Tanzania has either been destroyed 2003; A. Perkin unpubl. data), and G. z. udzungwensis is pres- or degraded (Burgess et al. 2004). Fortunately, all three species ent from 400 to 1,070 m a.s.l. at Kihanzi (which is c.24 km are able to persist in secondary forest and in mosaics of mixed to the east of Lulanda) (Honess 1996; Butynski et al. 1998; agriculture where some forest remains. N. Cordeiro pers. comm.). Kihanzi is the known southwestern As a recently resurrected species (Grubb et al. 2003), limit for G. z. udzungwensis, and Lulanda is the known north- G. cocos does not yet appear on the IUCN Red List. It is the western limit for G. granti. most abundant and widespread galago in the coastal forests of David Moyer and E. Mulungu (pers. comm.) tape Kenya, with densities of approximately 170 –180 individuals/ recorded the loud call of a Galagoides sp. at three sites in km² both at Gedi and Diani (Harcourt and Nash 1986a). Gala- extreme western Tanzania: Mbala Forest, Sitebe-Sifuta goides cocos is found at elevations from sea level to at least 210 Mountains (6º04′40′′S, 30º32′10′′E, 1,700 m a.s.l., 16 August m a.s.l. over the coastal zone of Kenya (T. Butynski and Y. de 2005), Mahale Mountains National Park at Mfitwa Mountain Jong unpubl. data), and to at least 350 m a.s.l. in the foothills (6º07′55′′S, 29º47′38′′E, 2,440 m a.s.l., 20 November 2005), of the East Usambara Mountains, Tanzania (A. Perkin unpubl. and at Pasagulu Mountain (6°03′47′′S, 29°45′14′′E, 1,500 m data). The information available indicates that G. cocos, when a.s.l., 6 February 2006). The species recorded may have been assessed for the IUCN Red List using the 2001 criteria (IUCN G. granti. If so, this extends its geographic range c.700 km to 2001), will be placed in the Least Concern category. the northwest (Fig. 15). G. zanzibaricus is listed as Lower Risk/Near Threatened Groves (2001) identified three specimens collected in sub- on the 2006 IUCN Red List (IUCN 2006), but assessed only montane forest at Bagilo (800 –1,000 m a.s.l.) in the Uluguru using the 1994 criteria (IUCN 1994). G. zanzibaricus is the Mountains (south of Morogoro, Fig. 15) as G. cf. granti (Grubb most abundant and widespread galago in the coastal forests et al. 2003). If these are G. granti, then the Uluguru Mountains of Tanzania, including ‘coastal’ forest sites located at least represent the northernmost site for this species, and the only 370 km inland (for example, Udzungwa Mountains) to c.1,100 known site for G. granti that is north of the Rufiji River. These m a.s.l. The density of G. zanzibaricus varies greatly from site three specimens had earlier been identified by Lawrence and to site. In the Udzungwa Mountains (for example, Matundu Washburn (1936) as G. senegalensis zanzibaricus. A. Perkin Forest Reserve), G. z. udzungwensis is estimated to occur at has since examined them at Harvard University’s Museum densities of more than 500 individuals/km² (Butynski et al. of Comparative Zoology (specimen numbers: 22450, 22449, 1998), whereas <100 individuals/km² occur at many other sites 22451) and also ascribes them to G. zanzibaricus. In 1993, (T. Butynski and A. Perkin pers. obs.). Although G. z. zanzi- P. E. Honess and S. K. Bearder (Honess 1996) visited the forest baricus is confined to Unguja Island, it is widespread over the at Bagilo but did not findG. granti there — only G. orinus was eastern and southern parts of the island (Lumsden and Masters present. Unfortunately, there is no longer any forest at Bagilo 2001), and is common in at least some places (e.g., >200 indi- (Perkin 2000; Doggart et al. 2004) but A. Perkin (unpubl. data) viduals/km² in Jozani–Chwaka Bay National Park (T. Butynski found G. zanzibaricus at 900 m a.s.l. on the edge of the Ulu- and Y. de Jong pers. obs). The information available indicates guru North Forest Reserve (c.2 km west of Bagilo Village) and that G. zanzibaricus, when reassessed for the IUCN Red List, up to 700 m a.s.l. in the Uluguru Mountains. should be placed in the ‘Least Concern’ category.

74 Galagoides in Eastern Africa

Mombasa Same - 4° Kigoma Tanga Tabora Pemba

Zanzibar (Unguja) - 6° Kabala L ? ? a k Dodoma e Dar es Salaam T a TANZANIA n g Morogoro a n Ru y fiji ik Ri a ver Mafia Iringa Mang'ula - 8° Lulanda Mahenge DRC Mbeya

Lindi

- 10° i

Kasama w

a l

Ma Songea

e k

Lubumbashi a Ruvuma

L River I

W - 12°

ZAMBIA A

L A

Ndola M Chipata MOZAMBIQUE

Lilongwe - 14° Cuamba Nacala N Songo " A E Lusaka Blantyre C O - 16° N

A I Z a D m b N e I z i Quelimane R iv e Harare r - 18° Tambarara ZIMBABWE Chimoio Beira - 20° Bulawago Masvingo

Francistown LEGEND - 22° G. granti distribution L im p o p o G. granti locality R Pembe iv e r Coguno " G. granti unconfirmed Pietersburg - 24°

SOUTH AFRICA 0 200 400

Maputo kilometers - 26° 28° 30° 32° 34° 36° 38° 40° 42° 44°

Figure 15. Approximate geographic distributions of the Mozambique galago (Galagoides granti). The shaded area extends out 20 km from the center of each locality point. The distribution of this species remains poorly known. Names of the sites plotted on this map, and the sources of these data, are available from Yvonne de Jong (e-mail: ).

75 Butynski et al.

The two subspecies, G. z. zanzibaricus and G. z. udzun- Literature Cited gwensis, have not as yet been assessed for the IUCN Red List. While G. z. udzungwensis is a widespread subspecies that Allen, G. M. 1939. A checklist of African mammals. Bull. will likely be assessed as Least Concern, G. z. zanzibaricus Mus. Comp. Zool. Harvard College 83: 1–763. has a far smaller ‘extent of occurrence’, being endemic to Allen, G. M. and A. Loveridge. 1927. Mammals from the Unguja Island, Zanzibar (c.2,000 km²) where rates of habi- Uluguru and Usambara Mountains, Tanganyika Territory. tat degradation, loss, and fragmentation are particularly high. Proc. Boston Soc. Nat. Hist. 38: 425– 426. As such, G. z. zanzibaricus is expected to be assessed as an Ambrose, L. 1999. Species diversity in West and Central Afri- Endangered taxon. can galagos (Primates, Galagonidae): The use of acous- Galagoides granti, assessed under the 1994 criteria tic analysis. PhD thesis, Oxford Brookes University, (IUCN 1994), is listed as Data Deficient in the 2006 IUCN Oxford. Red List (IUCN 2006). Galagoides granti is present in coastal Ambrose, L. 2003. Three acoustic forms of Allen’s galagos and submontane evergreen forest, gallery forest, and species- (Primates: Galagonidae) in the Central African region. rich woodlands, including some hilly miombo woodlands Primates 44: 25–39. (e.g., Mahenge foothills). In the southern part of its range, G. Anderson, M. J. 1999. The use of hand morphology in the granti is present between the coast and about 200 km inland taxonomy of galagos. Primates 40: 469– 478. (i.e., eastern Zimbabwe) up to 360 m a.s.l. (Smithers and Anderson, M. J. 2000. Penile morphology and classification Wilson 1979; Skinner and Smithers 1990). According to the of bush babies (subfamily Galagoninae). Int. J. Primatol. specimen tags, C. H. B. Grant collected this species up to 400 21: 815–836. m a.s.l. at Tambarara in central Mozambique. In the northern Bearder, S. K. 1999. Physical and social diversity among noc- part of its range, G. granti occurs from the coast to at least turnal primates: A new view based on long-term research. 310 km inland (i.e., Lulanda, Tanzania) up to at least 1,800 m Primates 40: 267–282. a.s.l. Galagoides granti occurs over a much greater range of Bearder, S. K., P. E. Honess and L. Ambrose. 1995. Species habitat types, altitudes, and climates than does G. cocos or G. diversity among galagos with special reference to mate zanzibaricus. There is now enough information available for recognition. In: Creatures of the Dark: The Nocturnal an IUCN Red List assessment of this species. The available Prosimians, L. Alterman, G. A. Doyle and M. K. Izard data indicate that G. granti, once reassessed, will be placed in (eds.), pp.331–352. Plenum Press, New York. the Least Concern category. Galagoides cocos, G. zanzibari- Bearder, S., L. Ambrose, C. Harcourt, P. Honess, A. Perkin, cus, and G. granti are all currently CITES Appendix II species E. Pimley, S. Pullen and N. Svoboda. 2003. Species-typi- (http://www.cites.org). cal patterns of infant contact, sleeping site use and social cohesion among nocturnal primates in Africa. Folia Pri- Acknowledgments matol. 74: 337–354. Bearder, S. K., K. A. I. Nekaris and D. J. Curtis. 2006. A re- We thank David Moyer and Esteban Sarmiento for their evaluation of the role of vision in the activity and com- unpublished data, the curatorial staff at the United States munication of nocturnal primates. Folia Primatol. 77: National Museum, British Museum of Natural History, and 50 –71. American Museum of Natural History for providing access Burgess, N, J. Salehe, N. Doggart, G. P. Clark, I. Gordon, to their collections; Harald Schuetz for the photograph in Fig- P. Sumbi and A. Rodgers. 2004. Coastal forests of east- ure 1; and Nike Doggart for the photograph in Figure 3. We ern Africa. In: Hotspots Revisited, R. A. Mittermeier, P. appreciate the support given to our research on primates in Robles Gil, M. Hoffmann, J. Pilgrim, T. Brooks, C. G. eastern Africa by the Margot Marsh Biodiversity Foundation, Mittermeier, J. Lamoreux and G. A. B. da Fonseca (eds.), Conservation International, Zoo Atlanta, the National Muse- pp.230 –239. CEMEX, Agrupación Serra Madre, SC, ums of Kenya, the IUCN Eastern Africa Regional Office, the México. Kenya Institute of Primate Research, the Critical Ecosystem Butynski, T. M., C. L. Ehardt and T. T. Struhsaker. 1998. Partnership Fund, the Primate Action Fund, Primate Conser- Notes on two dwarf galagos (Galagoides udzungwensis vation Inc., the Leverhulme Trust, Oxford Brookes University, and Galagoides orinus) in the Udzungwa Mountains, WWF-Tanzania, the Tanzanian Commission for Science and Tanzania. Primate Conserv. (18): 69–75. Technology, Tanzania Forestry Division, Tanzania National Butynski, T. M. and Y. A. de Jong. 2004. Natural history of the Parks, Kenya Wildlife Service, and Kenya Ministry of Edu- Somali lesser galago (Galago gallarum). J. E. Afr. Nat. cation, Science and Technology. Norbert Cordeiro, Nadine Hist. 93: 24–38. Svoboda, Alex Kempson, and Stephen Gregory provided tape Courtenay, D. O. and S. K. Bearder. 1989. The taxonomic sta- recordings of the advertising calls of G. cocos. Norbert Cor- tus and distribution of bushbabies in Malawi with empha- deiro assisted with vegetation descriptions and unpublished sis on the significance of vocalizations. Int. J. Primatol. data. Julie Anderson and the Colobus Trust helped with sur- 10: 17–34. veys funded by the Born Free Foundation. DelPero, M., J. C. Masters, D. Zuccon, P. Cervella, S. Crov- ella and G. Ardito. 2000. Mitochondrial sequences as

76 Galagoides in Eastern Africa

indicators of generic classification in bush babies. Int. J. Honess, P. E. 1996. Speciation among galagos (Primates: Primatol. 21: 889–904. Galagidae) in Tanzanian forests. PhD Thesis, Oxford Doggart, N., J. Lovett, B. Mhoro, J. Kiure, A. Perkin and N. D. Brookes University, Oxford, UK. Burgess. 2004. Biodiversity surveys in the forest reserves Honess, P. E. and S. K. Bearder. 1996. Descriptions of the of the Uluguru Mountains: Descriptions of the biodiver- dwarf galago species of Tanzania. African Primates 2: sity of individual forest reserves. Report, Tanzania Forest 75–79. Conservation Group, Dar es Salaam, Tanzania. IUCN. 1994. IUCN Red List Categories. IUCN Species Sur- Elliot, D.G. 1913. A Review of the Primates. Vol. 3. American vival Commission, Gland, Switzerland. Museum of Natural History, New York. IUCN. 2001. IUCN Red List Categories and Criteria: Version Groves, C. P. 1977. Taxonomy and phylogeny of prosim- 3.1. IUCN Species Survival Commission, Gland, Swit- ians. In: Prosimian Anatomy, Biochemistry and Evolu- zerland. tion, R. D. Martin, G. A. Doyle and A. C. Walker (eds), IUCN. 2006. 2006 IUCN Red List of Threatened Species. pp.449– 473. Duckworth, London. Website: www.iucnredlist.org. Accessed 15 May 2006. Groves, C. P. 1993. Order Primates. In: Mammal Species of Jenkins, P. D. 1987. Catalogue of Primates in the British the World: A Taxonomic and Geographic Reference, 2nd Museum (Natural History) and Elsewhere in the British Ed., D. E. Wilson and D. M. Reeder (eds.), pp.243 –277. Isles. Part IV. Suborder Strepsirrhini, Including the Sub- Smithsonian Institution Press, Washington, DC. fossil Madagascan Lemurs and Family Tarsiidae. British Groves, C. P. 2001. Primate Taxonomy. Smithsonian Institu- Museum (Natural History), London. tion Press, Washington, DC. Kingdon, J. 1971. East African Mammals. Vol. 1. Academic Groves, C. P. 2005. Order Primates. In: Mammal Species Press, London. of the World: A Taxonomic and Geographic Refer- Kingdon, J. 1997. The Kingdon Field Guide to African Mam- ence, 3rd edi. D. E. Wilson and D. M. Reeder (eds.), mals. Academic Press, London. pp.111–184.. Johns Hopkins University Press, Baltimore, Lawrence, B. and S. L. Washburn. 1936. A new eastern race Maryland. of Galago demidovii. Occ. Pap. Boston Soc. Nat. Hist. 8: Grubb, P., T. M. Butynski, J. F. Oates, S. K. Bearder, T. R. 255–266. Disotell, C. P. Groves and T. T. Struhsaker. 2003. Assess- Lumsden, W. H. R. and J. C. Masters. 2001. Galago (Galago- ment of the diversity of African primates. Int. J. Primatol. nidae) collections in East Africa (1953 –1955): Ecology 24: 1301–1357. of the study areas. African Primates 5: 37– 42. Harcourt, C. S. 1984. The behaviour and ecology of galagos Masters, J. C. 1991. Loud calls of Galago crassicaudatus and in Kenyan coastal forest. PhD thesis, University of Cam- G. garnettii and their relation to habitat structure. Pri- bridge, Cambridge, UK. mates 32: 149–174. Harcourt, C. S. 1986. Galago zanzibaricus: Birth seasonal- Masters, J. C. 1998. Speciation in the lesser galagos. Folia ity, litter size and perinatal behaviour of females. J. Zool., Primatol. 69(Suppl.1): 357–370. Lond. 210: 451– 457. Masters, J. C. and N. P. Bragg. 2000. Morphological corre- Harcourt, C. S. and S. K. Bearder. 1989. A comparison of lates of speciation in bush babies. Int. J. Primatol. 21: Galago moholi in South Africa with Galago zanzibaricus 793 –813. in Kenya. Int. J. Primatol. 10: 35– 45. Masters, J. C. and D. J. Brothers. 2002. Lack of congruence Harcourt, C. S. and L. T. Nash. 1986a. Species differences in between morphological and molecular data in recon- substrate use and diet between sympatric galagos in two structing the phylogeny of the Galagonidae. Am. J. Phys. Kenyan coastal forests. Primates 27: 41–52. Anthropol. 117: 79–93. Harcourt, C. S. and L. T. Nash 1986b. Social organization of Nash, L. T., S. K. Bearder and T. R. Olson. 1989. Synopsis galagos in Kenyan coastal forests. 1. Galago zanzibari- of galago species characteristics. Int. J. Primatol. 10: cus. Am. J. Primatol. 10: 339–355. 57–80. Heller, E. 1912. New races of insectivores, bats, and lemurs Perkin, A. W. 2000. The taxonomic status of bushbabies (gala- from British East Africa. Smithson. Misc. Coll. 60: 1–13. gos) in the Uluguru Mountains. Report for the Wildlife Hill, W. C. O. 1953. Primates: Comparative Anatomy and Conservation Society of Tanzania, Dar es Salaam. Web- Taxonomy. I. Strepsirhini. University of Edinburgh Press, page: www.africanconservation.com/uluguru. Accessed 8 Edinburgh, UK. May 2006. Hill, W. C. O. and J. Meester. 1977. Suborder Prosimii, Infra- Perkin, A. W. In press. Comparative penile morphology of order Lorisformes. In: The Mammals of Africa: An Iden- East African galagos of the genus Galagoides (Family tification Manual, Part 3.2., J. Meester and H. W. Setzer Galagidae): Implications for taxonomy. Amer. J. Prima- (eds.). Smithsonian Institution Press, Washington, DC. tol. Hollister, N. 1924. East African mammals in the United States Perkin, A. W., S. K. Bearder, T. M. Butynski, B. Agwanda National Museum. Part III. Primates, Artiodactyla, Pro- and B. Bytebier. 2002. The Taita mountain dwarf galago boscidea, and Hyracoidea. Bull. U. S. Natl. Mus. 99: 1– Galagoides sp.: A new primate for Kenya. J. E. Afr. Nat. 164. Hist. 91: 1–13.

77 Butynski et al.

Table B. Measurements for live specimens of Galagoides cocos. These data Schwarz, E. 1931. On the African long-tailed lemurs or gala- derive from the following two sources: (1) Harcourt and Nash (1986b) for G. gos. Ann. Mag. Nat. Hist. 7: 41– 66. cocos at Gedi (c.100 km northeast of Mombasa) and Diani (30 km south of Skinner, J. D. and R. H. N. Smithers. 1990. The Mammals of Mombasa) on the coast of Kenya; (2) A. Perkin (unpubl. data) for G. cocos at the Southern African Subregion. University of Pretoria, Arabuko-Sokoke Forest (c.100 km northeast of Mombasa, n = 2), Diani (n = 3), and East Usambara Mountains (c.150 km southwest of Mombasa (n = 1). Pretoria. Because adult male and adult female body linear measurements for G. cocos Smithers, R. H. N. and J. L. P. Lobão Tello. 1976. Check list are not significantly different, the data for males and females are combined in and atlas of the mammals of Mocambique. Museum Mem- this sample (Harcourt and Nash 1986b). Because body weights of adult male oir, Natl. Museums and Monuments of Rhodesia No. 8. and adult (non-pregnant) female G. cocos are significantly different (Harcourt and Nash 1986b), the body weight data are presented separately for each sex. Smithers, R. H. N. and V. J. Wilson. 1979. Check list and atlas Sample size of the mammals of Zimbabwe Rhodesia. Museum Mem- Measurement Mean Range (n) oir, Natl. Museums and Monuments of Rhodesia No. 9. Zimmermann, E. 1990. Differentiation of vocalisations in bush- Head + body length 159 mm 142–183 mm 38 Tail length 212 mm 182–230 mm 30 babies (Galaginae, Prosimiae, Primates) and the signifi- Hindfoot length 54 mm 40 –57 mm 49 cance for assessing phylogenetic relationships. Z. Zoolo- Ear length 30 mm 23 – 40 mm 32 gische Systematik und Evolutionsforschung 28: 217–239. Body weight (males) 150 g 135–167 g 36 Zimmermann, E., S. K. Bearder, G. A. Doyle and A. B. Ander- Body weight (females) 138 g 117–172 g 42 sson. 1988. Variations in vocal patterns of lesser bushba- bies (Galago senegalensis and Galago moholi) and their implications for taxonomic relationships. Folia Primatol. Table C. Combined measurements for male and female Galagoides zanzibari- cus zanzibaricus from Unguja Island, Zanzibar, Tanzania, the type locality for 51: 87–105. G. zanzibaricus. Measurements taken from one specimen captured and released by A. Perkin and from the tags of 10 specimens housed at the British Muse- Authors’ addresses: um of Natural History. Nine of these collected by W. H. R. Lumsden (speci- Thomas M. Butynski, Eastern Africa Regional Program, men numbers 1964.971, 1964.972, 1964.974, 1964.975, 1964.977, 1964.978, 1964.979, 1964.980, and 1964.981) and one obtained by an unknown collector Conservation International, c/o IUCN EARO, P.O. Box 68200, (specimen number: 1955.331). 00200 Nairobi, Kenya. Sample size Measurement Mean SD Range Yvonne A. de Jong, Kenya Institute of Primate Research, P.O. (n) Box 38, Ol Kalou 20303, Kenya. Head+body length 143 mm 8 125–150 mm 11 Andrew W. Perkin and Simon K. Bearder, Nocturnal Pri- Tail length 214 mm 12 198–235 mm 11 mate Research Group, Anthropology Department, School of Hindfoot length 56 mm 3 51–59 mm 11 Social Sciences and Law, Oxford Brookes University, Oxford Ear length 32 mm 2 30 –35 mm 11 OX3 0BP, UK. Body weight 127 g 20 104–172 g 10 Paul E. Honess, Department of Veterinary Services, Univer- sity of Oxford, Oxford OX1 3PT, UK. Table D. Combined measurements for male and female Galagoides zanzibari- cus udzungwensis obtained from the following sites in Tanzania; two from Received for publication: February 2006 Matundu Forest Reserve (Honess 1996); two from Kissarawe (housed at the Revised: May 2006 British Museum of Natural History); two from Pugu Forest Reserve, three from Pande Game Reserve (A. Perkin, unpubl. data); four from Bagilo, Uluguru Appendix I Mountains; and four from Amboni, near Tanga. These last eight specimens were collected by A. Loveridge and are housed at the Museum of Comparative Body measurements for adults of the Kenya coast galago (Gala- Zoology, Harvard University. goides cocos), the Zanzibar galago (Galagoides zanzibaricus), and the Sample size Measurement Mean SD Range Mozambique (galago (Galagoides granti). All measurements are either (n) known to be, or believed to be, from living or fresh adult specimens. Head+body length 162 mm 11 139–180 mm 17 Tail length 222 mm 16 202–270 mm 17 Table A. Measurements for eight Galagoides cocos collected at Mazeras, Ke- Hindfoot length 58 mm 6 nya, the type locality for G. cocos (Hollister 1924). All measurements taken 50 –70 mm 17 from the tags of specimens collected by E. Heller and housed at the United Ear length 31 mm 3 25–37 mm 17 States National Museum (specimen numbers: 181810, 184218, 184219, Body weight 145 g 27 118–195 g 6 184220, 184221, 184222, 184223, and 184225). Because adult male and adult female body linear measurements for G. cocos are not significantly different (Harcourt and Nash 1986b), the data for males and females are combined in this sample. Sample Measurement Mean SD Range size (n) Head+body length 155 mm 6 145–165 mm 8 Tail length 213 mm 11 200 –230 mm 8 Hindfoot length 57 mm 3 53 – 60 mm 7 Ear length 35 mm 2 33 –38 mm 7

78 Galagoides in Eastern Africa

Table E. Combined measurements for male and female Galagoides granti collected at Coguno (type locality) and Tambarara, Mozambique, by C. H. B. Grant during the Rudd Expedition. Coguno is the type locality. Measurements taken from specimen tags. All 12 specimens housed at the British Museum of Natural History (specimen numbers: 906.11.8.5, 1906.11.8.6, 1906.11.8.7, 1906.11.8.8, 1906.11.8.9, 1906.11.8.10, 1908.1.1.12, 1908.1.1.13, 1908.1.1.14, 1908.1.1.15, 1908.1.1.16, and 1908.1.1.129). Sample size Measurement Mean SD Range (n) Head+body length 153 mm 6 140 –160 mm 12 Tail length 230 mm 6 216 –237 mm 12 Hindfoot length 58 mm 3 54– 63 mm 12 Ear length 38 mm 2 36 – 43 mm 12

Table F. Combined measurements for male and female Galagoides granti from eastern Zimbabwe (Smithers & Wilson 1979). Sample size Measurement Mean Range (n) Head+body length 162 mm Not provided 10 Tail length 232 mm 214–254 mm 10 Hindfoot length 62 mm 59– 63 mm 10 Ear length 40 mm 38– 41 mm 9 Weight 165 g 139–178 mm 6

Table G. Combined measurements for two male and one female Galagoides granti from Tanzania, two from Kichi Hills Forest Reserve, and one from Lu- lunda, Udzungwa Mountains (A. Perkin unpubl. data). Sample size Measurement Mean SD Range (n) Head+body length 164 mm 15 154–181 mm 3 Tail length 214 mm 7 208–222 mm 3 Hindfoot length 58 mm 1 58–59 mm 3 Ear length 38 mm 1 37–38 mm 2 Body weight 136 g 25 110 –160 g 3

79 TsavoA new pig for

By James Culverwell, Jim Feely, be Africa’s least-known, non-forest, Uganda and Kenya). Their find- Sheila Bell-Cross, Yvonne A. de large mammal. The morphological ings were summarized in 2002 by Jong and Thomas M. Butynski differences between the two species Boy in Swara 25—1: 20-21. The of warthog are described by Grubb authors found the desert warthog here are only two spe- (1993) and by d’Huart & Grubb to occur from Puntland (northern cies of warthog, and (2005). The most diagnostic and Somalia) southwestward through both occur in Kenya; noticeable characteristics for both Somalia and southeastern Ethio- the common warthog species are summarized in Box 1. pia to central and eastern Kenya. Phacochoerus afri- d’Huart & Grubb (2001) The only sites shown on the map canus and the desert compiled a map depicting the for Kenya were Moyale, El Wak, warthogT Phacochoerus aethiopicus. geographical range of both the Merelle (‘midway between Arch- Little is known about the natural common warthog and the desert er’s Post and Mt Marsabit’), and history of the desert warthog—in- warthog in the Horn of Africa (Er- Mkokoni (in the Kiunga Ma- deed, the desert warthog might well itrea, Djibouti, Somalia, Ethiopia, rine Reserve). The southernmost

How to most readily differentiate the desert warthog from the common warthog in the field (d’Huart & Grubb 2005). See Figures 1 and 2. 1. Warts—Adult desert warthog has a hook- shaped wart under the eye. Adult common warthog has a cone-shaped wart under the eye. 2. Ears—Ear tips of desert warthog are bent backwards. Ear tips of common warthog are erect. 3. Suborbital area—Suborbital area (ie, under the eye) appears swollen in adult desert warthog. In adult common warthog this swelling is absent. 4. Head—Head of desert warthog is ‘egg- shaped’ when viewed from the front. Head of common warthog is slightly ‘diabolo- shaped’ when viewed from the front.

Figure 1. Adult male common warthog Phacochoerus africanus on the open plains of the Laikipia Plateau, central Kenya. Note the cone shaped warts, pointed ears, and the diabolo- shaped head. Photograph by T. M. Butynski & Y. A. de Jong.

Figure 2. Adult male desert warthog Phacochoerus aethiopicus in medium dense shrub in Tsavo West National Park, southeast Kenya. Note the flipped-back ears, hooked warts, the broad, egg-shaped head, and the swollen suborbital area (area under the eyes). Photograph by T. M. Butynski & Y. A. de Jong.

50 SWARA July – September 2008 FINDINGS

Figure 3. Desert warthog warthog sympatric? How far south for more photographs to be taken. Phacochoerus aethiopicus and does the desert warthog occur? What struck JC, JF and SBC im- common warthog Phacochoerus On 7 June 2007, JC, JF and mediately during these encounters africanus localities in Tsavo East SBC traversed Tsavo East Na- was the overall shape of the ani- National Park and Tsavo West tional Park. Briefed a few days mals’ heads, the hooked warts, and National Park, southeast Kenya. earlier about desert warthog char- the flipped-back tip of the ears (see acteristics by TMB and YDJ, they Box 1), all of which are diagnos- record for Kenya was from the travelled through the high-density tic, easily-seen field characters of Kiunga Marine Reserve, about 60 tourism areas south of the Voi the desert warthog. During this kilometres northeast of Lamu. All River, far to the southwest of the trip, no common warthogs were records were from north of the nearest confirmed desert warthog seen. All photographs were sent to Ewaso Ng’iro River and east of the locations. Late that afternoon they experts for confirmation. Some of Tana River. They speculated that were startled to find two adult these, together with some of our the two species would be found desert warthogs in low scrub on other warthog photographs, are to overlap in some parts of their the edge of the Dika Plains, just available on a digital map at www. range, but found no evidence for 13 kilometres north of Buchuma tinyurl.com/warthogMap. this; their data show the shortest Gate, and managed to take a few distance between common warth- photographs before the warthogs n July 2007, JC returned briefly og and desert warthog records as turned tail. They had just recorded to Tsavo East National Park 25 kilometres (in Puntland). a new large mammal species for Ifor two days. He observed two During a primate survey in Tsavo East National Park, and a sounders of desert warthogs—each 2005, TMB and YDJ encountered major range extension of over 300 of four animals—in open shrub- two solitary desert warthogs, 15 kilometres for the desert warthog! land north and south of the Voi and 80 kilometres southwest of The next day they returned River in the vicinity of Satao Safari Garissa respectively, in medium to the area and located four more Camp. Once again, no common dense acacia bushland. These en- desert warthogs only nine kilo- warthogs were seen. counters are important, as they are metres from the Kenya Wildlife In October 2007, JC visited the first records of desert warthog Service headquarters at Voi; one Tsavo West National Park for two west of the Tana River. Two ques- adult and three subadults stood days, where desert warthogs were tions remained, however. Are their ground long enough for posi- found to be fairly common around the desert warthog and common tive identification to be made and Severin Safari Camp and Kitani

SWARA July – September 2008 51 The East African Wild Life Society’s

Safari Lodge, just to the north areas where desert warthogs were References of the Tsavo River. Three desert seen during the earlier trip by JC). • Boy, G. 2002. The whole hog. 2009 CALENDARS warthogs were seen feeding in an About six kilometres of Finch Hat- Swara 25 (1): 20-21. open area along a major tourism ton’s Camp, in low bush on the • D’Huart, J. & Grubb, P. 2001. Dis- tribution of the common warthog circuit, permitting lengthy obser- edge of riverine forest, one group (Phacochoerus africanus) and the vation. One of these animals was of desert warthogs (one adult male desert warthog (Phacochoerus a subadult male. Subadult desert and three subadults) was encoun- aethiopicus). African Journal of warthogs, like adults, have ears tered only 150 metres from six Ecology 39: 156-169. that are flipped-back at the tips, common warthogs. This sight- • Grubb, P. 1993. The Afrotropical and broader-based genal warts, al- ing is not only the farthest range suids Phacochoerus, Hylochoerus, lowing them to be differentiated in extension for desert warthog (390 and Potamochoerus: taxonomy the field from common warthogs kilometres) from Mikoni, the and description. In: Pigs, Peccaries and Hippos. Status Survey and despite them lacking the hooked most southern record of d’Huart Conservation Action Plan (ed. W. warts of adult male desert warth- & Grubb, but it is also the first I. R. Oliver). IUCN/SSC, Gland, ogs. During this visit JC also found evidence that the two species of Switzerland common warthogs in Tsavo West warthogs are sympatric. • iucn.org/themes/ssc/sgs/pphsg/ National Park only a few hundred Finding desert warthog in Contents.htm metres from where he observed Tsavo, and finding them sympatric • D’Huart, J. & Grubb, P. 2005. A desert warthogs. with common warthog, is excit- photographic guide to the differenc- In April, 2008, TMB and YDJ ing news for the species and for es between the common warthog (Phacochoerus africanus) and the visited Tsavo West National Park Kenya. Tsavo West National Park desert warthog (Ph. aethiopicus). for three days, and encountered and Tsavo East National Park can Suiform Soundings 5 (2): 4-8 common warthogs around Kilagu- add one more species to their list • iucn.org/themes/ssc/sgs/pphsg/ ni Serena Safari Lodge, Komboyo of large mammals. What further Suiform%20soundings/Newslet- Camp, Kitani Safari Lodge and surprises can we expect from this ter%205(2)pdf Severin Safari Camp (including long-overlooked pig?

Have you seen warthogs in Africa?

To better understand the distribution of Africa’s warthogs, we would like to know if you have seen desert warthogs Desk Calendar - retail Wall Calendar - retail or common warthogs in Eritrea, Ethiopia, Member Non-Member Extra fee for Member Non-Member Extra fee for Djibouti, Somalia, Uganda, Kenya or Retail Prices Retail Prices Postage Retail Prices Retail Prices Postage Ksh 250 Ksh 270 - Ksh 300 Ksh 350 - Tanzania. Details and/or a photograph of US$ 5.00 US$ 5.80 US$ 3.40 US$ 5.00 US$ 5.80 US$ 6.30 UK£ 3.0 UK£ 3.20 your record would be highly appreciated. UK£ 2.50 UK£ 3.00 UK£ 2.00 UK£ 3.40 € 4.00 € 4.60 € 4.90 € 3.20 € 3.50 € 2.70 The following information is most important • date of sighting March 2009 species Su Mo Tu We Fr Sa • 6 7 name of nearest village or town 1 2 3 4 5 • 8 9 UG 10 11 12 13 14 • latitude and longitude 15 16 17 18 19 20 21 • elevation 22 23 24 25 26 27 28 • habitat 29 30 31 FEBRUARY 2009 APRIL 2009 We Fr Sa Su Mo Tu We Fr Sa Su Mo Tu 1 2 3 4 1 2 3 4 5 6 7 8 9 10 11 8 9 10 11 12 13 14 5 6 7 FEBRUAR 15 16 17 18 Y 2009 15 16 17 18 19 20 21 12 13 14 Please send your information and/or 25 SUNDAY MONDAY 28 19 20 21 22 23 24 TUESDAY WEDNESDAY 22 23 24 25 26 27 THURSDAY FRIDAYSATURDAY 26 27 28 29 30 1 2 3 4 5 6 photograph to: 7 APRIL 2009 Elephants in the Mara headed Y THURSDAY FRIDAY SATURDAY Yvonne de Jong for a sun downer of their own SUNDAY MONDAY TUESDAY WEDNESDA 8 MAY 2009 9 Th Fr Sa Ja 10 11 12 Su Mo Tu We 3 4 nu 13 1 2 2 1 ary 2009 14 31 3 4 5 6 7 8 9 Su 10 11 12 13 14 15 16 Mo 20 21 22 23 Tu We 17 18 19 PO Box 149 26 27 28 29 30 Fr Sa 24 25 15 16 17 Good Friday Nanyuki 10400, Kenya OVERPRINT AREA 1 2 3 18 19 20 21 5 6 7 8 9 10 11 4 5 6 7 8 9 10 [email protected] 22 23 24 25 26 27 Easter Monday 16 17 18 11 TZ 28 12 13 14 15 12 13 14 15 16 17 MARCH 2009 Thank you! Su Mo Tu 18 19 We Th Fr Sa 20 1 2 3 21 22 4 5 6 7 23 8 9 10 11 24 12 13 14 25 15 24 16 17 18 23 19 20 21 20 21 22 22 19 23 24 25 26 27 28 Your help is much appreciated! 25 26 UG 27 28 29 30 29 30 31 31 These Giraffe in the mag nificent evening Mara light FEBRU ARY 2009 DECEMBER 2008 TZ 28 29 30 Su Mo Tu We 26 27 Fr Sa Su Mo Tu We Fr Sa 1 2 3 4 5 6 7 1 2 3 4 5 6 8 9 10 11 Zebra in flight on the Mara Plains 12 13 14 7 8 9 10 11 12 13 15 16 17 18 19 20 21 14 15 16 17 18 19 20 22 23 24 25 26 27 28 21 22 23 24 25 26 27 52 SWARA July – September 2008 28 29 30 31 OVERPRINT AREA

Flamingo at sunset in Lake Nakuru National Park

Papers and

communications

Desert warthog Phacochoerus aethiopicus found in Tsavo East ational Park and Tsavo West ational Park, southern Kenya

Yvonne A. de Jong¹, James Culverwell² and Thomas M. Butynski³

¹P.O. Box 149, 10400 6anyuki, Kenya, [email protected] ²8 Manga Gardens, Kitisuru Rd, 6airobi, Kenya, [email protected] ³P.O. Box 149, 10400 6anyuki, Kenya, [email protected]

Both species of warthog, the common warthog Phacochoerus africanus and the desert warthog Phacochoerus aethiopicus, occur in Kenya. The desert warthog may be Africa’s least known non-forest large mammal as its distribution is poorly understood and it has never been the focus of an ecological or behavioural study. None of the earlier books and field guides on the mammals of eastern Africa mention the desert warthog (e.g., Dorst & Dandelot 1969, Haltenorth & Diller 1977, Kingdon 1979), and no game laws recognise this taxon (I. Parker, pers. comm.). Kingdon (1997) is the first major work to recognize the desert warthog as a full species and to bring this species to the attention of a large audience.

The preliminary distribution map for the desert warthog compiled by d’Huart & Grubb (2001) presents only four localities for Kenya. They show the southern-most locality as Mkokoni, 60 km northeast of Lamu Island (north coast of Kenya). d’Huart & Grubb found no evidence for desert warthog south of the Ewaso Ng’iro River in central Kenya or west of the Tana River in eastern Kenya. They questioned whether the common warthog and the desert warthog might be sympatric at some sites. d’Huart & Grubb (2005) produced a photographic guide that highlights the diagnostic differences be- tween the common warthog and the desert warthog. Some of the main diagnostic phenotypic characters used to identify the two species of warthog in the field are as follows: common warthogs have pointed ear tips, cone-shaped genal warts, a ‘diabolo-shaped’ head (when viewed from the front), and the suborbital areas are not swollen (fig. 1); desert warthogs have ear tips that are lax and flipped back, hook-shaped genal warts, an ‘egg-shaped’ head (when viewed from the front), and swollen suborbital areas (fig. 2)

In 2005, we started to opportunistically collect distribution data for both species of warthog in Kenya. TMB and YdJ found desert warthogs 15 km and 80 km west of Garissa town in 2005 when they encoun- tered two solitary individuals in medium-dense Acacia bushland during a primate survey. These are the first records west of the Tana River and extend the geographic range to ca. 265 km northwest of Mkok- oni, the nearest locality mentioned by d’Huart & Grubb (2001).

In 2007, JC, J. Feely, and S. Bell-Cross visited Tsavo East National Park south of the Voi River. Al- though they encountered no common warthogs during this trip, they did observe two sounders of desert warthogs in low bush on the edge of the Dika Plains, ca 13 km north and northwest of Buchuma Gate.

SUIFORM S OUNDINGS V OL 8(2) P AGE 4 Figure 1: Adult male common warthog Figure 2: Adult male desert warthog Phacochoe- Phacochoerus africanus on the plains of the Lai- rus aethiopicus in medium-dense shrub in Tsavo kipia Plateau, central Kenya. Note the pointed West National Park, southern Kenya. Note the ears, the cone-shaped warts, the ‘diabolo-shaped’ flipped-back ear tips, the hooked warts, the broad, head, and the lack of swelling of the suborbital ‘egg-shaped’, head, and the swollen suborbital area. area

Photographs were taken and sent to experts for confirmation. Some of these photographs, together with some of our other warthog photographs, are available for viewing on an online digital map at: http://picasaweb.google.com/wildsolutions/WarthogSightingsInKenya/photo? authkey=WigT2oFY78k#map .

These observations considerably extend the known geographical range for the desert warthog (ca. 310 km south from the nearest Garissa sighting and ca. 320 km southwest from Mkokoni) (Fig. 3).

JC made two further visits to the Tsavos in 2007. He found both species of warthog in Tsavo West Na- tional Park and desert warthog north of the Voi River in Tsavo East National Park.

In 2008, TMB and YdJ visited Tsavo West National Park and observed several sounders both of common warthogs and desert warthogs. In the northwest of the Park, in low bush on the edge of riverine forest, they found a sounder of six common warthogs only 150 m from a sounder of four desert warthogs. This locality represents not only the south western-most site in the range for desert warthog (ca. 390 km from Mkokoni, the southern-most point of d’Huart & Grubb 2001), but it also provided the first evidence that common warthog and desert warthog are at least narrowly sympatric over this part of their geographic ranges.

SUIFORM S OUNDINGS V OL 8(2) P AGE 5 Although we have yet to find the common warthog in Tsavo East National Park, it would be surprising if this species were not present there. If not present, however, the common warthog would need to be de- leted from the list of large mammals known for Tsavo East National Park. As concerns the desert wart- hog, Tsavo East National Park and Tsavo West National Park can now add one more species to their al- ready impressive list of large mammals.

Figure 3: Locations of common warthog Phacochoerus africanus and desert warthog Phacochoerus aethiopicus sightings in Tsavo East National Park and Tsavo West National Park, southern Kenya.

References d’Huart J and Grubb P. 2001. Distribution of the common warthog ( Phacochoerus africanus ) and the de- sert warthog ( Phacochoerus aethiopicus ). African Journal of Ecology 39: 156-169. d’Huart J & Grubb P. 2005. A photographic guide to the differences between the common warthog (Phacochoerus africanus ) and the desert warthog ( Ph. aethiopicus ). Suiform Soundings 5: 4-8. < http://iucn.org/themes/ssc/sgs/pphsg/Suiform%20soundings/Newsletter%205(2)pdf > Dorst J & Dandelot P. 1969. A Field Guide to the Larger Mammals of Africa . Houghton Mifflin Co., Boston, USA. Haltenorth T & Diller H. 1977. A Field Guide to the Mammals of Africa including Madagascar . Collins, London, UK. Kingdon J. 1979. East African Mammals . An Atlas of Evolution in Africa . Vol.III B. University of Chi- cago Press, Chicago, USA. Kingdon J. 1997. The Kingdon Field Guide to African mammals . Academic Press, New York, USA.

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