Primates in Peril
e World’s 25 Most Endangered Primates
Christoph Schwitzer, Russell A. Mittermeier,
Anthony B. Rylands, Federica Chiozza, Elizabeth A. Williamson,
Janette Wallis and Alison Cotton
Illustrations by Stephen D. Nash
IUCN SSC Primate Specialist Group (PSG)
International Primatological Society (IPS)
Conservation International (CI)
Bristol Zoological Society (BZS)
Published by: IUCN SSC Primate Specialist Group (PSG), International Primatological Society (IPS),
Conservation International (CI), Bristol Zoological Society (BZS)
Copyright: ©2015 Conservation International
All rights reserved. No part of this report may be reproduced in any form or by
any means without permission in writing from the publisher.
Inquiries to the publisher should be directed to the following address:
Russell A. Mittermeier, Chair, IUCN SSC Primate Specialist Group,
Conservation International, 2011 Crystal Drive, Suite 500, Arlington, VA 22202, USA.
Citation (report): Schwitzer, C., Mittermeier, R.A., Rylands, A.B., Chiozza, F., Williamson, E.A., Wallis, J. and Cotton, A.
(eds.). 2015. Primates in Peril: e World’s 25 Most Endangered Primates 2014–2016. IUCN SSC Primate
Specialist Group (PSG), International Primatological Society (IPS), Conservation International (CI), and
Bristol Zoological Society, Arlington, VA. iv+93pp.
Citation (species): Butynski, T.M. and Hamerlynck, O. 2015. Tana River red colobus Piliocolobus rufomitratus (Peters,
1879). In: C. Schwitzer, R.A. Mittermeier, A.B. Rylands, Chiozza, F., E.A. Williamson, J. Wallis and A.
Cotton (eds.), Primates in Peril: e World’s 25 Most Endangered Primates 2014–2016, pp. 20–22.
IUCN SSC Primate Specialist Group (PSG), International Primatological Society (IPS), Conservation
International (CI), and Bristol Zoological Society, Arlington, VA.
illustrations: © Stephen D. Nash, Conservation International, Arlington, VA, and Department of
Anatomical Sciences, Health Sciences Center, Stony Brook University, Stony Brook, NY, USA.
Available from: Jill Lucena, Conservation International, 2011 Crystal Drive, Suite 500, Arlington, VA 22202, USA.
Printed by: Tray, Glen Burnie, MD, USA
Front cover photo: Chamba sacred langur (Semnopithecus ajax). Photo © WILD / Himalayan Langur Project
Back cover photo: Ecuadorian brown-headed spider monkey (Ateles fusciceps fusciceps). Tesoro Escondido, Esmeraldas,
Ecuador. Photo © Juan de Dios Morales
e World’s 25 Most Endangered Primates: 2014–2016..........................................1
Rondo dwarf galago (Galagoides rondoensis) ....................................................................11
Roloway monkey (Cercopithecus diana roloway ................................................................14
Preuss’s red colobus (Piliocolobus preussi) .........................................................................17
Tana River red colobus (Piliocolobus rufomitratus) .......................................................20
Grauer’s gorilla (Gorilla beringei graueri) .........................................................................23
Madagascar ................................................................................................................ 26
Lavasoa Mountains dwarf lemur (Cheirogaleus lavasoensis) .........................................27
Lac Alaotra bamboo lemur (Hapalemur alaotrensis) ........................................................30
Red rued lemur (Varecia rubra) ....................................................................................33
Northern sportive lemur (Lepilemur septentrionalis) ....................................................36
Perrier’s sifaka (Propithecus perrieri) ...................................................................................38
Philippine tarsier (Carlito syrichta) ......................................................................................43
Javan slow loris (Nycticebus javanicus) ................................................................................45
Pig-tailed snub-nosed langur (Simias concolor) ...............................................................50
Delacour’s langur (Trachypithecus delacouri) .....................................................................53
Golden-headed langur or Cat Ba langur (Trachypithecus poliocephalus) .......................56
Tonkin snub-nosed monkey (Rhinopithecus avunculus) ..................................................58
Chamba sacred langur (Semnopithecus ajax).....................................................................60
Western purple-faced langur (Semnopithecus vetulus nestor) .......................................63
Hainan gibbon (Nomascus hainanus) ..................................................................................67
Sumatran orangutan (Pongo abelii) ......................................................................................70
Brown spider monkey (Ateles hybridus)..............................................................................74
Ecuadorian brown-headed spider monkey (Ateles fusciceps fusciceps) .........................76
Ka’apor capuchin (Cebus kaapori) .......................................................................................78
San Martín titi monkey (Callicebus oenanthe) ....................................................................81
Northern brown howler (Alouatta guariba guariba) .........................................................83
Editors’ addresses .....................................................................................................87
Contributors’ addresses .............................................................................................87
Here, we present the 2014–2016 iteration of the list of the World’s 25 Most Endangered
Primates, drawn up during an open meeting held during the XXV Congress of the International
Primatological Society (IPS), Hanoi, 13 August 2014.
We have updated the species proles from the 2012–2014 edition (Schwitzer et al. 2014) for
those species remaining on the list, and added additional proles for newly listed species.
is publication is a joint eort of the IUCN SSC Primate Specialist Group, the International
Primatological Society, Conservation International, and the Bristol Zoological Society.
We are most grateful to the Margot Marsh Biodiversity Foundation for providing signicant
support for research and conservation eorts on these endangered primates through the direct
provision of grants and through the Primate Action Fund, administered by Ms. Ella Outlaw, of
the Executive Vice Chair’s Oce at Conservation International. Over the years, the foundation
has provided support for the training workshops held before the biennial congresses of the
International Primatological Society and helped primatologists to attend the meetings to discuss
the composition of the list of the world’s 25 most endangered primates.
We would like to thank all of the authors who contributed to the nal 2014–2016 version:
Vishal Ahuja, Martina Anandam, Nary R. J. Andrianjaka, Matthew A. Banks, omas M.
Butynski, Bosco P. L. Chan, Lounès Chikhi, Fay Clark, Ana Gabriela de Luna, Marcos de Souza
Fialho, Dong anh Hai, Andrew Dunn, Leonardo Gomes Neves, Sharon Gursky, Olivier
Hamerlynck, Andreas Hapke, Leandro Jerusalinsky, Daphne Kerhoas, Le Khac Quyet, Neahga
Leonard, Joshua M. Linder, Andrés Link, Edward E. Louis Jr., W. Scott McGraw, Fabiano R.
Melo, Stefan Merker, Russell A. Mittermeier, Sanjay Molur, Alba Lucia Morales-Jiménez, Bethan
J. Morgan, Paola Moscoso-R., Tilo Nadler, K. Anna I. Nekaris, Vincent Nijman, Stuart Nixon,
Matthew G. Nowak, John F. Oates, Perry S. Ong, Lisa M. Paciulli, Richard J. Passaro, Erik R.
Patel, Andrew Perkin, Phan Duy uc, Martina Rael, Christian Roos, Rasanayagam Rudran,
Anthony B. Rylands, Jordi Salmona, Daniela Schrudde, Christoph Schwitzer, Myron Shekelle,
Ian Singleton, Roswitha Stenke, Pablo Stevenson, Jatna Supriatna, Maurício Talebi, Dana iele,
Diego G. Tirira, Graham Usher, Jan Vermeer, Serge A. Wich, Elizabeth A. Williamson, Lance
Woolaver, John R. Zaonarivelo.
Schwitzer, C., Mittermeier, R.A., Rylands, A.B., Taylor, L.A., Chiozza, F., Williamson, E.A.,
Wallis, J. and Clark, F.E. (eds.). 2014. Primates in Peril: e World’s 25 Most Endangered Primates
2012–2014. IUCN SSC Primate Specialist Group (PSG), International Primatological Society
(IPS), Conservation International (CI), and Bristol Zoological Society, Arlington, VA. 87pp.
Here we report on the eighth iteration of the biennial
listing of a consensus of the 25 primate species considered
to be among the most endangered worldwide and the
most in need of conservation measures.
e 2014–2016 list of the world’s 25 most endangered
primates has ve species from Africa, ve from
Madagascar, ten from Asia, and ve from the
Neotropics (Table 1). Madagascar tops the list with ve
species. Indonesia and Vietnam both have three, Brazil
two, and Cameroon, China, Colombia, Côte d’Ivoire,
the Democratic Republic of Congo, Ecuador, Ghana,
India, Kenya, Nigeria, Peru, the Philippines, Sri Lanka,
Tanzania and Venezuela each have one.
e changes made in this list compared to the previous
iteration (2012–2014) were not because the situation
of the eight species that were dropped (Table 2)
has improved. In some cases, such as, for example,
Microcebus berthae, the situation has in fact worsened,
due to ongoing deforestation in this species’ small
distribution range in western Madagascar. By making
these changes we intend rather to highlight other,
closely related species enduring equally bleak prospects
for their survival. One species for which the situation
may have improved since it was rst added to the list in
2008 is Eulemur avifrons, Sclater’s black lemur. While
severe threats to this species remain in large parts of its
range, some populations inside the Sahamalaza – Îles
Radama National Park are now under more eective
protection, mainly owing to a long-term research and
monitoring programme that has been active in this
protected area since 2004.
e World’s 25 Most Endangered Primates: 2014–2016
Eight of the primates were not on the previous (2012–
2014) list (Table 3). Four of them are listed as among the
world’s most endangered primates for the rst time. e
Lac Alaotra bamboo lemur, Perrier’s sifaka, the Hainan
gibbon and the Sumatran orangutan had already been
on previous iterations, but were subsequently removed
in favour of other highly threatened species. e 2014–
2016 list contains two members each of the genera
Piliocolobus, Trachypithecus, Semnopithecus and Ateles,
thus particularly highlighting the severe threats that
large primates are facing in all of the world’s primate
During the discussion of the 2014–2016 list at the XXV
Congress of IPS in Hanoi in 2014, a number of other
highly threatened primate species were considered for
inclusion (Table 4). For all of these, the situation in the
wild is as precarious as it is for those that nally made
it on the list.
Tabl e 1. e World’s 25 Most Endangered Primates 2014–2016.
Galagoides rondoensis Rondo dwarf galago Tanzania
Cercopithecus roloway Roloway monkey Côte d’Ivoire, Ghana
Piliocolobus preussi Preuss’s red colobus Cameroon, Nigeria
Piliocolobus rufomitratus Tana River red colobus Kenya
Gorilla beringei graueri Grauer’s gorilla DRC
Cheirogaleus lavasoensis Lavasoa Mountains dwarf lemur Madagascar
Hapalemur alaotrensis Lac Alaotra bamboo lemur Madagascar
Varecia rubra Red rued lemur Madagascar
Lepilemur septentrionalis Northern sportive lemur Madagascar
Propithecus perrieri Perrier’s sifaka Madagascar
Carlito syrichta Philippine tarsier Philippines
Nycticebus javanicus Javan slow loris Indonesia (Java)
Simias concolor Pig-tailed snub-nosed langur Indonesia (Mentawai Is.)
Trachypithecus delacouri Delacour’s langur Vietnam
Trachypithecus poliocephalus Golden-headed or Cat Ba langur Vietnam
Rhinopithecus avunculus Tonkin snub-nosed monkey Vietnam
Semnopithecus ajax Chamba sacred langur India
Semnopithecus vetulus nestor Western purple-faced langur Sri Lanka
Nomascus hainanus Hainan gibbon China
Pongo abelii Sumatran orangutan Indonesia (Sumatra)
Ateles hybridus Brown spider monkey Colombia, Venezuela
Ateles fusciceps fusciceps Ecuadorian brown-headed
Cebus kaapori Ka’apor capuchin Brazil
Callicebus oenanthe San Martín titi monkey Peru
Alouatta guariba guariba Northern brown howler Brazil
Table 2. Primate species included on the 2012–2014 list that were removed from the 2014–2016 list.
Piliocolobus pennantii Bioko red colobus Equatorial Guinea (Bioko Is.)
Microcebus berthae Madame Berthe’s mouse lemur Madagascar
Eulemur avifrons Sclater’s black lemur Madagascar
Propithecus candidus Silky sifaka Madagascar
Indri indri Indri Madagascar
Tarsius pumilus Pygmy tarsier Indonesia (Sulawesi)
Pygathrix cinerea Gray-shanked douc langur Vietnam
Nomascus nasutus Cao Vit or Eastern black-crested
Table 3. Primate species that were added to the 2014–2016 list. e Lake Alaotra bamboo lemur, Perrier’s sifaka,
the Hainan black-crested gibbon and the Sumatran orang-utan were on previous lists. e other four species,
marked with an asterisk, are new to the list.
Piliocolobus preussi* Preuss’s red colobus Cameroon, Nigeria
Cheirogaleus lavasoensis* Lavasoa Mountains dwarf lemur Madagascar
Hapalemur alaotrensis Lac Alaotra bamboo lemur Madagascar
Propithecus perrieri Perrier’s sifaka Madagascar
Carlito syrichta* Philippine tarsier Philippines
Semnopithecus ajax* Chamba sacred langur India
Nomascus hainanus Hainan gibbon China
Pongo abelii Sumatran orangutan Indonesia (Sumatra)
Table 4. Primate species considered during the discussion of the 2014–2016 list at the IPS Congress in Hanoi
that did not make it onto the list, but are also highly threatened.
Piliocolobus epieni Niger Delta red colobus Nigeria
Cheirogaleus sibreei Sibree’s dwarf lemur Madagascar
Lepilemur sahamalazensis Sahamalaza sportive lemur Madagascar
Daubentonia madagascariensis Aye-aye Madagascar
Nycticebus coucang Sunda slow loris Indonesia, Malaysia, Singapore,
Loris tardigradus nycticeboides Horton Plains slender loris Sri Lanka
Trachypithecus hatinhensis Hatinh langur Lao PDR, Vietnam
Cebus aequatorialis Ecuadorian white-fronted capuchin Ecuador, Peru
Photos of some of the Top 25 Most Endangered Primates. From top to bottom, le to right: 1. Carlito syrichta (photo by Russell A. Mittermeier);
2. Nomascus hainanus (juvenile)(photo by Zhao Chao); 3. Ateles hybridus (photo by Russell A. Mittermeier); 4. Alouatta guariba guariba (photo
by Russell A. Mittermeier); 5. Cercopithecus roloway (photo by S. Wolters, WAPCA); 6. Propithecus perrieri (photo by Russell A. Mittermeier);
7. Simias concolor (juvenile)(photo by Richard Tenaza); 8. Callicebus oenanthe (photo by Russell A. Mittermeier); 9. Varecia rubra (photo by
Russell A. Mittermeier); 10. Lepilemur septentrionalis (photo by Edward E. Louis, Jr.); 11. Trachypithecus poliocephalus (photo by Tilo Nadler); 12.
Gorilla beringei graueri (photo by Russell A. Mittermeier); 13. Pongo abelii (photo by Russell A. Mittermeier); 14. Galagoides rondoensis (photo
by Andrew Perkin); 15. Hapalemur alaotrensis (juvenile) (photo by Russell A. Mittermeier); 16. Trachypithecus delacouri (photo by Tilo Nadler).
Gulf of Guinea
Tropic of C
Tropic of C
0 500 1.000 1.500 2.000250
Gorilla beringei graueri
Weighing approximately 60 g, this is the smallest of all
galago species (Perkin et al. 2013). It is distinct from
other dwarf galagos in its diminutive size, a bottle-
brush-shaped tail, its reproductive anatomy, and its
distinctive “double unit rolling call” (Perkin and Honess
2013). Current knowledge indicates that this species
occurs in two distinct areas, one in southwest Tanzania
near the coastal towns of Lindi and Mtwara, the other
approximately 400 km further north, above the Ruji
River, in pockets of forest around Dar es Salaam. One
further population occurs in Sadaani National Park,
approximately 100 km north of Dar es Salaam. Rondo
dwarf galagos have a mixed diet of insects and fruit,
oen feed close to the ground, and move by vertical
clinging and leaping in the shrubby understorey.
ey build daytime sleeping nests, which are oen in
the canopy (Bearder et al. 2003). As with many small
primates, G. rondoensis is probably subject to predation
by owls and other nocturnal predators. Among these,
genets, palm civets and snakes invoke intense episodes
of alarm calling (Perkin and Honess 2013).
Over the last decade, the status of G. rondoensis on the
IUCN Red List has changed from Endangered in 2000
to Critically Endangered in 2008 (Perkin et al. 2008). In
fact, based on a comparative ranking of the 27 primate
species of Tanzania, the Taxon conservation score of
Galagoides rondoensis was the second highest (7.13 out
of 8; Davenport et al. 2014), thus, making this species one
of particular conservation concern. It has an extremely
limited and fragmented range in a number of remnant
patches of Eastern African Coastal Dry Forest (sensu
Burgess and Clarke 2000; p.18) in Tanzania, namely
those at Zaraninge forest (06°08’S, 38°38’E) in Sadaani
National Park (Perkin 2000), Pande Game Reserve
(GR) (06°42’S, 39°05’E), Pugu/Kazimzumbwi (06°54’S,
39°05’E) (Perkin 2003, 2004), Rondo (NR) (10°08’S,
39°12’E), Litipo (10°02’S, 39°29’E) and Ziwani (10°20’S,
40°18’E) forest reserves (FR) (Honess 1996; Honess and
Bearder 1996). New sub-populations were identied in
2007 near Lindi town in Chitoa FR (09°57’S, 39°27’E)
and Ruawa FR (09°44’S, 39°33’E), and in 2011 in Noto
Village Forest Reserve (09°53’S, 39°25’E) (Perkin et al.
2011, 2013) and in the northern population at Ruvu
South Forest Reserve (06°58’S, 38°52’E). Specimens of G.
rondoensis, originally described as Galagoides demidovii
phasma, were collected by Ionides from Rondo Plateau
in 1955, and Lumsden from Nambunga, near Kitangari,
(approximately 10°40’S, 39°25’E) on the Makonde
Plateau in Newala District in 1953. Doubts surround
the persistence of this species on the Makonde Plateau,
which has been extensively cleared for agriculture.
Rondo Dwarf Galago
Galagoides rondoensis Honess in Kingdon, 1997
Andrew Perkin & Daphne Kerhoas
Rondo dwarf galago (Galagoides rondoensis)
(Illustration: Stephen D. Nash)
Surveys there in 1992 failed to detect any extant
populations (Honess 1996). e areas most critical to
their long-term conservation are Kazimzumbwi Forest
Reserve (9 km²), Zaraninge Forest (20 km²) in Sadaani
National Park, Pugu Forest Reserve (24 km²), and
Rondo Forest Reserve (25 km²), eastern Tanzania (De
Jong and Butynski 2012).
No detailed surveys have been conducted to assess
population sizes of G. rondoensis. Distribution surveys
have been conducted, however, in the southern (Honess
1996; Perkin et al. in prep.) and northern coastal forests
of Tanzania (29 surveyed) and Kenya (seven surveyed)
(Perkin 2000, 2003, 2004; Perkin et al. 2013). Absolute
population sizes remain undetermined but recent
surveys have provided estimates of density (3–6/ha at
Pande Game Reserve (Perkin 2003) and 8/ha at Pugu
Forest Reserve (Perkin 2004)) and relative abundance
from encounter rates (3–10/hr at Pande Game Reserve
and Pugu/Kazimzumbwi Forest Reserve (Perkin
2003, 2004) and 3.94/hr at Rondo Forest Reserve
(Honess 1996)). ere is a clear and urgent need for
further surveys to determine population sizes in these
dwindling forest patches.
In 2008, it was reported that the total area of forest in
which G. rondoensis is currently known to occur does
not exceed 101.6 km² (Pande GR: 2.4 km², Rondo FR:
25 km², Ziwani FR: 7.7 km², Pugu/Kazimzumbwi FR:
33.5 km², Litipo FR: 4 km², Zaraninge forest: 20 km²,
Chitoa FR: 5 km², and Ruawa FR 4 km²) (Minimum area
data source: Burgess and Clarke 2000; Doggart 2003;
Perkin et al. in prep.). New data on forest area change
indicates that while two new sub-populations have been
discovered; the overall area of occupancy hovers around
100 km². 2008 and 2014 forest-area estimations are as
follows: Zaraninge 2008: 20 km², 2014: 15 km²; Pande
2008: 2.4 km², 2014: 2.4 km²; Pugu/Kazimzumbwe
2008: 33.5 km², 2014: 9 km²; Ruvu South 2008: 20 km²,
2014: 10 km²; Ruawa 2008: 4 km², 2014: 4 km²; Litipo
2008: 4 km², 2014: 3 km²; Chitoa 2008: 4 km², 2014: 5
km²; Noto 2008: 21 km², 2014: 20 km²; Rondo 2008: 25
km², 2014: 25 km²; Ziwani 2008: 7.7 km², 2014: 1 km².
e total forest area estimates are as follows: 2008 101.6
km²; 2014 94.4 km².
e major threat facing this species is loss of habitat.
All sites are subject to some level of agricultural
encroachment, charcoal manufacture and/or logging.
All sites, except Pande (Game Reserve), Zaraninge
(within Saadani National Park) and Rondo (Nature
Reserve), are national or local authority forest reserves
and as such nominally, but in practice minimally,
protected. Since 2008, there have been changes
resulting in the increase in protection of two forests.
e Noto plateau forest, formerly open village land, is
part of a newly created village forest reserve, and the
Rondo Forest Reserve has now been declared a new
Nature Reserve, both are important for Rondo galago
conservation given their relatively large size. Given
current trends in charcoal production for nearby Dar es
Salaam, the forest reserves of Pugu and Kazimzumbwi
have been predicted to disappear over the next 10–15
years (Ahrends 2005). Pugu/Kazimzumbwe as well as
Ruvu South have seen continued and predicted losses to
the rampant charcoal trade since Ahrends (2005) study.
Pande, as a Game Reserve, is perhaps more secure,
and Zareninge forest, being in a National Park, is the
most protected part of the range of G. rondoensis. In the
south, the Noto, Chitoa and Rondo populations are the
most secure, as they are buered by tracts of woodland.
e type population at Rondo is buered by woodland
and Pinus plantations managed by the Rondo Forestry
Project, and is now a Nature Reserve. Litipo, and Ruawa
FRs are under threat from bordering village lands.
Ziwani is now mostly degraded scrub forest, thicket and
Conservation action is urgently needed by monitoring
rates of habitat loss, surveying new areas for remnant
populations, estimating population size, reassessing
the phylogenetic relationships of the sub-populations
and increasing awareness. ere is emerging data
(vocal and penile morphology) that the northern and
southern populations may be phylogenetically distinct
with important taxonomic implications. As such the
conservation of all sub-populations is important.
Across its known range, the Rondo galago can be
found in sympatry with a number of other galagos,
including two much larger species in the genus
Otolemur: Garnett’s galago O. garnettii (Least Concern,
Butynski et al. 2008a), and the thick-tailed galago, O.
crassicaudatus (Least Concern, Bearder 2008). e
Rondo galago is sympatric with the Zanzibar galago,
Galagoides zanzibaricus (Least Concern, Butynski et al.
2008b), in the northern parts of its range (for example,
in Zaraninge forest, Pugu/Kazimzumbwi FR and Pande
GR). In the southern parts of its range (for example, in
Rondo, Litipo and Noto), the Rondo galago is sympatric
with Grant’s galago, Galagoides granti (Least Concern,
Honess et al. 2008).
A new project to address these conservation and
research issues has been implemented since 2012.
Targeted conservation initiatives are taking place in
Ruvu South FR, Chitoa FR and Noto VFR.
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Journal 17: 23.
Bearder, S.K. 2008. Otolemur crassicaudatus. In: IUCN
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Bearder, S. K., L. Ambrose, C. Harcourt, P. E. Honess,
A. Perkin, S. Pullen, E. Pimley and N. Svoboda. 2003.
Species-typical patterns of infant care, sleeping site
use and social cohesion among nocturnal primates in
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Burgess, N. D. and G. P. Clarke. 2000. Coastal Forests
of Eastern Africa. IUCN – e World Conservation
Union, Gland, Switzerland, and Cambridge, UK.
Butynski, T. M., S. K. Bearder and Y. de Jong. 2008a.
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reatened Species. Version 2013.2. <www.iucnredlist.
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Butynski, T. M., Y. de Jong, A. Perkin, S. K. Bearder and
P. Honess. 2008b. Galagoides zanzibaricus. In: IUCN
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2013.2. <www.iucnredlist.org>. Accessed 16 March
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Doggart, N. (ed.). 2003. Pande Game Reserve: A
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Galagidae) in Tanzanian Forests. Doctoral thesis,
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Perkin, A., S. K. Bearder and J. Karlsson. In prep. Galago
surveys in Rondo, Litipo, Chitoa, Ruawa, Ndimba
and Namatimbili forests, Lindi Region, southeastern
Perkin, A., B. Samwel and J. Gwegime. 2011. Going for
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Considered by some to be subspecies of Cercopithecus
diana, the Diana monkey and the roloway monkey are
highly attractive, arboreal primates that inhabit the
Upper Guinean forests of West Africa (Grubb et al.
2003). Groves (2001) considers the two subspecies to
be suciently distinct to be regarded as full species.
e roloway monkey is distinguished by its broad white
brow line, long white beard and yellow thighs. Of the
two forms, the roloway, which is known from Ghana
and central and eastern Côte d’Ivoire, is more seriously
threatened with extinction; it is rated as Endangered in
the current IUCN Red List (Oates et al. 2008), but its
status should be upgraded to Critically Endangered.
Roloway monkeys are upper-canopy specialists that
prefer undisturbed forest habitat. Destruction and
degradation of their habitat and relentless hunting for
the bushmeat trade have reduced their population to
small, isolated pockets. Miss Waldron’s red colobus
(Procolobus badius waldroni) once inhabited many of
the same forest areas as the roloway, but is now almost
certainly extinct (Oates 2011). Unless more eective
conservation action is taken, there is a strong possibility
that the roloway monkey will also disappear in the near
Over the last 40 years roloway monkeys have been
steadily extirpated in Ghana. Several recent surveys
have failed to conrm the presence of these monkeys in
any reserves in western Ghana, including Bia National
Park, Krokosua Hills Forest Reserve, Subri River Forest
Reserve and Dadieso Forest Reserve (Oates 2006;
Gatti 2010; Buzzard and Parker 2012; Wiafe 2013),
although it is possible that the Ankasa Conservation
Area still contains a few individuals (Magnuson 2003;
Gatti 2010). Community-controlled forests along
the Tano River (referred to as the “Kwabre Forest”)
in the far southwestern corner of the country are the
only place in Ghana at which any roloways have been
reported as seen by scientists or conservationists in
the last decade. Kwabre consists of patches of swamp
forest along the lower Tano River, adjacent to the Tanoé
forest in Côte d’Ivoire. Surveys of these forests have
been conducted under the auspices of the West African
Primate Conservation Action organization since 2011,
and several sightings of roloway groups have been
made, along with mona monkeys, spot-nosed monkeys,
white-naped mangabeys and olive colobus (WAPCA
2012, 2014; Dempsey 2014). WAPCA has launched a
community-based conservation project with villages
around these forests with the aim of establishing a
Kwabre Community Resource Management Area.
Meanwhile, further eorts should be made to ascertain
whether any roloway monkeys still survive in the
Ankasa, because this site has signicant conservation
Cercopithecus roloway (Schreber, 1774)
Ghana, Côte d’Ivoire
(2002, 2006, 2008, 2010, 2012, 2014)
W. Scott McGraw & John F. Oates
Roloway monkey (right) (Cercopithecus roloway) and Diana monkey (le) (Cercopithecus diana)
(Illustrations: Stephen D. Nash)
potential and roloways have been reported there in the
relatively recent past.
In neighbouring Côte d’Ivoire, the Roloway guenon’s
status is perhaps even direr. Less than ten years ago
roloways were known or strongly suspected to exist in
three forests: the Yaya Forest Reserve, the Tanoé forest
adjacent to the Ehy Lagoon, and Parc National des
Îles Ehotilé (McGraw 1998, 2005; Koné and Akpatou
2005). Surveys of eighteen areas between 2004 and 2008
(Gonedelé Bi et al. 2008, 2012) conrmed the presence
of roloways only in the Tanoé forest suggesting that
the roloway monkey may have been eliminated from
at least two forest areas (Parc National des Îles Ehotilé,
Yaya Forest Reserve) within the last decade. Subsequent
surveys carried out in southern Côte d’Ivoire suggest
a handful of roloways may still survive in two forest
reserves along the country’s coast. On 21 June, 2011,
Gonedelé bi Sery observed one roloway individual in
the Dassioko Sud Forest Reserve (Gonedelé Bi et al.
in review; Bitty et al. 2013). In 2012, Gonedelé Bi and
E. A. Bitty observed roloways in Port Gauthier Forest
Reserve, and in October 2013, Gonedelé Bi obtained
photographs of monkeys poached inside this reserve,
including an image purported to be a roloway. e
beard on this individual appears short for a roloway,
raising the possibility that surviving individuals in this
portion of the interuvial region may in fact be hybrids.
In any case, no sightings of roloways have been made
in the Dassioko Sud or Port Gauthier Forest Reserves
since 2012, despite regular patrols there. ese reserves
are described as coastal evergreen forests and both are
heavily degraded due to a large inux of farmers and
hunters from the northern portion of the country (Bitty
et al. 2013). Gonedelé Bi and colleagues, in cooperation
with SODEFOR (Société de Développement des Forêts)
and local communities, have organized regular foot
surveys aimed at removing illegal farmers and hunters
from both reserves; however, the most recent surveys
(August 2015) revealed that a logging company (SIDB)
has begun clearing a portion of the Port Gauthier
reserve. Eorts are underway to work with SODEFOR
at stopping logging and other illegal activities in these
reserves (Gonedelé Bi 2015).
us, the only forest in Côte d’Ivoire where roloways
are conrmed to exist is the Tanoé forest adjacent to
the Ehy Lagoon. is wet forest also harbors one of the
few remaining populations of white-naped mangabeys
in Côte d’Ivoire. Eorts led by I. Koné and involving
several organizations (CEPA, WAPCA) helped stop a
large palm oil company from causing further habitat
degradation, and a community-based conservation
eort has helped slow poaching within this forest (Koné
2015). Unfortunately, hunting still occurs in Tanoé and
the primate populations within it are undoubtedly
decreasing (Gonedelé Bi et al. 2013).
As the potential last refuge for roloways and white-
naped mangabeys, the protection of the Tanoé forest in
Côte d’Ivoire and the adjacent Kwabre Forest in Ghana
should be the highest conservation priority. By any
measure, the roloway monkey must be considered one
of the most critically endangered monkeys in Africa and
appears to be on the verge of extinction (Oates 2011).
Bitty, E. A., S. Gonedelé Bi, W. S. McGraw. 2013.
Accelerating deforestation and hunting in protected
reserves jeopardize primates in southern Côte d’Ivoire.
American Journal of Physical Anthropology Supp 56: 81-
Buzzard, P. J. and A. J. A. Parker. 2012. Surveys from the
Subri River Forest Reserve, Ghana. African Primates 7:
Dempsey, A. 2014. Save our Forest, Save our Future: A
Survey of the Tanoe‐Ankasa Community Owned Forest
for Primate Presence and Illegal Activity, Introducing
SMART Technology. Master’s thesis, Oxford Brookes
University, Oxford, England.
Gonedelé Bi, S. 2015. Report on primate surveys of
Dassioko Sud and Port Gauthier forest reserves in
southern Côte d’Ivoire. Unpublished report.
Gonedelé Bi, S., I. Koné, J-C. K. Béné, A. E. Bitty,
B. K. Akpatou, Z. Goné Bi, K. Ouattara and D. A.
Ko. 2008. Tanoé forest, south-eastern Côte-d’Ivoire
identied as a high priority site for the conservation of
critically endangered Primates in West Africa. Tropical
Conservation Science 1: 265-278.
Gonedelé Bi, S. and A. E. Bitty. 2013. Conservation of
threatened primates of Dassioko Sud and Port Gauthier
forest reserves in coast Côte d’Ivoire. Final report to
Primate Conservation Inc., Charlestown, Rhode Island,
USA. February, 2013.
Gonedelé Bi, S., J-C. K. Béné, A. E. Bitty, A. N’Guessan,
A. D. Ko, B. Akptatou and I. Koné. 2013. Roloway
guenon (Cercopithecus diana roloway) and white-naped
mangabey (Cercocebus atys lunulatus) prefer mangrove
habitats in Tanoé Forest, southeastern Ivory Coast.
Journal of Ecosystems and Ecography 3: 126.
Gonedelé Bi, S. I. Koné, A. E. Bitty, J-C. K. Béné,
B. Akptatou and D. Zinner. 2012. Distribution and
conservation status of catarrhine primates in Côte
d’Ivoire (West Africa). Folia Primatologica 82: 11–23.
Gonedelé Bi S., E. A. Bitty, W. S. McGraw. 2014.
Conservation of threatened primates in Dassioko Sud
and Port Gauthier forest reserves: use of eld patrols to
assess primate abundance and illegal human activities.
American Journal of Physical Anthropology Suppl. 58:
Groves, C. P. 2001. Primate Taxonomy. Smithsonian
Institution Press, Washington, DC.
Grubb, P., T. M. Butynski, J. F. Oates, S. K. Bearder, T.
R. Disotell, C. P. Groves and T. T. Struhsaker. 2003.
An assessment of the diversity of African primates.
International Journal of Primatology 24: 1301–1357.
Koné, I. 2015. Community empowerment for the
conservation of critically endangered primates and
their habitat in south-eastern Côte d’Ivoire. Twenty-
Two Stories of Conservation in Africa: Key Elements
for Eective and Well-Governed Protected Areas in
Sub-Saharan Africa. IUCN e IUCN Programme for
African Protected Areas and Conservation (PAPACO)
(ed.), pp.36-39. IUCN, Gland, Switzerland. URL: http://
Koné, I. and K. B. Akpatou. 2005. Recherche en Côte
d’Ivoire de trois singes gravement menaces d’extinction.
CEPA Magazine 12: 11–13.
Magnuson, L. 2003. Final Brief: Ecology and
Conservation of the Roloway Monkey in Ghana.
Unpublished report to the Wildlife Division of Ghana,
Forestry Commission, Ghana.
McGraw, W. S. 1998. Surveys of endangered primates
in the forest reserves of eastern Côte d’Ivoire. African
Primates 3: 22–25.
McGraw, W. S. 2005. Update on the search for Miss
Waldron’s red colobus monkey (Procolobus badius
waldroni). International Journal of Primatology 26: 605–
Oates, J. F. 2006. Primate Conservation in the Forests
of Western Ghana: Field Survey Results, 2005–2006.
Report to the Wildlife Division, Forestry Commission,
Oates, J. F. 2011. Primates of West Africa: A Field Guide
and Natural History. Conservation International,
Oates, J. F., S. Gippoliti and C. P. Groves, C.P. 2008.
Cercopithecus diana ssp. roloway. In: IUCN 2008. IUCN
Red List of reatened Species. Version 2013.2 <www.
iucnredlist.org>. Accessed 16 March 2014.
WAPCA. 2012. Annual Report. West African Primate
Conservation Action, Accra, Ghana.
WAPCA. 2014. Annual Report. West African Primate
Conservation Action, Accra, Ghana.
Wiafe, E. 2013. Status of the Critically Endangered
roloway monkey (Cercopithecus diana roloway) in the
Dadieso Forest Reserve, Ghana. African Primates 8:
Preuss’s red colobus (Piliocolobus preussi) is endemic to
western Cameroon and southeastern Nigeria where it is
found in dense, moist, high canopy forests (Oates 2011).
e taxonomic arrangement of this monkey has changed
considerably in recent years; some classications place
it as a subspecies of badius or pennantii and others
recognize it as a distinct species (Oates and Ting 2015).
Mittermeier et al. (2013) place preussi in the genus
Piliocolobus, following Groves (2007). Since 2008,
the IUCN Red List has listed P. preussi as a Critically
Although a comprehensive assessment of the
distribution and abundance of Preuss’s red colobus has
never been conducted, it is evident that populations
of this monkey have disappeared from much of their
original range since the beginning of the 20th century
(Struhsaker 1999). e largest populations are now
mostly found in Cameroon in the forests in and around
Korup National Park (Linder and Oates 2011; Forboseh
et al. 2007; Kupsch et al. 2014) and within the Ebo-
Makombe-Ndokbou forest block (Morgan et al. 2013).
In Nigeria, Preuss’s red colobus is restricted to a small
area of the Oban Division of Cross River National Park
close to the boundary with Cameroon, and contiguous
with Korup National Park; ranger patrols facilitated
by the Wildlife Conservation Society conrmed its
presence in this area in early 2015.
Although, as for other red colobus species, predation
by chimpanzees may be a threat to the viability of
some populations (Watts and Amsler 2013; Morgan et
al. 2013), it is clear that for P. preussi the threats from
bushmeat hunting and deforestation through logging,
agriculture and infrastructure development are the
major factors leading to its decline. Fa and colleagues
(2006) conducted point-of-sale bushmeat surveys
between August 2002 and January 2003 in the Cross-
Sanaga region of Nigeria and Cameroon and estimated
that 8,589 individual Preuss’s red colobus monkeys were
sold annually, over three-quarters of which originated
in Cameroon. In Korup, bushmeat hunting appears to
Preuss’s Red Colobus
Piliocolobus preussi Matschie, 1900
Joshua M. Linder, Bethan J. Morgan, John F. Oates, & Andrew Dunn
Preuss’ Red Colobus monkey
(Illustration: Stephen D. Nash)
be driving the decline of this species. Transect surveys
conducted between 2001 and 2015 in southern Korup
National Park indicate increasing hunting intensity
and declining sighting frequency (groups sighted/
km walked) of Piliocolobus preussi from 0.06 to 0.01
(Dunn and Okon 2003; Linder and Oates 2011; Linder
unpublished data). Piliocolobus preussi is also becoming
increasingly rare in northeast Korup, where sighting
frequency of this species along transects has declined
from 0.07 in 1990 (Edwards 1992) to 0.05 in 2004–2005
(Linder 2008) to 0.03 in 2014 (Robinson unpublished
data). Hunter harvest surveys in Korup also suggest that
the proportional representation of P. preussi has declined
signicantly between 1990 and 2005 (Linder and Oates
2011). Although temporal data on P. preussi abundance
and distribution are lacking for the Ebo forest, recent P.
preussi encounter rates suggest that numbers are now
Preuss’s red colobus is one of the most endangered
of all of the red colobus species, which are probably
more threatened than any other taxonomic group
of primates in Africa (Oates 1996; Struhsaker 2005,
2010). Elsewhere in the Gulf of Guinea region, the
Bioko red colobus is now restricted to a very small area
in the southwest of the island, where it is still hunted
(Cronin et al. 2014), and the Niger Delta red colobus
is in a precarious situation (Ikemeh 2015). On the
other hand, a population of Bouvier’s red colobus of
Congo Republic, long feared to be extinct, was located
in March 2015 in the Ntokou-Pikounda National Park
To secure the long term conservation of P. preussi,
we recommend the following actions: (i) bushmeat
hunting in the forests of Cameroon’s Korup National
Park and the contiguous Oban Division of Cross River
National Park in Nigeria must be drastically reduced
through improved law-enforcement and community-
based initiatives; (ii) the status of the Ebo forest in
Cameroon should be upgraded to a national park and
a results-driven law-enforcement regime implemented;
(iii) eld surveys are urgently needed to determine the
current distribution and abundance of P. preussi outside
of protected areas, and plans should be developed to
work with local communities in these areas to help
secure the populations. For example, the species was
encountered in what is now known as the Ndokbou
forest, north of the Ebo forest, in 2001, but since then
no surveys have been conducted; it is not clear whether
the species remains in this area, which is increasingly
isolated from the Ebo forest due to industrial logging
activities and acute pressure from the development of
large-scale oil-palm plantations in the intervening area;
and (iv) actions to raise the awareness of the existence
of the species (although it may be known to hunters and
bushmeat dealers the presence of the species remains
largely unacknowledged by park authorities, at least
Ultimately, conservationists must nd ways to convince
the Nigerian and Cameroon governments and local
communities that Preuss’s red colobus is worth saving.
Cronin, D. T., G. W. Hearn and J. F. Oates. 2014.
Bioko red colobus (Piliocolobus pennantii pennantii).
In: Primates in Peril: e World’s 25 Most Endangered
Primates 2012–2014. C. Schwitzer, R. A. Mittermeier, A.
B. Rylands, L. A. Taylor, F. Chiozza, E. A. Williamson,
J. Wallis and F. E. Clark (eds.), pp.17–19. IUCN SSC
Primate Specialist Group, International Primatological
Society, Conservation International, and Bristol
Zoological Society, Arlington, VA.
Devreese, L. 2015. Preliminary survey of the current
distribution and conservation status of the poorly
known and critically endangered Piliocolobus bouvieri
in the Republic of Congo. Unpublished report.
Dunn, A. and D. Okon. 2003. Monitoring the abundance
of diurnal primates and duikers in Korup National Park,
Cameroon, 2001–2003. Korup Project, Mundemba,
Edwards, A. E. 1992. e Diurnal Primates of Korup
National Park, Cameroon: Abundance, Productivity
and Polyspecic Associations. MSc esis, University
of Florida, USA.
Fa, J. E., S. Seymour, J. Dupain, R. Amin, L. Albrechtsen
and D. Macdonald. 2006. Getting to grips with the
magnitude of exploitation: Bushmeat in the Cross–
Sanaga rivers region, Nigeria and Cameroon. Biological
Conservation 129: 497–510.
Forboseh, P. F., T. C. H. Sunderland and M. Eno-Nku.
2007. Priority setting for conservation in south-west
Cameroon based on large mammal surveys. Oryx 41:
Groves, C. P. 2007. e taxonomic diversity of the
Colobinae of Africa. Journal of Anthropological Sciences
Ikemeh, R. A. 2015. Assessing the population status
of the Critically Endangered Niger Delta red colobus
(Piliocolobus epieni). Primate Conservation (29).
Kupsch, D., B. S. Kadiri and M. Waltert. 2014. Biodiversity,
carbon stock and market value assessment for the
SGSOC project area, Southwest region, Cameroon.
Report to World Wide Fund for Nature, Germany, and
Greenpeace International. Georg-August-Universität,
Göttingen, Germany, and University, of Dschang,
Dschang, Cameroon. 41pp.
Linder, J. M. 2008. e Impact of Hunting on Primates
in Korup National Park: Implications for Primate
Conservation. PhD esis, City University of New
York, New York.
Linder, J. M. and J. F. Oates. 2011. Dierential impact of
bushmeat hunting on monkey species and implications
for primate conservation in Korup National Park,
Cameroon. Biological Conservation 144: 738–745.
Mittermeier, R. A., A. B. Rylands, and D. E. Wilson
(eds.). 2013. Handbook of the Mammals of the World: 3.
Primates. Lynx Edicions, Barcelona, Spain.
Morgan, B. J., J. Ngu Suh and E. E. Abwe. 2012. Attempted
predation by Nigeria-Cameroon chimpanzees (Pan
troglodytes ellioti) on Preuss’s red colobus (Procolobus
preussi) in the Ebo forest, Cameroon. Folia Primatologica
Oates, J. F. 1996. African Primates: Status Survey and
Conservation Action Plan. IUCN, Gland, Switzerland.
Oates, J. F. 2011. Primates of West Africa: A Field Guide
and Natural History. Conservation International,
Oates, J. and N. Ting. 2015. Conservation consequences
of unstable taxonomies: e case of the red colobus
monkeys. In: Taxonomic Tapestries: e reads of
Evolutionary, Behavioural and Conservation Research,
A. M. Behie and M. F. Oxenham (eds.), pp.321−343.
Australian National University Press, Acton, Australia.
Struhsaker, T. T. 1999. Primate communities in Africa:
the consequence of longterm evolution or the artifact of
recent hunting. In: Primate Communities, J. G. Fleagle,
C. Janson and K. E. Reed (eds.), pp.289–294. Cambridge
University Press, Cambridge.
Struhsaker, T. T. 2005. Conservation of red colobus and
their habitats. International Journal of Primatology 26:
Struhsaker, T. T. 2010. e Red Colobus Monkeys:
Variation in Demography, Behavior, and Ecology of
Endangered Species. Oxford University Press, Oxford.
Watts, D. P. and S. J. Amsler. 2013. Chimpanzee–red
colobus encounter rates show a red colobus population
decline associated with predation by chimpanzees at
Ngogo. American Journal of Primatology 75: 927–937.
On the current IUCN Red List, the Tana River red
colobus is presented as one of four assessed subspecies
of Procolobus rufomitratus (i.e., as P. r. rufomitratus).
e other three are Oustalet’s red colobus Procolobus r.
oustaleti (Trouessart, 1906), ashy red colobus Procolobus
r. tephrosceles (Elliot, 1907), and Tshuapa red colobus
Procolobus r. tholloni (Milne-Edwards, 1886). Here,
however, we follow Groves (2001, 2005, 2007) and
Groves and Ting (2013) in placing all red colobus in the
genus Piliocolobus, and rufomitratus and the other three
subspecies mentioned above as full species.
Gallery forests along the Lower Tana River, Kenya, are
part of the East African Coastal Forests Biodiversity
Hotspot. e Lower Tana River forests and some forest
patches in the Tana Delta are the only habitat for two
endemic primates; the Tana River red colobus and the
Tana River mangabey, Cercocebus galeritus Peters, 1879.
Piliocolobus rufomitratus is classied as ‘Endangered’
on the current IUCN Red List (Butynski et al. 2008a).
Cercocebus galeritus is also classied as ‘Endangered’
(Butynski et al. 2008b).
Both the Tana River red colobus and the Tana River
mangabey inhabit forest fragments (size range about
1–500 ha) along a 60-km stretch of the Lower Tana
River (Butynski and Mwangi 1995; Mbora and Meikle
2004). In 2009, small populations of both species were
discovered in the Tana Delta (Hamerlynck et al. 2012).
e area of occurrence of the red colobus is <13 km², and
that of the mangabey <26 km² (Butynski and Mwangi
1995). e population of the Tana River red colobus is
about 1,000 individuals and declining (Butynski and
Mwangi 1995; Karere et al. 2004). e population of
the Tana River mangabey is roughly 2,000 individuals
and also declining (Karere et al. 2004; Wieczkowski and
Butynski 2013). Genetic analyses show that the eective
population sizes of the two species are less than 100
individuals (Mbora and McPeek 2015).
Tana River Red Colobus
Piliocolobus rufomitratus (Peters, 1879)
(2002, 2004, 2006, 2008, 2012, 2014)
omas M. Butynski & Olivier Hamerlynck
Tana River red colobus (Piliocolobus rufomitratus)
(Illustration: Stephen D. Nash)
ere are six (perhaps seven) other species of nonhuman
primate in the forests of the Lower Tana River, including
the ‘Vulnerable’ Pousargues’s monkey Cercopithecus
mitis albotorquatus Pousargues, 1896, and 3–4 species
of strepsirrhines (De Jong and Butynski 2012). Given
the small area of remaining forest (<37 km²), the serious
threats (see below), and the occurrence of two endemic
‘Endangered’ primates, the forests of the Lower Tana
River are the top priority for actions to conserve East
Africa’s primate diversity (De Jong and Butynski 2012;
Butynski and De Jong in press).
Several factors render the long-term survival of the
Tana River red colobus and Tana River mangabey
bleak. First, forest continues to be degraded, cleared,
and fragmented as a result of expanding agriculture
and the taking of building materials and other products
(Butynski and Mwangi 1995; Mbora and Meikle
2004; Moinde-Fockler et al. 2007; Duvail et al. 2012;
Hamerlynck et al. 2012; Butynski and De Jong in press).
Second, there has been considerable alteration of river
ow volume and the ood cycle by ve hydroelectric
power dams up-river (Butynski 1995; Maingi and
Marsh 2002), and a sixth dam, the High Grand Falls
Dam, is under construction (Hamerlynck et al. 2012).
is will be the second largest dam in Africa and will
be accompanied by large-scale irrigation schemes and
water transfer to the ‘Lamu Port and Lamu-Southern
Sudan-Ethiopia Transport Corridor’ (LAPSET).
LAPSET is, potentially, the largest infrastructure
project in Africa. us, the High Grand Falls Dam will
have additional negative impacts on the oods and
groundwater recharge required for the establishment
and maintenance of the forests of the Lower Tana River
and Tana Delta, on those people with ood-dependent
livelihoods (crop farmers, livestock-raisers, and
sherman), and on the biodiversity of this biologically-
sensitive region (Duvail et al. 2012). Construction of
the US$ 2 billion High Grand Falls Dam is now a top
priority of the Kenyan Government. Finance is from
rms in the People’s Republic of China and the Export-
Import Bank of China.
ird, in January 2007, the High Court of Kenya ordered
the annulment of the Tana River Primate National
Reserve (TRPNR; 171 km²) because the court found
that the Reserve had not been established in accordance
with the law. About half of the remaining forests gained
some protection from the TRPNR. As a result of this
court decision, no habitat of the Tana River red colobus,
or of the Tana River mangabey, is legally protected at
the present time.
Fourth, habitat loss has increased as a result of the
failure of the Tana Delta Irrigation Project (TDIP) to
protect forest. TDIP, nanced by the Japan International
Cooperation Agency (JICA), is a large rice-growing
scheme managed by the Tana and Athi Rivers
Development Authority (TARDA). e TDIP site had
some of the most important forests for the red colobus
and mangabey (Butynski and Mwangi 1994; Moinde-
Fockler et al. 2007).
In 2012, the Tana Delta became a Ramsar Site. is
internationally recognized protected area status
was expected to help maintain, if not enhance, the
conservation values of the Tana Delta, including those
small forests in which Tana River red colobus and Tana
River mangabey occur. e latest news, however, is that
the most important of these forests, Onkolde (60 ha), has,
since 2012, been decimated by mass loss of its dominant
tree canopy species, the ‘Vulnerable’ Oxystigma msoo.
is is probably the result of the lowering of groundwater
caused by a TDIP embankment and, in 2015, by massive
logging by people claiming to have “legal” documents
(typically a euphemism for documents signed by people
so high up that Kenya Forest Service sta on the ground
dare not challenge them) (O. Hamerlynck pers. obs).
As is oen the case for Ramsar Sites, the various land-
use plans for the Tana Delta have not been eectively
implemented, and established user-rights and public
interests have been disregarded (Butynski and De Jong
in press, O. Hamerlynck and S. Duvail pers. obs).
Despite the troubles highlighted above, there is reason
for hope for the Tana River forests and their primates.
Partly galvanized by the participatory nature of research
in the forests of the Lower Tana River, an organization
called ‘Ndera Community Conservancy’ (NCC)
has been established (Mbora and Allen 2011). e
mission of this formally registered community-based
organisation is to protect and conserve about half of the
forest patches within the former TRPNR, and improve
the viability of particular forest patches outside the
former reserve. e NCC is working with government
and NGO conservation initiatives and is making
progress. For the NCC to make signicant progress,
however, the support of international conservation
agencies is needed. With the NCC, government, and
the international conservation community working
together, the prospects for the long-term viability of the
Tana River primates can be greatly improved.
Butynski, T. M. 1995. Report says dam could threaten
Kenya’s endangered primates. African Primates 1: 14–
Butynski, T. M. and G. Mwangi. 1995. Census of Kenya’s
endangered red colobus and crested mangabey. African
Primates 1: 8–10.
Butynski, T. M., T. T. Struhsaker and Y. de Jong. 2008a.
Procolobus rufomitratus ssp. rufomitratus. In: IUCN
2015. IUCN Red List of reatened Species. Version
2015.3. <www.iucnredlist.org>. Accessed 9 October
Butynski, T. M., T. T. Struhsaker, J. Kingdon and Y. de
Jong 2008b. Cercocebus galeritus. In: IUCN 2015. IUCN
Red List of reatened Species. Version 2015.3. <www.
iucnredlist.org>. Accessed 9 October 2015.
Butynski, T. M. and Y. A. de Jong. In press. Primates
of Africa’s coastal deltas and their conservation. In:
Primates in Flooded Habitats: Ecology and Conservation,
A. A. Barnett, I. Matsuda and K. Nowak (eds.).
Cambridge University Press, Cambridge.
De Jong, Y. A. and T. M. Butynski. 2012. e primates
of East Africa: country lists and conservation priorities.
African Primates 7: 135–155.
Duvail, S., C. Médard, O. Hamerlynck and D. W. Nyingi.
2012. Land and water ‘grabbing’ in an East African
coastal wetland: the Tana Delta case study. Water
Alternatives 5: 322–343.
Groves, C. P. 2005. Order Primates. In: Mammal Species
of the World, D. E. Wilson and D. M. Reeder (eds.),
pp.111–184. e Johns Hopkins University Press,
Groves, C. P. 2007. e taxonomic diversity of the
Colobinae of Africa. Journal of Anthropological Sciences
Groves, C. P. and N. Ting. 2013. Tana River red colobus
Piliocolobus rufomitratus. In: Handbook of the Mammals
of the World. Volume 3. Primates, R. A. Mittermeier, A.
B. Rylands and D. E. Wilson (eds.), p.709. Lynx Edicions,
Hamerlynck, O., Q. Luke, T. M. Nyanger, S. Suvail
and C. Leauthaud. 2012. Range extension, imminent
threats and conservation options for two Endangered
primates: the Tana River red colobus Procolobus
rufomitratus rufomitratus (Peters, 1879) and the Tana
River mangabey Cercocebus galeritus (Peters, 1879) in
the Lower Tana Floodplain and Delta, Kenya. African
Primates 7: 211–217.
Karere, G.M., N.O. Oguge, J. Kirathe, P.K. Muoria, N.N.
Moinde and M.A. Suleman. 2004. Population sizes
and distribution of primates in the Lower Tana River
Forests, Kenya. International Journal of Primatology 25:
Maingi, J.K. and S. F. Marsh 2002. Quantifying
hydrologic impacts following dam construction along
the Tana River, Kenya. Journal of Arid Environments 50:
Mbora, D. N. M. and D. B. Meikle. 2004. Forest
fragmentation and the distribution, abundance and
conservation of the Tana River red colobus (Procolobus
rufomitratus). Biological Conservation 118: 67–77.
Mbora, D. N. M. and L. Allen. 2011. e Tana Forests
‘People for Conservation and Conservation for People’
Initiative (PCCP): Preserving the Habitat of the Tana
River Red Colobus (Procolobus rufomitratus) and the
Tana River Mangabey (Cercocebus galeritus) rough
Community Conservation and Development in
Tana River District, Kenya. Final Report on Phase 1,
Mohamed bin Zayed Species Conservation Fund, Abu
Mbora, D. N. M and M. A. McPeek. 2015. How monkeys
see a forest: population genetic structure in two forest
monkeys. Conservation Genetics 16: 559–569.
Moinde-Fockler, N. N., N. O. Oguge, G. M. Karere, D.
Otina and M. A. Suleman. 2007. Human and natural
impacts on forests along lower Tana River, Kenya:
implications towards conservation and management of
endemic primate species and their habitat. Biodiversity
and Conservation 16: 1161–1173.
Wieczkowski, J. A. and T. M. Butynski. 2013. Cercocebus
galeritus Tana River mangabey. In: Mammals of Africa.
Volume II: Primates, T. M. Butynski, J. Kingdon and J.
Kalina (eds.), pp.167–170. Bloomsbury, London.
Grauer’s gorilla (Gorilla beringei graueri), one of
two subspecies of eastern gorilla, is listed on CITES
Appendix I and expected to be upgraded to Critically
Endangered when the IUCN Red List is updated in
2016. It is endemic to the mixed lowland and montane
forests of the Albertine Ri escarpment in eastern
Democratic Republic of Congo (DRC), and although
formerly known as the eastern lowland gorilla, the
name is misleading as it ranges between 600 m and
2,900 m above sea level. e diet of Grauer’s gorillas is
rich in herbs, leaves, bark, lianas and vines, seasonally-
available fruit, bamboo (at higher altitudes) and insects,
and they show a preference for regenerating forests
associated with abandoned elds and villages (Schaller
1963; Yamagiwa et al. 2005; Nixon et al. 2006).
e rst surveys of Grauer’s gorillas were conducted in
1959 (Emlen and Schaller 1960). is landmark study
observed that eastern gorillas were rare west of the
Great Lakes, had a highly discontinuous distribution
and were severely threatened by hunting and habitat
destruction. During the 1960s, through to the late
1980s, habitat conversion in the eastern part of their
range intensied, destroying almost all montane forest
outside of protected areas and exterminating a number
of important high-altitude populations. Widespread
killing of gorillas for bushmeat and in retaliation for
crop raiding is likely to have impacted populations
across their entire range during this period.
Gorilla beringei graueri Matschie, 1914
Democratic Republic of Congo
(2010, 2012, 2014)
Stuart Nixon & Elizabeth A. Williamson
Grauer’s gorilla (Gorilla beringei graueri) (Illustration: Stephen D. Nash)
In the 1990s, eorts were made to determine the status of
Grauer’s gorilla in Maiko National Park (MNP; Hart and
Sikubwabo 1994), Kahuzi-Biega National Park (KBNP)
and adjacent forests (Hall et al. 1998a) and the Itombwe
Massif (Omari et al. 1999). From these surveys, Hall et
al. (1998b) concluded that Grauer’s gorillas remained
highly threatened across their range, and estimated
the total number surviving to be approximately 16,900
individuals, with KBNP and MNP supporting the
reats to the gorillas intensied enormously throughout
the 1990s and 2000s, due to persistent civil conict in
the eastern border regions of DRC. Refugees, internally
displaced people and armed groups settled throughout
the east of the country, putting enormous pressure on
natural resources, national parks included. KBNP and
MNP have been at the epicentre of this intense and illegal
resource extraction for the past 20 years. e isolated
KBNP highland population was decimated in the early
2000s (Amsini et al. 2008) following occupation by
rebel forces. e status of gorillas in northern MNP is
unknown as rebels control the illegal gold mines and
consequently much of the park remains inaccessible.
DRC’s rst community-managed protected areas—the
Tayna Nature Reserve and the Kisimba Ikoba Nature
Reserve (created in the mid 2000s specically to protect
their gorilla populations)—remain o limits due to land
tenure disputes and occupation by several rebel groups.
e Congolese Institute for Nature Conservation
(ICCN) faces continuing conicts with armed groups,
and highly dedicated ICCN personnel have been killed
in the line of duty while attempting to protect Grauer’s
gorilla populations and their habitat. Destruction of
forest for timber, charcoal production and agriculture
continue to threaten the isolated gorilla populations
that persist in the North Kivu highlands and Itombwe
Massif, while poaching presents a serious and immediate
threat to these gorillas across their entire range. Large
numbers of military personnel stationed in rural areas
and numerous rebel groups still active throughout the
region have been implicated in illegal mining activities
and have facilitated access to the rearms that fuel both
the ongoing insecurity and an illegal bushmeat trade
on a commercial scale. Since 2003, ICCN and partners
have conscated 15 Grauer’s gorilla infants—casualties
Conservation challenges are likely to increase as the DRC
government continues its eorts to stabilize the east.
Security will favour industrial extraction, large-scale
agriculture and infrastructure. While development will
increase the country’s ability to support its population
and participate in the global economy, it will also result
in increased human settlement in forest areas critical for
gorillas. Targeted conservation action in priority sites
will be vital to slow further demise of this subspecies.
To address the critical situation faced by Grauer’s
gorillas, international and local NGOs are working with
the government authorities to support protected area
rehabilitation and reinforce conservation programmes.
A conservation strategy with clear priorities for Grauer’s
gorillas has been published by IUCN (Maldonado et al.
2012). is action plan recognises four, broadly-dened
population centres: Maiko-Tayna-Usala (including
MNP and adjacent forests, Tayna Nature Reserve,
Kisimba-Ikoba Nature Reserve and the Usala forest),
Kahuzi-Kasese (including the lowland sector of KBNP
and adjacent forests), KBNP highlands and the Itombwe
Massif. In collaboration with ICCN, a consortium of
NGOs led by the Wildlife Conservation Society (WCS)
and Fauna & Flora International is completing a two-
year project to assess the status of Grauer’s gorilla
across its range. Until the results of ongoing surveys are
available, our best guesstimate from data collated during
the past 14 years is that Grauer’s gorilla numbers have
been reduced to 2,000–10,000 individuals (Nixon et al.
2012). Further evidence for such a decline comes from
an analysis of ape habitat across Africa, which estimates
that suitable environmental conditions for Grauer’s
gorillas been have halved since the 1990s (Junker et al.
ICCN and partners made signicant progress during
2014 and 2015, largely regaining control of KBNP
and the southern sector of MNP and re-establishing a
conservation presence. Signicant gains have also been
made in the Itombwe Nature Reserve, which will help
protect the core of the Itombwe Massif and its highly
fragmented gorilla population. Outside protected areas,
regular community-based gorilla monitoring has been
set up in Lubutu, Kasese and Biruwe/Nkuba, and may
be expanded to other remote regions, such as the Usala
forest. Conscated gorilla orphans are now cared for at
the Gorilla Rehabilitation and Conservation Education
(GRACE) Centre near Lubero in North Kivu, and
the possibility of reintroducing these gorillas at Mt.
Tshiaberimu in Virunga National Park oers hope for
small yet isolated subpopulations in well-protected sites.
e steady recovery of the KBNP highland population
(WCS unpublished data) is encouraging evidence that
highly-targeted conservation eorts can be successful
even in the face of acute and sustained human pressures.
Amsini, F., O. Ilambu, I. Liengola, D. Kujirakwinja, J.
Hart, F. Grossman and A. J. Plumptre. 2008. e Impact
of Civil War on the Kahuzi-Biega National Park: Results
of Surveys between 2000–2008. Wildlife Conservation
Society and Institut Congolais pour la Conservation de
la Nature, Kinshasa, DRC.
Emlen, J. T. and G. B. Schaller. 1960. Distribution and
status of the mountain gorilla (Gorilla gorilla beringei).
Zoologica 45: 41–52.
Hall, J. S., L. J. T. White, B. I. Inogwabini, I. Omari,
H. S. Morland, E. A. Williamson, K. Saltonstall, P.
Walsh, C. Sikubwabo, D. Bonny, P. Kiswele, A. Vedder
& K. Freeman. 1998a. Survey of Grauer’s gorillas
(Gorilla gorilla graueri) and eastern chimpanzees (Pan
troglodytes schweinfurthii) in the Kahuzi-Biega National
Park lowland sector and adjacent forest in eastern
Democratic Republic of Congo. International Journal of
Primatology 19: 207–235.
Hall, J. S., K. Saltonstall, B. I. Inogwabini and I. Omari.
1998b. Distribution, abundance and conservation status
of Grauer’s gorilla. Oryx 32: 122–130.
Hart, J. A. and C. Sikubwabo. 1994. Exploration of
the Maiko National Park of Zaire 1989–1992. Wildlife
Conservation Society, New York, USA.
Junker, J., S. Blake, C. Boesch, G. Campbell, L. du
Toit, C. Duvall, G. Etoga, A. Galat-Luong, J. Gamys,
J. Ganas-Swaray, S. Gatti, A. Ghiurghi, N. Granier, J.
Hart, J. Head, I. Herbinger, T. C. Hicks, B. Huijbregts,
I. S. Imong, N. Kumpel, S. Lahm, J. Lindsell, F. Maisels,
M. McLennan, L Martinez, B. Morgan, D. Morgan, F.
Mulindahabi, R. Mundry, K. P. N’Goran, E. Normand,
A. Ntongho, D. T. Okon, C. A. Petre, A. Plumptre, H.
Rainey, S. Regnaut, C. Sanz, E. Stokes, A. Tondossama,
S. Tranquilli, J. Sunderland-Groves, P. Walsh, Y. Warren,
E. A. Williamson and H. S. Kuehl. 2012. Recent decline
in suitable environmental conditions for African great
apes. Diversity and Distributions 18: 1077–1091.
Maldonado, O., C. Aveling, D. Cox, S. Nixon, R. Nishuli,
D. Merlo, L. Pintea and E. A. Williamson. 2012. Grauer’s
Gorillas and Chimpanzees in Eastern Democratic
Republic of Congo (Kahuzi-Biega, Maïko, Tayna and
Itombwe Landscape): Conservation Action Plan 2012–
2022. IUCN/SSC Primate Specialist Group, Ministry
of Environment, Nature Conservation and Tourism,
Institut Congolais pour la Conservation de la Nature,
and Jane Goodall Institute, Gland, Switzerland.
Nixon, S., E. Emmanuel, K. Mufabule, F. Nixon, D.
Bolamba and P. Mehlman. 2006. e Post-conict
Status of Grauer’s Eastern Gorilla (Gorilla beringei
graueri) and Other Wildlife in the Maïko National
Park Southern Sector and Adjacent Forests, Eastern
Democratic Republic of Congo. Unpublished report,
Institut Congolais pour la Conservation de la Nature
and Dian Fossey Gorilla Fund International, Goma,
Nixon. S., A. J. Plumptre, L. Pintea, J. A. Hart, F. Amsini,
E. Bahati, E. Delattre, C. K. Kaghoma, D. Kujirakwinja,
J. C. Kyungu, K. Mufabule, R. Nishuli and P. Ngobobo.
2012. e forgotten gorilla; historical perspectives
and future challenges for conserving Grauer’s gorilla.
Abstract #641. XXIV Congress of the International
Primatological Society, Cancun, Mexico.
Omari, I., J. A. Hart, T. M. Butynski, N. R. Birhashirwa,
A. Upoki, Y. M’keyo, F. Bengana, M. Bashonga and N.
Bagurubumwe. 1999. e Itombwe Massif, Democratic
Republic of Congo: biological surveys and conservation,
with an emphasis on Grauer’s gorilla and birds endemic
to the Albertine Ri. Oryx 33: 301–322.
Schaller, G. B. 1963. e Mountain Gorilla: Ecology and
Behav ior. University of Chicago Press, Chicago, IL.
Yamagiwa, J., A. K. Basabose, K. Kaleme and T Yumoto.
2005. Diet of Grauer’s gorillas in the montane forest of
Kahuzi, Democratic Republic of Congo. International
Journal of Primatology 26: 1345–1373.
0 100 200 300 40050
e Lavasoa Mountains Dwarf Lemur, Cheirogaleus
lavasoensis, was discovered in the Lavasoa-
Ambatotsirongorongo Mountains in 2001 and rst
assigned to Cheirogaleus crossleyi (Hapke et al. 2005).
Genetic data for comparison to other Cheirogaleus
populations were largely lacking at that time and became
available later (Groeneveld et al. 2009, 2010). iele et
al. (2013) nally described Cheirogaleus lavasoensis as a
new species. ey assessed its distinctiveness based on
analyses of mitochondrial and nuclear genetic data.
Cheirogaleus lavasoensis has a head-body-length of
22.2–28.5 cm and a body weight of 248–297 g. It has
black eye rings, dark furry ears, and a darkly pigmented,
pointed nose. e coloration on the crown, forehead, and
neck is intensely reddish-brown and changes gradually
to grey-brown towards the tail. e light creamy ventral
coloration extends into a sharply delimited lateral stripe
on the neck (iele et al. 2013). Cheirogaleus lavasoensis
is nocturnal. Data about its life-history, ecology, and
behavior are not available.
e Lavasoa-Ambatotsirongorongo Mountains are
situated south of the north-southward directed Anosy-
and Vohimena Mountain chains in extreme southern
Madagascar. ese mountains act as a climatic barrier
between rainforest on their eastern anks and dry
spiny bush in their western rain shadow. is sharp
ecological rupture is contrasted by a wide climatic
gradient from dry spiny bush over transitional forest
into humid littoral forest south of the mountain
chains. e Lavasoa-Ambatotsirongorongo Mountains
are situated in the center of this gradient. ey are
surrounded by water and lowlands and isolated from
the Anosy Mountains by a plain at 20–30 m above
sea level. e three main summits are aligned in
west eastern direction: Grand Lavasoa (823 m), Petit
Lavasoa (617 m), and Ambatotsirongorongo (438). A
topographic map based on aerial photographs from
1957 (Foiben-Taosarintanin’i Madagasikara, 1979)
displays one large, continuous forest on their southern
and eastern anks. is forest was isolated from the
huge, continuous humid forest of the Anosy Mountain
Chain by the forestless northern ank of the Lavasoa-
Lavasoa Mountains Dwarf Lemur
Cheirogaleus lavasoensis iele, Razamahatratra and Hapke, 2013
Andreas Hapke, Nary R. J. Andrianjaka & Dana iele
Lavasoa Mountains Dwarf Lemur (Cheirogaleus lavasoensis)
(Illustration: Stephen D. Nash)
Ambatotsirongorongo Mountains and the forestless
plain. Most of this forest has disappeared and only small
fragments remain on the southern anks of the three
main summits. eir oristic composition has been
characterized as a predominantly humid, transitional
mixture (Andrianarimisa et al. 2009; Ramanamanjato
et al. 2002).
Until recently, three of these forest fragments were the
only known habitat of Cheirogaleus lavasoensis (iele
et al. 2013). e three forest fragments had sizes of 50 ha,
30 ha, and 25 ha in 2009 (Andrianarimisa et al. 2009).
Extensive eldwork in nearby transitional, humid, and
littoral forest yielded no evidence for the occurrence
of the species (Hapke et al. 2005; Hapke, Gligor, and
Andrianjaka pers. obs). Lei et al. (2014) reported the
occurrence of Cheirogaleus lavasoensis at Kalambatritra,
170 km north of the Lavasoa-Ambatotsirongorongo
Mountains at an altitude of approximately 1200 m.
e Kalambatritra forest is Madagascar’s westernmost
rainforest and situated on the islands west-eastern
drainage divide in an elevation range of 1200–1680 m
(Irwin et al. 2001). It is separated from the eastern humid
forest by 16 km of open grassland and extends over an
area with a north-south diameter of approximately 60
iele et al. (2013) reported data from 17 individuals
of Cheirogaleus lavasoensis that had been captured and
released between 2001 and 2006. e habitat of the
species in the Lavasoa-Ambatotsirongorongo Mountains
is biogeographically isolated, small, and fragmented. A
survey in 2015 conrmed the persistence of Cheirogaleus
lavasoensis in this area but also a further decline of forest
cover (Andrianjaka and Hapke unpublished). e forest
of Kalambatritra could harbour a considerably greater
population of Cheirogaleus lavasoensis. At present, it is
not possible to estimate its size because Lei et al. (2014)
report data from a single individual that they could
assign to the species. With respect to the dierences
in latitude and elevation, it is probable that there are
strong ecological dierences between Kalambatritra
and the Lavasoa-Ambatotsirongorongo Mountains.
Further research is necessary to gather information
about the population size at Kalambatritra and about
possible local adaptations of Cheirogaleus lavasoensis in
the two areas. e population at Ambatotsirongorongo
thus deserves high conservation priority, at least until
more data become available about the population at
e entire remaining forest in the Lavasoa-
Ambatotsirongorongo Mountains is situated within
the Nouvelle Aire Protégée Ambatotsirongorongo.
e greatest threats for the survival of Cheirogaleus
lavasoensis in this area are further habitat loss and
degradation. e main pressures are wood extraction,
slash-and-burn cultivation, and accidental res, which
are frequent due to the practice of regular grassland
burning. e forest fragments inhabited by Cheirogaleus
lavasoensis are apparently remnants of the most humid
patches within a formerly larger, continuous forest.
Until recently, the local population relied on dryer
forest remnants as main sources of construction timber
and rewood. ese fragments have now mostly
disappeared, which generates the risk of increasing
pressure onto the remaining habitat of Cheirogaleus
lavasoensis. At least two forest fragments inhabited by
Cheirogaleus lavasoensis harbor sources that are used by
the local population for paddy irrigation and drinking
water. A successful conservation program should thus
integrate water management, reforestation and habitat
Andrianarimisa, A., V. Andrianjakarivelo, Z.
Rakotomalala, Z. and M. Anjeriniaina 2009. Vertébrés
terrestres des fragments forestiers de la Montagne
d’Ambatotsirongorongo, site dans le Système des Aires
Protégées de Madagascar de la Région Anosy, Tolagnaro.
Malagasy Nature 2: 30–51.
Foiben-Taosarintanin’i Madagasikara. 1979. Ranopiso,
Feuille M-62, Carte Topographique au 100000. Institut
National de Géodésie et Cartographie, Antananarivo,
Groeneveld, L.F., D. W. Weisrock, R. M. Rasoloarison,
A. D. Yoder and P. M. Kappeler. 2009. Species
delimitations in lemurs: multiple genetic loci reveal
low levels of species diversity in the genus Cheirogaleus.
BMC Evolutionary Biology 9: 30.
Groeneveld, L.F., M. B. Blanco, J. L. Raharison, V.
Rahalinarivo, R. M. Rasoloarison, P. M. Kappeler,
P.M., L. R. Godfrey and M.T. Irwin. 2010. MtDNA and
nDNA corroborate existence of sympatric dwarf lemur
species at Tsinjoarivo, eastern Madagascar. Molecular
Phylogenetics and Evolution 55: 833–845.
Hapke, A., J. Fietz, S. D. Nash, D. Rakotondravony,
B. Rakotosamimanana, J. B. Ramanamanjato, G. F. N.
Randria and H. Zischler. 2005. Biogeography of dwarf
lemurs: genetic evidence for unexpected patterns in
southeastern Madagascar. International Journal of
Primatology 26: 873–901.
Irwin, M.T., K. E. Samonds and J. L. Raharison. 2001. A
biological inventory of the lemur community of Réserve
Spéciale de Kalambatritra, south-central Madagascar.
Lemur News 6: 24–28.
Lei, R., C. L. Frasier, A. T. McLain, J. M. Taylor, C. A. Bailey,
S. E. Engberg, A. L. Ginter, R. Randriamampionona, C.
P. Groves, R. A. Mittermeier and E. E. Louis Jr. 2014.
Revision of Madagascar’s dwarf lemurs (Cheirogaleidae:
Cheirogaleus): designation of species, candidate species
status and geographic boundaries based on molecular
and morphological data. Primate Conservation (28):
Ramanamanjato, J. B., P. B. McIntyre and R. A.
Nussbaum. 2002. Reptile, amphibian, and lemur
diversity of the Malahelo Forest, a biogeographical
transition zone in southeastern Madagascar. Biodiversity
and Conservation 11: 1791–1807.
iele, D., E. Razamahatratra and A. Hapke. 2013.
Discrepant partitioning of genetic diversity in mouse
lemurs and dwarf lemurs - Biological reality or
taxonomic bias? Molecular Phylogenetics and Evolution
Hapalemur alaotrensis exclusively inhabits the dense
papyrus and reed beds surrounding Lac Alaotra,
Madagascar’s largest lake, located on the western edge
of the eastern rain forest region (Mutschler and Feistner
1995). e species occurs as two subpopulations, a
small fragmented one in the northern part of the lake
around the Belempona Peninsula and a larger one in
contiguous marshland along the lake’s southwestern
shore. Its entire range appears to be less than 5,800
ha and it occurs only up to elevations of 750 m. e
lake is surrounded by a vast wetland area consisting
of 14,000 ha of marshes and 175,000 ha of rice elds,
and constitutes an important biodiversity area (Pidgeon
1996). Guillera-Arroita et al. (2010) reported that
diculties in monitoring this species arise because the
wetland is dicult to survey. It can only be accessed via
canals cut by shermen, resulting in limited transects
for monitoring, and visibility is typically restricted to a
e Alaotra watershed is of international importance
under the Ramsar Convention of 2003. e government
of Madagascar also recognized the conservation value
of this area by classifying it as a Temporary New
Protected Area within national law N°381-2007/
MINENVEF/MAEP in January 2007, which was given
permanent protection status in July 2015. However,
the human population in the Alaotra watershed has
rapidly increased in the last few decades, from 109,000
in 1960 to approximately 550,000 in 2003 (Pidgeon
1996; PRD 2003). People rely heavily on rice cultivation
and shing for their livelihoods (Andrianandrasana
et al. 2005; Copsey et al. 2009a; Wallace et al. 2015),
leading to severe loss, degradation and fragmentation
of the Alaotran marshes (Mutschler et al. 1995, 2005).
Conversion of marsh habitat to rice elds has been the
most signicant historical and continuing threat to the
survival of H. alaotrensis. From 2001–2007 there was a
decrease in the coverage of marsh vegetation of 29.7%,
from 19,000 to 14,000 ha (Durrell Wildlife Conservation
Trust unpublished data). Marsh burning inhibits the
regeneration of H. alaotrensis habitat, which also
declined during that same period from 9,400 to 5,800 ha.
Unfortunately, marsh burning increased considerably in
2013 (3,000 ha) and 2014 (2,600 ha) due to a lack of law
enforcement and an increase in coordinated pressure by
people in power organising conversion of the marsh to
riceelds for their own nancial benet (Ratsimbazafy
et al. 2013). Some of this burned marsh will regenerate
if not converted to rice but the trend in annual burning
has accelerated. Hunting for food and capture for pets
has signicantly reduced lemur numbers in the past
but has been mitigated somewhat in more recent years
due to intervention by conservation organizations
(Razamananahaka et al. in prep). Various methods
Lac Alaotra Bamboo Lemur
Hapalemur alaotrensis Rumpler, 1975
Fay Clark and Lance Woolaver
Lac Alaotra bamboo lemur (Hapalemur alaotrensis)
(Illustration: Stephen D. Nash)
of hunting and trapping are employed by local people.
Direct pursuit by dogs is the most common, but they
may also be captured by using a harpoon, a snare, a
stick to knock them out or into the water, by burning
their reed bed habitat or just by chasing them down
(Copsey et al. 2009a,b). Commercial drainage projects
represent a potential threat. Regular burning to increase
cattle pasture and facilitate local shing reduces suitable
lemur habitat and also promotes the invasion of exotic
plant species that may choke the remaining marshes.
According to the most recent IUCN Red List
assessment, H. alaotrensis is now Critically Endangered
(Andriaholinirina et al. 2014). eir numbers have
decreased from >10,000 individuals in 1994 (Mutschler
and Feistner 1995) to <3,000 in 2002 (Ralainasolo 2004).
e most recent population estimates for H. alaotrensis
range from 2,500–5,000 individuals, representing a
decline of approximately 30% in just over a decade
(Ralainasolo 2004; J. Ratsimbazafy, unpublished data).
Existing habitat is so fragmented that gene ow between
existing populations is greatly reduced, aecting the
long-term viability of populations (Ralainasolo et
al. 2006). anks to eorts of the Durrell Wildlife
Conservation Trust and local community associations,
the Lac Alaotra New Protected Area was recently
granted permanent protection status, providing an
ocial legal framework for the co-management of the
Protected Area by local communities living around the
wetland. is includes both a strict conservation area
of 8,000 ha, and a surrounding 5,200 ha zone where
controlled activities (e.g. shing) are permitted. In
addition, public awareness campaigns continue to focus
on the benets of habitat conservation to the half million
or more people who live by the lake-erosion control,
the biological ltering of agricultural pollutants, and
ood prevention. Hapalemur alaotrensis is currently
being used as a agship species by Madagascar Wildlife
Conservation (MWC), where the economic benets
from its ‘Camp Bandro’ (Bandro is the vernacular word
for H. alaotrensis) are invested back into community
development (Ratsimbazafy et al. 2013; Rendigs et al.
More work is needed to secure the future of H. alaotrensis,
a unique wetland lemur experiencing severe conict
with humans (Waeber et al. in press). Ratsimbazafy et
al. 2013 suggest more stringent policy and management
mechanisms to halt marsh conversion around the lake,
and the general need to improve the socio-economic
status of local people in order for conservation measures
to be eective. e eective management of the New
Protected Area (NAP) will provide the legal framework
by which marsh protection and wise management
of the lake and marshes can be linked to sustainable
rural development that has a positive impact on the
environment. Immediate actions to ensure that the
New Protected Area is operational include the physical
delimitation of the NAP and capacity-building and
support for the management structure of the NAP
which will be co-managed by local communities and
resource users around Lac Alaotra. A more eective
system for censusing H. alaotrensis, potentially using
drone technology, is required to improve monitoring
and rene annual population estimates (Guillera-
Arroita et al. 2010).
Andriaholinirina et al. 2014. Hapalemur alaotrensis.
IUCN Red List of reatened Species 2014. <www.
iucnredlist.org>. Accessed 26 October 2015.
Andrianandrasana, H. T., J. Randriamahefasoa, J.
Durbin, R. E. Lewis and J. H Ratsimbazafy. 2005.
Participatory ecological monitoring of the Alaotra
wetlands in Madagascar. Biodiversity Conservation 14:
Copsey, J. A., L. H. Rajaonarison, R. Randriamihamina
and L. J. Rakotoniaina. 2009a. Voices from the marsh:
livelihood concerns of shers and rice cultivators in
the Alaotra wetland. Madagascar Conservation and
Development 4: 25–30
Copsey, J. A., J. P. G. Jones, H. Andrianandrasana, L. H.
Rajaonarison and J. E. and Fa. 2009b. Burning to sh:
local explanations for wetland burning in Lac Alaotra,
Madagascar. Oryx 43: 1–4.
Guillera-Arroita, G., J. J. Lahoz-Monfort, E. J. Milner-
Gulland, R. P. Young and E. Nicholson. 2010. Using
occupancy as a state variable for monitoring the
Critically Endangered Alaotran gentle lemur Hapalemur
alaotrensis. Endangered Species Research 11: 157–166.
Mutschler T., A. J. Randrianarisoa and A. T. C. Feistner.
2001. Population status of the Alaotran gentle lemur
(Hapalemur griseus alaotrensis). Oryx 35: 152–157.
Mutschler, T. and A. T. C. Feistner. 1995. Conservation
status and distribution of the Alaotran gentle lemur
Hapalemur griseus alaotrensis. Oryx 29: 267–274.
Mutschler, T., C. Nievergelt and A. T. C. Feistner. 1995.
Human-induced loss of habitat at Lac Alaotra and its
eect on the Alaotran gentle lemur. In: Environmental
Change in Madagascar, B. D. Patterson, S. M. Goodman
and J. L. Sedlock (eds), pp.35–36, 103–104. Field
Museum of Natural History, Chicago.
Pidgeon, M. 1996. An ecological survey of Lake Alaotra
and selected wetlands of central and eastern Madagascar
in analysing the demise of Madagascar pochard Aythy a
innotata. Unpublished report for World Wide Fund for
Nature and Missouri Botanical Gardens, Antananarivo.
PRD (Plan Régional de Développement). 2003. Zone
de Développement Rural Intégré de l’Alaotra (Z.D.R.I),
Ralainasolo, F. B. 2004 Inuence des eets anthropiques
sur la dynamique de population de Hapalemur griseus
alaotrensis ou ‘bandro’ dans son habitat naturel. Lemur
News 9: 32–35.
Ralainasolo, F. B., P. O. Waeber, J. Ratsimbazafy, J.
Durbin and R. E. Lewis. 2006. e Alaotra gentle lemur:
Population estimation and subsequent implications.
Madagascar Conservation & Development 1: 9–10.
Ratsimbazafy, J.H., F. B. Ralainasolo, A. Rendigs,
J. Mantilla Contreras, H. Andrianandrasana, A. R.
Mandimbihasina, C. M. Nievergelt, R. Lewis and P. O.
Waeber. 2013. Gone in a pu of smoke? Hapalemur
alaotrensis at great risk of extinction. Lemur News 17:
Razamananahaka, J. H., N. Piludu, J. P. G. Jones, L.
Woolaver, H. Andrianandrasana and R. Jenkins. In
prep. Assessment of hunting pressure on the Critically
Endangered Alaotran bamboo lemur Hapalemur
Rendigs, A., L. M. Reibelt, F. B. Ralainasolo, J. H.
Ratsimbazafy and P. O. Waeber. 2015. Ten years into the
marshes – Hapalemur alaotrensis conservation, one step
forward and two steps back? Madagascar Conservation
& Development 10: 13–20.
Waeber, P. O., J. H. Ratsimbazafy, H. Andrianandrasana,
F. B. Ralainasolo and C. M. Nievergelt. In press.
Hapalemur alaotrensis, a conservation case study from
the swamps of Alaotra, Madagascar. In: Primates in
Flooded Habitats: Ecology and Conservation. Cambridge
University Press, Cambridge, UK.
Wallace, A. P. C., E. J. Milner-Gulland, J. P. G. Jones,
N. Bunnefeld, R. Young and E. Nicholson. 2015.
Quantifying the short-term costs of conservation
interventions for shers at Lake Alaotra, Madagascar.
PLoS ONE 10(6), e0129440.
e red rued lemur is conned to the Masoala
Peninsula and the region immediately north of the Bay
of Antongil in northeastern Madagascar (Petter and
Petter-Rousseaux 1979; Tattersall 1982). It may have
occurred as far north as Antalaha in the past, but this
is not certain (Tattersall 1977). e Antainambalana
River appears to separate it from V. variegata subcincta,
and recent surveys have shown that the westernmost
distribution of V. r ub ra is near the conuence of the
Antainambalana and Sahantaha rivers (Hekkala et al.
2007). Variations in colour pattern are well known in this
species, but have not been attributed to clear geographic
regions. It may intergrade with V. variegata subcincta;
the conuence of the Vohimara and Antainambalana
rivers has been investigated as a possible contact or
hybrid zone between the two, but without conclusive
results (Tattersall 1982; Lindsay and Simons 1986; Vasey
and Tattersall 2002; Hekkala et al. 2007).
With a head-body-length of 50–55 cm and a body mass
of 3.0–3.6 kg (Vasey 2003), Varecia rubra is a large
member of the Lemuridae. It inhabits primary and some
secondary moist lowland forest (up to 1200 m above sea
level) and prefers tall forest, where it is oen observed
in the crowns of large feeding trees. e species feeds
mainly on fruit, supplemented with owers, nectar,
and leaves. In one study conducted between May and
November (Rigamonti 1993), red rued lemurs fed on
ripe fruits for 73.9% of their feeding time, owers for
5.3%, and leaves for 20.9% (18.3% of these mature).
Only a few plant species were used as food resources:
72.5% of the observed feeding bouts occurred in only
seven tree species. e animals fed on 42 plant species
altogether, compared to 106 species that would have
been available to them in their home range area. e
composition of the diet varied from month to month,
but fruits were consistently the main item, even when
they were hard to nd. e core areas used within their
territories always correlated with large, fruit-bearing
trees. In the cold-wet season, when few fruits are
available, groups split up into subgroups to use dierent
core areas. Females are reported to eat more low-bre,
high-protein items (young leaves and owers) prior to
giving birth and during lactation, presumably to meet
the higher energy demands of reproduction (Vasey
2000a, 2002). At Andranobe, 132 dierent plant species
from 36 families were eaten over the course of a year
is species has been studied in the forests of
Ambatonakolahy (Rigamonti 1993) and Andranobe
(Vasey 1997a) on the Masoala Peninsula. Social
organization is described as ssion/fusion, and
communities are usually multimale-multifemale and
number 5–31 individuals. Home ranges cover 23–58
ha and appear to be defended (Rigamonti 1993; Vasey
Red Rued Lemur
Varecia rubra (É. Georoy Sainte-Hilaire, 1812)
Christoph Schwitzer, Russell A. Mittermeier, Edward E. Louis Jr. & Matthew Richardson
Red Rued Lemur
(Illustration: Stephen D. Nash)
2006). In one study at Andranobe, V. r u br a spent 28%
of its time feeding, 53% resting, and 19% traveling.
Females fed more and rested less than males (Vasey
2005). e species is most active during the hot rainy
season. Mating occurs in early July, and infants are born
in October and fully weaned by February (Vasey 2007).
e red rued lemur is classied as Critically Endangered
(Andriaholinirina et al. 2014) based on a suspected
population reduction of ≥80% over a 3-generation time
period of 24 years. e principal threats to the species
are habitat loss and hunting (Simons and Lindsay
1987; Rigamonti 1996; Vasey 1996, 1997b; Borgerson
2015). Because of their large size and evident need
for tall primary forest, these animals are particularly
susceptible to human encroachment (Borgerson 2015),
and hunting and trapping for food still takes place.
Furthermore, remaining populations are concentrated
on the Masoala Peninsula, where they are threatened by
the frequent cyclones that hit this part of Madagascar.
e only protected area where Varecia rubra is known
to occur is Masoala National Park (Kremen 1998).
Masoala was the national park most aected by the very
rapid upsurge of illegal logging aer the political events
of early 2009, and this logging continued well into
2010. Population density has been variously estimated
at 6 individuals/km² (Rakotondratsima and Kremen
2001; Borgerson 2015 [at an unnamed village site]),
17 individuals/km² (Borgerson 2015 [at an unnamed
forest site away from villages]), 21–23 individuals/
km² in Ambatonakolahy (Rigamonti 1993), and 31–54
individuals/km² in Andranobe (Vasey 1997b).
e IUCN lemur conservation strategy 2013–2016
(Schwitzer et al. 2013) proposes a suite of conservation
measures for Masoala National Park to ensure the
conservation of the red rued lemur: further patrols and
surveillance; campaigns of environmental education
and awareness; and support for small-scale husbandry
of domestic animals as a source of protein. As of 2015,
there were 600 red rued lemurs reported in captivity
worldwide (ISIS 2015). Such populations in American
and European zoos represent a safeguard against
extinction, but they are unfortunately very limited in
their genetic diversity (Schwitzer 2003).
Andriaholinirina et al. 2014. Varecia rubra. In:
e IUCN Red List of reatened Species 2014:
e.T22920A16121712. Accessed 09 September 2015.
Borgerson, C. 2015. e eects of illegal hunting
and habitat on two sympatric endangered primates.
International Journal of Primatology 36: 74–93.
Hekkala, E. R., M. Rakotondratsima and N. Vasey.
2007. Habitat and distribution of the rued lemur,
Varec ia, north of the Bay of Antongil in north-eastern
Madagascar. Primate Conservation (22): 89–95.
ISIS. 2015. Zoological Information Management System
(ZIMS) version 2.2, released 29 June 2015. ISIS, Apple
Kremen, C. 1998. Madagascar creates its largest
protected area on the Masoala Peninsula. Lemur News
Lindsay, N. B. D. and H. J. Simons. 1986. Notes on
Varec ia in the northern limits of its range. Dodo 23:
Petter, J.-J. and A. Petter-Rousseaux. 1979. Classication
of the prosimians. In: e Study of Prosimian Behavior,
G. A. Doyle and R. D. Martin (eds.), pp.359–409.
Academic Press, New York.
Rakotondratsima, M. and C. Kremen. 2001. Suivi
écologique de deux espèces de lémuriens diurnes
Varecia variegata rubra et Eulemur fulvus albifrons dans
la presqu’île de Masoala (1993–1998). Lemur News 6:
Rigamonti, M. M. 1993. Home range and diet in red
rued lemurs (Varecia varigata rubra) on the Masoala
Peninsula, Madagascar. In: Lemur Social Systems and
their Ecological Basis, P. M. Kappeler and J. U. Ganzhorn
(eds.), pp.25–39. Plenum Press, New York.
Rigamonti, M. M. 1996. Red rued lemur (Varecia
variegata rubra): a rare species from the Masoala rain
forests. Lemur News 2: 9–11.
Schwitzer, C. 2003. Energy Intake and Obesity in
Captive Lemurs (Primates, Lemuridae). Doctoral
thesis, Universität zu Köln. Schüling Verlag, Münster,
Schwitzer, C., R. A. Mittermeier, N. Davies, S. Johnson,
J. Ratsimbazafy, J. Razandramanana, E. E. Louis Jr. and
S. Rajaobelina (eds.). 2013. Lemurs of Madagascar: A
Strategy for eir Conservation 2013–2016. IUCN SSC
Primate Specialist Group, Bristol Conservation and
Science Foundation, and Conservation International,
Bristol, UK. 185pp.
Simons, H. J. and N. B. D. Lindsay. 1987. Survey work
on rued lemurs (Varecia variegata) and other primates
in the northeastern rain forests of Madagascar. Primate
Conservation (8): 88–91.
Tattersall, I. 1977. Distribution of the Malagasy lemurs
part 1: e lemurs of northern Madagascar. Annals of
the New York Academy of Sciences 293: 160–169.
Tattersall, I. 1982. e Primates of Madagascar. Columbia
University Press, New York.
Vasey, N. 1996. Clinging to life: Varecia variegata rubra
and the Masoala coastal forests. Lemur News 2: 7–9.
Vasey, N. 1997a. Community Ecology and Behavior
of Varecia variegata rubra and Lemur fulvus albifrons
on the Masoala Peninsula, Madagascar. PhD thesis,
Washington University, St. Louis, MO.
Vasey, N. 1997b. How many red rued lemurs are le?
International Journal of Primatology 18: 207–216.
Vasey, N. 2000a. Niche separation in Varecia variegata
rubra and Eulemur fulvus albifrons: I. Interspecic
patterns. American Journal of Physical Anthropology
Vasey, N. 2000b. Plant species composition of diet in
two sympatric lemurs: Varecia variegata rubra and
Eulemur fulvus albifrons. American Journal of Physical
Anthropology 30 (suppl.): 309–310.
Vasey, N. 2002. Niche separation in Varecia variegata
rubra and Eulemur fulvus albifrons: II. Intraspecic
patterns. American Journal of Physical Anthropology
Vasey, N. 2003. Vare ci a , rued lemurs. In: S. M.
Goodman and J. P. Benstead (eds), e Natural History
of Madagascar, pp.1332–1336. Aldine de Gruyter, New
Yor k .
Vasey, N. 2005. Activity budgets and activity rhythms
in red rued lemurs (Varecia rubra) on the Masoala
Peninsula, Madagascar: seasonality and reproductive
energetics. American Journal of Primatology 66: 23–44.
Vasey, N. 2006. Impact of seasonality and reproduction
on social structure, ranging patterns, and ssion-
fusion social organization in red rued lemurs. In: L.
Gould and M. A. Sauther (eds), Lemurs: Ecology and
Adaptation, pp.275–304. Springer, New York.
Vasey, N. 2007. e breeding system of wild red rued
lemurs (Varecia rubra): a preliminary report. Primates
Vasey, N. and I. Tattersall. 2002. Do rued lemurs
form a hybrid zone? Distribution and discovery of
Varec ia, with systematic and conservation implications.
American Museum Novitates (3376): 1–26.
Originally described based on cytogenetic and
morphometric characteristics (Rumpler and Albignac
1975), the taxonomic status of the northern sportive
lemur (Lepilemur septentrionalis) has since been
supported by more detailed cytogenetic, morphogenetic
and especially molecular data (Ravoarimanana et al.
2004; Andriaholinirina et al. 2006; Louis et al. 2006),
and subsequently accepted in recent taxonomic
revisions of primates (Groves 2001, 2005) and lemurs
(Mittermeier et al. 2008; Mittermeier et al. 2010). With
the taxonomic revision conrming L. septentrionalis
and L. ankaranensis as distinct species, the perceived
range of the northern sportive lemur was drastically
reduced; limited to a few degraded patches of dry
forest in the Sahafary region just south of Antsiranana.
e number of animals observed during surveys has
decreased dramatically over the past ten years. e rst
signicant survey of the northern sportive lemur was
performed in 2001 by I. Ravoarimanana, following by
another one in 2007 by A. Zaramody in the Andrahona,
Ankarakataova, and Sahafary regions. e population
was then estimated at about 120 individuals, mainly
concentrated in the Sahafary area.
Expeditions by Omaha’s Henry Doorly Zoo and
Aquarium (OHDZA) and the Madagascar Biodiversity
Partnership (MBP) in 2010 and 2011 veried the
continued existence of the northern sportive lemur,
albeit noting a tremendous decline in the Sahafary
classied forest. Furthermore, the surveys did not
detect a single animal in the Analalava forest where it
had previously been seen in 2005. A follow-up survey
in July 2012 to Analalava did however document one
individual (Ranaivoarisoa et al. 2013). Fortunately,
Ranaivoarisoa et al. (2013) also conrmed the presence
of the northern sportive lemur in Montagne des
Français (MDF) in 2010, but could only identify 19
individuals across its range based on capture and direct
visual observations. Further surveys of the Montagne
des Français region in 2012–2013 by OHDZA and MBP,
including the previously known habitats of Sahafary
and Analalava classied forests through to its northern
extent in MDF, documented only 52 L. septentrionalis
individuals, with 95% of these lemurs located in MDF.
e most recent population estimates based on only
capture surveys in 2013 in the Montagne des Français
area provided the following population estimates:
1) Abatoire - 7 individuals; 2) Andranonakomba - 2
individuals; 3) Ampamakiampafana - 11 individuals; 4)
Ambatobe - 2 individuals; and Berambo - 5 individuals.
is is a total of 27 individuals. e species was also
documented at Anketrakala and Ampitsinjozatsambo
in 2012, which were not recently surveyed. Surveys in
the region continue to estimate the population around
50 individuals in 2015.
Northern Sportive Lemur
Lepilemur septentrionalis Rumpler and Albignac, 1975
(2008, 2010, 2012, 2014)
Edward E. Louis Jr. & John R. Zaonarivelo
Northern sportive lemur (Lepilemur septentrionalis)
(Illustration: Stephen D. Nash)
In 2008, the Service d’Appui à la Gestion de
l’Environnement promoted the designation of
Montagne des Français as a newly protected area,
and supported the development of a Vondron’Olona
Ifototra (VOI) in Andavakoera, the primary village
of this mountain forest. However, sustained human
encroachment from the city of Antsiranana continues
to nance the production of charcoal and collection of
sand, activities that are threatening this last remaining
northern sportive lemur population (Ranaivoarisoa et
al. 2013). us, habitat loss from uncontrolled long-term
charcoal practices is the primary challenge to overcome.
e northern sportive lemur is nocturnal, spending
the day sleeping in tree holes, and very little is known
about its ecology and behaviour. However, recent work
has shown that its folivorous diet and predilection for
new-growth leaves complicates any attempts or plans
to maintain it in captivity. Currently, there is no record
of any sportive lemur held in any zoological park, as
all known attempts to maintain them in captivity
have failed; on average within one week of capture.
In situ conservation programs and community-based
interactions are, therefore, the only viable solutions. e
combination of a very small range composed of rapidly
deteriorating suitable habitat with high pressure from
charcoal production continues to place the Critically
Endangered northern sportive lemur (Andrainarivo et
al. 2011) on the cusp of extinction.
Andrainarivo, C., V. N. Andriaholinirina, A. T. C.
Feistner, T. Felix, J. U. Ganzhorn, N. Garbutt, C.
Golden, W. B. Konstant, E. E. Louis Jr., D. M. Meyers,
R. A. Mittermeier, A. Perieras, F. Princee, J. C.
Rabarivola, B. Rakotosamimanana, Rasamimanana, H.,
J. Ratsimbazafy, G. Raveloarinoro, A. Razamanantsoa,
Y. Rumpler, C. Schwitzer, R. Sussman, U. almann, L.
Wilmé and P. C. Wright. 2011. Lepilemur septentrionalis.
In: IUCN 2013. IUCN Red List of reatened Species.
Version 2013.2. <www.iucnredlist.org>. Accessed 16
Andriaholinirina, N., J.-L. Fausser, C. Roos., D. Zinner,
U. almann, C. Rabarivola, I. Ravoarimanana, J.
U. Ganzhorn, B. Meier, R. Hilgartner, L. Walter., A.
Zaramody, C. Langer, T. Hahn, E. Zimmermann, U.
Radespiel, M. Craul, J. Tomiuk, I. Tattersall and Y.
Rumpler. 2006. Molecular phylogeny and taxonomic
revision of the sportive lemurs (Lepilemur, Primates).
BMC Evolutionary Biology 6: 17.
Groves, C. P. 2001. Primate Taxonomy. Smithsonian
Institution Press, Washington, DC.
Groves, C. P. 2005. Order Primates. In: Mammal Species
of the World: A Taxonomic and Geographic Reference, 3rd
edition, D. E. Wilson and D. M. Reeder (eds.), pp.111–
184. Johns Hopkins University Press, Baltimore, MD.
Louis Jr., E. E., S. E. Engberg, R. Lei, H. Geng, J. A.
Sommer, R. Randriamampionona, J. C. Randriamanana,
J. R. Zaonarivelo, R. Andriantompohavana, G. Randria,
R. B. Prosper, G. Rakotoarisoa, A. Rooney and R.
A. Brenneman. 2006. Molecular and morphological
analyses of the sportive lemurs (Family Megaladapidae:
genus Lepilemur) reveals 11 previously unrecognized
species. Texas Tech University Museum, Special
Publications 49: 1–47.
Mittermeier, R. A., J. U. Ganzhorn, W. R. Konstant, K.
Glander, I. Tattersall, C. P. Groves, A. B. Rylands, A.
Hapke, J. Ratsimbazafy, M. I. Mayor, E. E. Louis Jr., Y.
Rumpler, C. Schwitzer and R. M. Rasoloarison. 2008.
Lemur diversity in Madagascar. International Journal of
Primatology 29: 1607–1656.
Mittermeier, R. A., E. E. Louis Jr., M. Richardson, C.
Schwitzer, O. Langrand, A. B. Rylands, F. Hawkins, S.
Rajaobelina, J. Ratsimbazafy, R. Rasoloarison, C. Roos,
P. M. Kappeler and J. Mackinnon. 2010. Lemurs of
Madagascar, 3rd edition. Conservation International
Tropical Field Guide Series, Arlington, VA.
Ranaivoarisoa, J. F., J. R. Zaonarivelo, R. Lei, S. E.
Johnson, T. M. Wyman, R. A. Mittermeier, and E. E.
Louis Jr. 2013. Rapid survey and assessment of the
northern sportive lemur, Lepilemur septentrionalis, in
northern Madagascar. Primate Conservation (27): 23–
Ravoarimanana, I. B., R. Tiedemann, D. Montagnon
and Y. Rumpler. 2004. Molecular and cytogenetic
evidence for cryptic speciation within a rare endemic
Malagasy lemur, the northern sportive lemur (Lepilemur
septentrionalis). Molecular Phylogenetics and Evolution
Rumpler, Y. and R. Albignac. 1975. Intraspecic
chromosome variability in a lemur from north of
Madagascar: Lepilemur septentrionalis, species nova.
American Journal of Physical Anthropology 42: 425–429.
Perrier’s sifaka (Propithecus perrieri) is intermediate in
size relative to other members of the genus Propithecus
(Ranaivoraisoa et al. 2006; Lehman et al. 2005) and is
characterized by an all-black pelage, naked black face
and striking orange-red eyes (Mittermeier et al. 2010).
At present, Perrier’s sifaka occurs only in dry deciduous
forests on limestone karst and semi-evergreen
transitional forests on sandstone soils and has a diet of
predominantly leaves and owers (Lehman and Mayor
2004). Little is known about many details of its biology
including its behaviour and past distribution. Its current
distribution is the smallest of all Propithecus species and
it has been recognized as Critically Endangered since
1996 (IUCN 2015; Mittermeier et al. 2010; Salmona
et al. 2013). Its geographic range is restricted to the
extreme northeast of Madagascar, some 50 km to the
south of Antsiranana (Diego Suarez). It extends from
the eastern edges of the Analamerana limestone massif,
along the Indian Ocean coast to the sandstone forests
of the Andraamena mountains as far west as the
peaks of Ambohibe northeast of the rural commune
of Marivorahona. It is bound in the north by the Irodo
River and in the south by the Andraamena mountain
range (Banks 2012; Zaonarivelo et al. 2007). Despite
evidence of the species’ presence in the Ankarana
National Park in the 80s and 90s (Hawkins et al. 1990;
Meyers 1996), three recent surveys in 2003, 2004 and
2012 (Banks et al. 2007; Rasoloharijaona et al. 2005;
Salmona et al. 2013) failed to nd Perrier’s sifakas there.
Furthermore, suggestions that its distribution might
also extend south of the Andraamena mountains and
into the Andavakoera forest (Schwitzer et al. 2006)
could not be conrmed during two surveys to the area
in 2006 and 2012 (Zaonarivelo et al. 2007; Salmona et
Earlier estimates of its total population size based
on total remaining habitat within the species’ range
suggested that less than 1000 individuals persist in the
wild. Estimates of the eective population size from
eld data (about 230 individuals; Banks et al. 2007)
and from genetic data (Ne ~50–100; Salmona et al.
submitted), further support that the population is
small. However, doctoral thesis work by Banks between
2007 and 2012 provided revised estimates of population
density that addressed dierences in habitat quality in
dry and semi-evergreen forest types. Aer modelling
detection in this elusive species as a function of
behaviour, observer, and habitat dierences, the study
reached a population size estimate of 2133 (95% CI:
1761–2584) individuals. Although a small number of
forest patches within the range are not included in these
estimates, the area sampled (252 km2) comprises 85%
of the total remaining habitat available estimated (296.6
Propithecus perrieri Lavauden, 1931
(2000, 2002, 2004, 2014)
Matthew A. Banks, Erik R. Patel, Lounès Chikhi & Jordi Salmona
Perrier’s sifaka (Propithecus perrieri).
(Illustration: Stephen D. Nash)
km2) by Banks et al. (2007), for this species. Population
densities were found to be up to an order of magnitude
higher in sandstone forests. Despite representing only
a small portion of the habitat (about 12%), sandstone
forests likely host nearly 40% of the remaining Perrier’s
sifaka population (Banks 2013). Ominously, a Landsat
7 imagery analysis of the region between 1994 and 2003
reveals that sandstone forests were the most susceptible
to forest loss, with a total loss of more than 60% over
Although its habitat is fragmented, Perrier’s sifaka can
cross open areas for distances of up to 600 m (Mayor
and Lehman 1999). Other sifaka species are known to
disperse over much larger distances of open habitat
(Meyers and Wright 1993; Richard et al. 1993). is
and evidence from studies of Perrier’s sifaka population
levels in eleven forest fragments (range: 1.1 and 124
km2) and occupancy patterns in 45 fragments less
than 1 km2 in area indicate that the population is not
strongly inuenced by the fragmentation of forests and
the matrix of open grassland habitats. However, with
fewer than 2,200 known individuals le in the wild
(Banks 2013), not all of which are reproductive, and a
long generation time of about 18 years, the viability of
the population is at a high risk of being compromised
(generation time between 6 and 20 years, based on data
from Verreaux’s sifaka, P. verreauxi; Lawler et al. 2009;
Morris et al. 2011). Genetic data are extremely limited.
Mayor et al. (2002) identied relatively high levels of
genetic diversity compared to other sifaka species and
useful genetic markers for the genus Propithecus. Some
of these markers together with markers from other
sifaka species were used for the rst population study
conducted by Salmona et al. (submitted). e results of
this study suggest that the three main forest fragments
do not “behave” as dierentiated populations. ey
also suggest that, in the best interests of safeguarding
the species, measures to maintain connectivity between
forest fragments should be implemented (Salmona
et al. submitted). Even though Perrier’s sifaka may
have the ability to cross open grassland, most sifakas
encountered on the ground were elusive and ed from
humans (Salmona et al. submitted). Furthermore,
studies of occupancy patterns show that Perrier’s sifakas
avoid forest patches in proximity to human settlements
with ≥ 10 households where they are susceptible to
attacks from dogs, particularly when attempting to
cross matrix habitat (Banks 2013). e combined eects
of deforestation, fragmentation and human activity
could prevent them from routinely crossing open land,
thereby decreasing gene ow and further fragmenting
the remaining population (Salmona et al. submitted).
Decades ago, several authors reported Perrier’s sifaka
presence outside of its current distribution (Hawkins
et al. 1990; Meyers 1996). Moreover, subfossils of P. cf.
diadema (most probably P. perrieri) were reported in
Ankarana cave (Jungers et al. 1995; Godfrey et al. 1996)
and far north from its current distribution in Montagne
des Français, Andavakoera cave (Dewar et al. 2013 and
reference therein). Perrier’s sifaka paleodistribution
and population size may have been larger than today. In
addition recently found genetic signatures of population
decline suggest that the population underwent a major
decline in the past 5,000 years (Salmona et al. in prep.)
similar to the decline detected in the neighbouring
golden crowned sifaka (Quéméré et al. 2012). Although
it is not clear which events brought Perrier’s sifaka to
its current restricted distribution and small population
size, it is likely due to the conjugated eects of climatic
and human driven forest size uctuations.
ere is also emerging evidence that pressures on
lemur populations in northern Madagascar are on
the rise (Rakotodravony 2006, Reuter et al. 2014) and
local guides indicate that the absence of regular surveys
across the region since the Banks’ studies ended in 2012
has substantially exacerbated the impact from threats
such as hunting, re, land conversion and habitat loss
(J. R. Zaonarivelo pers. comm). Given the small total
population size, persistence of pressures from local
threats and the paucity of wildlife patrols in remote
forests critical for Perrier’s sifaka survival, a return to
monitoring activities and an appraisal of its population
levels are urgently needed. Moreover, a unied regional
management plan is required to safeguard this species
from extinction. e species’ natural range and potential
areas of migration/seasonal presence overlap with three
protected areas all with dierent protected statuses,
independently managed by Madagascar National
Parks (Analamerana and Ankarana) and Fanamby
(Andraamena). Stakeholders involved must operate
at dierent levels of the decision making process (e.g.
park services, ministries, universities, tour operators,
local businesses, farmers, etc.) making the integration
of all perspectives a real challenge. Reaching targets for
conservation with this species given the diverse group
of stakeholders involved requires a clearly dened
institution, committed to leading a species conservation
plan with incentives for action that are inclusive and take
advantage of the strengths of the dierent participants.
Banks, M. A., E. R. Ellis, A. Wright and P. C. Wright.
2007. Global population size of a critically endangered
lemur, Perrier’s sifaka. Animal Conservation 10: 254–
Banks, M. A. 2013. Determinants of Abundance and
the Distribution of Primates in Northern Madagascar.
Unpublished PhD thesis. Stony Brook University, Stony
Brook, New York.
Banks M. A. 2012. Propithecus perrieri (Perrier’s sifaka).
In: All the World’s Primates, N. Rowe, M. Myers (eds.),
Primate Conservation Inc., Charlestown, Rhode Island.
Dewar, R. E., C. Radimilahy, H. T. Wright, Z. Jacobs, G.
O. Kelly and F. Berna. 2013. Stone tools and foraging
in northern Madagascar challenge Holocene extinction
models. Proceedings of the National Academy of Sciences
Godfrey, L. R., W. L. Jungers, E. L. Simons, P. S.
Chatrath and B. Rakotosamimanana. 1999. Past and
present distributions of lemurs in Madagascar. In: New
Directions in Lemur Studies, B. Rakotosamimanana et
al. (eds.), pp.19–53. Springer, New York.
Hawkins, A. F. A., P. Chapman, J.U. Ganzhorn, Q. M. C.
Bloxam, S.C. Barlow and S. J. Tonge. 1990. Vertebrate
conservation in Ankarana special reserve, northern
Madagascar. Biological Conservation 54: 83–110.
IUCN 2015. e IUCN Red List of reatened
Species. Version 2015. <http://www.iucnredlist.org>.
Downloaded on 7 October 2015.
Jungers, W. L., L. R. Godfrey, E. L. Simons and P. S.
Chatrath. 1995. Subfossil Indri indri from the Ankarana
Massif of northern Madagascar. American Journal of
Physical Anthropology 97: 357–366.
Mayor M.I., J.A. Sommer, R.M. Huebinger, C. Barber
and E.E. Louis Jr. 2002. Characterization of seven
microsatellite marker loci in a genus of Malagasy lemurs
(Propithecus). Molecular Ecology Notes 2: 385-388.
Meyers D. M. and P. C. Wright. 1993. Resource tracking:
food availability and Propithecus seasonal reproduction.
In: Lemur Social Systems and eir Ecological Basis, P.
M. Kappeler and J. U. Ganzhorn (eds.), pp.179−192.
New York: Plenum Press.
Meyers D.M. 1996. Update on the endangered sifaka of
the north. Lemur News 2: 13–14.
Mittermeier R. A., E.E. Louis Jr, M. Richardson, C.
Schwitzer, O Langrand, A. B. Rylands, F. Hawkins, S.
Rajaobelina, J. Ratsimbazafy, R. Rasoloarison, C. Roos,
P. M. Kappeler P. M. and J. Mackinnon. 2010. Lemurs
of Madagascar. 3rd edition. Conservation International,
Morris, W. F., J. Altmann, D. K. Brockman, M. Cords, L.
M. Fedigan, A. E. Pusey, T. S. Stoinski, A. M. Bronikowski,
S. C. Alberts and K.B. Strier. 2011. Low demographic
variability in wild primate populations: tness impacts
of variation, covariation, and serial correlation in vital
rates. e American Naturalist 177: E14–28.
Quéméré, E., X. Amelot, J. Pierson, B. Crouau-Roy and L.
Chikhi. 2012. Genetic data suggest a natural prehuman
origin of open habitats in northern Madagascar and
question the deforestation narrative in this region.
Proceedings of the National Academy of Sciences 109:
Rasoloharijaona, S., B. Randrianambinina, B.
Rakotosamimanana and E. Zimmermann. 2005.
Inventaires des lémuriens dans la forêt d’Andranovelona/
Madirovalo (nord ouest de Madagascar), les “savoka” dé
Manehoko, la Réserve de Lokobe, la Réserve Spéciale
de l’Ankarana, et le Réserve Spéciale d’Analamerana, au
nord de Madagascar. Lemur News 10: 8–11.
Rakotodravony H. A. 2006. Communautés locales
et gibiers dans la region de Daraina, extrême Nord-
Est de Madagascar. Madagascar Conservation and
Development 1: 19–21.
Reuter K. E., H. Gilles, A. R. Willis and B. J. Sewall. 2014.
Live capture and ownership of lemurs in Madagascar:
extent and conservation implications. Oryx 1–11.
Richard, A. F., P. Rakotomanga and M. Schwartz.
1993. Dispersal by Propithecus verreauxi at Beza
Mahafaly, Madagascar 1984–1991. American Journal of
Primatology 30: 1–20.
Salmona, J., H. Teixeira, E. Rasolondraibe, I. Pais, C.
Kun-Rodrigues, A. Rakotonanahary, et al. Submitted.
Genetic diversity, population size and conservation of
the Critically Endangered Perrier’s sifaka (Propithecus
Salmona, J., R. Heller, E. Quéméré, E. and L. Chikhi.
In press. Comparative demographic history of northern
Madagascar lemurs using Approximate Bayesian
Salmona, J., F. Jan, E. Rasolondraibe, R. Zaranaina, D.
Saïd Ousseni, I. Mohamed ani, A. Rakotonanahary,
T. Ralantoharijaona, C. Kun-Rodrigues, M. Carreira,
S. Wohlhauser, P. Ranirison, J. R. Zaonarivelo, J. C.
Rabarivola and L. Chikhi. 2013. Survey of the critically
endangered Perrier’s sifaka (Propithecus perrieri) across
most if its distribution range. Lemur News 17: 9–12.
Salmona J., J. R. Zaonarivelo and M. A. Banks. 2013.
Analamerana and Andraamena. In: Lemurs of
Madagascar: A Strategy for their Conservation 2013–
2016, Schwitzer, C., R. A. Mittermeier, N. Davies, S.
Johnson, J. Ratsimbazafy, J. Razandramanana, E. E.
Louis Jr. and S. Rajaobelina (eds.), pp.140–141. IUCN
SSC Primate Specialist Group, Bristol Conservation and
Science Foundation, and Conservation International,
Schwitzer, C., O. Arnoult and B. Rakotosamimanana.
2006. An international conservation and research
programme for Perrier’s sifaka (Propithecus perrieri
Lavauden, 1931) in northern Madagascar. Lemur News
Zaonarivelo J. R., R. Andriantompohavana, A.
Razandrakoto, J. Andrianasolo, S. Rajaobelina, R.
A. Brenneman, S. Wolhauser and E. E. Louis Jr. 2007.
Preliminary survey of Andavakoera Classied Forest
and regional forest fragments of Madagascar. Lemur
News 12: 26–28.
Modern genetic tools reveal that Philippine tarsiers,
Carlito syrichta, are a group of three distinct genetic
lineages: the Bohol-Samar-Leyte lineage, the Dinagat-
Caraga lineage and the Mindanao-Zamboanga lineage
(Brown et al. 2014). However, these distinct genetic
lineages do not perfectly match the three possible
subspecies, T. syrichta syrichta from Leyte and Samar, T.
s. carbonarius from Mindanao, and T. s. fraterculus from
Bohol, creating a very uncertain situation in which
some populations might be threatened with extinction.
Furthermore, we don’t know which one, or ones, are
threatened, and with deforestation having removed
nearly all of the Philippine tarsiers’ original habitat in
many places, there is urgent need to act now and identify
these unique primate taxa as a conservation priority.
e history of the Philippine tarsier’s IUCN threat status
has changed through the years, indicating a profound
lack of knowledge. Originally listed as Endangered (EN),
it was downgraded in 1996 to Lower Risk/Conservation
Dependent (LR/CD). Subsequently it was listed as
Data Decient (DD), and is currently listed as Near
reatened (NT) (Shekelle & Arboleda 2008). What this
reects is that the metapopulation of all tarsiers found
within the Philippines, scattered over 4 large islands
(Samar, Leyte, Bohol, and Mindanao) and an unknown
number of smaller islands (e.g., Dinagat, Basilan) might
not be in imminent danger of going extinct across its
entire range. However, this merely masks the true state
of some populations, representing genetically distinct
lineages, which could more likely be in trouble.
e three genetically distinct lineages do not correspond
with the distribution of the three subspecies described,
albeit recognized as such by only a few taxonomists.
Brandon-Jones et al. (2004) inferred that the lack of
acceptance is more due to the limited morphological
evidence needed for taxonomic separation. As
with many taxonomically cryptic nocturnal species,
morphological variation is minimal. is is further
complicated by museum specimens of Philippine
tarsiers being greatly skewed toward just two sites on
the Gulf of Davao on Mindanao Island. us, taxonomic
clarity of the three lineages hinges on acquiring more
morphological data as well as genetic data across its
known range, quickly, before these distinct lineages
disappear and the opportunity to reveal the real species
status within the metapopulation is lost forever.
Carlito syrichta* (Linnaeus, 1758)
Myron Shekelle, Sharon Gursky, Stefan Merker & Perry Ong
(Carlito syrichta subspecies)
(Illustrations: Stephen D. Nash)
*Carlito syrichta is used as it is the most recent
published name, however some authors prefer using
Tarsius syrichta when referring to Philippine tarsiers.
e Philippines have been described as the region
where “megadiversity meets mega deforestation”
(Panela 2014). On the list of e World’s 10 Most
reatened Forest Hotspots, the Philippines rank 4th,
ahead of Madagascar, a country that is infamous for its
conservation crisis (CI 2011). Indeed, perhaps as little
as 7% of the original forests remain in the Philippines.
is is particularly distressing since whether tarsiers
are obligate forest species or not remains unclear. It
is yet another issue that must be resolved quickly.
Further compounding the threats to Philippine tarsier
survival are the seeming increase in frequency and
intensity of typhoons due to climate change. In 2013,
supertyphoon Haiyan swept directly over critical tarsier
habitats in Leyte and Samar. e direct eects of this
supertyphoon and other expected typhoons, as well as
other natural disasters of similar magnitude, on tarsier
survival are still being studied, but are most likely
devastating (Gursky unpublished data).
Finally, while tarsiers are used as an ecotourism mascot
on the island of Bohol, the regulation of this practice is
weak. Many tarsiers are on display as roadside attractions
in conditions that can be heartbreaking, especially
as these are nocturnal animals on display during the
daytime. Given the diculties of keeping tarsiers alive
in captivity, it is assumed that mortality among these
animals is high, and that replacements are collected
from the wild when they die. Furthermore, there is
a burgeoning illegal trade in tarsiers as pets, which
unfortunately, is probably caused to some degree by
the tarsiers’ status as tourism mascots. While the use of
tarsiers as tourism mascots is laudable and proves their
ability to promote tourism and thereby conservation,
the industry in the Philippines needs to improve how
tarsiers are managed for tourism and conservation.
erefore, the inclusion of the Philippine tarsier among
the world’s 25 most endangered primates, even though
its current IUCN threat status is just Near reatened, is
due to the following: First, the NT status was accorded
to the metapopulation of all Philippine tarsiers on all
islands. With the identication of three distinct genetic
lineages of tarsiers, the extent of occurrence of each
genetic lineage is now considerably reduced and the
population status of each lineage is unknown, thus the
threat of extinction for some distinct genetic lineage
is now a real possibility. Second, the Philippines are
a land of mega deforestation and it remains unclear
whether non-forest habitats, though reported to be
potential alternative habitats, could sustain the three
distinct genetic lineages of tarsiers in perpetuity.
ird, climate change might lead to the increased
frequency and intensity of typhoons, particularly in
a highly vulnerable country like the Philippines, thus
combined with the rst two factors described above,
this might further exacerbate the extinction risk of
tarsiers. Finally, tarsiers possess great power to promote
conservation through their role as tourism mascots, but
unfortunately, owing to weak management, current
practices could exacerbate the risk of extinction, and
this needs correction. e reasons enumerated above
were based on the best available information and
should be further validated on the ground. Experts with
direct experience of these tarsiers need to be mobilized
immediately to determine if the three genetic lineages
are indeed separate species, as well as the population
level of each lineage; assess the suitability of non-forest
habitats to sustain tarsier populations; study how the
extinction risk of tarsiers is impacted by catastrophic
disturbances; and examine ways by which tarsiers as a
agship species for conservation through ecotourism
can be enhanced without causing them harm.
Brandon-Jones, D., A. A. Eudey, T. Geissmann, C.
Groves, D. J. Melnick, J. C. Morales, M. Shekelle and
C.–B. Stewart. 2004. Asian primate classication.
International Journal of Primatology 25: 97-164.
Brown, R.M., et al. 2014. Conservation genetics of the
Philippine tarsier: cryptic genetic variation restructures
conservation priorities for an island archipelago
primate. PLoS ONE 9(8): e104340. doi:10.1371/journal.
Conservation International. 2011. e World’s 10 Most
reatened Forest Hotspots.
Panela, S. 2014. e Philippines: where ‘megadiversity’
meets mega deforestation. Mongabey. http://news.
Shekelle, M. and I. Arboleda. 2008. Tar siu s
syrichta. e IUCN Red List of reatened Species:
e.T21492A9289252. Accessed on 11 November 2015.
All Asian lorises are imperiled by the devastating loss
of their habitat. An even greater immediate threat to
Asian slow lorises, however, is their high demand in the
rampant Asian pet and traditional medicine trades and
their use as tourist photo props (Schulze and Groves
2004; Das et al. 2009; Nekaris et al. 2010; Osterberg and
Nekaris 2015). eir wide availability as pets has led to
new threats, including their body parts made available
as fashionable key rings (Nijman et al. 2014) and a
rising Internet trade fuelled by multiple viral Internet
videos of pet lorises (Nekaris et al. 2013). Easy to catch
due to their slow locomotion, numbers of slow lorises
in animal markets far outstretch the ability of these
slow-reproducing primates to recover their population
numbers in the wild (Nekaris and Starr 2015). Indeed,
this threat raised international concern, resulting in
the transfer of all members of the genus Nycticebus to
CITES Appendix I in 2007 (Nekaris and Nijman 2007).
Eight species of slow loris are now recognized: N.
coucang (greater), N. pygmaeus (pygmy), N. bengalensis
(Bengal), N. menagensis (Philippine), N. borneanus
(Bornean), N. kayan (Kayan), N. bancanus (Sody’s),
and N. javanicus (Javan) (Chen et al. 2007; Munds et
al. 2013; Pozzi et al. 2014). All slow lorises suer from
trade throughout their range, but when combined with
tremendous habitat loss, no other species has been
harder hit than the Javan slow loris.
Recognized by the IUCN as a species in 2006, and
currently listed as Critically Endangered (IUCN 2014),
the Javan slow loris is distinguished easily from its
congeners in several respects. Both morphologically
and genetically, it is most similar to, yet still distinct
from, the largest slow loris, N. bengalensis of mainland
Asia (Roos 2003; Groves and Maryanto 2008). Weighing
about 1 kg, the most distinctive feature of the Javan slow
loris is its facial mask, comprised of bold fork marks
leading from the eyes and ears to the crown of the head,
revealing a white diamond pattern on the forehead
(Nekaris and Jae 2007). Despite being legally protected
since 1973, with its creamy neck, bold dorsal stripe, and
panda-like face, it is no wonder that Indonesian pet
traders in the 1990s targeted Javan slow lorises above
other endemic loris species. Since 2002, however, the
numbers of Javan slow lorises in trade have decreased,
with a stark rise in numbers of (Sumatran) greater slow
lorises (N. coucang), a species whose threat status must
Javan Slow Loris
Nycticebus javanicus É. Georoy Saint-Hilaire, 1812
(2008, 2012, 2014)
K. Anna I. Nekaris & Vincent Nijman
Javan slow loris (Nycticebus javanicus)
(Illustration: Stephen D. Nash)
also be carefully monitored. Indeed, over three years of
markets surveys on Java from 2012 to 2015, quadruple
the number of greater than Javan slow lorises were
counted, with traders claiming that Javan slow lorises
could no longer be found (Nijman et al. in press). In
November 2013 alone, nearly 300 greater slow lorises
were conscated in two raids. e smaller raid, yielding
76 individuals, was followed by the almost immediate
death toll of 31 individuals. All of these animals were
conscated before ever making it to markets, showing
the dramatic extent of this trade.
e Javan slow loris is found only on the Indonesian
island of Java. Java has a long history of cultivation
and deforestation that already started c.1000 AD, but
took o in 1830 when the Dutch colonial government
imposed the so-called ‘cultuurstelsel’. To support this
agro-economic system, farmers were forced to grow
export crops on communal grounds, which were oen
forest (Whitten et al. 1996). By the end of the 19th
century the natural forest was severely fragmented,
and at the beginning of the last century the remaining
forest, especially in West and Central Java, showed a
fragmentation pattern very similar to that seen today.
Over the last few decades, the decrease in forest area
has been slow. At present, less than 10% of the original
forest remains, most of it covering the higher slopes of
the central mountains.
GIS models made available by orn et al. in 2009
suggested that historic forest loss and continued
degradation mean that less than 20% of habitat suitable
for Javan slow loris remains and that only 17% of the
potential distribution of Javan slow loris is currently
within the protected area network of Java. Voskamp et
al. (2014) and Nekaris et al. (2014) investigated nine
of these areas along with an additional six unprotected
areas. eir results concurred with those obtained by
three separate research groups, with animals occurring
at 0.02 to 0.20 animals per km, when they could be
found at all, meaning 5-50 km must be walked to see
a single slow loris (Nekaris and Nijman 2008; Winarti
2008). Roads and human disturbance have been shown
to correlate negatively with Javan slow loris abundance
(Winarti 2008). During Voskamp et al.’s study, numbers
of slow lorises were higher in agroforest that in some
cases was extremely disturbed by humans. Nekaris et al.
(2014) found that walking speed signicantly inuenced
the number of slow lorises spotted.
Also urgently required are programmes to mitigate
trade in all species of slow loris. A number of studies
have found that slow lorises are not always a targeted
group, but that they do have economic value throughout
their range. Rather than seeking a slow loris, villagers
moving through the forest simply pick up a loris when
they happen to see it (Starr et al. 2008). Similarly, when
forests are clear cut (for agriculture or cash crops),
villagers pick through the felled trees and collect the
slow lorises; with a defense mechanism to cling to
branches rather than to ee, and with their nocturnal
senses stunned by bright daylight, lorises are an easy
target (Ratjacsek 1998). Nijman and Nekaris (2014)
showed that traditional beliefs about slow lorises may
hinder people from hunting them, particularly beliefs
regarding their being venomous or poisonous.
Slow lorises are oen targeted, with, in Java, specialized
collectors searching the countryside. In addition, locals
who come across a slow loris may collect it, and pass
it on to middlemen. Most slow lorises collected in this
manner end up in the so-called bird markets that are
found in most major towns in Java. Once they arrive at
a market, slow lorises face other threats. To avoid being
bitten by slow lorises, which are one of few venomous
mammals, traders habitually cut or pull out an animal’s
lower front teeth (Nekaris et al. 2013). Most of these
slow lorises die due to general infection, dental abscess
or pneumonia (Nekaris and Starr 2015). ose that do
survive are no longer able to eat their preferred food
(gum) (Wiens et al. 2006), or to engage in the important
behavior of social grooming with the toothcomb,
meaning that any conscated animals are unlikely to
survive if released to the wild.
Reintroduction itself is a threat to the Javan slow loris;
the major trade hubs on Java and the neighboring island
of Sumatra receive dierent species of slow loris from
throughout the region. e similar appearance of slow
lorises to the untrained eye results in release of other
slow loris species into Java, with potential for disastrous
eects from hybridization or displacement of native
species by introduced ones (Nekaris and Starr 2015).
e ability for slow lorises to persist in human habitat if
le undisturbed means that well-meaning people may
translocate animals to habitat that is unknown to the
animals, causing potential death, disease transmission,
and invasive species issues (Kumar et al. 2014).
Moore et al. (2014) assessed the success of reintroduction
of Javan slow lorises, nding up to a 90% death rate.
Illness, hypothermia and exhaustion were all implicated
in the death of lorises. Sadly, reintroductions were
started before anything was known about the behavior,
ecology or wild distribution of slow lorises. No habitat
assessment could be made since it was not even
known in what type of habitat the species occurred.
Subsequently it is reported that success is improving,
but no published data are available. A related study
of pygmy slow loris in Vietnam found that the season
in which lorises are released as well as the age of the
releases is vital for success (Kenyon et al. 2014).
In 2011, the rst long-term study of Javan slow loris
behavioural ecology was instigated by the Little
Fireface Project in Garut District, West Java, Indonesia
(Nekaris 2014; Rode-Margono et al. 2014). is multi-
disciplinary project has obtained the rst data about
slow loris behavior in an agroforest matrix, including
home range size, social organization, infant dispersal,
and feeding ecology. Some notable discoveries have
been that both sexes disperse from their natal range at
about 18 months old, that dispersal distances are some
1–2 km from the natal range, that home range sizes are
large relative to the size of the animal (5–10 ha), and that
the diet of lorises comprises mainly gum, supplemented
with nectar and insects. Several initiatives have been
put into place to conserve slow lorises in the area and
in other parts of Java. National workshops have been
held for law enforcement ocers and rescue center
employees to feed essential data into a national slow
loris action plan. At the local level, slow lorises are
totally dependent on local people for their protection,
feeding on human-planted tree species and residing
on farmland. us a major conservation program,
combining empowerment activities, conservation
education and village events, has been launched, and it
is hoped that it can be used as a model for other key
slow loris sites in Indonesia (Nekaris and Starr 2015).
For a long time, slow lorises were thought to be common
throughout Indonesia, and the presence of animals in
trade was believed to be an indicator of their abundance.
We are only beginning to unravel the complexity of
their taxonomy and distribution, leading to an overall
bleak picture. While Java has an impressive and
comprehensive protected area network, encompassing
over 120 terrestrial conservation areas covering some
5,000 km², enforcement of environmental laws and
active protection of forest is lacking in most of these
parks. Besides curbing the illegal trade, it is paramount
that these conservation areas, and indeed all other
remaining forest areas on the island, be eectively
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