· Y.A. de Jong & T.M. Butynski, 2009. Primates of the coastal forests of Kenya. 2 Primate Biogeography, Diversity, Taxonomy and Conservation of the Coastal Forests of Kenya. Yvonne

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.

Primates of the coastal forests of Kenya. 2

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.



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

Cercopithecus mitis albogularis 84

5. Discussion 87 Acknowledgements 89 References 90



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.



Tables 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.

22

2 Rates of encounter with Otolemur garnettii lasiotis from 2005 – 2008 in the coastal forests of Kenya.

39

3 Rates of encounter with Galagoides cocos during 2005 – 2008 in the coastal forests of Kenya.

47

4 Encounter rates of Cercopithecus mitis albogularis during 2005 – 2008 in the coastal forests of Kenya and outside the Hotspot.

52

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.

53

6 Encounter rates of Cercopithecus mitis albotorquatus during 2005 – 2008 in the Coastal Forests Kenya and while driving to the Hotspot.

59

7 Field descriptions of C. m. albogularis, Gedi Ruins, Kenya, C. m. albotorquatus, Witu Forest Reserve, Kenya and C. m. albogularis, Unguja Island, Tanzania.

60

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.

65

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.

66

10 Encounter rates of Papio cynocephalus ibeanus during 2005 – 2008 in the Kenya coastal forest Hotspot and while driving to the Hotspot.

73

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).

76

12 Encounter rates (groups/hour and groups/km) of Colobus angolensis palliatus during 2005 – 2008 in the coastal forests of Kenya.

78

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.

85



Figures 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. 16

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.

17

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.

18

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

lasiotis from Diani Beach. 39

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

coastal area of Kenya. 44

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


18 Adult male C. m. albogularis at Jozani-Chwaka Bay National Park, Unguja Island, Tanzania.

55

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

albotorquatus. 58

22 Chlorocebus pygerythrus hilgerti male at Diani Beach, Kenya. 63 23 Approximate geographic distribution of Chlorocebus pygerythrus hilgerti in the


24 Suspected geographic distribution and evidence for Chlorocebus pygerythrus excubitor.

68

25 Adult male Papio cynocephalus ibeanus foraging on the beach of Manda Toto Island, Lamu Archipelago, Kenya.

71

26 Approximate geographic distribution of Papio cynocephalus ibeanus over the coast of Kenya.

72

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

in the coastal forests of Kenya. 77

29 Cercocebus galeritus at Mchelelo Research Station, Tana River Primate National Reserve, Kenya.

82

30 Adult male Chlorocebus pygerythrus hilgerti x Cercopithecus mitis albogularis hybrid, Diani, Kenya.

84

31 Encounter site of C. p. hilgerti x C. m. albogularis hybrid. Diani, Kenya. 84



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



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



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.



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?



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



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.



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).



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.



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).



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.



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.



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



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



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



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/ChlorocebusPygerythrusHilgertiXCercopithecusMitisAlbogularisHybrid?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.



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

encounters in the coastal forests of Kenya

Number of encounters outside the coastal forestsof Kenya

Population trend (IUCN, 2008)

Degree of threat category (IUCN, 2008)

Abundance in the coastal forest of Kenya

Nocturnal primates

1 Otolemur garnettii lasiotis

>40 individuals

Present, not counted

Stable Least Concern Very common

2 Galago senegalensis braccatus

0 Present, not counted

Stable Least Concern Uncommon

3 Galagoides cocos >20 individuals

0 Stable Least Concern Common

Diurnal primates

4 Cercopithecus mitis albogularis

24 groups 3 groups Decreasing Least Concern Common

5 Cercopithecus mitis albotorquatus

16 groups

0 Decreasing Vulnerable Common in along the Lower Tana River and Witu area.

6 Chlorocebus pygerythrus hilgerti

11 groups 33 groups Stable Not assessed Common outside of the coastal forests.

7 Chlorocebus pygerythrus excubitor

0 0 ? Not assessed Rare

8 Papio cynocephalus ibeanus

32 groups

39 groups Stable Least Concern Very common

9 Colobus angolensis palliatus

11 groups 0 ? Least Concern Common in the coastal forest south of Mombasa

10 Procolobus rufomitratus rufomitratus

2 groups 0 Decreasing Endangered Endemic to the Lower Tana River



where it is common.

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

Other Chlorocebus

pygerythrus hilgerti x Cercopithecus mitis albogularis hybrid

1 individual 0 - - Rare

Cercopithecus mitis 4 groups Total 99 groups, 61

solitary individuals

79 groups, solitary 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



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).



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, these islands are not particularly isolated. 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.

Figure 5. Manda Toto Island on the left, Manda Island on the right. During low tide the two islands are almost connected.



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). 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

Figure 6. Cercopithecus mitis albotorquatus in Witu Forest Reserve



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.



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).



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.



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.



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)



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



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



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)



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)



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



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



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 a

Documents

· Y.A. de Jong & T.M. Butynski, 2009. Primates of the coastal forests of Kenya. 2 Primate Biogeography, Diversity, Taxonomy and Conservation of the Coastal Forests of Kenya. Yvonne