Primate Biogeography, Diversity, Taxonomy and ... - Wild Solutions

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
yvonne@wildsolutions.nl & 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
Cercopithecus mitis albogularis
84
5. Discussion 87
Acknowledgements 89
References 90
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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,
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
coastal forests of Kenya.
3 Rates of encounter with Galagoides cocos during 2005 – 2008 in the coastal
forests of Kenya.
4 Encounter rates of Cercopithecus mitis albogularis during 2005 – 2008 in
the coastal forests of Kenya and outside the Hotspot.
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.
6 Encounter rates of Cercopithecus mitis albotorquatus during 2005 – 2008 in
the Coastal Forests Kenya and while driving to the Hotspot.
7 Field descriptions of C. m. albogularis, Gedi Ruins, Kenya, C. m.
albotorquatus, Witu Forest Reserve, Kenya and C. m. albogularis, Unguja
Island, Tanzania.
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.
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.
10 Encounter rates of Papio cynocephalus ibeanus during 2005 – 2008 in the
Kenya coastal forest Hotspot and while driving to the Hotspot.
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).
12 Encounter rates (groups/hour and groups/km) of Colobus angolensis
palliatus during 2005 – 2008 in the coastal forests of Kenya.
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.
22
39
47
52
53
59
60
65
66
73
76
78
85
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
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
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
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
coastal area of Kenya.
51
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
coastal area of Kenya.
64
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
16
17
18
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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

3. METHODS
Y.A. de Jong & T.M. Butynski, 2009.
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
encounters
in the coastal
forests of
Nocturnal primates
Kenya
Number of
encounters
outside the
coastal forests
of Kenya
Population
trend (IUCN,
2008)
Degree of
threat
category
(IUCN, 2008)
Abundance in
the coastal
forest of
Kenya
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
individuals
0 Stable Least Concern Common
Diurnal primates
4 Cercopithecus mitis
albogularis
5 Cercopithecus mitis
albotorquatus
6 Chlorocebus
pygerythrus hilgerti
24 groups 3 groups Decreasing Least Concern Common
16 groups
0 Decreasing Vulnerable Common in
along the
Lower
Tana River
and Witu area.
11 groups 33 groups Stable Not assessed Common
outside of the
coastal
forests.
0 0 ? Not assessed Rare
7 Chlorocebus
pygerythrus excubitor
8 Papio cynocephalus
ibeanus
32 groups 39 groups Stable Least Concern Very common
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
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
Primates of the coastal forests of Kenya. 23

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

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
Y.A. de Jong & T.M. Butynski, 2009.
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
Figure 5. Manda Toto Island on the left, Manda Island on the right.
During low tide the two islands are almost connected.
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

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

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

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

28 April 2008
Papio: 0.7 groups/h, 0.05 groups/km (n=2)
Y.A. de Jong & T.M. Butynski, 2009.
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

Altitude: 700 – 1300 m asl
Y.A. de Jong & T.M. Butynski, 2009.
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

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
white.
Y.A. de Jong & T.M. Butynski, 2009.
Figure 7. Otolemur garnettii lastiotis present in good
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.
pinkish. Finger tips brownish. Ear length ca. 41 mm.
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
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

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
pinkish or pale grey. Interocular stripe white.
Eye rings dark grey. Sides of the face and lower
Y.A. de Jong & T.M. Butynski, 2009.
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
Figure 12. Galago senegalensis braccatus in Euphorbia
tirucalli hedge around human settlement, Isiolo, central
Kenya.
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

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 buffybrown
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,
extending to slightly above eyes. Eye rings grey.
Throat, cheeks and ventrum off white to pale grey.
Y.A. de Jong & T.M. Butynski, 2009.
Figure 14. Galagoides cocos in Acacia – baobab
woodland at Takwa Ruins, Manda Island, Kenya.
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,
Mazeras
- - Foot No counts
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

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
fringe narrow, pale grey. Brow band grizzled
Y.A. de Jong & T.M. Butynski, 2009.
Figure 16. Cercopithecus mitis albogularis adult male at
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
Road Surveys
0.4 (n=3) 0.02 (n=3) Vehicle
Kiboko to Lunga
Lunga (excluding
0 0 Vehicle Highway
Mrima Hill)
Lunga Lunga to
Mazeras (including
Shimba Hills NR,
excluding Mrima
Hill)
Mtito Andei to
Kilimangodo North
Kilimangodo North
to Kaya Sega (Vanga,
Lunga Lunga, Kaya
Gonja, Godo)
Kaya Sega to Diani
(Vanga, Kaya
Dzombo, Kaya
Gonja, Buda Forest)
0 0 Vehicle Main road and
bush tracks
0 0 Vehicle Bush track
0.6 (n=3) 0.03 (n=3) Vehicle Using main roads
0.5 (n=4) 0.02 (n=4) Vehicle Using tracks and
main roads. Three
encounters in
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
(including Malindi)
0.2 (n=1) 0.01 (n=1) Vehicle Using main roads
Watamu to Tsavo
East NP (including
Malindi, Arabuko
Sokoke Forest, Sala
Gate Tsavo East NP)
0 0 Vehicle
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

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.
Y.A. de Jong & T.M. Butynski, 2009.
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
Unguja Island,
Altitude
(m asl)
Photo
C. m. albogularis C. m. albogularis Cercopithecus C. m.
Usa River, Mrima Hill, mitis albogularis albogularis Gedi
Tanzania Tanzania Kenya
Diani, Kenya Ruins, Kenya
20 1195 80 16 20
Ears Narrow, pale grey,
fringe
Pale grey tufts Narrow, pale grey,
fringe
Narrow, pale grey,
fringe (some
whitish)
Primates of the coastal forests of Kenya. 53
Fringe absent or
narrow, pale
grey.
Face Grizzled pale grey. Grizzled pale grey. Grizzled grey. Bare Grizzled grey. Bare Grizzled grey.

Bare black skin
around eyes.
Cheeks Whiskers full,
grizzled pale grey.
No collar line
crossing the lower
cheeks.
Brow band Projects forward
and upwards.
Grizzled pale grey.
Crown Grizzled olive-grey
with rufus wash.
Collar Whitish-pale grey,
ca 55-60%
complete.
Bare black skin
around eyes
Whiskers full,
grizzled olive-grey.
No collar line
crossing the lower
cheeks.
Projects forward
and upwards.
Grizzled olive-pale
grey.
Grizzled olive-grey
with rufus wash.
Absent. White
throat patch
present.
Y.A. de Jong & T.M. Butynski, 2009.
black skin around
eyes.
Whiskers not full,
grey. Collar line
through the cheeks.
Projects forward
and upwards.
Grizzled grey.
Grizzled olive-grey
with pale rufus
wash.
White, ca 75-80%
complete. Collar
line sharp, running
from upper jaw,
across cheeks ca 4
cm below the ears,
onto neck, down
the shoulders to
include the throat
and upper chest.
black skin around
eyes.
Whiskers not full,
grey. Collar line
through the cheeks.
Projects forward
and upwards.
Grizzled grey.
Grizzled olive-grey
with pale rufus
wash.
Not sharply
defined. Around
lower ca. 60% of
neck. Throat offwhite.
Sides of
neck light grey.
Primates of the coastal forests of Kenya. 54
Bare black skin
around eyes.
Whiskers not
full, grey. Collar
line through the
cheeks.
Projects forward
and upwards.
Grizzled grey.
Grizzled olivegrey.
White, ca 75%
complete. Collar
line sharp,
running from
upper jaw, across
cheeks to ca 4
cm below the
ears onto neck,
down shoulders,
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
arms
Black Black Black Black Black
Inner arms Dark grey Black Dark grey Blackish Dark grey
Outer legs Grizzled grey, feet
blackish
Dark grey Dark grey Grizzled grey Dark grey
Inner legs Grizzled grey Dark grey Upper part white,
lower part dark
grey
Pale grey Pale 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
ventrum pale grey.
Pale grey
Sides Grizzled reddish- Grizzled reddish- Grizzled grey Grizzled grey Grizzled grey
greygrey
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

characters dark grey ventrum,
rich red dorsum,
full cheek
whiskers.
blackish ventrum,
rich red dorsum,
full cheek
whiskers.
white upper inner
legs.
Y.A. de Jong & T.M. Butynski, 2009.
ventrum grey ventrum.
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

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
from upper jaw, across the cheeks to ca 2 cm below the
ears then down onto the sides of the neck, across the top
Y.A. de Jong & T.M. Butynski, 2009.
Figure 20. Cercopithecus mitis albotorquatus male
from Witu Forest, Kenya, where this subspecies is
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
Reserve
7.14, n=3 0.6 n=3 Vehicle Using existing roads
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 semiarid
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
Foto
20 m asl 20 m asl
Ears Narrow pale grey fringe Absent or narrow pale
grey fringe.
Face Grizzled pale grey. Bare black Grizzled grey. Bare black skin
skin around eyes.
around eyes.
Cheeks Full, grizzled pale grey. Grizzled grey, slender
appearance. Collar line
crossing the lower cheeks
Brow band Projecting forward and upwards.
Grizzled pale grey
Projecting forward and
upwards. Grizzled grey.
Narrow fringe, grey.
Triangular shape with
short hair. Grizzled grey.
Bare black skin around eyes.
Grizzled grey, slender
appearance. Collar line
crossing the lower cheeks
Projects forward and
upwards. Grizzled greyisholive
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
blackish
Dark grey. Grey.
Inner legs Grizzled grey Pale grey Pale grey to off white.
Back of hind
limbs
Grizzled grey. Pale grey with rufus wash Long, off white, hairs with
distinguishing rusty wash.
Rust darkest around base
of tail.
Dorsum Grizzled grey between
shoulders. Half way the back
onto the upper tail rich red.
Grizzled grey between
shoulders. Below
shoulders grizzled rusty
red extending onto the upper
tail.
Grizzled grey between
shoulders. Behind shoulders
grizzled olive-grey-rusty
becoming increasingly rusty
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

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
contigua (Heller, 1920), Changamwe, 10 W
Mombasa, Kenya
luteus (Elliot, 1910), Wambugu, SW Mt Kenya, Kenya
rubellus (Elliot, 1909), Fort Hall, Kenya.
tumbili (Heller, 1913), Ndi, Taita District, S Kenya.
Y.A. de Jong & T.M. Butynski, 2009.
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.
Figure 22. Chlorocebus pygerythrus hilgerti male
at Diani Beach, Kenya.
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
Conservancy
/Mpeketoni/ Witu
Forest Reserve
0.1 (n=2) 0.02 (n=2) Vehicle
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,
south coast
Tana River PNR –
Kipini
Conservancy
Garissa – Tana
River PNR
Watamu – N.
Malindi – W.
Malindi - Watamu
Tanga (Tz) – Kaya
Rabai, Mazeras
Kaya Rabai,
Mazeras - Nairobi
Mwea NR –
Mwingi East
Mwingi East –
Tana River PNR
0.4 (n=1) 0.02 (n=1) Vehicle, 25.2 km/h Travelling slowly
on highway
0.2 (n=1) 0.01 (n=1) Vehicle, 27.5 km/h
0.4 (n=2) 0.01 (n=2) Vehicle, 38.3 km/h
0.2 (n=1) 0.01 (n=1) Vehicle, 23.5 km/h
0.4 (n=3) 0.01 (n=3) Vehicle Through Mrima
Hill and Shimba
Hills NR
0.1 (n=1) 0.002 (n=1) Vehicle Highway
0.2 (n=1) 0.01 (n=1) Vehicle
0.2 (n=2) 0.10 (n=2) Vehicle
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.
Ears Black
eyes
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.
Under tail slightly lighter
grey.
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.
Long whiskers stand out.
p. hilgerti.
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

4.2.7 Chlorocebus pygerythrus excubitor
(Schwarz, 1926)
Y.A. de Jong & T.M. Butynski, 2009.
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

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). Males have a trace of mane, are
larger and more robust than females. Muzzle
Y.A. de Jong & T.M. Butynski, 2009.
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
Figure 25. Adult male Papio cynocephalus ibeanus
foraging on the beach of Manda Toto Island, Lamu
Archipelago, Kenya.
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
River PNR
0.1, n=2 0.004 Vehicle, ca 45 km/h
Tana River PNR –
Garissa
0.2, n=1 0.01, n=1 Vehicle, ca 45 km/h
Tana River Primate
Reserve –Garsen –
Witu
0.3, n=1 0.01, n=1 Vehicle, 30 km/h
Witu – Garsen – Tana
River PNR
1.1, n=4 0.03, n=3 Vehicle, 40.2 km/h
Nairobi – Makindu –
Mrima Hill
0.7, n=5 0.13, n=1 Vehicle, ca 65 km/h
Mazeras – Nairobi 0.4, n=3 0.01, n=1 Vehicle, 43.8 km/h
Tana River PNR –
Garissa – Nanyuki
0.3, n=3 0.01, n=3 Vehicle, ca 50 km/h
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 –
Kilimangodo
0.2, n=1 0.004, n=1 Vehicle, 41.5 km/h
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
NP
0.3, n=2 0.01, n=2 Vehicle, 31.8 km/h
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
description Manda Toto Island
Foto
P. c. ibeanus
Tana River PNR
P. c. ibeanus
Diani Beach
Cheeks Yellowish to off-white Yellowish-white to pale
grey
Pale yellowish-grey
Nostrils Directed upwards Directed upwards Directed upwards
Pale grey
muzzle
patch
Present Present but vague Present
Head
shape
Pointed Pointed Pointed
Mane Trace Absent Trace
Dorsum Grizzled yellowish- grey Grizzled yellowish- grey.
Some with brighter yellow
than others
Grizzled yellowish- grey
Ventrum Greyish to off-white with Pale yellowish-white to pale Pale yellowish-grey
and inner
limbs
yellow wash
grey
Callosite
shape
Medium U-shape Medium U-shape Medium U-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.
Also, more phenotypic
variation seen within groups
than observed for other
populations of Papio in East
Africa.
Island (close to type locality)
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

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).
Extent of occurrence: ca 272,000 km². Area of
occupancy in Kenya ca. 249 km² (Anderson et al., 2007).
Y.A. de Jong & T.M. Butynski, 2009.
Figure 27. Adult male Colobus angolensis palliatus
at Mrima Hill, south coast of Kenya, where this
subspecies is common.
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 -
Diani, road
survey
0.3, n=2 0.01 (n=2) Vehicle, 27.9 km/h
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.
Primates of the coastal forests of Kenya. 78

Y.A. de Jong & T.M. Butynski, 2009.
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/km 2 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/km 2 ; in the Mwache F. R., ca. 6 individuals/km 2 . 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.
Primates of the coastal forests of Kenya. 79

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)
Y.A. de Jong & T.M. Butynski, 2009.
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.
Primates of the coastal forests of Kenya. 80

Y.A. de Jong & T.M. Butynski, 2009.
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.
Primates of the coastal forests of Kenya. 81

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
Tana River, SE Kenya, from Kanjonja in the north
to Garsen in the south (01.40S-02.15S, 40.10E,
20-40 m asl; Butynski & Mwangi 1994, 1995; Butynski
et al. 2008d).
Y.A. de Jong & T.M. Butynski, 2009.
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)
Figure 29. Cercocebus galeritus at Mchelelo
Research Station, Tana River Primate
National Reserve, Kenya.
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).
Primates of the coastal forests of Kenya. 82

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
Y.A. de Jong & T.M. Butynski, 2009.
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.
Primates of the coastal forests of Kenya. 83

4.2.12 Hybrid
Chlorocebus pygerythrus hilgerti x Cercopithecus mitis
albogularis
Y.A. de Jong & T.M. Butynski, 2009.
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
light grey. Hands and feet dark grey. Crown, dorsum, sides
and proximal ca. 10% upper part of tail grizzled greyish-olive
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 (14 th 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
Figure 30. Adult male Chlorocebus pygerythrus hilgerti
x Cercopithecus mitis albogularis hybrid on a large
beach hotel compound in Diani, Kenya.
Primates of the coastal forests of Kenya. 84

Y.A. de Jong & T.M. Butynski, 2009.
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
characteristics
Photo
pygerythrus x mitis Chlorocebus pygerythrus
hilgerti
Cercopithecus mitis
albogularis
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
upwards and backwards
covering part of the ear.
grizzled pale grey.
Face Triangular-shaped area of Jet black, hairless Grizzled grey. Bare black
bare grey skin around eyes
and nose. Grizzled pale
grey pelage around the
mask.
skin around eyes.
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
Primates of the coastal forests of Kenya. 85

Collar Vague and not sharply
defined. Around lower ca.
50% of the neck. Throat
off white. Sides of neck
mouse grey.
Y.A. de Jong & T.M. Butynski, 2009.
Absent Present but not distinct or
sharply defined. Around
lower ca. 50% of the neck.
Throat off-white. Sides of
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
arms
Grizzled mouse grey.
Lower arms slightly darker
grey.
Grizzled mouse grey Black.
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
legs
Hair slightly longer. Off
white to pale grey. Rusty
wash around ischial
callosities.
Dorsum Grizzled olive-grey with
rufus wash extending unto
the upper tail.
Off white to pale grey.
Red patch under base of
tail in males.
Hair longer. Off white
with rufus wash.
Grizzled olive-grey. Grizzled grey between
shoulders. Below
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
grey then blackish distally.
the tail in males
tail.
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
intermediate in colour
between C. p. hilgerti and
C. m. albogularis.
hilgerti.
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’?
Primates of the coastal forests of Kenya. 86

Y.A. de Jong & T.M. Butynski, 2009.
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?
Primates of the coastal forests of Kenya. 87

Y.A. de Jong & T.M. Butynski, 2009.
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.
Primates of the coastal forests of Kenya. 88

Y.A. de Jong & T.M. Butynski, 2009.
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.
Primates of the coastal forests of Kenya. 89

Y.A. de Jong & T.M. Butynski, 2009.
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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
Face
- Colour cheek
- Colour naked
skin
P. anubis P.a. furax P.a. neumanni
Purple-black (1), naked, dark grey to black (3), ruffs of hair at cheeks (3),
dark grey (4), black (5), hair on cheeks prominent (5)
Mane Short to medium (1), male large manes (4),
Upper parts Olive-brown (1), dark grey to olive-brown sometimes grading into olive or
light brown (khaki or grey) (3),
Eastern Uganda and western Kenya
(above the Chanler Falls on the Northern
Ewaso Ngiro).
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)
Below the Chanler Falls on the Ewaso
Ngiro, then southward through Kenya
and Tanzania.
Darker than neumanni (6) Pattern and colour generally similar (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
Face
- Colour cheek
- Colour naked
skin
P. cynocephalus P.c. typical P.c. ibeanus P.c. kinda
Purple-black (1), muzzle greyish to black (2), cheeks almost white and hair sparse (2),
cheeks white (4), cheeks very pale approaching a yellow-white(5), hair short on top of head
and lays flat (5)
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
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
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
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.
Mikumi (T), Ruaha (T) Amboseli (K), Tana River (K) Angola, Zambia (west of
Typical and Ibean)
pelage (2)
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
wavier and coarser pelage
(2)
reddish-brown (2)
Curved (4)
straight and soft, silky
pelage (2)

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
district.
Witu Islands, Kenya Sudan & Ethiopia NE Tanzania & Kenya
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
black and the tight white (5)
Legs sometimes washed with yellow (6)
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
tinge. Lighter bands of dorsal hairs
Tail Creamy white tip, white basal tuft (1). Long shiny
hairs (5). Whitish/yellow upper side and lightly
speckled with black tips underside. Terminal end
creamy white (5) white tuft at root (5)
Face Black (1) Black line from corner eye running back
Brow band & frontal
band
Cheek
whiskers
along temples (1)
White (1). Frontal band distinct, narrow in the
centre and white (5) separate from the whiskers
(5). Long black brow hairs border it in front (5)
Long, stiff and yellowish with short black tips (1).
White with yellow at tips, directed upwards and
backwards conceiling the ears (5) Ends of longer
hairs speckled black & yellow, tips black (5)
Above dark grey with basal part
showing extension of back colour,
terminal end black, beneath yellowish
white with red patch at root (5)
between dark olive-buff and old gold (5)
Washed out tawny grey (5)(6)
End of tail whitish (4) Grey above, pure black at tip, rusty red at
base (5). Grey above, scarcely washed
with yellow black at tip (6) Large rusty
rusty-red basal patch
Whitish moustache Jet black (5)(6)
Distinct but narrow (5) Frontal band greyish white to tawny white
(5). Greyish-white or tawny-white
browband (6)
Whiskers long, falciform, pure white (4) Long whiskers passing gradually above
into colour of crown (5). Greyish-white or
tawny-white extending from browband (6)
Long cheek hairs overlaying ears (6)
Crown
Neck
Ears
Grizzled, gold to greenish (1). Golden green (5) Washed out tawny grey (5)
Body size Smaller than hilgerti (2)(4)
Scrotum Sky blue (1) Turquoise blue
Overall Sides golden green (5) rusty patch on pubic and
perineal area and around scrotum (5)
Differs from hillgerti in that there is no
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
Ethiopia
Limbs Deep mouse grey, upper arms tinge of dorsal
colour (5)
Tanzania, Kenya between L. Victoria and Rift
Valley to Mt Elgon and Mt Debasien; Elgeyo and
Mau esc.
NW Tanzania, SW Uganda, Rwanda, Burundi, DRC.
Laterally iron grey (5) Light iron grey laterally, sharp demarcation with dorsal (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
with greenish tinge; below white or yellowish white.
Top black (5)
Grey speckled above. Greenish of back continues
on tail (5). Terminal end black (5)
Grey with yellow speckling near root dorsally. Terminal end black
(5). Few rusty hairs at root (5)(6)
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,
Taitas, Mazeras
W. Kenya & Uganda W. Kenya & Highlands & U Mau (Kenya) Patta & Witu Island (K) Tutha, Kenya district ?
Under base of tail Red patch extending to thighs
sometimes present (2)(3)
Russet tuft (2)
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
white (9)
Blackish chest (10)
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
yellow (9)
faintly greenish (7)
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).
Speckled and sides of neck as brow
band(10)
Face dark olive (2)
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
white half collar (9)
than kolbi (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
(good indicator)(8)
white patch (10)
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
becoming sooty grey towards wrists.
Outer limbs speck smokey grey (9).
Inner limbs pale grey to white(9)
kolbi (8)
black, dark grey (2), partly black (3)
black (2)
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
inferiorly, rest dark speckled. Term end
black (9) Rusty red at root above and
outside (9)
(6)
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
yellowish-grey with black (little greyer
than head) (9)
(10)

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,
younger animals do show this patch (8)
Russet tuft (3) Reddish (3)
Under parts speckled grey (2) Dark (1), dark slate grey (7).
Smokey grey (9)
Dark (7) Not black (1)
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).
Cheeks speckled greyish yellow
(9)
Cheek whiskers yellowish grey (3),
cheeks yellowish-olive (7)
Collar No collar (10) Creamy (3) No whitish half colar (9) Nearly complete (3), broad snow-white
(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
size (7)
Shoulders Speckled with greyish yellow (9).
In front of shoulders black (9)
Limbs slightly speckled (2) Thighs speckled grey (9). Black
upperarms (9). Grey speckled (9)
H/F black speckled (9)
Tail 66% of distal half black (3), ventral
surface reddish (7) Black,
speckled towards base, reddish
above root (9)
Other Speckled gray with dark reddish
saddle (3), nape dark grey (3),
sides and thighs light grey (7)(1).
Sides speckled grey (9)
Yellowish-olive (7). Behind
shoulders speckled and rich
reddish orange-yellow (9)
Greyish (2), cheek whiskers (3) Long grizzled whiskers (5)
Nearly complete (3)
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.
White (2) (3)
Black (5) Greyish (2), speckled yellowish grey
(3)
Thighs dark grey speckled (9) Hind legs speckled, looking dark grey (8)
Reddish tinge on hindlegs
40% of terminal end black (3) No red on on anal region and base tail
(9)
Neck yellowish-olive (7). Neck
thickly speckled with black (9)
Closely resembling C. a.
albogularis (9)
Dark grey, shiny black (5), hindlimbs
blackish grey (7)
60% distal part black 30% terminal end black
Longer fur than albotorquatus (9) Nape black (3)(7). Buff speckled
frontal band (3). Possibly the closest
resemblance with ancestral type (5)

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
Cap
- tip
- base
- brush
Tail tip white , bushy, occupying about 33% of tail
length (1), no distinct brush at end of tail, the distal
third being white but the area up to the middle
being distinctly speckled (4)
Cap hairs lying flat toward back with nuchal parting
(3), hairs of occiput lengthened into a distinct tuft
(4)
Distal 5-10cm of tail greyish (1), tail almost entirely
black, distal 5% grizzled (2), tail almost entirely black
(3)(4)
Distal 25% of tail grizzled (2), tail brush rather full,
white and 33% of tail length but the speckling
extends farther up to about the middle of the tail (4)
Hairs of the occiput broadly and evenly lengthened
(4)
Brow band White brow band fairly broad, broadly connected
via full cheek whiskers with epaulettes (1)
Narrow brow band (2)(4) Distinct white brow band (2)
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,
broadly connected with shoulder mane (4)
(1), large epaulettes (3).
epaulettes (2)
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
patch small (4)
region (1), pubic patch small and grizzled (2)(3)
Pelage Coat long, thick and soft (4) Very long pelage (3), Long pelage (2), coat very long, thick and soft but
rather short (4)
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.
Literature

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
Lakes and Kahe.
Tail
- Length
- Tuft
- Base
Veil
- length
White tail tuft making up over
80% of tail length, only extreme
proximal part of tail black (1)(6).
Base of tail black, slightly shorter
than in kikuyensis (3). Tail tuft
longer than kikuyensis (3), tail
completely white, often yellow in
colour (4). 3 to 4 inches of base
black, the rest white tuft (20-21
inch)(7)
Flank veil longer than kikuyensis
(1)(3). Veil covers black part of
tail completely
Kenya, west of Rift and in the Rift,
west to Mt Elgon, south to
Ngorongoro Crater and the
Grumeti River Tanzania
White tail tuft short (45% of tail
length), rest black (1)(6), tail
longer than head and body (1),
tuft copious, 50% of the length of
tail (3)(4). Tail base black, tail tuft
full covering 33% of total length
(5).
Flank veil much yellowed, not
extending up onto back but
covering base of tail (1), posterior
portion of veil about as long as the
tail tuft, extending to the basal
portion of the tail (3)(4)(5),
occasionally as far down as the
beginning of the tuft (3).
Shoulders White hairs sprinkled over
shoulders, nearly linking veil to
white of throat (1)
Throat White (1)
Under parts Fur less woolly than kikuyensis
(1)(3)
Other Longer and distinct thigh-band
(compared to kikuyensis) (3)
From Ngong Escarpment to Mt
Kenya and Aberdares, Kenya
White tail tuft very bushy, extending
over 72% of tail length (1), tail
proximally gray (1), tail short, about
equal to head plus body (1), Base of
tail grey, slightly longer than in
caudatus (4). Tail completely white
with tuft almost from base (4). Base
of tail black, white tuft voluminous
forming about 66% of the length of
tail (5)(6).
Flank veil very long extending well up
on back and beyond base of tail tuft
(1)(3), veil covering 30% of the tail
(4). Veil covering up to a quarter of
the tail (5)
Hair short (1) Hair long; thigh stripe short (1),
Thigh-band short and pale (3), fur
soft and woolly (3)
Mount Gargues, Mathews Range,
Kenya
White tail tuft extends over 63% of
tail (1), tail as long as head plus
body (1), tuft full (3)(6) and bushy,
occupies 66% of the whole tail (3).
Tail tuft smaller than caudatus, longer
tail, larger body than caudatus (8)
Flank veil very long, creamy yellow,
only just reaching tail base (1)(3).
No white sprinkling on shoulders
below veil (1)
Hair very long (1). Temporal ridges
into sagittal crest (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.
Mau Forest (8)
More extensively black tail than
C. g. matschiei (8). White
terminal end of tail 25%, other
75% black short hair (8)
Resembling C. g. matschiei but
smaller. Sagittal cres on the
skull when aged (8)

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
baobab cocos palm
07-02-06 17.16
190 C. m. albogularis Mrima Hill -4.48444 39.27057 110 Secondary forest edge, coastal and shamba mango cashew
baobab cocos palm
07-02-06 17.27
191 C. m. albogularis Mrima Hill -4.48361 39.27133 82 Secondary forest edge, coastal and shamba mango cashew
baobab cocos palm
07-02-06 17.29
194 C. m. albogularis Mrima Hill -4.49069 39.25439 101 Secondary forest edge, coastal and shamba mango cashew
baobab cocos palm
07-02-06 17.55
195 C. m. albogularis Mrima Hill -4.48881 39.25431 124 Secondary forest edge, coastal and shamba mango cashew
baobab cocos palm
07-02-06 18.07
197 C. m. albogularis Mrima Hill -4.48656 39.25418 130 Secondary forest edge, coastal and shamba mango cashew
baobab cocos palm
07-02-06 18.19
198 C. m. albogularis Mrima Hill -4.48428 39.25394 146 Secondary forest edge, coastal and shamba mango cashew
baobab cocos palm
07-02-06 18.20
248 C. m. albogularis Mrima Hill -4.48694 39.27056 79 Secondary forest edge, coastal and shamba mango cashew
baobab cocos palm
20-02-06 11.13
- 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,
Finch Hatton camp
-2.93328 37.90464 798 Ground water forest 20-4-08 12.10
333 C. m. albogularis Tsavo West NP,
Finch Hatton camp
-2.93171 37.90537 806 Ground water forest 20-4-08 13.40
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
highway
22-4-08 10.30
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
road in town
23-4-08 16.32
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
including Acacia, Commiphora, wild fruit trees on coral rag
7-7-08 13.15
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
Campsite, riverine forest and dry
09-08-05 11.00
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)
Tana River Primate
-1.78122 40.10539 25 Acacia bushland, sandy soil, close to village 20-06-06 10.08
252A C. p. hilgerti NR N
Tana River Primate
-1.84272 40.06139 42 Acacia bushland, sandy soil, close to village 20-06-06 10.47
256 C. p. hilgerti NR S -1.93767 40.08487 46 Acacia bushland sandy soil
Grassland with doumpalm, greenbush whith some
21-06-06 10.48
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
Makindu, Hunters
0.16298 38.21313 590 13-2-07 8.07
324 C. p. hilgerti lodge
Tsavo West NP,
-2.21298 37.71455 Garden of a lodge, around pond 19-4-08 12.45
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
Tsavo West NP,
-2.98758 37.94847 751 Riverine Forest 20-4-08 11.24
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.
On the edge of the beach, medium dense bushland
20-4-08 17.25
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
(including Acacia, commiphora)
13-10-05
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
NR
-1.87700 40.13878 35 Dry bushland, near riverine forest 06-08-05 11.29
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
woodland
21-06-06 12.54
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
some taller trees
22-06-06 10.45
261 P. c. ibeanus Kipini Conservancy -2.47582 40.55717 13 Grassland with doumpalm, greenbush with some
taller trees, on edge with agricultural land
22-06-06 10.55
262 P. c. ibeanus Kipini Conservancy -2.47265 40.55675 Grassland, open bush/woodland with doumpalm
bordering agricultural land
22-06-06 10.58
263 P. c. ibeanus Kipini Conservancy -2.42517 40.61950 Grassland, open bush/woodland with doumpalm
bordering agricultural land
22-06-06 14.35
264 P. c. ibeanus Kipini Conservancy -2.42258 40.62205 16 Grassland, open bush/woodland with doumpalm
bordering agricultural land
22-06-06 14.39
266 P. c. ibeanus Witu forest E -2.34858 40.59468 16 Coastal forest clearing, moving into agricultural land and
tall grass
23-06-06 07.48
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
NR S
-1.98548 40.09865 Town 25-06-06 13.40
282 P. c. ibeanus Tana River Primate
NR
-1.87642 40.13892 Riverine gallery forest edge with dry bushland 25-06-06 14.35
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
to no undergrowth
26-06-06 9.18
340 P. c. ibeanus Mwardimu -4.17304 39.01811 251 Close to agricultural, outside medium dense Acacia
bushland
21-4-08 17.06
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,
Kiboko camp
-2.19331 37.67158 927 Highway 06-02-06 13.10
176 Papio Makindu,
Hunters lodge
-2.20999 37.71606 Lodge compound with large yellow fever 06-02-06 15.54
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,
Chyulu Gate
-2.90508 38.03984 787 Acacia - Commiphora bushland with dense tall grass 19-4-08 16.16
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,
Sala Gate E
-3.12033 39.36525 145 Acacia bushland, medium 26-4-08 11.08
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
bushland and agricultural land
28-4-08 10.45
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
Tana River Primate
Latitude Longitude Altitude m Vegetation Date Time
17B P. r. rufomitratus NR
Tana River Primate
-1.87641 40.13891 46 Riverine gallery forest
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
(23 individuals) of C. a. palliates in the forest.
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.
Figure 32. Dzombo Hill Forest Reserve, surrounded by
degraded bushland, agriculture and human settlements.
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).

Taxonomy, Distribution, and Conservation Status of Three Species
of Dwarf Galagos (Galagoides) in Eastern Africa
Thomas M. Butynski 1 , Yvonne A. de Jong 2 , Andrew W. Perkin 3 , Simon K. Bearder 3 , and Paul E. Honess 4
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.
Key Words: Dwarf galagos, Galagoides, cocos, granti, zanzibaricus, udzungwensis, taxonomy, conservation
Introduction
1 Eastern Africa Regional Program, Conservation International, Nairobi, Kenya
2 Kenya Institute of Primate Research, Ol Kalou, Kenya
3 Nocturnal Primate Research Group, Anthropology Department, Oxford Brookes University, Oxford, UK
4 Department of Veterinary Services, University of Oxford, Oxford, UK
Many of the species and subspecies of the family Galagidae
(galagos or bushbabies) have been subjected to repeated
taxonomic revisions and name changes over the past century
(for example, Elliot 1913; Allen 1939; Hill 1953; Groves 1977,
1993, 2001, 2005; Grubb et al. 2003). The Kenya coast galago
(or Diani small galago), Galagoides cocos, is no exception
(Figs. 1 and 2). Galagoides cocos is a recently revived, highly
cryptic species of the coastal forest of eastern Africa.
In this paper we (1) review the nomenclatural history for
the Kenya coast galago, (2) provide new information that confirms
that ‘cocos’ is the valid name for this recently revived
species, (3) summarize the phenotypic and the main qualitative
vocal differences among G. cocos, the Zanzibar galago (Galagoides
zanzibaricus), and the Mozambique or Grant’s galago
(Galagoides granti), (4) review their geographic ranges, and
(5) examine their conservation status (Figs. 1–7).
63
Primate Conservation 2006 (21): 63 –79
Nomenclatural History for the Kenya Coast Galago
On 16 December 1911, Edmund Heller (1912) collected
an adult male dwarf galago (one of 10 specimens)
at Mazeras, Kenya, which he named Galago moholi cocos.
This taxon was raised to species status (Galago cocos) by
Elliot (1913), but later placed as a subspecies of the Somali
galago (Galago gallarum cocos), as a subspecies of the
Zanzibar galago (Galago zanzibaricus cocos), or simply not
recognized as a valid taxon and placed as a junior synonym
of the Zanzibar galago (Galago senegalensis zanzibaricus
or Galago zanzibaricus zanzibaricus or Galago zanzibaricus
or Galagoides zanzibaricus) (Table 1). Most recently,
this taxon has been provisionally referred to as ‘Galagoides
cf. cocos’ (Bearder et al. 2003; Grubb et al. 2003). This
binomial is ‘provisional’ because the validity of the use of
the name ‘cocos’ requires confirmation.

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 3. Adult male Zanzibar galago (Galagoides zanzibaricus udzungwensis)
from Pande Game Reserve, Tanzania (near Dar es Salaam). Note the
absence of muzzle patches. Photograph by Nike Doggart.
64
Figure 4. Adult (sex not known) Udzungwa (or Matundu) galago (Galagoides
zanzibaricus udzungwensis) from Matundu Forest Reserve, Udzungwa Mountains,
south-central Tanzania (near Ifakara). Note that the hairs of the tail are of
even length, sparse, and wiry, that the bone of the tail is visible, and that the tip
of the tail is dusky. Photograph from Honess (1996).

Many of the more recent taxonomic studies on Galagidae
make no mention of cocos, but presumably they consider
cocos to be a synonym of Galago zanzibaricus (for example,
Groves 1977; Nash et al. 1989; Masters 1998; Zimmermann
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 6. Typical adults of three Galagoides spp. at the British Museum of
Natural History, London. Bottom to top: Kenya coast galago (Galagoides
cocos) from Gande, Kenya, Udzungwa galago (Galagoides zanzibaricus udzungwensis)
from Kissarawe, Tanzania, and Mozambique galago (Galagoides
granti) from Coguno, Mozambique. Note the great similarity in the color of the
dorsum, and that G. granti is the largest of the three species and has a wider,
fuller, tail. Photograph by Tom Butynski.
65
Galagoides in Eastern Africa
1990) or of Galagoides zanzibaricus (for example, Honess
1996; Anderson 1999, 2000; DelPero et al. 2000; Masters and
Bragg 2000; Masters and Brothers 2002).
The Need for Confirmation of the Name ‘cocos’ for the
Kenya Coast Galago
Until recently, G. zanzibaricus was considered a polytypic
species of the coastal forests from southern Somalia
through Kenya and Tanzania (including Unguja Island, Zanzibar,
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
(1939), Hill and Meester (1977)
Jenkins (1987)
Kingdon (1971, 1997), Groves (2005)
Groves (1993)
cocos a synonym of Galago
senegalensis zanzibaricus
cocos a synonym of Galago
zanzibaricus zanzibaricus
cocos a synonym of Galago
zanzibaricus
cocos a synonym of Galagoides
zanzibaricus
Groves (2001) Galago zanzibaricus cocos
Bearder et al. (2003), Grubb et al.
(2003)
Galagoides cocos
Figure 7. Two adult Zanzibar galagos (Galagoides zanzibaricus zanzibaricus)
(left) from Unguja Island, Zanzibar, Tanzania, and two adult Udzungwa galagos
(Galagoides zanzibaricus udzungwensis) (right) from Kissarawe, Tanzania. All
four specimens are at the British Museum of Natural History (BMNH), London.
The two specimens of G. z. zanzibaricus represent the extremes in pelage
coloration among the 10 adult specimens at the BMNH. Note that the dorsum,
tail, and outer front limbs are medium to bright cinnamon in G. z. zanzibaricus
and buffy-brown in G. z. udzungwensis. Photograph by Tom Butynski.

Butynski et al.
Tanzania, Malawi, and extreme eastern Zimbabwe (Hill 1953;
Groves 1977; Hill and Meester 1977; Smithers and Wilson
1979; Jenkins 1987; Courtenay and Bearder 1989; Skinner
and Smithers 1990; Groves 2001, 2005; Bearder et al. 2003;
Grubb et al. 2003). The ecology, behavior, and vocal repertoire
of the mainland subspecies, G. z. cocos, is well known, having
been the focus of detailed field studies at Diani and Gedi
Forests, Kenya (Harcourt 1986; Harcourt and Nash 1986a,
1986b). Far less well known is the nominotypical subspecies,
G. z. zanzibaricus, an endemic of Unguja Island, Zanzibar.
It was not until A. Perkin visited Unguja Island in 1998 and
recorded the species-specific advertising call of topotypical
G. zanzibaricus (Fig. 8) that it became clear that these two
forms were different:
(1) The species-specific advertising call of G. z. zanzibaricus
is very different from that of G. z. cocos. Galagoides
z. zanzibaricus has a ‘single unit rolling call’ (Fig. 8),
and G. z. cocos has an ‘incremental call’ (Fig. 9) (Honess
1996; Honess and Bearder 1996; Perkin et al. 2002;
Grubb et al. 2003).
(2) The species-specific advertising call of G. z. zanzibaricus
is identical, or nearly so, to the species-specific advertising
call of the recently named Udzungwa (or Matundu)
galago, Galagoides udzungwensis (see A. Perkin unpubl.
data, cited in Bearder 1999). This led to the realization
that G. udzungwensis may not be a new species, but rather
synonymous with, or a subspecies of, G. zanzibaricus
(see Perkin et al. 2002; Bearder et al. 2003; Grubb et al.
2003). Here we treat the Udzungwa galago as a distinct
mainland subspecies (G. z. udzungwensis), but emphasize
that the taxonomic status of the Udzungwa galago is far
from resolved (see below).
Based on a considerable body of knowledge concerning
the species-specific advertising calls of the Galagidae, and
their wide use and acceptance as a robust species recogni-
66
tion and taxonomic tool (Zimmermann et al. 1988; Courtenay
and Bearder 1989; Harcourt and Bearder 1989; Nash et al.
1989; Zimmermann 1990; Masters 1991; Bearder et al. 1995,
2003; Honess 1996; Honess and Bearder 1996; Butynski et
al. 1998; Ambrose 1999, 2003; Bearder 1999; Groves 2001;
Perkin et al. 2002), it was judged that the level of difference
between the advertising calls of G. z. cocos and G. z. zanzibaricus/G.
z. udzungwensis is far greater than can be accommodated
at the subspecies level. In fact, the advertising call of
G. z. cocos is far more similar to the advertising ‘incremental’
call of G. granti (formerly G. zanzibaricus granti) (Fig. 10)
than it is to the ‘single unit rolling’ call of G. z. zanzibaricus
(see Bearder et al. 1995). As such, G. z. cocos was reassigned
species status, G. cf. cocos (Bearder et al. 2003; Grubb et al.
2003). Interestingly, the geographic range of G. zanzibaricus
is located between the geographic ranges of G. cocos and
G. granti (see below).
Grubb and co-authors give a succinct overview of this
complex situation.
“Galagos at Diani, which were thought to be
Galagoides zanzibaricus (Harcourt and Nash, 1986a,
b) are vocally distinct from true or topotypical G. zanzibaricus
of Zanzibar. They are provisionally identified
as Galagoides cf. cocos, and we assign them to
the G. granti group. Galagos from the Udzungwa
Mtns and other localities in Tanzania have been
named Galagoides udzungwensis (Honess, 1996)
but, on the basis of their vocalization, do not differ
from those of topotypical G. zanzibaricus of Zanzibar
(A. Perkin unpubl. data, cited in Bearder, 1999). The
form udzungwensis may prove to be a valid taxon at
the subspecific level but until the systematics is clarified,
we relegate it to the synonymy of Galagoides
zanzibaricus.” (Grubb et al. 2003, pp.1315–1316).
And below
“The galago recorded from Diani is vocally distinct
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 Kingdon
(1997).

67
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).

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 northeast
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 of G. 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
A. Perkin (in litt.) reported it from the northern tip
of the East Usambara Mtns, seemingly close to G.
zanzibaricus (Table IV). The form Galago moholi
cocos Heller, 1912, was described from Mazeras
(Manzeras), relatively close to Diani (Table IV).
Groves (2001) recognized cocos as a mainland form
of Galagoides zanzibaricus. Vocalizations typical of
G. zanzibaricus have not been recorded in Kenya.
Therefore, it seems highly likely that the Diani galago
is a separate species, Galagoides cf. cocos. Nevertheless
it is important to confirm this. Vocalizations
recorded from as near to the type locality of Galagoides
cocos as possible should be compared with
the voice of the Diani galago. The forest at the type
locality (Kaya Mazeras) has been destroyed, but forests
5 km and 20 km distant (Kaya Mtswakana and
68
Kaya Fungo respectively) and some others might be
visited (Butynski unpubl., including information from
Q. Luke). Museum specimens of Galagoides cocos
should be compared with the Diani galago further to
ensure that we are dealing with a single taxon. We provisionally
consider the Diani galago to be conspecific
with Galagoides cocos, under the vernacular name
Kenya coast galago.” (Grubb et al. 2003, p.1317).
In other words, in considering the Kenya coast galago
once again a valid species, one important action remains: the
confirmation of ‘cocos’ as the species name. This is necessary
because the tape recordings used to describe the loud call for
the Kenya coast galago were made at Diani, Kenya (04°18′S,
39°35′E) (Zimmermann 1990; Bearder et al. 1995), c.40 km
south of the type locality for G. cocos (i.e., Mazeras). There
are three rivers between these two sites that are potential barri-

ers to dispersal (Maji ya Chumvi, Mambome, and Pemba/Cha
Shimba). A visit to Mazeras was, therefore, required to determine
whether the species-specific loud call of G. cocos at the
type locality is the same as that of the dwarf galago at Diani.
Note that, in the above quotation, Grubb et al. (2003)
assume that the holotype of G. cocos was collected in the forest
of ‘Kaya Mazeras’, which has since been destroyed. What
Heller (1912) actually says about the type locality is:
“The Mazeras specimens were all obtained on
the brushy borders of the cocoa-palm groves. These
groves are the dominant feature in the landscape of
the coast belt, and extend almost unbroken from the
sea beaches inland a distance of ten or fifteen miles.
They mark the tropical littoral zone more precisely
than any other plant growth.” (Heller 1912, p.2).
In short, Heller obtained the holotype (and nine other
specimens) of G. cocos in an area of coconut palms (Cocos
nucifera) and bushland, and not in forest.
Confirmation of the Name ‘cocos’ for the Kenya Coast
Galago
T. Butynski and Y. de Jong visited the Mazeras area on
10 –12 February 2004. The first night was spent searching
for G. cocos in the Mazeras Botanical Garden (03°57′58′′S,
39°33′05′′E, 134 m a.s.l.) in Mazeras town. No dwarf galagos
were heard or seen, although the small-eared greater galago,
(Otolemur garnettii), was common (about eight individuals
seen or heard).
The second night was spent searching for G. cocos
on the edge of Kaya Chijembeni (Rabai) (03°56′42′′S,
39°34′54′′E, 210 m a.s.l.), a relatively large coastal forest
located about 4 km northeast of Mazeras town, 17 km from
the Indian Ocean, and 40 km north of Diani. Coconut palms
are an extremely common species there on the forest edge
and, with cashew nut (Anacardium occidentale) and mango
(Mangifera indica), are scattered throughout the farmlands
and bush lands that surround Kaya Chijembeni. Many dwarf
galagos were heard and seen at this site (both on the forest
edge and inside the forest), and tape recordings were acquired
of ‘incremental’ advertising calls and other vocalizations. To
our ears, the advertising call recorded in Kaya Chijembeni
matched that recorded for the dwarf galago at Diani, as well
as the advertising call that T. Butynski has heard many hundreds
of times (and recorded) at others sites on the coast of
Kenya (for example, Kilifi, Watamu, Gedi), and in forests
along the lower Tana River (02°30′S, 40°30′E), c.150 km to
the north of Mazeras.
That the incremental advertising call recorded from
G. cocos at Kaya Chijembeni is the same as that recorded
from the dwarf galago at Diani is confirmed through comparisons
of the sonograms and oscillograms of these calls (Figs.
9 and 11). The advertising calls from these two sites have a
very similar incremental structure pattern, frequency range,
and call duration.
69
Galagoides in Eastern Africa
T. Butynski and Y. de Jong heard about 100 G. cocos
incremental calls during one night at Kaya Chijembeni. As at
other sites where C. cocos is similarly common, there was a
distinct ‘dusk chorus’ (c. 19:05–19:15h) of incremental calls, a
much lower rate of incremental calls throughout the night, and
a slight ‘dawn chorus’ (c. 05:45–05:55h) of incremental calls.
The only other species of galago heard at Kaya Chijembeni
was O. garnettii. Thus, only two species of galagos
were seen or heard in the Mazeras/Kaya Chijembeni/Rabai
area — G. cocos and O. garnettii. Y. de Jong and T. Butynski
returned to Kaya Chijembeni on 20 February 2006 to obtain
better (digital) recordings of the dwarf galago (Fig. 11). Again,
the only galagos heard were G. cocos and O. garnettii.
Incremental calls identical to those of topotypical G. cocos
at Mazeras have been recorded (by A. Perkin, T. Butynski,
Y. de Jong, S. Bearder, N. Cordiero, N. Svoboda, A. Kempson
and S. Gregory) at several localities along the Kenya coast
both north and south of Mazeras, as well as in the northern
lowland coastal forests of the East Usambara Mountains in
northeastern Tanzania. These calls were analyzed by A. Perkin
and S. Bearder.
Visual examination of the type G. cocos and eight other
adult specimens of G. cocos from Mazeras (housed at the
United States National Museum, Washington, DC) reveal
that they are not phenotypically different from the three specimens
of G. cocos at the National Museums of Kenya that
were collected along the Tana River (NMK 992), and on the
Kenya coast at Gedi (NMK 5351) and Kipendi (no specimen
number). One of the specimens at the National Museums of
Kenya (MR14) was collected at Mrima Hill, Kenya, very near
the border with Tanzania. This specimen has an intact penis,
the morphology of which helps to identify this population as
G. cocos (see below).
We conclude that there is now no doubt that the type
of G. cocos that Heller (1912) described from Mazeras is
conspecific with the dwarf galago found along much of the
coast of Kenya and into northern Tanzania. In short, the name
‘cocos’ can correctly be used as the species name of the Kenya
coast galago.
This clarification of the correct name for the Kenya coast
galago has at least three important implications for previous
research conducted on G. cocos, G. zanzibaricus, and
G. granti. First, most of the field research that has been conducted
on the distribution, abundance, behavior, and ecology
of ‘G. zanzibaricus’ was, in fact, conducted on G. cocos (for
example, Harcourt 1984, 1986; Harcourt and Nash 1986a,
1986b; Harcourt and Bearder 1989). Second, the distinctive
rolling calls of galagos on mainland Tanzania that led
Honess (1996) and Honess and Bearder (1996) to name this
form G. udzungwensis, belong to G. zanzibaricus (i.e., G. z.
udzungwensis). Third, those researchers who obtained data
from specimens initially assigned to ‘G. zanzibaricus’ have
sometimes, unknowingly, combined data from two species
(G. cocos and G. zanzibaricus), and, on occasion, from a third
species (G. granti). This is especially the case for specimens
collected from coastal Kenya, from coastal Tanzania south of

Butynski et al.
the East Usambara Mountains, and from Unguja Island, Zanzibar.
This means that (1) the results and conclusions of some
previous studies of ‘G. zanzibaricus’ may need to be reviewed
and reevaluated, and (2) that the providence of each and every
specimen labeled ‘G. zanzibaricus’ must be known in order
to help ensure that the specimen is not, in fact, G. cocos or
G. granti.
Morphological Differences Among G. cocos, G. zanzibaricus
and G. granti
Galagoides cocos, G. zanzibaricus, and G. granti are
among the most cryptic of primate species. That they are similar
phenotypically and morphologically is demonstrated by the
inability of some of the foremost primate taxonomists of their
time to differentiate among them. For example, Schwarz (1931),
in reference to G. senegalensis zanzibaricus, states (p. 56):
“There can be no doubt that Heller’s cocos is
identical with this race. The size, coloration, and the
large upper M³ are found both in the series at Berlin
and the one of cocos studied by Heller and Hollister.
By the identification of the two the known range of
zanzibaricus is considerably extended. There is no
difference between the island and coast specimens.”
What is needed next is a detailed comparison of large
numbers of G. cocos and G. zanzibaricus specimens to determine
their morphological differences, coupled with ecological,
behavioral, acoustic, and molecular studies. Other than
the highly distinctive species-specific advertising call, one of
the differences noted thus far is that G. cocos is slightly larger
than G. z. zanzibaricus. For example, the mean length of the
head+body for G. cocos from southeast Kenya and northeast
Tanzania is 158 mm (n = 46, range = 142–183 mm) (Appendix
I, Tables A and B), while the mean length of the head+body for
G. z. zanzibaricus from Unguja Island, Zanzibar, is 143 mm
(n = 11, range 125–150) (Appendix I, Table C). This size difference
extends to body weight; G. cocos has a mean body
weight of 144 g (n = 78, range = 117–172), whereas G. z.
zanzibaricus has a mean body weight of 127 g (n = 10, range
= 104–172).
Of the two subspecies of G. zanzibaricus, it appears that
the island form, G. z. zanzibaricus, is smaller than the mainland
form, G. z. udzungwensis (Appendix I, Tables C and
D). Mean length of the head+body for G. z. udzungwensis is
162 mm (n = 17, range = 139–180) and mean body weight is
145 g (n = 6, range = 118–105). The data available suggest
that G. granti is larger than G. z. zanzibaricus, and very similar
in size to G. z. udzungwensis and G. cocos (Appendix I,
Tables E, F and G).
Color of the nose stripe, chin, throat, cheeks and ventrum,
length of the nose stripe, and length of the tail relative
to length of the head+body, have all been proposed as useful
for distinguishing among G. cocos, G. z. zanzibaricus, G. z.
udzungwensis, and G. granti (for example, Elliot 1913; Nash
et al. 1989; Honess 1996; Groves 2001). However, our stud-
70
ies lead us to conclude that there is (1) too much intraspecific
variation and, especially, (2) too much interspecific overlap for
these characters to serve as diagnostic features.
The full range of phenotypic variation present in G. cocos,
G. z. zanzibaricus, G. z. udzungwensis, and G. granti remains
unknown. Although they need to be examined quantitatively,
and with sample sizes far larger than those currently available,
the following phenotypic characters hold promise for distinguishing
among G. cocos, G. z. zanzibaricus, G. z. udzungwensis,
and G. granti, especially when taken together:
Muzzle patches
G. cocos – patch on either side of muzzle dark, blackish, and prominent
(Figs. 1 and 2).
G. z. zanzibaricus – patch on either side of muzzle less dark, grayish, and less
prominent.
G. z. udzungwensis – patch on either side of muzzle less dark, grayish, and
less prominent (Figs. 3 and 4).
G. granti – patch on either side of muzzle less dark, grayish, and less prominent
(Fig. 5).
Ears
G. cocos – seldom longer than 38 mm, dusky behind.
G. z. zanzibaricus – seldom longer than 35 mm, dusky behind.
G. z. udzungwensis – seldom longer than 33 mm, dusky behind.
G. granti – seldom shorter than 37 mm, blackish behind. Ears not only long
but also relatively broad (Fig. 5).
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
(Fig. 7).
G. z. udzungwensis – hairs c.9 mm long, tipped buffy-brown (Figs. 6 and 7).
G. granti – hairs c.12 mm long, tipped buffy-brown with slight pinkish tint
(Figs. 5 and 6).
Tail
G. cocos – even length hairs over tail; hairs dense, c.14 mm long, soft.
Proximal c.25% of tail same color as dorsum (i.e., buffy-brown); distal
c.33% dark buffy-brown in some (Mazeras) specimens, but same color as
dorsum in other specimens (Fig. 6).
G. z. zanzibaricus – even length hairs over tail; hairs sparse, c.13 mm long,
wiry, rufous-cinnamon, cinnamon, or dusky-cinnamon (highly variable). Tail
darker/brighter cinnamon than dorsum, and either evenly colored or with
gradual darkening to reddish or dusky toward tip (Fig. 7).
G. z. udzungwensis – even length hairs over tail; hairs sparse, c.11 mm, wiry.
Proximal c.75% of tail same color as dorsum (i.e., buffy-brown); distal c.25%
slightly darker brown or dusky. Some with tail tipped white (Figs. 4, 6, and
7).
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
tail tipped white (Fig. 6).
Of these four taxa, G. z. zanzibaricus and G. granti are
phenotypically the most distinctive. The dorsum of G. z. zanzibaricus
is cinnamon and the tail has at least some rufous,
whereas the dorsum of the other three taxa is buffy-brown and
all lack rufous in the tail (Fig. 7). The color of the dorsum of G.
cocos, G. z. udzungwensis, and G. granti is virtually identical,
although there is a slight pinkish tint to the dorsum of G. granti
when seen in good light (Fig. 6).
When observed at close range in the field, G. granti is
distinguished from G. cocos, G. z. zanzibaricus, and G. z.
udzungwensis by its relatively large, broad, round, and blackish
(behind) ears, and by the very full, bottlebrush-shaped tail,
which is blackish-brown over the distal part (Figs. 5 and 6).

The particularly large ears of G. granti have been noted previously
(for example, Honess 1996; Masters and Bragg 2000).
As a species, G. zanzibaricus is probably best distinguished
phenotypically from G. cocos and G. granti by the
relatively short, wiry, stiff hairs over the tail. This hair type
makes it relatively easy to see the skin of the tail through the
pelage (Fig. 4). Galagoides cocos and G. granti have relatively
long, soft, lax hairs over the tail.
Of the four taxa considered here, G. cocos and G. z.
udzungwensis are, phenotypically, the most difficult to distinguish
from one another (Fig. 6). The presence in G. cocos of
a prominent dark, blackish patch on either side of the muzzle
is probably the best phenotypic character available for distinguishing
C. cocos and G. z. udzungwensis in the field (Figs.
1–4) (A. Perkin pers. obs.). As already stated, however, all of
the characters listed above are in need of detailed quantitative
study in order to determine their reliability, both in the field
and in the museum.
The penile morphology of G. cocos, G. zanzibaricus,
and G. granti is diagnostic (Fig. 12). For details, see Hon-
Figure 12. Schematic drawings showing the penile morphology of: 1 – Galagoides
cocos; 2 – Galagoides granti; 3 – Galagoides zanzibaricus udzungwensis.
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
Perkin (in press).
71
Galagoides in Eastern Africa
ess (1996), Honess and Bearder (1996), Anderson (2000), and
Perkin (in press). See sketches in Kingdon (1997).
Vision plays an important role in the life histories of all
galagos, perhaps especially for species recognition. Detailed
study of the light and dark facial markings of these four taxa
may reveal that they are species-specific and, therefore, a
useful diagnostic tool. This is not only a priority topic for
research related to the search for species-typical differences
among G. cocos, G. z. zanzibaricus, G. z. udzungwensis, and
G. granti, but also among the many other cryptic taxa within
the Galagidae (Bearder 1999; Bearder et al. 2006).
The differences noted here among G. cocos, G. zanzibaricus,
and G. granti in their species-specific advertising calls,
body measurements, and phenotypic characters are consistent
with the species-level differences observed for other species
in the Galagidae (Honess 1996; Honess and Bearder 1996;
Masters and Bragg 2000; Masters and Brothers 2002).
In contrast to the great similarity among G. cocos, G. zanzibaricus,
and G. granti, these three species are readily distinguished
from the other seven species of galagos with which
one or all are sympatric or parapatric. These are O. garnettii,
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 identified
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 northeastern
Tanzania (Figs. 13 and 14) (A. Perkin in litt. in Grubb
et al. 2003). Although there is a complex mosaic of habitat
types in this region, preliminary observations indicate that
G. cocos is present in the dry mixed coastal forests and mixed
woodland of the northernmost forests of Tanzania’s coastal
strip (for example, Bombo East I and Bombo East II For-

Butynski et al.
Nairobi
Arusha
Machakos
TANZANIA
0 50
0
Athi River
Kiboko
Same
kilometers
G. cocos locality
kilometers
Galana River
LEGEND
Mwingi
Kitui
G. cocos distribution
100
G. cocos unconfirmed
50 100
Voi
Amani
Pangani River
KENYA
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
Sabaki River
Tanga
Mazeras
Zanzibar
(Unguja)
Diani
Tana River
Garsen
Mombasa
37° 38° 39° 40°
Pemba
Hola
Gedi
Malindi
INDIAN OCEAN
- 1°
- 2°
- 3°
- 4°
- 5°

Farkwa
Babati
Kondoa
Dodoma
Iringa
Kihansi
TANZANIA
Ifakara
Kilosa
Great Ruaha River
Mahenge
Mang'ula
Mikumi
Morogoro
Same
Pangani River
Rufiji River
73
Galagoides in Eastern Africa
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: ).
Handeni
0
Amani
Wami River
Rufiji River
Ruvu River
Utete
LEGEND
50 100
kilometers
KENYA
Tanga
Zanzibar
(Unguja)
Dar es Salaam
Kisiju
G. zanzibaricus distribution
G. zanzibaricus locality
Kilwa Kivinje
G. zanzibaricus unconfirmed
Mombasa
36° 37°
38° 39° 40°
Pemba
INDIAN OCEAN
Mafia
- 3°
- 4°
- 5°
- 6°
- 7°
- 8°

Butynski et al.
est Reserves), as well as in the more moist (but relatively
tree-species and bird-species poor) groundwater forests and
adjacent woodlands on the lower northern slopes of the East
Usambara Mountains (for example, Mgambo Forest Reserve)
(A. Perkin and N. Cordeiro unpubl. data). In this region, G. z.
udzungwensis appears to be confined to the more moist (and
relatively tree-species and bird-species rich) forest on the
eastern slopes of the East Usambara Mountains (for example,
Kambai, Segoma, Manga, and Marimba forest reserves) (A.
Perkin unpubl. data).
Galagoides z. udzungwensis and G. granti appear to be
parapatric at the Kilombero-Rufiji River with G. z. udzungwensis
reaching its southern limit on the north (left) bank
(Fig. 14) and G. granti reaching its northern limit on the
south (right) bank (Fig. 15) (Honess 1996; A. Perkin in litt.
in Grubb et al. 2003). In addition, the geographic ranges of
G. z. udzungwensis and G. granti either approach one another
or meet at the southern end of the Udzungwa Mountains in
south-central Tanzania. Here, G. granti is present at 1,500 to
1,800 m a.s.l. in the Lulanda Forest Reserve at the headwaters
of the Kilombero River (A. Perkin in litt. in Grubb et al.
2003; A. Perkin unpubl. data), and G. z. udzungwensis is present
from 400 to 1,070 m a.s.l. at Kihanzi (which is c.24 km
to the east of Lulanda) (Honess 1996; Butynski et al. 1998;
N. Cordeiro pers. comm.). Kihanzi is the known southwestern
limit for G. z. udzungwensis, and Lulanda is the known northwestern
limit for G. granti.
David Moyer and E. Mulungu (pers. comm.) tape
recorded the loud call of a Galagoides sp. at three sites in
extreme western Tanzania: Mbala Forest, Sitebe-Sifuta
Mountains (6º04′40′′S, 30º32′10′′E, 1,700 m a.s.l., 16 August
2005), Mahale Mountains National Park at Mfitwa Mountain
(6º07′55′′S, 29º47′38′′E, 2,440 m a.s.l., 20 November 2005),
and at Pasagulu Mountain (6°03′47′′S, 29°45′14′′E, 1,500 m
a.s.l., 6 February 2006). The species recorded may have been
G. granti. If so, this extends its geographic range c.700 km to
the northwest (Fig. 15).
Groves (2001) identified three specimens collected in submontane
forest at Bagilo (800 –1,000 m a.s.l.) in the Uluguru
Mountains (south of Morogoro, Fig. 15) as G. cf. granti (Grubb
et al. 2003). If these are G. granti, then the Uluguru Mountains
represent the northernmost site for this species, and the only
known site for G. granti that is north of the Rufiji River. These
three specimens had earlier been identified by Lawrence and
Washburn (1936) as G. senegalensis zanzibaricus. A. Perkin
has since examined them at Harvard University’s Museum
of Comparative Zoology (specimen numbers: 22450, 22449,
22451) and also ascribes them to G. zanzibaricus. In 1993,
P. E. Honess and S. K. Bearder (Honess 1996) visited the forest
at Bagilo but did not find G. granti there — only G. orinus was
present. Unfortunately, there is no longer any forest at Bagilo
(Perkin 2000; Doggart et al. 2004) but A. Perkin (unpubl. data)
found G. zanzibaricus at 900 m a.s.l. on the edge of the Uluguru
North Forest Reserve (c.2 km west of Bagilo Village) and
up to 700 m a.s.l. in the Uluguru Mountains.
74
The known southern limit for G. granti is the Limpopo
River in southern Mozambique. The western limit in the southern
part of the range appears to be extreme eastern Zimbabwe
(Smithers and Lobão Tello 1976; Smithers and Wilson 1979;
Skinner and Smithers 1990).
In summary, present information indicates that G. cocos,
G. zanzibaricus, and G. granti are parapatric, or narrowly sympatric,
species of the evergreen forests of the coastal strip of
eastern Africa from northern Kenya (perhaps southern Somalia)
to extreme southern Mozambique and extreme eastern Zimbabwe.
Galagoides cocos is the northern species, G. zanzibaricus
is the central species, and G. granti is the southern species.
Conservation Status of G. cocos, G. zanzibaricus, and
G. granti
Galagoides cocos, G. zanzibaricus, and G. granti now
survive in highly fragmented, probably declining, populations
as a result of the extensive (>65%) loss of eastern Africa’s
original coastal forest cover. More than 90% of the original
coastal forest of Kenya and Tanzania has either been destroyed
or degraded (Burgess et al. 2004). Fortunately, all three species
are able to persist in secondary forest and in mosaics of mixed
agriculture where some forest remains.
As a recently resurrected species (Grubb et al. 2003),
G. cocos does not yet appear on the IUCN Red List. It 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 and Nash 1986a). Galagoides
cocos is found at elevations from sea level to at least 210
m a.s.l. over the coastal zone of Kenya (T. Butynski and Y. de
Jong unpubl. data), and to at least 350 m a.s.l. in the foothills
of the East Usambara Mountains, Tanzania (A. Perkin unpubl.
data). The information available indicates that G. cocos, when
assessed for the IUCN Red List using the 2001 criteria (IUCN
2001), will be placed in the Least Concern category.
G. zanzibaricus is listed as Lower Risk/Near Threatened
on the 2006 IUCN Red List (IUCN 2006), but assessed only
using the 1994 criteria (IUCN 1994). G. zanzibaricus is the
most abundant and widespread galago in the coastal forests
of Tanzania, including ‘coastal’ forest sites located at least
370 km inland (for example, Udzungwa Mountains) to c.1,100
m a.s.l. The density of G. zanzibaricus varies greatly from site
to site. In the Udzungwa Mountains (for example, Matundu
Forest Reserve), G. z. udzungwensis is estimated to occur at
densities of more than 500 individuals/km² (Butynski et al.
1998), whereas 200 individuals/km²
in Jozani–Chwaka Bay National Park (T. Butynski
and Y. de Jong pers. obs). The information available indicates
that G. zanzibaricus, when reassessed for the IUCN Red List,
should be placed in the ‘Least Concern’ category.

Kabala
DRC
Lubumbashi
Francistown
Ndola
Lusaka
Bulawago
?
Lake Tanganyika
Pietersburg
Kigoma
?
ZIMBABWE
Kasama
Harare
Masvingo
SOUTH AFRICA
ZAMBIA
Chipata
Tabora
TANZANIA
Songo
Limpopo River
Mbeya
Chimoio
Maputo
MALAWI
Lake Malawi
Lilongwe
Coguno
Lulanda
Zambezi River
Tambarara
Songea
Beira
Dodoma
Iringa
Blantyre
Pembe
75
Rufiji River
Galagoides in Eastern Africa
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: ).
Mang'ula
Cuamba
Quelimane
Same
Morogoro
Mahenge
Ruvuma River
Tanga
MOZAMBIQUE
0 200
Mombasa
Dar es Salaam
Lindi
Nacala
Pemba
Zanzibar
(Unguja)
LEGEND
G. granti distribution
G. granti locality
G. granti unconfirmed
kilometers
400
INDIAN OCEAN
28° 30° 32° 34° 36° 38° 40° 42° 44°
Mafia
- 4°
- 6°
- 8°
- 10°
- 12°
- 14°
- 16°
- 18°
- 20°
- 22°
- 24°
- 26°

Butynski et al.
The two subspecies, G. z. zanzibaricus and G. z. udzungwensis,
have not as yet been assessed for the IUCN Red List.
While G. z. udzungwensis is a widespread subspecies that
will likely be assessed as Least Concern, G. z. zanzibaricus
has a far smaller ‘extent of occurrence’, being endemic to
Unguja Island, Zanzibar (c.2,000 km²) where rates of habitat
degradation, loss, and fragmentation are particularly high.
As such, G. z. zanzibaricus is expected to be assessed as an
Endangered taxon.
Galagoides granti, assessed under the 1994 criteria
(IUCN 1994), is listed as Data Deficient in the 2006 IUCN
Red List (IUCN 2006). Galagoides granti is present in coastal
and submontane evergreen forest, gallery forest, and speciesrich
woodlands, including some hilly miombo woodlands
(e.g., Mahenge foothills). In the southern part of its range, G.
granti is present between the coast and about 200 km inland
(i.e., eastern Zimbabwe) up to 360 m a.s.l. (Smithers and
Wilson 1979; Skinner and Smithers 1990). According to the
specimen tags, C. H. B. Grant collected this species up to 400
m a.s.l. at Tambarara in central Mozambique. In the northern
part of its range, G. granti occurs from the coast to at least
310 km inland (i.e., Lulanda, Tanzania) up to at least 1,800 m
a.s.l. Galagoides granti occurs over a much greater range of
habitat types, altitudes, and climates than does G. cocos or G.
zanzibaricus. There is now enough information available for
an IUCN Red List assessment of this species. The available
data indicate that G. granti, once reassessed, will be placed in
the Least Concern category. Galagoides cocos, G. zanzibaricus,
and G. granti are all currently CITES Appendix II species
(http://www.cites.org).
Acknowledgments
We thank David Moyer and Esteban Sarmiento for their
unpublished data, the curatorial staff at the United States
National Museum, British Museum of Natural History, and
American Museum of Natural History for providing access
to their collections; Harald Schuetz for the photograph in Figure
1; and Nike Doggart for the photograph in Figure 3. We
appreciate the support given to our research on primates in
eastern Africa by the Margot Marsh Biodiversity Foundation,
Conservation International, Zoo Atlanta, the National Museums
of Kenya, the IUCN Eastern Africa Regional Office, the
Kenya Institute of Primate Research, the Critical Ecosystem
Partnership Fund, the Primate Action Fund, Primate Conservation
Inc., the Leverhulme Trust, Oxford Brookes University,
WWF-Tanzania, the Tanzanian Commission for Science and
Technology, Tanzania Forestry Division, Tanzania National
Parks, Kenya Wildlife Service, and Kenya Ministry of Education,
Science and Technology. Norbert Cordeiro, Nadine
Svoboda, Alex Kempson, and Stephen Gregory provided tape
recordings of the advertising calls of G. cocos. Norbert Cordeiro
assisted with vegetation descriptions and unpublished
data. Julie Anderson and the Colobus Trust helped with surveys
funded by the Born Free Foundation.
76
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Authors’ addresses:
Thomas M. Butynski, Eastern Africa Regional Program,
Conservation International, c/o IUCN EARO, P.O. Box 68200,
00200 Nairobi, Kenya.
Yvonne A. de Jong, Kenya Institute of Primate Research, P.O.
Box 38, Ol Kalou 20303, Kenya.
Andrew W. Perkin and Simon K. Bearder, Nocturnal Primate
Research Group, Anthropology Department, School of
Social Sciences and Law, Oxford Brookes University, Oxford
OX3 0BP, UK.
Paul E. Honess, Department of Veterinary Services, University
of Oxford, Oxford OX1 3PT, UK.
Received for publication: February 2006
Revised: May 2006
Appendix I
Body measurements for adults of the Kenya coast galago (Galagoides
cocos), the Zanzibar galago (Galagoides zanzibaricus), and the
Mozambique (galago (Galagoides granti). All measurements are either
known to be, or believed to be, from living or fresh adult specimens.
Table A. Measurements for eight Galagoides cocos collected at Mazeras, Kenya,
the type locality for G. cocos (Hollister 1924). All measurements taken
from the tags of specimens collected by E. Heller and housed at the United
States National Museum (specimen numbers: 181810, 184218, 184219,
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.
Measurement Mean SD Range
Sample
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
Table B. Measurements for live specimens of Galagoides cocos. These data
derive from the following two sources: (1) Harcourt and Nash (1986b) for G.
cocos at Gedi (c.100 km northeast of Mombasa) and Diani (30 km south of
Mombasa) on the coast of Kenya; (2) A. Perkin (unpubl. data) for G. cocos at
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).
Because adult male and adult female body linear measurements for G. cocos
are not significantly different, the data for males and females are combined in
this sample (Harcourt and Nash 1986b). Because body weights of adult male
and adult (non-pregnant) female G. cocos are significantly different (Harcourt
and Nash 1986b), the body weight data are presented separately for each sex.
Measurement Mean Range
Sample size
(n)
Head + body length 159 mm 142–183 mm 38
Tail length 212 mm 182–230 mm 30
Hindfoot length 54 mm 40 –57 mm 49
Ear length 30 mm 23 – 40 mm 32
Body weight (males) 150 g 135–167 g 36
Body weight (females) 138 g 117–172 g 42
Table C. Combined measurements for male and female Galagoides zanzibaricus
zanzibaricus from Unguja Island, Zanzibar, Tanzania, the type locality for
G. zanzibaricus. Measurements taken from one specimen captured and released
by A. Perkin and from the tags of 10 specimens housed at the British Museum
of Natural History. Nine of these collected by W. H. R. Lumsden (specimen
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
(specimen number: 1955.331).
Measurement Mean SD Range
Sample size
(n)
Head+body length 143 mm 8 125–150 mm 11
Tail length 214 mm 12 198–235 mm 11
Hindfoot length 56 mm 3 51–59 mm 11
Ear length 32 mm 2 30 –35 mm 11
Body weight 127 g 20 104–172 g 10
Table D. Combined measurements for male and female Galagoides zanzibaricus
udzungwensis obtained from the following sites in Tanzania; two from
Matundu Forest Reserve (Honess 1996); two from Kissarawe (housed at the
British Museum of Natural History); two from Pugu Forest Reserve, three from
Pande Game Reserve (A. Perkin, unpubl. data); four from Bagilo, Uluguru
Mountains; and four from Amboni, near Tanga. These last eight specimens
were collected by A. Loveridge and are housed at the Museum of Comparative
Zoology, Harvard University.
Measurement Mean SD Range
Sample size
(n)
Head+body length 162 mm 11 139–180 mm 17
Tail length 222 mm 16 202–270 mm 17
Hindfoot length 58 mm 6 50 –70 mm 17
Ear length 31 mm 3 25–37 mm 17
Body weight 145 g 27 118–195 g 6

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).
Measurement Mean SD Range
Sample size
(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).
Measurement Mean Range
Sample size
(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 Lulunda,
Udzungwa Mountains (A. Perkin unpubl. data).
Measurement Mean SD Range
Sample size
(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
Galagoides in Eastern Africa

By James Culverwell, Jim Feely,
Sheila Bell-Cross, Yvonne A. de
Jong and Thomas M. Butynski
There are only two species
of warthog, and
both occur in Kenya;
the common warthog
Phacochoerus africanus
and the desert
warthog Phacochoerus aethiopicus.
Little is known about the natural
history of the desert warthog—indeed,
the desert warthog might well
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 hookshaped
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 ‘eggshaped’
when viewed from the front. Head
of common warthog is slightly ‘diaboloshaped’
when viewed from the front.
A new pig for
Tsavo
be Africa’s least-known, non-forest,
large mammal. The morphological
differences between the two species
of warthog are described by Grubb
(1993) and by d’Huart & Grubb
(2005). The most diagnostic and
noticeable characteristics for both
species are summarized in Box 1.
d’Huart & Grubb (2001)
compiled a map depicting the
geographical range of both the
common warthog and the desert
warthog in the Horn of Africa (Eritrea,
Djibouti, Somalia, Ethiopia,
Uganda and Kenya). Their findings
were summarized in 2002 by
Boy in Swara 25—1: 20-21. The
authors found the desert warthog
to occur from Puntland (northern
Somalia) southwestward through
Somalia and southeastern Ethiopia
to central and eastern Kenya.
The only sites shown on the map
for Kenya were Moyale, El Wak,
Merelle (‘midway between Archer’s
Post and Mt Marsabit’), and
Mkokoni (in the Kiunga Marine
Reserve). The southernmost
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 diaboloshaped
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

Figure 3. Desert warthog
Phacochoerus aethiopicus and
common warthog Phacochoerus
africanus localities in Tsavo East
National Park and Tsavo West
National Park, southeast Kenya.
record for Kenya was from the
Kiunga Marine Reserve, about 60
kilometres northeast of Lamu. All
records were from north of the
Ewaso Ng’iro River and east of the
Tana River. They speculated that
the two species would be found
to overlap in some parts of their
range, but found no evidence for
this; their data show the shortest
distance between common warthog
and desert warthog records as
25 kilometres (in Puntland).
During a primate survey in
2005, TMB and YDJ encountered
two solitary desert warthogs, 15
and 80 kilometres southwest of
Garissa respectively, in medium
dense acacia bushland. These encounters
are important, as they are
the first records of desert warthog
west of the Tana River. Two questions
remained, however. Are
the desert warthog and common
SWARA July – September 2008
warthog sympatric? How far south
does the desert warthog occur?
On 7 June 2007, JC, JF and
SBC traversed Tsavo East National
Park. Briefed a few days
earlier about desert warthog characteristics
by TMB and YDJ, they
travelled through the high-density
tourism areas south of the Voi
River, far to the southwest of the
nearest confirmed desert warthog
locations. Late that afternoon they
were startled to find two adult
desert warthogs in low scrub on
the edge of the Dika Plains, just
13 kilometres north of Buchuma
Gate, and managed to take a few
photographs before the warthogs
turned tail. They had just recorded
a new large mammal species for
Tsavo East National Park, and a
major range extension of over 300
kilometres for the desert warthog!
The next day they returned
to the area and located four more
desert warthogs only nine kilometres
from the Kenya Wildlife
Service headquarters at Voi; one
adult and three subadults stood
their ground long enough for positive
identification to be made and
for more photographs to be taken.
What struck JC, JF and SBC immediately
during these encounters
was the overall shape of the animals’
heads, the hooked warts, and
the flipped-back tip of the ears (see
Box 1), all of which are diagnostic,
easily-seen field characters of
the desert warthog. During this
trip, no common warthogs were
seen. All photographs were sent to
experts for confirmation. Some of
these, together with some of our
other warthog photographs, are
available on a digital map at www.
tinyurl.com/warthogMap.
In July 2007, JC returned briefly
to Tsavo East National Park
for two days. He observed two
sounders of desert warthogs—each
of four animals—in open shrubland
north and south of the Voi
River in the vicinity of Satao Safari
Camp. Once again, no common
warthogs were seen.
In October 2007, JC visited
Tsavo West National Park for two
days, where desert warthogs were
found to be fairly common around
Severin Safari Camp and Kitani
FINDINGS
51

Safari Lodge, just to the north
of the Tsavo River. Three desert
warthogs were seen feeding in an
open area along a major tourism
circuit, permitting lengthy observation.
One of these animals was
a subadult male. Subadult desert
warthogs, like adults, have ears
that are flipped-back at the tips,
and broader-based genal warts, allowing
them to be differentiated in
the field from common warthogs
despite them lacking the hooked
warts of adult male desert warthogs.
During this visit JC also found
common warthogs in Tsavo West
National Park only a few hundred
metres from where he observed
desert warthogs.
In April, 2008, TMB and YDJ
visited Tsavo West National Park
for three days, and encountered
common warthogs around Kilaguni
Serena Safari Lodge, Komboyo
Camp, Kitani Safari Lodge and
Severin Safari Camp (including
areas where desert warthogs were
seen during the earlier trip by JC).
About six kilometres of Finch Hatton’s
Camp, in low bush on the
edge of riverine forest, one group
of desert warthogs (one adult male
and three subadults) was encountered
only 150 metres from six
common warthogs. This sighting
is not only the farthest range
extension for desert warthog (390
kilometres) from Mikoni, the
most southern record of d’Huart
& Grubb, but it is also the first
evidence that the two species of
warthogs are sympatric.
Finding desert warthog in
Tsavo, and finding them sympatric
with common warthog, is exciting
news for the species and for
Kenya. Tsavo West National Park
and Tsavo East National Park can
add one more species to their list
of large mammals. What further
surprises can we expect from this
long-overlooked pig?
Have you seen
warthogs in Africa?
References
• Boy, G. 2002. The whole hog.
Swara 25 (1): 20-21.
• D’Huart, J. & Grubb, P. 2001. Distribution
of the common warthog
(Phacochoerus africanus) and the
desert warthog (Phacochoerus
aethiopicus). African Journal of
Ecology 39: 156-169.
• Grubb, P. 1993. The Afrotropical
suids Phacochoerus, Hylochoerus,
and Potamochoerus: taxonomy
and description. In: Pigs, Peccaries
and Hippos. Status Survey and
Conservation Action Plan (ed. W.
I. R. Oliver). IUCN/SSC, Gland,
Switzerland
• iucn.org/themes/ssc/sgs/pphsg/
Contents.htm
• 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 (2): 4-8
• iucn.org/themes/ssc/sgs/pphsg/
Suiform%20soundings/Newsletter%205(2)pdf
To better understand the distribution
of Africa’s warthogs, we would like to
know if you have seen desert warthogs
or common warthogs in Eritrea, Ethiopia,
Djibouti, Somalia, Uganda, Kenya or
Tanzania. Details and/or a photograph of
your record would be highly appreciated.
The following information is most
important
• date of sighting
• species
• name of nearest village or town
• latitude and longitude
• elevation
• habitat
Please send your information and/or
photograph to:
Yvonne de Jong
PO Box 149
Nanyuki 10400, Kenya
yvonne@wildsolutions.nl
Thank you!
Your help is much appreciated!
52 SWARA July – September 2008

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 anyuki, Kenya, yvonne@wildsolutions.nl
²8 Manga Gardens, Kitisuru Rd, airobi, Kenya, jc@lubombo.net
³P.O. Box 149, 10400 anyuki, Kenya, tbutynski@aol.com
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 between
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 encountered
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 Mkokoni,
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. Although
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 SOUNDINGS VOL 8(2) PAGE 4

Figure 1: Adult male common warthog
Phacochoerus africanus on the plains of the Laikipia
Plateau, central Kenya. Note the pointed
ears, the cone-shaped warts, the ‘diabolo-shaped’
head, and the lack of swelling of the suborbital
area.
Figure 2: Adult male desert warthog Phacochoerus
aethiopicus in medium-dense shrub in Tsavo
West National Park, southern Kenya. Note the
flipped-back ear tips, the hooked warts, the broad,
‘egg-shaped’, head, and the swollen suborbital
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 National
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 SOUNDINGS VOL 8(2) PAGE 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 deleted
from the list of large mammals known for Tsavo East National Park. As concerns the desert warthog,
Tsavo East National Park and Tsavo West National Park can now add one more species to their already
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 desert
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 Chicago
Press, Chicago, USA.
Kingdon J. 1997. The Kingdon Field Guide to African mammals. Academic Press, New York, USA.
SUIFORM SOUNDINGS VOL 8(2) PAGE 6