Primates in Peril

The World’s 25 Most Endangered Primates

2022–2023

Edited by
Russell A. Mittermeier, Kim E. Reuter, Anthony B. Rylands,
Leandro Jerusalinsky, Christoph Schwitzer, Karen B. Strier,
Jonah Ratsimbazafy and Tatyana Humle
PRIMATES IN PERIL
The World’s 25 Most Endangered Primates
2022–2023

Edited by
Russell A. Mittermeier, Kim E. Reuter, Anthony B. Rylands,
Leandro Jerusalinsky, Christoph Schwitzer, Karen B. Strier,
Jonah Ratsimbazafy and Tatyana Humle

Illustrations by
Stephen D. Nash

IUCN SSC Primate Specialist Group (PSG)

International Primatological Society (IPS)
Re:wild
Published by: IUCN SSC Primate Specialist Group (PSG), International Primatological
Society (IPS) and Re:wild.

Copyright: © 2022 Re:wild

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, Re:wild,
PO Box 129, Austin, TX 78767, USA.

Citation (report): Mittermeier, R.A., Reuter, K.E., Rylands, A.B., Jerusalinsky, L., Schwitzer, C.,
Strier, K.B., Ratsimbazafy, J. and Humle, T. (eds.), 2022. Primates in Peril:
The World’s 25 Most Endangered Primates 2022–2023. IUCN SSC Primate
Specialist Group, International Primatological Society, Re:wild, Washington,
DC. 163pp.

Citation (species): Schäffler, L., Markolf, M. and Kappeler, P.M. 2022. Madame Berthe’s Mouse
Lemur Microcebus berthae Rasoloarison et al. 2000. In: R.A. Mittermeier,
K.E. Reuter, A.B. Rylands, L. Jerusalinsky, C. Schwitzer, K.B. Strier, J.
Ratsimbazafy and T. Humle (eds.), Primates in Peril: The World’s 25 Most
Endangered Primates 2022–2023, pp. 8–10. IUCN SSC Primate Specialist
Group, International Primatological Society, Re:wild, Washington, DC.

Layout: Paula Katharina Rylands, Ubatuba, SP, Brazil.

Illustrations: © Stephen D. Nash, Re:wild, Washington DC, and Department of

Anatomical Sciences, Health Sciences Center, State University of New York
at Stony Brook, NY, USA.

Available from: Jill Lucena, Re:wild

E-mail: [email protected]
Website: www.primate-sg.org

Printed by: Tray Inc.

Glen Burnie, MD, USA.

ISBN: 978-1-7372851-5-1

Cover photo: The Buffy-headed Marmoset, Callithrix flaviceps, in the Brazilian Atlantic
Forest. Photograph by Geraldo Lucas Amaral.
 CONTENTS
Preface...........................................................................................................................................1
The World’s 25 Most Endangered Primates 2022–2023......................................................3

Madagascar
Madame Berthe’s Mouse Lemur, Microcebus berthae .............................................................9
Sahafary Sportive Lemur, Lepilemur septentrionalis ...............................................................13
Blue-eyed Black Lemur, Eulemur flavifrons ..............................................................................17
Coquerel’s Sifaka, Propithecus coquereli .................................................................................23

Africa
Rondo Dwarf Galago, Paragalago rondoensis ........................................................................31
Golden-bellied Mangabey, Cercocebus chrysogaster ............................................................35
Southern Patas Monkey, Erythrocebus baumstarki ..................................................................38
Roloway Monkey, Cercopithecus roloway ................................................................................43
Niger Delta Red Colobus, Piliocolobus epieni ........................................................................49
Nigeria-Cameroon Chimpanzee, Pan troglodytes ellioti ........................................................53

Asia
Javan Slow Loris, Nycticebus javanicus ....................................................................................61
Sangihe Tarsier, Tarsius sangirensis ...........................................................................................67
Purple-faced Langur, Semnopithecus vetulus ..........................................................................71
Golden-headed Langur, Trachypithecus poliocephalus ..........................................................77
Raffles’ Banded Langur, Presbytis femoralis .............................................................................81
Guizhou Snub-nosed Monkey, Rhinopithecus brelichi ............................................................85
Gaoligong Hoolock Gibbon, Hoolock tianxing .......................................................................89
Tapanuli Orangutan, Pongo tapanuliensis ...............................................................................93

Neotropics
Buffy-headed Marmoset, Callithrix flaviceps ...........................................................................101
Ka’apor Capuchin, Cebus kaapori ...........................................................................................107
Ecuadorian White-fronted Capuchin, Cebus aequatorialis ..................................................111
Groves’ Titi Monkey, Plecturocebus grovesi ..........................................................................117
Brown Howler Monkey, Alouatta guariba ...............................................................................121
Brown-headed Spider Monkey, Ateles fusciceps ...................................................................127
Geoffroy’s Spider Monkey, Ateles geoffroyi ...........................................................................135

Other Species Considered ......................................................................................................142

Affiliations .................................................................................................................................157
 PREFACE
We present here the 11th edition of The World’s 25 Most Endangered Primates, this one for 2022–
2023. The consultation for this list was held in an open meeting on the evening of 12 January 2022,
during the IPS-SLAPrim Joint Meeting (XXVIII Congress of the International Primatological Society -
IPS and IV Congress of the Latin American Society of Primatology - SLAPrim) in Quito, Ecuador (9–15
January 2022). This session was attended by more than 100 people and we subsequently consulted
with many experts to refine the information on those species that were selected. Note also that since
the Quito IPS Congress, originally planned for 2020, was not held until January 2022, the 2020–2022
list of the World’s 25 Most Endangered Primates wound up being this list for 2022–2023.

Using the information obtained over the past six months, we have updated the profiles for those
species remaining on the list from the 2018–2020 (2022) edition and for those from previous editions
that were returned to the list, and we have added additional profiles for newly listed species. Of note,
the 2018–2020 edition was the last that was published before the onset of the COVID-19 pandemic.

This publication is a joint initiative of the IUCN SSC Primate Specialist Group, the International
Primatological Society and Re:wild.

We are most grateful to the people and organizations who have provided major support for the
conservation of endangered primates over the years, both those listed in this report and many others
as well. In particular, we would like to thank Jon Stryker and Slobodan Randjelović for their support
through the Arcus Foundation and in other ways, to Annette Lanjouw of the Arcus Foundation for
major support of great ape and gibbon conservation, and to the Mohamed bin Zayed Species
Conservation Fund for supporting numerous primate projects since 2009. Special thanks also to the
Margot Marsh Biodiversity Foundation for their support of primate conservation since 1996, both
through the provision of grants from the foundation itself and through the Primate Action Fund of the
Primate Program at Re:wild. Besides providing the wherewithal publish this report, the foundation has
funded training workshops and field courses held at multiple biennial congresses of the International
Primatological Society, besides helping primatologists to attend the meetings to discuss The World’s
25 Most Endangered Primates lists. Much of the work on the administration of these grants is done
by Ella Outlaw, Anthony Rylands and Jill Lucena of the Re:wild Primate Program and by William R.
Konstant, a Re:wild Associate.

We thank all of those who contributed to the final 2022–2023 version. They are also listed as authors
on the individual species accounts to which they contributed.

Ekwoge Abwe, Felipe Alfonso-Cortes, André C. Alonso, Mónica Alzamora, Aubin Andriajaona,
Herizo T. Andrianandrasana, Aristide Andrianarimisa, Andie Ang, Márcia M. Assis Jardim, Renata B.
Azevedo, Carolyn A. Bailey, Hamilton F. Barreto, Raone Beltrão-Mendes, Richard Bergl, Hosnah Bezara
Hortensia, Júlio César Bicca-Marques, Jean P. Boubli, Gerson Buss, Thomas M. Butynski, Gustavo
Rodrigues Canale, Tatiane dos Santos Cardoso, Rodrigo Salles Carvalho, Laura Cervera, Óscar M.
Chaves, Susan M. Cheyne, Lounès Chikhi, Fanny Cornejo, Sam Cotton, Tim R.B. Davenport, Yvonne
A. De Jong, Stella de la Torre, Andrea Dempsey, Mary P. Dinsmore, Sarisha Trindade do Carmo, Osiris
Doumbe, Andrew Dunn, Paul Dutton, Timothy M. Eppley, María Fernanda Solórzano, Stephen F.
Ferrari, Chefor Fotang, Cynthia L. Frasier, Nathalia Fuentes, Paul A. Garber, Mary Katherine Gonder,
Sery Gonedelé Bi, Marcelo Gordo, Sidney Gouveia, Karol M. Gutiérrez-Pineda, Panut Hadisiswoyo,
Fei Hanlan, John A. Hart, Rio Heriniaina, Daniel Hending, Alma Hernández-Jaramillo, Zelinda B.
Hirano, Claudia Igayara-Souza, Rachel Ashegbofe Ikemeh, Inaoyom Imong, Sabrina Jabbar, Leandro

1
Jerusalinsky, Serge A. Kamgang, Peter M. Kappeler, Inza Koné, Martin Kowalewski, Wanda Kuswanda,
Diogo Lagroteria, Chien Lee, Zan Hui Lee, Neahga Leonard, Richard Lewis, Edward E. Louis Jr., Zhang
Lu, Matthias Markolf, Eduardo Marques, Jesus Martinez, Reiko Matsuda Goodwin, W. Scott McGraw,
Edward McLester, Fabiano R. de Melo, Sérgio Lucena Mendes, Ana Cristina Mendes-Oliveira, Erik
Meijaard, Pedro G. Méndez-Carvajal, Sebastián Montilla, Alba Lucía Morales-Jiménez, Citlalli Morelos-
Juárez, Bethan Jane Morgan, Jo A. Myers Thompson, Tilo Nadler, K. Anna I. Nekaris, Leonardo G.
Neves, Vincent Nijman, Amadou Ngouh, John F. Oates, Luciana I. Oklander, Richard L. Passaro,
Fan Pengfei, Andrew Perkin, George (PJ) Perry, Eduardo J. Pinel Ramos, Herinandrasana Pollensio,
Marcio Port-Carvalho, Gilberto Pozo-Montuy, Didik Prasetyo, Helder Queiroz, Martina Raffel, Brigitte
M. Raharivololona, Gilbert Rakotoarisoa, Felix Rakotondraparany, Valérie F. Rakotomalala, Jeannin
Nicolas Rakotondrazandry, Romeo Rakotondrina, Fidy Ralainasolo, Doménico R. Randimbiharinirina,
Guy H. Randriatahina, Noa E. Rasoanaivo, Lucile Rasoamazava, Solofonirina Rasoloharijaona, Jonah
Ratsimbazafy, Lucile Mialisoa Raveloarimalala, Faranky Ravelomandrato, Herimalala Raveloson,
André Luis Ravetta, Lena M. Reibelt, Baoping Ren, Jhon Rigobert, Esteban Rivera, Rizaldi, Melissa
E. Rodríguez-Menjívar, Noel Rowe, Rasanayagam Rudran, Nadine Ruppert, Ronald Sánchez-Porras,
Jordi Salmona, Livia Schäffler, Daniela Schrudde, Christoph Schwitzer, Timothy M. Sefczek, Tukuru
Seibokuro, Thiago B.F. Semedo, Brandon Semel, Meredith Semel, Arif Setiawan, Sam Shanee, Myron
Shekelle, Denise Spaan, Travis S. Steffens, Roswitha Stenke, Jatna Supriatna, Phan Duy Thuc, Jen
Tinsman, Diego G. Tirira, Mônica M. Valença-Montenegro, Orlando Vital, Sylviane Volampeno, Patrick
O. Waeber, Robert Wallace, Serge Wich, Leslie Wilmet, Karla P. Zaldaña-Orantes, John Zaonarivelo,
Jiang Zhou.

2
 THE WORLD’S 25 MOST
ENDANGERED PRIMATES
2022–2023
Here we report on the 11th edition of the biennial listing of a consensus of the 25 primate species
considered to be among the most endangered worldwide and most in need of conservation measures.
The previous edition (Schwitzer et al. 2019)1 – published in 2019 and spanning 2018-2020 – was the last
to be published before the onset of the global COVID-19 pandemic. However, since the Quito IPS
Congress, originally planned for 2020, was not held until January 2022, the 2018–2020 Top 25 actually
wound up being the list for 2018–2022. Following the publication of the 2018–2020 list, we conducted
an in-depth analysis of the traceable impacts of the 25 Most Endangered Primates lists since their
inception in 2000 (Reuter et al. 2021)2.

The 2022–2023 list of the World’s 25 Most Endangered Primates has six species from Africa, four
from Madagascar, eight from Asia, and seven from the Neotropics (Table 1). Madagascar and Brazil
both have four, Indonesia has three, China, Colombia, Ecuador, Nigeria, Panama, and Tanzania have
two, and Argentina, Belize, Cameroon, Costa Rica, Côte d’Ivoire, El Salvador, Ghana, Guatemala,
Honduras, Democratic Republic of the Congo, Mexico, Myanmar, Nicaragua, Peru, Singapore, Sri
Lanka, and Vietnam each have one.

Eight of these primates are listed as among the world’s most endangered primates for the first time
– those listed with an asterisk in Table 1. The remaining 17 species and subspecies have been listed
in previous versions of the 25 most endangered primates. Fourteen primates were dropped from the
previous list, 2018–2020 (see Table 2).

The changes made in this list compared to the previous iteration (2018–2020) were not because the
situation of the 14 species that were dropped (Table 2) has improved. In some cases, the situation
has in fact worsened. By making these changes we intend rather to highlight other, closely related
species enduring equally bleak prospects for their survival.

During the discussion of the 2022–2023 list at the IPS-SLAPrim Joint Meeting (XXVIII IPS Congress and
IV SLAPrim Congress) in Quito, a number of other highly threatened primates were considered for
inclusion. For all of these, the situation in the wild is as precarious as it is for those that finally made it
on the list, thus they have been included as ‘Other Species Considered’ (p. 142).

1 Schwitzer, C., Mittermeier, R.A., Rylands, A.B., Chiozza, F., Williamson, E.A., Byler, D., Wich, S., Humle, T., Johnson, C.,
Mynott, H., and McCabe, G. (eds.). 2019. Primates in Peril: The World’s 25 Most Endangered Primates 2018–2020. IUCN SSC
Primate Specialist Group (PSG), International Primatological Society (IPS), Global Wildlife Conservation (GWC), and Bristol
Zoological Society, Washington, DC. 130pp.

2 Reuter, K.E., Mittermeier, R.A., Schwitzer, C., McCabe, G.M., Rylands, A.B., Jerusalinsky, L., Konstant, W.R., Kerhoas, D.,
Ratsimbazafy, J., Strier, K.B., Webber, A.D., Williamson, E.A. and Wise, J. 2021. The 25 most endangered primates list: im-
pacts on conservation fundraising and policy. In: E. Freedman, S.S. Hiles and D. Sachsman (eds.), Communicating Endan-
gered Species: Extinction, News and Public Policy, pp.101–115. Routledge, Abingdon, UK.

3
Table 1. The World’s 25 Most Endangered Primates: 2022–2023. * First time on the list.

MADAGASCAR
Microcebus berthae Madame Berthe’s Mouse Lemur Madagascar
Lepilemur septentrionalis Sahafary Sportive Lemur Madagascar
Eulemur flavifrons Blue-eyed Black Lemur Madagascar
Propithecus coquereli* Coquerel’s Sifaka Madagascar
AFRICA
Paragalogo rondoensis Rondo Dwarf Galago Tanzania
Cercocebus chrysogaster* Golden-bellied Mangabey Democratic Republic of the Congo
Erythrocebus baumstarki* Southern Patas Monkey Tanzania
Cercopithecus roloway Roloway Monkey Côte d’Ivoire, Ghana
Piliocolobus epieni Niger Delta Red Colobus Nigeria
Pan troglodytes ellioti* Nigeria-Cameroon Chimpanzee Cameroon, Nigeria
ASIA
Nycticebus javanicus Javan Slow Loris Indonesia (Java)
Tarsius sangirensis* Sangihe Tarsier Indonesia (Sulawesi)
Semnopithecus vetulus Purple-faced Langur Sri Lanka
Trachypithecus poliocephalus Golden-headed Langur Vietnam
Presbytis femoralis* Raffles’ Banded Langur Peninsular Malaysia, Singapore
Rhinopithecus brelichi Gray Snub-nosed Monkey China
Hoolock tianxing Skywalker or Gaoligong Hoolock China, Myanmar
Pongo tapanuliensis Tapanuli Orangutan Indonesia (Sumatra)
NEOTROPICS
Callithrix flaviceps* Buffy-headed Marmoset Brazil
Cebus kaapori Ka’apor Capuchin Brazil
Cebus aequatorialis Ecuadorian Capuchin Ecuador, Peru
Plecturocebus grovesi* Groves’ Titi Monkey Brazil
Alouatta guariba Brown Howler Monkey Argentina, Brazil
Ateles fusciceps Brown-headed Spider Monkey Colombia, Ecuador, Panama
Belize, Costa Rica, El Salvador, Gua-
Ateles geoffroyi Geoffroy’s Spider Monkey temala, Honduras, Mexico, Nicara-
gua, Panama, Colombia (?)

4
Table 2. Primate species included on the 2018–2020 list that were removed from the 2022–2023 list.

MADAGASCAR
Microcebus manitatra Bemanasy Mouse Lemur Madagascar
Hapalemur alaotrensis Lac Alaotra Bamboo Lemur Madagascar
Lepilemur jamesorum Manombo Sportive Lemur Madagascar
Indri indri Indri Madagascar
Daubentonia madagascariensis Aye-aye Madagascar
AFRICA
Rungwecebus kipunji Kipunji Tanzania
Benin, Côte d’Ivoire, Ghana, Nigeria,
Colobus vellerosus White-thighed Colobus
Togo
Piliocolobus rufomitratus Tana River Red Colobus Kenya
Benin (?), Burkina Faso (?), Côte
d’Ivoire, The Gambia (Extinct), Gha-
Pan troglodytes verus Western Chimpanzee na, Guinea, Guinea-Bissau, Liberia,
Mali, Nigeria, Senegal, Sierra Leone,
Togo (?)
ASIA
Simias concolor Pig-tailed Langur Indonesia (Sumatera)
Trachypithecus geei Golden Langur Bhutan, India (Assam)
NEOTROPICS
Saguinus bicolor Pied Tamarin Brazil
Callithrix aurita Buffy-tufted-ear Marmoset Brazil
Plecturocebus olallae Olalla’s Titi Bolivia

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6
MADAGASCAR
MADAGASCAR

© Russell Mittermeier

@ Russell Mittermeier
 MADAME BERTHE’S MOUSE
LEMUR
Microcebus berthae Rasoloarison, Goodman and Ganzhorn, 2000

Madagascar
(2012, 2022)

Livia Schäffler, Matthias Markolf and Peter M. Kappeler

Madame Berthe’s Mouse Lemur (Microcebus It is categorized as Critically Endangered on the

berthae) was discovered in Kirindy Forest of IUCN Red List of Threatened Species – habitat
the Centre National de Formation, d’Études et loss and degradation due to slash-and-burn
de Recherche en Environnement et Forestier agriculture is causing a continuing decline in
(CNFEREF) in 1992 (Schmid and Kappeler 1994). the species’ area of occupancy and extent of
With a mean adult body mass of 31 g, it is the occurrence (Markolf et al. 2020). A population
smallest known primate (Rasoloarison et al. 2000). decrease of more than 80% over ten years is
Its range is confined to the central Menabe region expected as a direct consequence of ongoing
of western Madagascar (Rasoloarison et al. 2000), habitat destruction. Data on the species’
where it is restricted to the dry deciduous forests population size and distribution are currently
of Kirindy and Ambadira. It is absent from the being updated. There are no captive populations
Reserve Spécial Andranomena (Schäffler and of this species.
Kappeler 2014). Microcebus berthae is the
rarest lemur in the central Menabe region. Not As the protection of the remaining forest patches
all the remaining forest there is suitable due to and the restoration of habitat quality and
its specific habitat and dietary requirements connectivity are critical to preserve biodiversity in
(Dammhahn and Kappeler 2009), and it is the central Menabe region, several conservation
sensitive to anthropogenic activities (Schäffler and education programs are being implemented
and Kappeler 2014). by NGOs using M. berthae as a flagship species.
Development projects, reforestation, and
Microcebus berthae is sympatric with the more restoration campaigns have been initiated as
widely distributed M. murinus, and ecological well.
differentiation in the two mouse lemurs is
evidently insufficient to prevent competitive The Aire Protégée Menabe Antimena (APMA),
exclusion (Ganzhorn and Kappeler 1996, Schwab with the key core areas being the Kirindy and
and Ganzhorn 2004, Dammhahn and Kappeler Ambadira forests, unfortunately still lacks
2008, 2009, 2010). Interactions with two other effective law enforcement. Since the protected
sympatric cheirogaleids, Mirza coquereli area’s designation in 2015, slash-and-burn
and Cheirogaleus medius, however, stabilize practices have destroyed about 30% of its forests
mouse lemur coexistence on a landscape scale (Markolf et al. 2020). The loss of revenue from
by providing M. berthae with refuges from tourism during the COVID-19 pandemic has
competition with its congener. At the community hampered adequate forest protection, and the
level, the densities of the two mouse lemur absence of foreign researchers has further fueled
species are correlated in opposite directions with deforestation. Effective protection of the APMA
those of Mirza and Cheirogaleus – when they are will require addressing not only the needs of a
abundant, M. berthae does better. growing human population, but also the need
to reduce the exploitation of natural resources.

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MADAGASCAR

The region suffers from a lack of sustainable

agriculture, agroforestry, and livestock farming
(Frank and Schäffler 2019) and there are numerous
other problems to solve if we are to preserve the
remaining habitat of the iconic Madame Berthe’s
Mouse Lemur.

Dammhahn, M. and Kappeler, P.M. (2008). Small-scale

coexistence of two mouse lemur species (Microcebus
berthae and M. murinus) within a homogeneous
competitive environment. Oecologia 157: 473–483.
Dammhahn, M. and Kappeler, P.M. (2009). Females go where
the food is: does the socio-ecological model explain
variation in social organisation of solitary foragers? Behav.
Ecol. Sociobiol. 63: 393–352.
Dammhahn, M. and Kappeler, P.M. (2010). Scramble or
contest competition over food in solitarily foraging
mouse lemurs (Microcebus spp.): new insights from stable
isotopes. Am. J. Phys. Anthropol. 141: 181–189.
Frank, A.K. and Schäffler, L. (2019). Identifying key knowledge
gaps to better protect biodiversity and simultaneously
secure livelihoods in a priority conservation area.
Sustainability 11: 5695.
Ganzhorn, J.U. and Kappeler, P.M. (1996). Lemurs of the
Kirindy forest. Primate Report 46: 257–274.
Markolf, M., Schäffler, L. and Kappeler, P. (2020). Microcebus
berthae. The IUCN Red List of Threatened Species 2020:
e.T41573A115579496.
Rasoloarison, R.M., Goodman, S.M. and Ganzhorn, J.U.
(2000). Taxonomic revision of mouse lemurs (Microcebus)
in the western portions of Madagascar. Int. J. Primatol.
21: 963–1019.
Schäffler, L. and Kappeler, P.M. (2014). Distribution and
abundance of the world’s smallest primate, Microcebus
berthae, in central western Madagascar. Int. J. Primatol.
35: 57–72.
Schäffler, L., Saborowski, J. and Kappeler P.M. (2015).
Agent-mediated spatial storage effect in heterogeneous
habitat stabilizes competitive mouse lemur coexistence
in Menabe Central, Western Madagascar. BMC Ecol. 15:
7.
Schäffler, L., Kappeler, P.M. and Halley J.M. (2021). Mouse
lemurs in an assemblage of cheirogaleid primates in
Menabe Central, Western Madagascar – three reasons to
coexist. Front. Ecol. Evol. 9: 585781.
Schmid, J. and Kappeler, P.M. (1994). Sympatric mouse
lemurs (Microcebus spp.) in western Madagascar. Folia
Primatol. 63: 162–170.
Schwab, D. and Ganzhorn, J.U. (2004). Distribution,
population structure and habitat use of Microcebus
berthae compared to those of other sympatric
cheirogaleids. Int. J. Primatol. 25: 307–330.

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MADAGASCAR

© Edward E. Louis Jr.

12
 SAHAFARY SPORTIVE LEMUR
Lepilemur septentrionalis Rumpler and Albignac, 1975

Madagascar
(2008, 2010, 2012, 2014, 2022)

Edward E. Louis Jr., Carolyn A. Bailey, Cynthia L. Frasier, Leslie Wilmet, Aubin Andriajaona,
Lucile Rasoamazava, Valérie F. Rakotomalala, Mary P. Dinsmore and Timothy M. Sefczek

Lepilemur septentrionalis is a small sportive lemur forest fragments (Dinsmore et al. 2016). Due
with a body weight of just 600–750 g (Louis Jr. et to the topography of karst mountains and the
al. 2006) from Madagascar’s far northern region. deforestation from charcoal enterprises, forest
Originally believed to have been distributed habitat has become more and more disconnected
across the north of Madagascar, from Montagne in Montagne des Français, limiting migration and
d’Ambre south to the Mahavavy River close to gene flow in the current population.
Ambilobe (Hawkins et al. 1990, Rumpler et al.
2001), it is now evidently restricted to just a few Lepilemur septentrionalis is listed as Critically
patches of forest. Endangered on the IUCN Red List of Threatened
Species (Louis Jr. et al. 2020). This must be
In 2007, a survey in the forests of Sahafary and considered one of the most restricted and least
the Analalava Forest Reserve estimated a protected lemurs in Madagascar, and probably
population of 100 (Ravaorimanana et al. 2009). the one closest to extinction, perhaps with the
A follow-up survey in 2012 and 2013 conducted exception of Microcebus berthae. The principal
by Omaha’s Henry Doorly Zoo and Aquarium threat is forest loss due to illicit firewood and
and the Madagascar Biodiversity Partnership charcoal production. This species is also hunted
(MBP) found only three individuals in these for food incidentally at illegal charcoal enterprises.
forests, suggesting that these populations have Most of the species’ habitat is already gone, but at
since been completely extirpated in the two least it now benefits from some protection given
western parts of its range. As such, the species that Montagne des Français became a protected
is thought to exist now only in Montagne des site in 2015 (Dinsmore et al. 2021a, 2021b).
Français (Louis Jr. and Zaonarivelo 2015). In a
subsequent field survey conducted in August This species is on the verge of extinction and in
and September 2019, Omaha’s Henry Doorly Zoo need of special attention. Although undertaking
and Aquarium and MBP field surveys detected a captive breeding program would be an option,
87 individuals, all of them in the central-western sportive lemurs have thus far proven difficult to
portions of Montagne des Français. Whether this keep in zoos due to their specialized folivorous
increase is based on protective steps taken by diet. Continuing direct surveys linked to random
the Direction Régionale de l’Environnement et patrols by government authorities is considered
du Développement Durable to conduct ranger a minimum requirement for this species’ survival
patrols with the MBP field teams, or an artificial over the next ten years, prioritizing the forest
response to the collapse of the surrounding fragments of Vangisay, Andrananakomba, and
habitat is not known (Bailey et al. 2020). Ampamakiampafana, which contain 98% of the
known population. As of 2022, this species was
Prior to 2010, this species had not been studied not being held in captivity (ZIMS 2022).
in the wild, though more is known now about
its ecology (e.g., Rasoamazava et al. 2022). It Conservation priorities for this species center on
inhabits tropical dry, deciduous and gallery restoration and reforestation efforts to establish

13
MADAGASCAR

habitat corridors between the Andranonakoba,

Ambavala, and Ambatove forest fragments in
Montagne des Français to allow for immigration
between these lemur subpopulations.
Furthermore, ranger patrols by the regional
Eaux et Forêt gendarmerie, VOI (Vondron’Olona
Ifotany or Communauté de Base) and local non-
profit organizations should be increased in these
areas to eliminate all human-related disturbances.
Research priorities should include annual census
surveys of the species’ remaining population.
Research is also needed to confirm yearly
population fluctuations and whether population
increases detected between the two previous
censuses (Ranaivoarisoa et al. 2013 and Bailey et
al. 2020) were positive recruitment or simply the
concentration of individuals into the eastern half
of Montagne des Français due to habitat loss.

Bailey, C.A., Sefczek, T.M., Robertson, B.A.D., Rasoamazava,

L., Rakotomalala, V.F. Andriamadison, J.D.N.
Randrianasolo, F., Andriajaona, A. and Louis Jr., E.E.
(2020). A re-evaluation of the northern sportive lemur
(Lepilemur septentrionalis) population at Montagne des
Français, and a review of its current state of conservation
in the protected area. Primate Conserv. (34): 53–59.
Dinsmore, M.P., Louis Jr., E.E., Randriamahazomanana, D.,
Hachim, A., J. Zaonarivelo, J.R. and Strier, K.B. (2016).
Variation in habitat and behavior of the northern sportive
lemur (Lepilemur septentrionalis) at Montagne des
Français, Madagascar. Primate Conserv. (30): 73–88.
Dinsmore, M.P., Strier, K.B. and Louis Jr., E.E. (2021a).
Anthropogenic disturbances and deforestation of
northern sportive lemur (Lepilemur septentrionalis)
habitat at Montagne des Français, Madagascar. Primate
Conserv. (35): 125–138.
Dinsmore, M.P., Strier, K.B. and Louis, Jr., E.E. (2021b). The
influence of seasonality, anthropogenic disturbances,
and cyclonic activity on the behavior of northern sportive
lemurs (Lepilemur septentrionalis) at Montagne des
Français, Madagascar. Am. J. Primatol. 83(12): e23333.
Hawkins, A.F.A., Chapman, P., Ganzhorn, J.U., Bloxam,
Q.M.C., Barlow, S.C. and Tonge, S.J. (1990). Vertebrate
conservation in Ankarana Special Reserve, northern
Madagascar. Biol. Conserv. 54: 83–110.
Louis Jr., E.E., Engberg, S.E., Lei, R., Geng, H., Sommer,
J.A., Randriamampionona, R., Randriamanana, J.C.,
Zaonarivelo, J.R., Andriantompohavana, R., Randria, G.,
Prosper, Ramaromilanto, B., Rakotoarisoa, G., Rooney, A.
and Brenneman, R.A (2006). Molecular and morphological
analyses of the sportive lemurs (Family Megaladapidae:
Genus Lepilemur) reveals 11 previously unrecognized
species. Spec. Publ. Mus. Texas Tech Univ. 49: 1–47.
Louis Jr., E.E., Bailey, C.A., Frasier, C.L., Sefczek, T.M.,
Raharivololona, B., Schwitzer, C., Ratsimbazafy, J., Wilmet,
L., Andriajaona, A., Rasoamazava, L., Rakotomalala, V.F.

14
 and Dinsmore, M. (2020). Lepilemur septentrionalis.
The IUCN Red List of Threatened Species 2020:
e.T11622A115567059.
Louis, Jr., E.E. and Zaonarivelo, J.R. (2015). Northern sportive
lemur Lepilemur septentrionalis (Rumpler and Albignac,
1975). In: C. Schwitzer et al. (eds.), Primates in Peril:
The World’s 25 Most Endangered Primates 2014–2016,
pp. 36–37. Arlington, VA: IUCN SSC Primate Specialist
Group (PSG), International Primatological Society (IPS),
Conservation International (CI), and Bristol Zoological
Society.
Ranaivoarisoa, J.F., Zaonarivelo, J.R., Lei, R., Johnson, S.E.,
Wyman, T.M., Mittermeier, R.A. and Louis Jr., E.E. (2013).
Rapid Survey and Assessment of the northern sportive
lemur, Lepilemur septentrionalis, in northern Madagascar.
Primate Conserv. (27): 23–31.
Rasoamazava, L., Rakotomalala, V.F., Sefczek, T.M., Frasier,
C.L., Dinsmore, M.P., Rasoloharijaona, S. and Louis, E.E.,
Jr. (2022). Feeding ecology of Lepilemur septentrionalis
in the dry forest of Montagne des Français, northern
Madagascar. Folia Primatol. doi.org/10.1163/14219980–
20210702
Ravaorimanana, I., Zaramondy, A., Rabarivola, C. and
Rumpler, Y. (2009). Northern sportive lemur. In: R.A.
Mittermeier et al. (eds.), Primates in Peril: The World’s
25 Most Endangered Primates 2008–2010, pp. 21–22.
Arlington, VA: IUCN SSC Primate Specialist Group
(PSG), International Primatological Society (IPS), and
Conservation International (CI).
Rumpler, Y., Ravaoarimanana, B., Hauwy, M. and Warter, S.
(2001). Cytogenetic arguments in favour of a taxonomic
revision of Lepilemur septentrionalis. Folia Primatol. 72:
308–315.
ZIMS. (2022). Zoological Information Management Software.
Available online: https://zims.species360.org.

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MADAGASCAR

© Sam Cotton

16
 BLUE-EYED BLACK LEMUR
Eulemur flavifrons (Gray, 1867)

Madagascar
(2008, 2010, 2012, 2022)

Guy Randriatahina, Sam Cotton, Dan Hending, Christoph Schwitzer,

Jen Tinsman and Sylviane Volampeno

Eulemur flavifrons is a medium-sized lemur. It is forests to the south. It occurs from sea level to
sexually dichromatic but the eyes of both sexes 1,200 m (Randriatahina and Rabarivola 2004). Its
are blue to blue-gray. It is one of the very few home range size and the way it uses its range
primates in the world that consistently has blue differs between primary and secondary forest
eyes, hence the English common name. fragments. Schwitzer et al. (2007a) reported
an average home range of 6.8 ha in primary
Following its description in 1867, the taxonomic forest, and 13.0 ha in secondary forest. These
status of this lemur was debated until genetic larger home ranges and the lower densities of
analysis confirmed that it was indeed distinct E. flavifrons in secondary forest suggest that it is
(Rabarivola 1998, McLain et al. 2012). The Blue- less suitable habitat for the species than primary
eyed Black Lemur has a very limited distribution forest. Parasite prevalence seems to be higher
in northwestern Madagascar. Due to the lack in secondary than in primary forest and appears
of locality data for the few museum specimens high when compared to data from other lemurs,
collected, nothing was known of its status in the suggesting that E. flavifrons on the Sahamalaza
wild until its ‘rediscovery’ in November 1983, Peninsula is generally under pressure, possibly
when it was located in northwestern Madagascar, due to the high degree of fragmentation and
in forests north of Befotaka and south of degradation of the remaining forest habitat
Maromandia, just south of the Sambirano domain (Schwitzer et al. 2010).
(Tattersall 1982, Koenders et al. 1985). It is now
known to occur on the Sahamalaza Peninsula During a 12-month study, E. flavifrons ate parts of
as well as in a stretch of forest on the adjacent 72 different plant species from 35 families; 52.3%
mainland, extending from around Befotaka in were fruits, and 47.7% were leaves. The animals
the south to the Manongarivo mountains in also fed on flowers, insects, insect excretions and
the north. The Maevarano River serves as the fungi (Polowinsky and Schwitzer 2009). At certain
southern boundary of the species’ range, the times of the year, the species may feed on large
Sandrakota River as the eastern boundary, and quantities of cicadas.
the Manongarivo River anad its tributary, the
Antsahakolana River, as the northern boundary Population densities of 24 individuals/km² have
(Rabarivola et al. 1991, Andrianjakarivelo 2004, been estimated in the eastern part of the species’
Randriatahina and Rabarivola 2004, Tinsman et range (Andrianjakarivelo 2004) and 97 individuals/
al. 2020). A recent range revision by Tinsman et km² in the Ankarafa Forest on the Sahamalaza
al. (2020) estimated the Extent of Occurrence to Peninsula (Volampeno et al. 2011), but the latter
be 3,475 km². figure seems to be unusually high compared to
other sites throughout the species’ range. Group
This lemur inhabits more or less disturbed primary size ranges from 4 to 11 (Andrianjakarivelo 2004,
and secondary subtropical, sub-humid forests, in Randriatahina and Rabarivola 2004, Schwitzer
a transition zone between the Sambirano region 2004, Volampeno et al. 2011).
to the north and the western dry, deciduous

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MADAGASCAR

The Blue-eyed Black Lemur has a bimodal activity

pattern, with peaks during the morning and
evening twilight. It is cathemeral, with activity
bouts during the day and night year-round.
Nocturnal illumination and the proportion of
illuminated lunar disk are positively associated
with the amount of nocturnal activity. Total daily
activity, as well as nocturnal activity, is higher in
secondary forest than in primary forest (Schwitzer
et al. 2007b).

Eulemur flavifrons is listed as Critically Endangered

on the IUCN Red List of Threatened Species
(Volampeno et al. 2020). The principal threat to
its survival is forest destruction due to slash-and-
burn agriculture, selective logging, mining and
feu de la colère (deliberately-set fires to express
people’s discontent, usually with the authorities
or government policy) (Seiler et al. 2010). It is
hunted for food, especially in the eastern part of
its range, and captured live for the local pet trade
(Reuter and Schaefer 2017b). Andrianjakarivelo
(2004) found a density of up to 570 traps/km² in
certain areas. In recent surveys of E. flavifrons,
evidence of hunting or slash-and-burn agriculture
was present in all locations surveyed (Tinsman et
al. 2020). Brown and Yoder (2015) estimated that
there will be an 88% reduction in the species’
range from 2000 to 2080 due to climate change
alone. Moreover, Volampeno et al. (2015)
predicted the extinction of E. flavifrons from the
Ankarafa forest on the Sahamalaza Peninsula
within a few decades if habitat destruction and
fragmentation continue.

Parts of the range of E. flavifrons received official

protected area status in 2007 as the Sahamalaza–
Îles Radama National Park, including the
Sahamalaza Peninsula and some mainland
coastal forests to the north and east (Lernould
2002, Schwitzer and Lork 2004, Schwitzer et al.
2006, 2009). The Sahamalaza Peninsula is also a
UNESCO Biosphere Reserve.

As of 2022, there were 59 individuals in captivity

(ZIMS 2022). Breeding in captivity, however,
has been limited, with only a small number
of successful breeding pairs after some initial
success with the founder generation. In these
settings, obesity can also be a problem (Schwitzer
2003, Volampeno et al. 2020).

18
Research priorities include estimating the community projects. Mikajy Natiora also plans
number and density of Blue-eyed Black Lemurs to build a research camp in nearby Mahitsihazo
remaining, both on the mainland and on the village to better protect the Blue-eyed Black
Sahamalaza Peninsula, and determining if the Lemurs of Andilambologno forest (Mikajy Natiora
hybrid zone between E. flavifrons and its sister 2022).
species E. macaco still exists (Tinsman et al. 2020).
Additional conservation efforts are needed in
Conservation efforts are underway on the Manongarivo Special Reserve, which covers the
Sahamalaza Peninsula. The Association largest remaining parcel of forest in the Blue-eyed
Européenne pour l’Etude et la Conservation des Black Lemur’s range. The reserve’s rugged terrain
Lémuriens (AEECL) has run a community-based and remote location make patrolling difficult,
conservation program there since 2000. The and human incursion into the forest for hunting,
program promotes alternative livelihoods and agriculture, and cattle grazing are common
improved, sustainable farming techniques. It (Tinsman et al. 2020). Increased financial support
also provides primary and secondary education for Madagascar National Parks rangers and clear
in this very remote region of the country (AEECL boundary markers would help protect the Blue-
2022). AEECL has been promoting ecotourism to eyed Black Lemur. Measures to support the
the Sahamalaza–Îles Radama National Park and communities in and surrounding Manongarivo
has established a tourist camp near the Ankarafa are also critical for long-term success. A program
forest, where visitors can observe the Blue-eyed similar to AEECL’s work in Sahamalaza or Mikajy
Black Lemur. The Ankarafa forest is also home Natiora’s work in Andilambologno forest could
to a field station from which lemur monitoring reduce these communities’ reliance on the
and forest patrols are conducted along with Manongarivo Reserve. Further, a proposed
experimental research on different methods of expansion of Manongarivo Special Reserve
reforestation and forest restoration (Cotton 2021, (MEFT and MEM 2008) would place a good
Hending et al. 2021). Previous reforestation proportion of the Blue-eyed Black Lemur’s
initiatives to counter E. flavifrons habitat loss mainland distribution range under protection
in Sahamalaza have been largely unsuccessful (Tinsman et al. 2020).
(Saunders 2017). The recent construction of a
permanent seedling nursery in the field station
is, therefore, a key step in the reforestation and
restoration of the Ankarafa forest complex (and
beyond). The nursery serves as a research base AEECL. (2022). The Lemur Conservation Association.
Available online: https://aeecl.org.
in which to refine propagation protocols and
Andrianjakarivelo, V. (2004). Exploration de la zone en
provide seedling stock, as well as providing dehors de la péninsule Sahamalaza pour l’évaluation
permanent employment and alternative, rapide de la population d’E. m. flavifrons. Unpublished
sustainable, livelihoods for local people. Current report. Antananarivo: WCS Madagascar. 31pp.
reforestation locations are strategically prioritized Brown, J.L. and Yoder A.D. (2015). Shifting ranges and
conservation challenges for lemurs in the face of climate
towards areas that yield maximum benefits for change. Ecol. Evol. 5(6): 1131–1142.
Blue-eyed Black Lemurs in terms of increased Cotton, S. (2021). Growing links for lemurs: towards an
connectivity between fragments and forest core- effective reforestation of Sahamalaza-Iles Radama
to-edge ratios (Von Bülow et al. in prep). National Park. Final Report. Gland: IUCN.
Hending, D., Randrianarison, H., Holderied, M., McCabe, G.
and Cotton, S. (2021). The kapok tree (Ceiba pentandra
Outside the Sahamalaza–Îles Radama National
(L.) Gaertn, Malvaceae) as a food source for native
Park, Mikajy Natiora has been working to conserve vertebrate species during times of resource scarcity and
the Blue-eyed Black Lemurs of Andilambologno its potential for reforestation in Madagascar. Austral Ecol.
forest, located east of the Route Nationale 6, near 46: 1440–1444.
the town of Maromandia. The pressures there Koenders, L., Rumpler, Y., Ratsirarson, J. and Peyrieras,
A. (1985). Lemur macaco flavifrons (Gray, 1867): a
are higher than in the protected area and include rediscovered subspecies of primate. Folia Primatol. 44:
slash-and-burn agriculture. Mikajy Natiora has 210–215.
trained local rangers, who are now patrolling the Lernould, J.-M. (2002). Un programme international de
forest, and conducts environmental outreach and recherche et de conservation pour le lémur aux yeux

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MADAGASCAR

turquoise (Eulemur macaco flavifrons). Lemur News 7: Schwitzer, N., Randriatahina, G.H., Kaumanns, W.,
30–33. Hoffmeister, D. and Schwitzer, C. (2007a). Habitat
McLain, A.T., Meyer, T.J., Faulk, C., Herke, S.W., Oldenburg, utilization of blue-eyed black lemurs, Eulemur macaco
J.M., Bourgeois, M.G., Abshire, C.F., Roos, C. and flavifrons (Gray, 1867), in primary and altered forest
Batzer, M.A. (2012). An Alu-based phylogeny of lemurs fragments. Primate Conserv. (22): 79–87.
(Infraorder: Lemuriformes). PLoS One 7(8): e44035. Schwitzer, N., Kaumanns, W., Seitz, P.C. and Schwitzer, C.
Mikajy Natiora. (2022). Lemur Conservation Network. (2007b). Cathemeral activity patterns of the blue-eyed
Available online: https://www.lemurconservationnetwork. black lemur Eulemur macaco flavifrons in intact and
org/organization/mikajy-natiora/ degraded forest fragments. Endang. Species Res. 3: 239–
Polowinsky, S.Y. and Schwitzer, C. (2009). Nutritional ecology 247.
of the blue-eyed black lemur (Eulemur flavifrons): Schwitzer, C., Polowinsky, S.Y. and Solman, C. (2009). Fruits
integrating in situ and ex situ research to assist the as foods – common misconceptions about frugivory. In:
conservation of a critically endangered species. In: M. M. Clauss, A. Fidgett, G. Janssens, J.-M. Hatt, T. Huisman,
Clauss, A. Fidgett, G. Janssens, J.-M. Hatt, T. Huisman, J. J. Hummel, J. Nijboer and A. Plowman (eds.), Zoo Animal
Hummel, J. Nijboer and A. Plowman (eds.), Zoo Animal Nutrition. Volume IV, pp. 131–168. Fuerth: Filander
Nutrition, Volume IV, pp. 169–178. Fuerth: Filander Verlag.
Verlag. Schwitzer, N., Clough, D., Zahner, H., Kaumanns, W.,
Rabarivola, C. (1998) Etude génétique comparative de Kappeler, P. and Schwitzer, C. (2010). Parasite prevalence
populations insulaires et ‘continentales’ de Eulemur in blue-eyed black lemurs (Eulemur flavifrons) in differently
macaco. Utilisation simultanée des dermatoglyphes, de degraded forest fragments. Endang. Species Res. 12:
marqueurs sanguins et de l’ADN (RAPD) pour étudier 215–225.
la différenciation de E. macaco en deux sous-espèces: Seiler, M., Randriatahina, G.H. and Schwitzer, C. (2010).
E. m. macaco et E. m. flavifrons. Doctoral dissertation, Ongoing threats to lemurs and their habitat inside the
Université d’Antananarivo, Antananarivo. Sahamalaza–Iles Radama National Park. Lemur News 15:
Rabarivola, C., Meyers, D. and Rumpler, Y. (1991). Distribution 7–9.
and morphological characters of intermediate forms Tattersall, I. (1982). The Primates of Madagascar. New York,
between the black lemur (Eulemur macaco macaco) and NY: Columbia University Press.
Sclater’s lemur (Eulemur macaco flavifrons). Primates 32: Tinsman, J.C., Eschmann, C.L., Solofondranohatra, J.S.,
269–73. Ralainirina, J.R., Holderied, M. and McCabe, G. (2020).
Randriatahina, G.H. and Rabarivola, J.C. (2004). Inventaire Range and conservation updates for the Critically
des lémuriens dans la partie nord-ouest de Madagascar Endangered blue-eyed black lemur Eulemur flavifrons
et distribution d’Eulemur macaco flavifrons. Lemur News and the Vulnerable black lemur Eulemur macaco. Oryx
9: 7–9. 54(6): 819–827.
Reuter, K.E. and Schaefer, M.S. (2017). Illegal captive lemurs Volampeno, M.N., Masters, J.C. and Downs, C.T. (2011).
in Madagascar: comparing the use of online and in- A population estimate of blue-eyed black lemurs in
person data collection methods. Am. J. Primatol. 79: Ankarafa forest, Sahamalaza–Iles Radama National Park,
e22541. Madagascar. Folia Primatol. 81: 305–314.
Saunders, J.S. (2017). Is Reforestation an Effective Volampeno, M.S., Randriatahina, G.H., Kalle, R., Wilson, A.L.
Conservation Strategy for the Threatened Vertebrate and Downs, C.T. (2015). A preliminary population viability
Fauna of the Sahamalaza Peninsula, Northwest analysis of the critically endangered blue-eyed black
Madagascar? Masters dissertation, University of Bristol, lemur (Eulemur flavifrons). Afr. J. Ecol. 53(4): 419–427.
Bristol. Volampeno, S., Randriatahina, G., Schwitzer, C. and Seiler,
Schwitzer, C. (2003). Energy Intake and Obesity in Captive M. (2020). Eulemur flavifrons. The IUCN Red List of
Lemurs (Primates, Lemuridae). Doctoral dissertation, Threatened Species 2020: e.T8211A115563094.
Universität zu Köln, Köln. Von Bülow, J., Cotton, S. and Abrahams, M.I. (in prep).
Schwitzer, C. (2004). Sahamalaza field trip 28.09.-19.11.2003. Applying a Novel GIS-Based Method to Determine
Report to the board of AEECL. Köln: Zoologischer Optimal Reforestation Sites in the Ankarafa Fragment
Garten Köln. Complex, Sahamalaza-Îles Radama National Park,
Schwitzer, C. and Lork, A. (2004). “Projet Sahamalaza–Iles Madagascar.
Radama“ Ein internationales Schutzprojekt für den ZIMS. (2022). Zoological Information Management Software.
Sclater’s Maki (Eulemur macaco flavifrons Gray, 1867). Z. Available online: https://zims.species360.org.
des Kölner Zoo 47: 75–84.
Schwitzer, C., Schwitzer, N., Randriatahina, G.H., Rabarivola,
C. and Kaumanns, W. (2006). “Programme Sahamalaza”:
New perspectives for the in situ and ex situ study and
conservation of the blue-eyed black lemur (Eulemur
macaco flavifrons) in a fragmented habitat. In: C.
Schwitzer, S. Brandt, O. Ramilijaona, M. Rakotomalala
Razanahoera, D. Ackermand, T. Razakamanana and J. U.
Ganzhorn (eds.), Proceedings of the German-Malagasy
Research Cooperation in Life and Earth Sciences, pp.
135–149. Berlin: Concept Verlag.

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MADAGASCAR

© Russell Mittermeier

22
 COQUEREL’S SIFAKA
Propithecus coquereli (A. Grandidier, 1867)

Madagascar
(2022)

Jordi Salmona, Solofonirina Rasoloharijaona, Lounès Chikhi, Travis S. Steffens,

Timothy M. Eppley and Gilbert Rakotoarisoa

Propithecus coquereli is endemic to the tropical, groups of 3–10, using home ranges of 4–33 ha
dry, lowland forests of northwestern Madagascar. (Petter 1962a, 1962b, Albignac 1981, McGoogan
Historically, the species has been reported in 2011). Sexual maturity is reached at roughly 2.5
the Sofia region between the Betsiboka and years of age for both sexes. A gestation period
Maevarano watersheds (Tattersall 1982, Wilmé et of 162 days normally leads to a single offspring
al. 2006, Mittermeier et al. 2010). Its southeastern and births are clustered in the months of June
and eastern boundaries are, however, less clear and July (Richard 1978b, 1987). Infants become
and would need extensive field efforts to be completely independent by about six months of
properly defined (Salmona et al. 2014). To age and reach adult size by one year (Mittermeier
date, the southernmost known occurrence is et al. 2010 and references therein). Infanticide
southeast of Maevatanana along the east bank and cannibalism, the context of which remains
of the Betsiboka (Rakotonirina et al. 2014), and to be clarified, has been reported in Mariarano
the easternmost occurrence is located near (Ramsay et al. 2020).
Antetemasy (just west of Befandriana Nord).
Propithecus coquereli is known to occur in Coquerel’s Sifaka is listed as Critically Endangered
eleven protected areas: Anjajavy Reserve, on the IUCN Red List of Threatened Species
Anjiamangirana, Ankarafantsika National Park, (Louis Jr. et al. 2020) based on a suspected
Bongolava forest corridor, Bora Special Reserve, reduction of the population of 80% in the past
Mariarano, Marosakoa and Namakia, Mifoko, and upcoming 30 years. The main threats to
Narindra peninsula, and Bombetoka-Beloboka this species are habitat loss and fragmentation
(Salmona et al. 2014, Goodman et al. 2018). (Waeber et al. 2015) and hunting (Garcia and
Goodman 2003, Borgerson et al. 2019). High
This diurnal, vertical clinger-and-leaper is most recent (1996–2016) deforestation rates in the
commonly found in mixed deciduous, dry forests, Bongolava (39.0%) and Ankarafantsika (11.7%)
secondary forests, plantations, brush-and-scrub, protected areas (Goodman et al. 2018), and
and in mosaics of fragmented forests (Salmona the probability of increased deforestation in
et al. 2014). It has also been seen in coastal unprotected lowland forests (Schüßler et al.
mangroves (Donati et al. 2019, Chell et al. 2020). 2020) suggest alarming deforestation rates
Coquerel’s Sifaka feeds mostly on young leaves, across P. coquereli’s distribution. Slash-and-
flowers, fruit, bark and dead wood in the wet burn agriculture and annual grassland burning to
season, and on mature leaves and buds in the dry create fresh pasture for livestock are the principal
season (Richard 1974, McGoogan 2011). As many causes of forest loss in northwestern Madagascar,
as 98 different plant species have been recorded but trees are also cut to produce charcoal as
in its diet (Richard 1978a, 1978b, McGoogan well as planks for building (Waeber et al. 2015).
2011). As with other western sifaka species, Combined, these practices threaten all of its
P. coquereli regularly descends to the ground remaining habitats, even those that are officially
where its locomotion is often characterized as protected (Goodman et al. 2018). Wildlife
‘dancing’, a mixture of chasséing and leaping. hunting and consumption (including lemurs) is
In the forests of Ankarafantsika, it occurs in

23
MADAGASCAR

common around Ankarafantsika (Borgerson et al.

2019) and P. coquereli hunting has been reported
repeatedly (Nicoll and Langrand 1989, Garcia
and Goodman 2003) despite taboos on hunting
sifakas. In some areas, local traditions place
taboos on hunting or eating sifakas, but human
immigration from other regions likely change
such long-held customs (Nicoll and Langrand
1989, Borgerson et al. 2019). The species is
suspected to have been completely extirpated
from Bora Special Reserve after it was delisted
as a protected area by the Malagasy government
following high levels of degradation (Louis Jr. et
al. 2020).

The latest extensive field survey, conducted

between 2009 and 2011, confirmed the presence
of the species in many of the remaining forest
fragments in its range (Salmona et al. 2014). The
Extent of Occurrence is estimated to be less
than 28,000 km² (Louis Jr. et al. 2020). Its Area
of Occupancy is expected to be much smaller
due to the high fragmentation of its habitat. No
systematic surveys have been carried out across
its large range over the past 10 years. Kun-
Rodrigues et al. (2014) found highly variable
densities (between 5 and 100 individuals/
km²) and a negative edge effect, across the
Ankarafantsika National Park. The negative edge
effect penetrated up to 625 m into the forest and
was associated with home range sizes more than
double the size of those in the interior (McGoogan
2011). Steffens and Lehman (2018) found sifakas
only in the largest forest patches (19–118 ha)
of the Ambanjabe mosaic landscape (Western
Ankarafantsika) and reported their absence in
forests smaller than 12 ha. Ramilison et al. (2021),
on the other hand, reported Coquerel’s Sifaka in
forest patches in Mariarano that were as small
as 2.5 ha. The majority of encounters were in
habitat edge zones, most groups were sighted in
introduced Mango trees (Mangifera indica), and
many sightings were in villages and fields. The
latter result echoes the seeming ubiquity of P.
coquereli across its range (Salmona et al. 2014)
with its presence in all sites visited. Salmona et
al. (2014) also reported that they frequently use
secondary forests, plantations, and introduced
Mango and Tamarind trees.

24
In 2009–2010, the Ankarafantsika population was or population decline (Bailey et al. 2016, Guevara
considered to be the largest remaining across et al. 2021), which suggests that the demography
its range, estimated at around 47,000 individuals of P. coquereli is impacted by both the historical
(Kun-Rodrigues et al. 2014). Kun-Rodrigues et al. human shift to extensive agropastoralism
(2014) suggested that it had been declining prior practices in the region (Voarintsoa et al. 2017)
to their survey, and that it would likely continue and late Quaternary climate fluctuations. The
declining, especially following the uncontrolled viability of the population is at high risk of
fires in 2021. Steffens et al. (2020) recorded sifakas being compromised if habitat availability keeps
in only two of the nine sites they surveyed along decreasing, especially given the long generation
the periphery of Ankarafantsika National Park. time of 10–20 years and that a decade ago there
In a comparative analysis of how diurnal lemurs were only 47,000 individuals left in Ankarafantsika
cope with different landscape metrics, Eppley et (Kun-Rodrigues et al. 2014), not all of which are
al. (2020) found that P. coquereli relies on large reproductive. Generation time estimates are
forest blocks. Using species distribution models, tentative, however, and partly based on data
Brown and Yoder (2015) inferred that P. coquereli from Verreaux’s Sifaka, P. verreauxi (Lawler et al.
is unlikely to face drastic range contraction from 2009, Morris et al. 2011; see Salmona et al. 2017
climate change alone between 2000 and 2080. for details).

In 2022, 60 individuals of this species were being Except for Ankarafantsika National Park, most
held in a small number of zoos around the world forests where this species is found have not
(ZIMS 2022), more than 50 in registered captive been surveyed for at least a decade, and the
facilities in Madagascar, and even more held in south-eastern and eastern limits of P. coquereli’s
illegal facilities that are not registered with the distribution are still unclear (Salmona et al. 2014).
government (Reuter and Schaefer 2017, LaFleur Clarifying P. coquereli’s south-eastern range is an
et al. 2019). It is difficult to maintain healthy urgent prerequisite to inclusive conservation and
indriids in captivity (Zehr et al. 2014, Cassady et al. will require extended survey efforts in remote
2018). Sifakas in captivity outside of Madagascar areas. To date, population sizes have only been
(P. coquereli and P. coronatus) frequently suffer estimated in Ankarafantsika (Richard 1978b,
from gastrointestinal and diarrheal diseases, Albignac 1981, Kun-Rodrigues et al. 2014), and
decreased longevity, low infant survivorship, and systematic surveys of all known localities are
decreased reproductive success (Zehr et al. 2014, needed. Accurate estimates of habitat size and
Cassady et al. 2018). deforestation will allow scientists to estimate
the overall population size of the species and
Propithecus coquereli has the highest the demographic trends—critical for proper
mitochondrial (Dloop sequence) and nuclear conservation status assessments.
(microsatellites) genetic diversity among the
northern and northwestern sifakas, that includes The prioritization of sites suitable for conserving
P. perrieri and P. tattersalli (Bailey et al. 2016). It the species should be updated (considering
has the highest genome-wide heterozygosity especially the population genetic structure and
of all assembled sifaka species genomes (P. the potential barriers to dispersal) to enable the
tattersalli, P. coquereli, P. diadema, P. verreauxi; conservation of the species’ genetic diversity, and
Guevara et al. 2021). These estimates are in line to ensure and re-establish connectivity among
with its large geographic range and potentially historically connected populations. Further
large population size. It appears to be genetically fine-scale population genetic studies should
structured in at least four distinct populations be conducted, to ascertain the current level of
(Mariarano, Ankarafantsika, Anjiamangirana, connectivity between sites and the impacts on
and Bora), suggesting a strong effect of major the viability of the remaining sifaka populations
rivers (Mahajamba and Sofia) and large tracts of in the wild. Research efforts should also focus
grassland in limiting gene flow among the forests on the dichotomy of forest edge use – avoidance
where it is found (Bailey et al. 2016). These in Ankarafantsika and preference in Mariarano
populations also show genetic signals that imply (McGoogan 2011, Kun-Rodrigues et al. 2014,
changes in habitat connectivity (fragmentation) Ramilison et al. 2021). Local traditions placing

25
MADAGASCAR

taboos on hunting and eating sifakas may play a History, Description, and Biota. Volume III: Western
pivotal role, and should be leveraged (Lingard et and Southwestern Madagascar – Synthesis. Chicago:
University of Chicago Press.
al. 2003, Jones et al. 2008). Guevara, E.E, Webster, T.H., Lawler, R.R., Bradley, B.J.,
Greene, L.K., Ranaivonasy, J., Ratsirarson, J., Harris, R.A.,
Current conservation projects targeting Liu, Y., Murali, S., Raveendran, M., Hughes, D.S.T., Muzny,
Coquerel’s Sifakas and their habitat include the D.M., Yoder, A.D., Worley, K.C. and Rogers, J. (2021).
long-term endeavors of Madagascar National Comparative genomic analysis of sifakas (Propithecus)
reveals selection for folivory and high heterozygosity
Parks and the more recent efforts of Planet despite endangered status. Sci. Adv. 7(17): eabd2274.
Madagascar and Operation Wallacea. They Jones, J.P., Andriamarovololona, M.M. and Hockley, N.
include fire management, monitoring patrols, (2008). The importance of taboos and social norms to
forest restoration, and community development. conservation in Madagascar. Conserv. Biol. 22(4): 976–
986.
Kun-Rodrigues, C., Salmona, J., Besolo, A., Rasolondraibe,
E., Rabarivola, C., Marques, T.A. and Chikhi, L. (2014).
New density estimates of a threatened sifaka species
Albignac, R. (1981). Lemurine social and territorial (Propithecus coquereli) in Ankarafantsika National Park.
organization in a north-western Malagasy forest Am. J. Primatol. 76(6): 515–528.
(restricted area of Ampijoroa). In: A. Chiarelli and R. LaFleur, M., Reuter, K. and Schaefer, M. (2019). Tourism and
Corrucini (eds.), Primate Behavior and Sociobiology, pp. lemurs: the fate of diurnal indriids at popular tourist
25–29. Berlin: Springer Verlag. destinations. Lemur News 22: 54–56.
Bailey, C.A., McLain, A.T., Paquette, S.R., McGuire, S.M., Lawler, R.R., Caswell, H., Richard, A.F., Ratsirarson, J., Dewar,
Shore, G.D., Lei, R., Randriamanana, J.C., Rabekinaja, R.E. and Schwartz, M. (2009). Demography of Verreaux’s
J.D., Rakotoarisoa, G., Razafindrakoto, A. and sifaka in a stochastic rainfall environment. Oecologia
Brenneman, R. (2016). Evaluating the genetic diversity 161(3): 491–504.
of three endangered lemur species (genus: Propithecus) Lingard, M., Raharison, N., Rabakonandrianina, E.,
from northern Madagascar. J. Primatol. 5(1): 132. Rakotoarisoa, J.A., and Elmqvist, T. (2003). The role of
Borgerson, C., Randrianasolo, J.F., Anjaranirina, E.J.G. local taboos in conservation and management of species:
and Golden, C.D. (2019). Wildlife hunting in complex the radiated tortoise in southern Madagascar. Conserv.
human-environmental systems: how understanding Soc. 1(2): 223–246.
natural resource use and human welfare can improve Louis Jr., E.E., Bailey, C.A., Sefczek, T.M., King, T., Radespiel,
conservation in the Ankarafantsika National Park, U. and Frasier, C.L. (2020). Propithecus coquereli.
Madagascar. Madag. Conserv. Dev. 14(1): 37–45. The IUCN Red List of Threatened Species 2020:
Brown, J.L. and Yoder, A.D. (2015). Shifting ranges and e.T18355A115572275.
conservation challenges for lemurs in face of climate McGoogan, K.C. (2011). Edge Effects on the Behaviour
change. Ecol. Evol. 5(6): 1131–1142. and Ecology of Propithecus coquereli in Northwest
Cassady, K., Cullen, J.M. and Williams, C.V. (2018). Mortality Madagascar. Doctoral dissertation, University of Toronto,
in Coquerel’s sifakas (Propithecus coquereli) under Toronto.
human care: a retrospective survey from the Duke Lemur Mittermeier, R.A., Louis Jr., E.E., Richardson, M., Schwitzer,
Center 1990–2015. J. Zoo Wildl. Med. 49(2): 315–323. C., Langrand, O., Rylands, A.B., Hawkins, F., Rajaobelina,
Chell, C., Corral, M.E., Grant, I., Laskai, C.J., Gough, H., S., Ratsimbazafy, J., Rasoloarison, R., Roos, C., Kappeler,
Rakotomalala, S.D., Ward, S.J. and Thompson, C. P.M. and MacKinnon, J. (2010). Lemurs of Madagascar.
(2020). A brief observation of endangered Coquerel’s 3rd edition. Arlington, VA: Conservation International.
sifaka (Propithecus coquereli) feeding on red mangrove Morris, W.F., Altmann, J., Brockman, D.K., Cords, M., Fedigan,
(Rhizophora mucronata) vegetation in a mangrove L.M., Pusey, A.E., Stoinski, T.S., Bronikowski, A.M., Alberts,
environment, northwest Madagascar. Int. J. Primatol. 41: S.C. and Strier, K.B. (2011). Low demographic variability
562–565. in wild primate populations: fitness impacts of variation,
Donati, G., Eppley, T.M., Ralison, J., Youssouf, J. and covariation, and serial correlation in vital rates. Am. Nat.
Ganzhorn, J.U. (2019). Lemurs in mangroves and other 177(1): E14–E28.
flooded habitats. In: A.A. Barnett, I. Matsuda and K. Nicoll, M.E. and Langrand, O. (1989). Madagascar: Revue
Nowak (eds.), Primates in Flooded Habitats: Ecology de la Conservation et des Aires Protégées. Gland,
and Conservation, pp. 29–32. Cambridge: Cambridge Switzerland: World Wildlife Fund (WWF).
University Press. Petter, J.-J. (1962a). Recherches sur l’écologie et l’éthologie
Eppley, T.M., Santini, L., Tinsman, J.C. and Donati, G. (2020). des lémuriens malgaches. Mem. Mus. Natl. Hist. Nat.
Do functional traits offset the effects of fragmentation? Hist., Ser. A 27: 1–146.
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western Madagascar. Oryx 37(1): 115–118. populations de Propithecus coquereli dans les fragments
Goodman, S.M., Raherilalao, J. and Wohlhauser, S. (2018). forestiers d’Anjiamangirana, Ankarafantsika, Mariarano
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 et Bora, du Nord-Ouest de Madagascar. Doctoral reconstruct paleoenvironmental changes in northwestern
dissertation, Université d’Antananarivo, Antananarivo. Madagascar between 370 CE and 1300 CE. Palaeogeogr.
Rakotonirina, L.H.F., Randriatsara, F., Rakotoarisoa, A.H., Palaeoclimatol. Palaeoecol. 469: 138–155.
Rakotondrabe, A.R., Razafindramanana, J., Ratsimbazafy, Waeber, P.O., Wilmé, L., Ramamonjisoa, B., Garcia, C.,
J. and King, T. (2014). A preliminary assessment of sifaka Rakotomalala, D., Rabemananjara, Z.H., Kull, C.A.,
(Propithecus) distribution, chromatic variation and Ganzhorn, J.U. and Sorg, J.P. (2015). Dry forests in
conservation in western central Madagascar. Primate Madagascar: neglected and under pressure. Int. For. Rev.
Conserv. (28): 43–53. 17(2): 127–148.
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Rakotondravony, R., Radespiel, U. and Ramsay, M.S. (2021). Biogeographic evolution of Madagascar’s microendemic
Distribution of the critically endangered Coquerel’s sifaka biota. Science 312(5776): 1063–1065.
(Propithecus coquereli) across a fragmented landscape in Zehr, S.M., Roach, R.G., Haring, D., Taylor, J., Cameron,
NW Madagascar. Afr. J. Ecol. 59(2): 350–358. F.H. and Yoder, A.D. (2014). Life history profiles for 27
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Reuter, K.E. and Schaefer, M.S. (2017). Illegal captive lemurs Available online: https://zims.species360.org.
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Climate change and human colonization triggered
habitat loss and fragmentation in Madagascar. Mol. Ecol.
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specific metapopulation responses to habitat loss and
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Steffens, T.S., Maheritafika, H.M.R., Hildebrand, J. and
Aylward, M. (2020). Lemur distribution and resident
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Ankarafantsika National Park, Madagascar. Primate
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Columbia University Press.
Voarintsoa, N.R.G., Wang, L., Railsback, L.B., Brook,
G.A., Liang, F., Cheng, H. and Edwards, R.L. (2017).
Multiple proxy analyses of a U/Th-dated stalagmite to

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AFRICA
 AFRICA

© Andrew Perkin

30
 RONDO DWARF GALAGO
Paragalago rondoensis (Honess in Kingdon, 1997)

Tanzania
(2006–2014, 2018, 2022)

Andrew Perkin

Weighing approximately 60 g, the Rondo Dwarf the northern parts of its range (for example, in
Galago is the smallest of all galagos (Perkin and Zaraninge forest, the Pugu/Kazimzumbwi Forest
Honess 2013). It is distinct from other dwarf Reserve [FR] and the Pande Game Reserve
galagos in its diminutive size, a bottle-brush- [GR]), the Rondo galago is sympatric with the
shaped tail, its reproductive anatomy, and its Zanzibar Galago, Paragalago zanzibaricus (Near
distinctive “double unit rolling call” (Perkin and Threatened: Perkin et al. 2020) and in the southern
Honess 2013). Current knowledge indicates that parts of its range (for example, in Rondo, Litipo
P. rondoensis is endemic to the coastal forests and Noto), it is sympatric with Grant’s Galago, P.
of Tanzania. There are three spatially distinct granti (Least Concern: De Jong et al. 2019b).
sub-populations. One is in southeast Tanzania
near the coastal towns of Lindi and Mtwara. Paragalago rondoensis, considered Endangered
The second is approximately 400 km north, in on the IUCN Red List of Threatened Species
pockets of forest around the capital city of Dar es (Perkin 2020), has an extremely limited and
Salaam. The third sub-population is in Sadaani fragmented range across a number of remnant
National Park, approximately 100 km north of Dar patches of Eastern African Coastal Dry Forest
es Salaam. There is emerging data (vocal and (Burgess and Clarke 2000) in Tanzania. These are
penile morphological), however, that suggests at Zaraninge forest (06°08’S, 38°38’E) in Sadaani
the northern and southern populations may be National Park (Perkin 2000), Pande GR (06°42’S,
phylogenetically distinct. 39°05’E), Pugu/Kazimzumbwi (06°54’S, 39°05’E)
(Perkin 2003, 2004), Rondo Nature Reserve (NR)
Rondo Dwarf Galagos have a mixed diet of (10°08’S, 39°12’E), and the Litipo (10°02’S, 39°29’E)
insects and fruit. They often feed close to the and Ziwani (10°20’S, 40°18’E) forest reserves
ground, moving by vertical clinging and leaping (Honess 1996, Honess and Bearder 1996). New
in the shrubby understory. They build daytime sub-populations were identified in 2007 near
sleeping nests, usually in the canopy (Bearder Lindi town in Chitoa FR (09°57’S, 39°27’E) and the
et al. 2003). As with many small primates, P. Ruawa FR (09°44’S, 39°33’E) and, in 2011, in Noto
rondoensis is probably subject to predation Village FR (09°53’S, 39°25’E) (Perkin et al. 2011,
from owls and other nocturnal predators, such as 2013) and, of the northern population, at the
genets, palm civets and snakes. The presence Ruvu South FR (06°58’S, 38°52’E). Specimens of
of these predators invokes intense episodes of P. rondoensis, originally described as Galagoides
alarm calling (Perkin and Honess 2013). demidoffi phasma, were collected by Ionides
from the Rondo Plateau, southeast Tanzania in
Across its known range, the Rondo Dwarf Galago 1955, and by Lumsden from Nambunga, near
can be found in sympatry with a number of other Kitangari, (approximately 10°40’S, 39°25’E) on the
galagos, including two much larger species in the Makonde Plateau, Newala District in 1953. There
genus Otolemur: Garnett’s Greater Galago O. are doubts as to the persistence of the species on
garnettii (Least Concern: De Jong et al. 2019a) the Makonde Plateau, which has been extensively
and the Thick-tailed Galago, O. crassicaudatus cleared for agriculture. In 1992, surveys there
(Least Concern: Masters and Bearder 2019). In failed to detect any extant populations (Honess

31
AFRICA

1996). Distribution surveys have been conducted

in the southern (Honess 1996; Perkin et al. unpubl.
data) 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 density estimates
of: 3–6/ha at the Pande GR (Perkin 2003) and 8/ha
at the Pugu FR (Perkin 2004). Relative abundance
has also been estimated from encounter rates:
3–10/hr at Pande GR and Pugu/ Kazimzumbwi
FR (Perkin 2003, 2004), and 3.94/hr at Rondo FR
(Honess 1996). There is a clear and urgent need
for further surveys to determine population sizes
in these dwindling forest patches.

The major threat facing the Rondo Dwarf Galago

is loss of habitat. All sites are subject to some
level of agricultural encroachment, charcoal
manufacture, and/or logging. In 2008, the
known Area of Occupancy of P. rondoensis was
<101.6 km², but new data on forest area change
indicates this figure has fallen to 87.4 km². In the
Pande GR (2.4 km²), Chitoa (5 km²), and Rondo
(25 km²) forest reserves, forest cover remained
the same between 2008 and 2014. However,
forest cover between 2008 and 2014 declined in
the Zaraninge forest from 20 km² to 15 km², in
Pugu/Kazimzumbwi FR from 33.5 km² to 8 km², in
Ruawa FR and Litipo FR from 4 km² to 3 km², and
in Ziwani FR from 7.7 km² to 1 km². Two newly
discovered Areas of Occupancy are Ruvu South,
in which forest cover fell from 20 km² to 5 km²,
and Noto, in which forest cover fell from 21 km²
to 20 km² in the same time period (Burgess and
Clarke 2000, Doggart 2003, Perkin et al. unpubl.
data).

As habitat availability decreases, the population

trend must also be assumed to be declining, the
rate varying according to the level of protection
of each forest fragment. All sites, except the
Pande GR, Zaraninge (in Saadani National Park)
and the Rondo NR, are national or local authority
forest reserves and, as such, nominally, but
in practice minimally, protected. Since 2008,
protection of two forests has increased: the Noto
plateau forest, formerly open village land, as 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 Dwarf Galago conservation considering

32
their relatively large size. Given current trends Honess, P.E. (1996). Speciation Among Galagos (Primates,
in charcoal production for nearby Dar es Salaam, Galagidae) in Tanzanian Forests. Doctoral thesis, Oxford:
Oxford Brookes University, Oxford, UK.
the forest reserves of Pugu and Kazimzumbwi Honess, P.E. and Bearder, S. (1996). Descriptions of the dwarf
were predicted to disappear over the next 10– galago species of Tanzania. Afr. Primates. 2(2): 75–79.
15 years (Ahrends 2005). Recorded forest loss Masters, J. and Bearder, S. (2019). Otolemur crassicaudatus.
in Pugu/Kazimzumbwe and Ruvu South has The IUCN Red List of Threatened Species 2019:
e.T15643A17963699.
been attributed to the rampant charcoal trade. Perkin, A.W (2000). A Field Study of the Conservation Status
Pande, as a Game Reserve, is relatively secure, and Diversity of Galagos in Zaraninge Forest, Coast
and Zareninge forest, being in a national park, Region, Tanzania. Report, WWF-Tanzania, Dar es Salaam,
is the most protected part of the range of G. Tanzania.
rondoensis. In the south, the Noto, Chitoa and Perkin, A.W. (2003). Mammals. In: N. Doggart (ed.), Pande
Game Reserve: A Biodiversity Survey, p.95. Dar es Salaam,
Rondo populations are the most secure, as they Tanzania: Tanzania Forest Conservation Group.
are buffered by tracts of woodland. The type Perkin, A.W. (2004). Galagos of the Coastal Forests and
population at Rondo is buffered by woodland Eastern Arc Mtns. of Tanzania – Notes and Records.
and Pinus plantations managed by the Rondo Tanzania Forest Conservation Group, Technical Paper 8.
Dar es Salaam, Tanzania.
Forestry Project, and is now a nature reserve. Perkin, A.W. (2020). Paragalago rondoensis. The IUCN Red
Litipo and Ruawa forest reserves are under threat List of Threatened Species 2020: e.T40652A17962115.
from bordering village lands. Ziwani is now mostly Perkin, A.W. and Honess, P.E. (2013). Galagoides rondoensis
degraded scrub forest, thicket and grassland. Rondo Dwarf Galago. In: T.M. Butynski, J. Kingdon and J.
Kalina (eds.), Mammals of Africa. Volume II: Primates, pp.
The following conservation actions are 450–452. London: Bloomsbury Publishing.
Perkin, A.W, Samwel, B. and Gwegime, J. (2011). Going for
recommended to safeguard the future of this gold in the Noto Plateau, SE Tanzania. Arc Journal. 26:
species: 1) continued monitoring of habitat 14–16.
loss rates, 2) surveying new areas for remnant Perkin, A.W., Honess, P.E. and Butynski, T.M. (2013).
populations, 3) implementation of community- Galagoides orinus Mountain Dwarf Galago. In: T.M.
Butynski, J. Kingdon and J. Kalina (eds.), Mammals
based conservation and awareness, 4) assessment of Africa. Volume II: Primates, pp. 452–454. London:
of population status and phylogenetic Bloomsbury Publishing.
relationships between the sub-populations Perkin, A.W., Butynski, T.M. and de Jong, Y.A. (2020).
and confirmation of suspected phylogenetic Paragalago zanzibaricus (errata version published in
distinctions. Until such time that the latter has 2020). The IUCN Red List of Threatened Species 2020:
e.T8790A179843811.
been carried out, each subpopulation must be
considered to be of high conservation value.

Ahrends, A. (2005). Pugu Forest: going, going... The Arc

Journal. 17: 23.
Bearder, S.K., Ambrose, L., Harcourt, C., Honess, P., Perkin,
A.W., Pullen, S., Pimley, E. and Svoboda, N. (2003).
Species-typical patterns of infant care, sleeping site use
and social cohesion among nocturnal primates in Africa.
Folia Primatol. 74: 337–354.
Burgess, N.D. and Clarke G.P. (2000). Coastal Forests of
Eastern Africa. Switzerland and Cambridge, UK: IUCN
Gland.
De Jong, Y.A., Butynski, T.M., Perkin, A.W. and Svensson,
M. (2019a). Otolemur garnettii. The IUCN Red List of
Threatened Species 2019: e.T15644A17963837.
De Jong, Y.A., Butynski, T.M. and Perkin, A.W. (2019b).
Paragalago granti. The IUCN Red List of Threatened
Species 2019: e.T91970347A17962454.
Doggart, N. (ed.). 2003. Pande Game Reserve: A Biodiversity
Survey. Tanzania Forest Conservation Group, Technical
Paper 7. Dar es Salaam, Tanzania.

33
AFRICA

© Edward McLester

34
 GOLDEN-BELLIED MANGABEY
Cercocebus chrysogaster Lydekker, 1900

Democratic Republic of the Congo

(2022)

Edward McLester, John A. Hart and Jo A. Myers Thompson

More than 120 years after being first described, eating Blue Duikers (Philantomba monticola) at
the Golden-bellied Mangabey remains one LuiKotale (McLester 2022).
of Africa’s least-known primates. Previously
considered Data Deficient, it is now listed as Golden-bellied Mangabeys are endemic to
Endangered on the IUCN Red List of Threatened the Democratic Republic of the Congo (DRC).
Species (Hart and Thompson 2020). Found only The species is known to occur in two widely
in a limited range in the central Congo basin, separated subpopulations (a western and an
Golden-bellied Mangabeys are highly threatened eastern population – Ehardt and Butynski 2013,
throughout their distribution and face continuing, Hart and Thompson 2020). Surveys conducted
and potentially rapid, declines. between 1994 and 2007 (Inogwabini and
Thompson 2013) and 2016–2018 (Bessone et al.
The largest of the seven Cercocebus mangabeys, 2019, 2020) identified the western population
these monkeys are characterized by vivid yellow- as occurring within an area of 68,000 km²,
orange fur that covers the belly and abdomen. bordered by the Lokolo River to the north, the
Golden-bellied Mangabey ecology is poorly Luilaka River to the east, the Kasai and Sankuru
known and there have been no detailed behavioral rivers to the south, and the Congo River to the
studies of this species in the wild. Almost all insight west. The eastern range covers an area of about
is limited to anecdotes reported prior to the year 12,000 km² with identified populations known
2000 and preliminary data recorded in 2021 from between the Sankuru and Lubefu rivers in an
two habituated groups at LuiKotale (a study site area locally termed the Kipula Block (Thompson
of the Max Planck Institute of Animal Behavior) in and Hart 2015). Eastern and western populations
Lokolama sector, bordering the southern sector are separated by at least 300 km. There is no
of the Salonga National Park (SNP). Group sizes evidence to support the suggestion by Erhardt
typically number 50–70 individuals (Ehardt and and Butynski (2013) that the two populations are
Butynski 2013). Home ranges are also relatively connected by a forest corridor along the Lukenie
large for cercopithecine monkeys in tropical forest or Sankuru rivers. The total range for both the
(approximately 20–25 km²) and are comparable western and eastern populations is around
and may exceed those of sympatric bonobo 80,000 km². However, Golden-bellied Mangabey
(Pan paniscus) communities (McLester and Fruth occurrence is highly patchy within this large
in prep.) Individuals travel almost exclusively range, and their distribution and abundance
terrestrially and generally climb arboreally only to remain poorly known over many areas.
sleep, forage in large fruit trees, or during heavy
rainfall. The diet consists primarily of fruit, with The western sub-population includes portions
considerable time also spent feeding on seeds of the southern sector of the SNP, which is the
and insects that are obtained from the ground, only protected area in which Golden-bellied
particularly in large patches of leaf litter. Similar Mangabeys occur. Known occurrence within
to Agile Mangabeys (C. agilis) north of the Congo the southern sector of SNP is patchy, with
River, Golden-bellied Mangabeys are notable for observations restricted to the northern and
relatively high rates of mammal consumption, eastern areas of the park (n = 22 observations
with adult males observed regularly catching and from about 4,000 km of line transects and recces
35
AFRICA

covering 93.5% of the sector) (Bessone et al.

2019). There are no records of C. chrysogaster
from the southwestern area of SNP’s southern
sector or from the northern sector of SNP (J.
Eriksson, F. Maisels and G. Reinartz pers. comm.).

The typical elevational range for this species is

300–500 m and habituated groups at LuiKotale
almost exclusively use high and dry terra firma
vegetation at ≥400 m and rarely venture into
swamps or riparian areas (McLester and Fruth
in prep.). Approximately 60% of the habitat in
the western distribution is permanent swamp
or seasonally flooded and riparian forest, within
which areas of terra firma forest comprise an even
lower percentage (Inogwabini and Thompson
2013). In the absence of any detailed estimates
of population density, limited habitat choice and
large home range sizes suggest densities are
likely to be low.

Golden-bellied Mangabeys are in decline

throughout their distribution. In the past 20
years, at least 32% of the total potential habitat
has been lost and an estimated 40% of the
population extirpated (Hart and Thompson 2020).
A major threat is unsustainable hunting, with high
numbers of Golden-bellied Mangabeys killed
for the commercial bushmeat trade in both the
western and eastern populations. In the western
population, surveys from 2001 to 2007 found
Golden-bellied Mangabeys comprised as much
as 70% of bushmeat available in some markets
(Inogwabini and Thompson 2013). Surveys
of markets in the eastern range conducted in
2010 found this species represented about 10%
of all bushmeat recorded. That number fell
considerably by 2015, when the Golden-bellied
Mangabey population was also reported to
have declined significantly in some localities
(Thompson and Hart 2015). Hunting varies across
the distribution. Rates of human activity in the
northern half of the southern sector of SNP, where
the species were reported to be found, were
lower than in the south of the sector, where no
observations of Golden-bellied Mangabeys were
made (Bessone et al. 2019). In some Lokoloma
sector villages in the center of the species’
distribution, Golden-bellied Mangabeys are
rarely consumed as bushmeat and hunting may
be reduced because larger or more conspicuous
species (e.g., colobus monkeys) are easier targets
for hunters (N. Bondjengo and B. Fruth, unpubl.
36
data). In other areas, hunting is likely exacerbated in the SNP but illegal hunting remains a significant
because Golden-bellied Mangabeys are threat, especially associated with villages that do
considered to be agricultural pests (Inogwabini not practice agriculture and for which bushmeat
and Thompson 2013). is the only source of income. The park is also
threatened by an increase in uncontrolled
Golden-bellied Mangabeys are threatened by commercial hunting (Bessone et al. 2019). Across
habitat loss due to both small-scale and larger the distribution, there is a critical need to confirm
industrial logging operations. Inogwabini et where Golden-bellied Mangabeys still occur,
al. (2013) reported that west of the Lake Mai- along with an improved understanding of where
Ndombe, where the species no longer occurs, (and why) they are absent across their large
local communities reported its disappearance range. This will be essential to focus resources
over the course of two decades following the on an effective conservation strategy.
arrival of intensive logging. Currently, small-
scale logging is particularly widespread in the
western range, further decreasing the availability
of suitable habitat for this species. In total, Bessone, M., Bondjengo, N., Hausmann, A., Herbinger, I.,
more than 30% of the remaining habitat has Hohmann, G., Kühl, H., Mbende, M., N’Goran, P., Soliday,
V. and Fruth, B. (2019). Inventaire de la biodiversité
been ceded to logging concessions (Hart and
dans le bloc sud du parc national de la Salonga et
Thompson 2020). développement d’une stratégie de suivi écologique
pour améliorer la protection de la faune et de la flore
The illegal pet trade is also a threat and captive menacées dans le parc. Unpublished report produced
individuals are frequently seen in both the by L’Institut Congolais pour la Conservation de la Nature,
western and eastern subpopulations, as well World Wildlife Fund, Berlin, Germany: The Max Planck
Institute for Evolutionary Anthropology, and Ludwig-
as in Kinshasa (Hart and Thompson 2020). The Maximilians-Universität.
trade appears to extend internationally. In 2021, Bessone, M., Kühl, H.S., Hohmann, G., Herbinger, I., N’Goran
for example, a single shipment confiscated in K.P., Asanzi, P., Da Costa, P.B., Dérozier, V., Fotsing, E.D.B.,
Zimbabwe contained eleven individuals that were Beka, B.I., Iyomi, M.D., Iyatshi, I.B., Kafando, P., Kambere,
M.A., Moundzoho, D.B., Wanzalire, M.L.K. and Fruth, B.
being smuggled to South Africa. John Hart (pers.
(2020). Drawn out of the shadows: surveying secretive
obs.) observed captive infants for sale on the forest species with camera trap distance sampling. J.
streets of Kinshasa as recently as February 2022. Appl. Ecol. 57: 963– 974.
These observations are an alarming indicator of Ehardt, C.L. and Butynski, T.M. (2013). Cercocebus
the threat this species is under, given that captive chrysogaster Golden-bellied Mangabey. In: T.M.
Butynski, J. Kingdon and J. Kalina (eds.), Mammals of
infants are likely a by-product of adult individuals Africa. Volume 2: Primates, pp. 174–177). London, UK:
killed for bushmeat. Bloomsbury Publishing.
Hart, J.A. and Thompson, J. (2020). Cercocebus
Golden-bellied Mangabeys are one of Africa’s chrysogaster. The IUCN Red List of Threatened Species
most threatened but least known primates. To 2020: e.T4207A17956177.
Inogwabini, B.-I. and Thompson, J.A.M. (2013). The golden-
effectively address ongoing declines, research is
bellied mangabey Cercocebus chrysogaster (Primates:
needed on their distribution, population sizes, Cercopithecidae): distribution and conservation status. J.
and ecology as well as the impact of hunting and Threat. Taxa. 5: 4069–4075.
habitat transformation. Recently initiated studies Inogwabini, B.-I., Nzala, A.B. and Bokika, J.C. (2013). People
of habituated groups at LuiKotale should provide and bonobos in the southern Lake Tumba landscape,
Democratic Republic of Congo. Am. J. Hum. Ecol. 2:
the first detailed insight into the behavioral 44–53.
ecology of this species and can directly inform McLester, E. (2022). Golden-bellied mangabeys (Cercocebus
conservation planning by providing data on chrysogaster) consume and share mammalian prey at
habitat use, ranging behavior, and dispersal LuiKotale, Democratic Republic of the Congo. J. Trop.
Ecol. 5pp. doi.10.1017/S026646742200013X
patterns.
Thompson, J.A.M. and Hart, J. (2015). Biological Inventory
and Survey of Bonobos within the Lukenie-Sankuru-
Effective protection of SNP is a high conservation Lusambo Forest Block, Democratic Republic of Congo.
priority given that it is the only protected area in Unpublished report to USFWS. Kinshasa: Lukuru
the species’ range. At present, land conversion Foundation.
and commercial logging are relatively uncommon

37
 AFRICA

SOUTHERN PATAS MONKEY

Erythrocebus baumstarki Matschie, 1906

Kenya (Extirpated), Tanzania

(2022)

Thomas M. Butynski and Yvonne A. de Jong

A detailed review of the taxonomic arrangement

of the patas monkeys (Erythrocebus) is long over-
due. The Southern Patas Monkey (Erythrocebus
baumstarki) was described by Matschie in 1906
from east of Ikoma in central-northern Tanzania.
Elliot (1913) appears to be the last to recognize
baumstarki as a species. Subsequently, this
taxon has been treated as either a synonym or
subspecies of Erythrocebus patas. Only recently
has baumstarki been reinstated as a species
(Gippoliti 2017) based on its unique pelage
coloration and pattern, and geographic isolation
(De Jong and Butynski 2020, 2021, De Jong et al.
2020).

Erythrocebus baumstarki is a large, slender, long-

limbed, semi-terrestrial guenon that typically
lives in one-male, multi-female groups. The
natural history of E. baumstarki is poorly known.
Its geographically closest relative, the Eastern
Patas Monkey (Erythrocebus patas pyrrhonotus),
has been studied in Uganda and Kenya and, at
this time, is used as a proxy for the natural history
of E. baumstarki.

In East Africa, E. p. pyrrhonotus prefers open,

short grass, acacia woodlands and wooded
savannas, where it occurs at low densities (0.03–
1.50 individuals/km²). This monkey rarely sleeps
in the same area on successive nights and has
long day ranges (1,380–7,500 m) and large home
ranges (23–52 km²; Hall 1965, Chism and Rowell
1988, Isbell 1998, Isbell and Chism 2007, Isbell
2013). These characteristics, together with its
typically shy and flighty behavior and ability to
run at high speed (55 km/hour; Hall 1965), makes
Erythrocebus especially difficult to locate and
observe (Makacha and Sirolli 2005, De Jong
et al. 2008, Loishooki et al. 2016). Like E. p.

38
pyrrhonotus in central Kenya, E. baumstarki is primarily due to agricultural expansion and
an ecological specialist, being highly dependent intensification (both crops and livestock), charcoal
on large areas of healthy Whistling Thorn Acacia production, fire, and development activities
(Acacia drepanolobium), its primary food plant, (settlements, roads, dams, power-lines), which
and probably also upon the on-going mutualistic have resulted in widespread habitat degradation,
interactions between ants (Crematogaster spp.) loss, and fragmentation, and extreme declines
and A. drepanolobium. in wildlife populations (Homewood et al. 2001,
Makacha and Sirolli 2005, BurnSilver et al. 2008,
In the early 20th century, E. baumstarki occupied Ogutu et al. 2014, 2016, Loishooki et al. 2016).
large parts of the Serengeti-Mara Ecosystem
and Amboseli Ecosystem of southern Kenya and Throughout the historic range of E. baumstarki,
northern Tanzania (De Jong et al. 2008, 2009, A. drepanolobium woodlands continue to rapidly
2020, De Jong and Butynski 2020, 2021). It seems disappear due to over-use by livestock and
that, at present, E. baumstarki remains only in conversion to cropland. Other major concerns
the protected areas of the western Serengeti are competition with people and livestock for
(Serengeti National Park [14,750 km²], Grumeti habitat and water, particularly during droughts,
Game Reserve [428 km²], Ikorongo Game Reserve hunting by poachers and domestic dogs (Canis
[567 km²], Ikona Wildlife Management Area [255 familiaris), climate change, and loss of genetic
km²]), with the western Serengeti National Park variation. Although these threats apply mostly to
being the stronghold (De Jong and Butynski regions outside protected areas, pastoralists now
2021). move livestock illegally into the protected areas
that support E. baumstarki (African BioServices
The geographic distribution of E. baumstarki in 2019, Veldhuis et al. 2019).
the early 20th century was about 66,000 km². This
has declined roughly 85% to around 9,700 km² Poaching, primarily through the use of wire
at present (post-2009). It was extirpated from snares, is a widespread and serious problem in
Kenya in about 2015 and from the Kilimanjaro western Serengeti (Loibooki et al. 2002, Holmern
Region of Tanzania in about 2011. The present et al. 2007, Nyahongo et al. 2009), the region
Extent of Occurrence (EOO) is roughly 2,150 where the remaining E. baumstarki population
km². The total number of individuals remaining occurs, and on ranches that border the Maasai-
in the wild is probably between 100 and 200, Mara National Reserve (Ogutu et al. 2011, 2016).
including between 50 and 100 mature individuals Although E. baumstarki is not a target species
(De Jong and Butynski 2021). There is no captive for poachers, it is likely that some individuals are
population. captured in snares (Makacha and Sirolli 2005,
Loishooki et al. 2016, De Jong and Butynski
Erythrocebus baumstarki is listed as Critically 2020, 2021). This monkey is probably hunted in
Endangered on the IUCN Red List of Threatened retaliation for raiding crops. The meat is eaten
Species based on its small EOO, fragmented and the pelt used in traditional ceremonies and
distribution, rapid decline in distribution and witchcraft (Makacha and Sirolli 2005, Loishooki et
abundance, small population size, and small al. 2016).
effective population size. All of these parameters
are expected to continue to worsen as the causes Patas monkeys require perennial sources of
are ongoing and unlikely to be reversed in the drinking water (Chism and Rowell 1988, Isbell
foreseeable future (De Jong and Butynski 2020, and Chism 2007, De Jong et al. 2008). The all-day
2021). presence of herders and livestock at increasingly
scarce sources of water appears to be a serious
The ultimate threat to Erythrocebus, and to the problem for E. baumstarki, particularly because
other primates in Tanzania and Kenya, is the rapidly of the attacks by herders and dogs.
growing human population, which is doubling
about every 25–30 years. The main proximate Although data are lacking, it is likely that E.
threats are the widespread unsustainable baumstarki experiences increased exposure to
exploitation of natural resources by humans, parasites and diseases at water sources as they

39
wait, forage, and drink in an environment that BurnSilver, S.B., Worden, J. and Boone, R.B. (2008). Processes
is densely populated by humans and livestock. of fragmentation in the Amboseli Ecosystem, southern
Kajiado District, Kenya. In: K.A. Galvin, R.S. Reid, R.H.
Data are also lacking on the impacts of climate Behnke Jr. and N.T. Hobbs (eds.), Fragmentation in Semi-
change and loss of genetic diversity. Although it arid and Arid Landscapes: Consequences for Human
seems inevitable that these impacts are negative, and Natural Systems, pp. 225–253. Dordrecht, Germany:
they pale against the more immediate threats Springer.
Chism, J. and Rowell, T.E. (1988). The natural history of patas
posed by human population growth and the monkeys. In: A. Gautier-Hion, F. Bourlière, J.P. Gautier
related degradation, loss, and fragmentation of and J. Kingdon (eds.), A Primate Radiation: Evolutionary
A. drepanolobium woodlands and water sources. Biology of the African Guenons (pp. 412–438). Cambridge,
UK: Cambridge University Press.
Erythrocebus baumstarki has never been the focus De Jong, Y.A. and Butynski, T.M. (2020). Erythrocebus
of conservation activities and no conservation baumstarki. The IUCN Red List of Threatened Species
2020: e.T92252436A92252442.
actions are planned to secure the long-term De Jong, Y.A. and Butynski, T.M. (2021). Is the southern
survival of this charismatic species. Indeed, with patas monkey Erythrocebus baumstarki Africa’s next
fewer than 200 individuals remaining in the wild, primate extinction? Reassessing taxonomy, distribution,
an EOO of only about 2,150 km², and the absence abundance, and conservation. Am. J. Primatol. e23316.
10.1002/ajp.23316.
of focused conservation actions, it appears that De Jong, Y.A., Butynski, T.M. and Nekaris, K.A.-I. (2008).
E. baumstarki will be among the first primate Distribution and conservation of the patas monkey
extinctions for continental Africa in historic times. Erythrocebus patas in Kenya. J. East Afr. Nat. Hist. 97:
83–102.
De Jong and Butynski (2021) recommended the De Jong, Y.A., Butynski, T.M., Isbell, L.A. and Lewis, C. (2009).
following conservation actions for E. baumstarki: Historic and current distribution of the southern patas
monkey Erythrocebus patas baumstarki in Tanzania. Oryx
(1) Establish a network of people who will help 43: 267–274.
locate all groups and then closely monitor group De Jong, Y.A., Rylands, A.B. and Butynski, T.M. (2020).
size and age/sex composition, home ranges, Erythrocebus patas. The IUCN Red List of Threatened
and threats; (2) Conduct detailed surveys every Species 2020: e.T174391079A17940998.
Elliot, D.G. (1913). A Review of the Primates. Volume 3:
two years to re-assess geographic distribution, Anthropoidea (Miopithecus to Pan). Monograph Series.
abundance, population structure, conservation New York, USA: American Museum of Natural History.
status, and threats; (3) Undertake a detailed, Gippoliti, S. (2017). On the taxonomy of Erythrocebus with a
long-term, ecological and behavioral study; re-evaluation of Erythrocebus poliophaeus (Reichenbach,
(4) Implement molecular research projects to 1862) from the Blue Nile region of Sudan and Ethiopia.
Primate Conserv. (31): 53–59.
assess the level of genomic erosion; (5) Establish Hall, K.R.L. (1965). Behavior and ecology of the wild patas
dedicated, reliable, wildlife water sources where monkey (Erythrocebus patas) in Uganda. J. Zool. 148:
E. baumstarki occurs; (6) Stop poaching and 15–87.
illegal livestock grazing, and keep domestic dogs Holmern, T., Muya, J. and Røskaft, E. (2007). Local law
enforcement and illegal bushmeat hunting outside the
out of the protected areas; (7) Study and monitor Serengeti National Park, Tanzania. Environ. Conserv. 34:
the impacts of browsing on A. drepanolobium 55–63.
by livestock, Savanna Elephant (Loxodonta Homewood, K., Lambin, E.F., Coast, E., Kariuki, A., Kikula,
africana), Black Rhinoceros (Diceros bicornis), I., Kivelia, J., Said, M., Serneels, S. and Thompson, M.
and Rothschild’s Giraffe (Giraffe camelopardalis), (2001). Long-term changes in Serengeti-Mara wildebeest
and land cover: pastoralism, population, or policies.
and assess how this affects E. baumstarki; (8) PNAS 98: 12544–12549.
Bring the plight of E. baumstarki to wide national Isbell, L.A. (1998). Diet for a small primate: Insectivory and
and international attention; and (9) Produce an gummivory in the (large) patas monkey (Erythrocebus
‘Erythrocebus baumstarki Conservation Action patas pyrrhonotus). Am. J. Primatol. 45: 381–398.
Isbell, L.A. (2013). Erythrocebus patas Patas Monkey (Hussar
Plan’ and ensure that this plan is implemented by Monkey, Nisnas Monkey, nisnas). In: T.M. Butynski, J.
those authorities responsible for the conservation Kingdon and J. Kalina (eds.), Mammals of Africa. Volume
of Tanzania’s biodiversity. II: Primates, pp. 257–264. London, UK: Bloomsbury
Publishing.
Isbell, L.A. and Chism, J. (2007). Distribution and abundance
of patas monkeys (Erythrocebus patas) in Laikipia, Kenya,
1979–2004. Am. J. Primatol. 69: 1223–1235.
African BioServices (2019). Serengeti Squeeze – Interactive Loibooki, M., Hofer, H., Campbell, K.L.I. and East, M.L.
Maps. Available online: https://arcg.is/0851br. (2002). Bushmeat hunting by communities adjacent to

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the Serengeti National Park, Tanzania: The importance

of livestock ownership and alternative sources of protein
and income. Environ. Conserv. 29: 391–398.
Loishooki, A.G., Kihwele, E.S., Nasari, D.S. and Mafuru, G.
(2016). Assessment of Distribution and Conservation
Mitigations of the Patas Monkey (Erythrocebus patas) in
Serengeti Ecosystem. Unpublished report by the Unit of
Ecological Monitoring, Tanzania National Parks, Arusha,
Tanzania.
Makacha, S. and Sirolli, A. (2005). Status and Distribution
of Ikoma Patas Monkeys Cercopithecus (Erythrocebus)
patas baumstarki in the Serengeti National Park.
Unpublished report by the Serengeti Biodiversity Project,
Arusha, Tanzania.
Nyahongo, J.W., Holmern, T., Kaltenborn, B.P. and Røskaft,
E. (2009). Spatial and temporal variation in meat and fish
consumption among people in the western Serengeti,
Tanzania: The importance of migratory herbivores. Oryx
43: 258–266.
Ogutu, J.O., Owen-Smith, N., Piepho, H.-P. and Said, M.Y.
(2011). Continuing wildlife population declines and
range contraction in the Mara region of Kenya during
1977–2009. J. Zool., Lond. 285: 99–109.
Ogutu, J.O., Reid, R.O., Piepho, H.-P., Hobbs, N.T., Rainy,
M.E., Kruska, R.L., Worden, J.S. and Nyambenge, M.
(2014). Large herbivore responses to surface water
and land use in an East African savanna: Implications
for conservation and human-wildlife conflict. Biodiv.
Conserv. 23: 573–596.
Ogutu, J.O., Piepho, H.-P., Said, M.Y., Ojwang, G.O., Njino,
L.W., Kifugo, S.C. and Wargute, P.W. (2016). Extreme
wildlife declines and concurrent increases in livestock
numbers in Kenya: What are the causes? PLoS One 11:
e0163249.
Veldhuis, M.P., Ritchie, M.E., Ogutu, J.O., Morrison, T.A.,
Beale, C.M., Estes, A.B., Mwakilema, W., Ojwang, G.O.,
Parr, C.L., Probert, J., Wargute, P.W., Hopcraft, J.G.C.
and Olff, H. (2019a). Cross-boundary human impacts
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363: 1424–1428.

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 AFRICA

© Andrew Gooch

42
 ROLOWAY MONKEY
Cercopithecus roloway (Schreber, 1774)

Côte d’Ivoire, Ghana

(2002, 2006–2022)

Sery Gonedelé Bi, Andrea Dempsey, Inza Koné, W. Scott McGraw and John F. Oates

Cercopithecus roloway and its close relative converted to agricultural land annually between
Cercopithecus diana are very attractive, arboreal 2000 and 2015 (O’Sullivan et al. 2018); conversion
monkeys that inhabit the Upper Guinean forests to cocoa production, in particular, accounted for
of West Africa. The Roloway Monkey, which 54.6% and 77.8% of closed and open forest loss,
once occurred in many of the southern forests respectively (Benefoh et al. 2018). Hunting has
of Ghana and south-central and south-eastern very likely been the major cause of the crash in
Côte d’Ivoire, is distinguished from the Diana roloway populations. Wild meat is a major food
Monkey by its broad white brow line, long white source for Ghanaians, with an estimated 80% of
beard, and yellow thighs. Because individuals the rural population dependent on it as their main
with intermediate coat patterns are known from source of animal protein (Trench 2000, Dempsey
near the Sassandra River in Côte d’Ivoire, some 2014). Any conservation measures to secure the
scientists treat roloway and diana as subspecies future of the Roloway Monkey in Ghana must,
of one species, C. diana (for example, Oates therefore, consider the socio-economic needs
2011). Of the two species, roloway is the more of the local people and address them using an
seriously threatened, and it is now classified as inclusive and sustainable model.
Critically Endangered on the IUCN Red List of
Threatened Species (Koné et al. 2019). Surveys undertaken in Ghana in the early 2000’s
have failed to confirm the presence of Roloway
Roloway Monkeys are upper-canopy specialists Monkeys in many reserves, including the Ankasa
that prefer undisturbed forests. Destruction Conservation Area, Bia National Park, Krokosua
and degradation of their forest habitat due to Hills Forest Reserve, Subri River Forest Reserve,
illegal lumbering and clearing for agriculture and Dadieso Forest Reserve, Atewa Range Forest
mining, together with relentless hunting for the Reserve, and Tano Offin Forest Reserve (Oates
bushmeat trade over many years, have reduced 2006, Gatti 2010, Buzzard and Parker 2012, Wiafe
their population to small, isolated pockets. Miss 2013, 2021). The only two localities in Ghana
Waldron’s Red Colobus (Piliocolobus waldroni) where Roloway Monkeys have been recorded
once inhabited many of the same forests as the by scientists or conservationists in the last
roloway but is now almost certainly extinct (Oates decade are the community-owned forest along
2011). Despite existing conservation efforts, the Tano River (referred to as the “Ankasa-Tano
there is a strong possibility that the roloway Community Rain Forest”) and the Cape Three
monkey will also disappear in the near future. Points Forest Reserve, in the southwest of the
country. The Ankasa-Tano Community Rainforest
Ghana is experiencing a 2% rate of annual consists of patches of swamp forest along the
deforestation and forest degradation, which lower Tano River, adjacent to the Tanoé-Ehy forest
translates into approximately 135,000 ha/year of in Côte d’Ivoire. Surveys of these forests have
forest cover loss due to anthropogenic causes been conducted since 2011 under the auspices
(Kyere-Boateng et al. 2021). A main driver of of West African Primate Conservation Action
forest cover loss is agricultural conversion. In the (WAPCA), a non-government organization,
high forest zone, close to 110,000 ha of forest were and several sightings of Roloway groups have

43
AFRICA

been made, along with Mona Monkeys, Spot-

nosed Monkeys, White-naped Mangabeys and
Olive Colobus (WAPCA 2014, Dempsey 2014,
Osei et al. 2015). WAPCA, with support from
Noè, the Critical Ecosystem Partnership Fund,
the French Development Agency, the Sofi
Tucker Foundation and IUCN’s Programme de
Petites Initiatives, has supported a community-
based conservation project with communities
around these forests, establishing an Ankasa-
Tano Community Resource Management Area
(CREMA), which works to protect the forest
through the sustainable management of natural
resources. In 2018, a survey of the Cape Three
Points Forest Reserve recorded a single sighting
of C. roloway and, since 2019, WAPCA has
been working with the local communities and
government to replicate the methodologies
which were successfully implemented around
the Ankasa-Tano Community Rainforest. Future
conservation action should be focused around
Ankasa-Tano CREMA, supporting patrols,
reforestation and the effective running of the
elected community management committees, as
well as developing additional income revenues
(e.g., from palm oil and cassava), in order to
complement existing organic cocoa and coconut
oil businesses. Around Cape Three Point Reserve,
similar support is required to maintain the newly
elected CREMA, including associated activities
such as patrolling, and the development of
sustainable and improved community livelihoods
(for instance, honey production and ecotourism).
WAPCA has an ongoing project to survey the
historical range of the Roloway in Ghana, and
assess their distribution and abundance more
accurately. In the next two years, these surveys
should be conducted in Dadieso Forest Reserve,
Krokosua Hills Forest Reserve, and Yoyo River
Forest Reserve using camera traps and audio
recorders, as well as more traditional methods.

The Roloway Monkey’s status in neighboring Côte

d’Ivoire is also dire. In the late 1990s, Roloway
Monkeys were known or strongly suspected to
exist in three forests: the Yaya Forest Reserve,
the Tanoé-Ehy forest adjacent to the Ehy Lagoon,
and Parc National des Îles Ehotilé (McGraw 1998,
2005, Koné and Akpatou 2005, Gonedelé Bi et
al. 2013). Surveys of 18 areas between 2004 and
2008 (Gonedelé Bi et al. 2008, 2012) confirmed

44
the presence of Roloway Monkeys only in the vellerosus) in Côte d’Ivoire. An intensive camera
Tanoé forest, suggesting that the Roloway trapping campaign in the Tanoé-Ehy Forest
Monkey may have been eliminated from at least initiated to ‘rediscover’ Miss Waldron’s Red
two forest areas (Parc National des Îles Ehotilé Colobus began in late 2019. This initiative,
and Yaya Forest Reserve) in the last dozen years. supported by Re:wild, has yielded videos
Subsequent surveys carried out in southern Côte and photographs of Roloways, White-naped
d’Ivoire suggest a handful of Roloway Monkeys Mangabeys, White-thighed Black-and-white
may still survive in two forest reserves along Colobus, but not Miss Waldron’s Red Colobus,
the country’s coast. In June 2011, Gonedelé Bi together with other arboreal vertebrates,
observed one Roloway individual in the Dassioko including the Tree Pangolin (Phataginus
Sud Forest Reserve (Bitty et al. 2013, Gonedelé Bi tricuspis). Efforts led by I. Koné involving several
et al. 2014). In 2012, Gonedelé Bi and A.E. Bitty organizations, including the NGOs West African
observed Roloway Monkeys in Port Gauthier Primate Conservation Action (WAPCA), Action
Forest Reserve, and in October 2013, Gonedelé pour la Conservation de la Biodiversité en Côte
Bi obtained photographs of monkeys poached d’Ivoire (ABC-CI), Mulhouse Zoo, and SOS
inside this reserve, including an image purported Forêts, helped stop a large palm oil company
to be a Roloway. The beard on this individual from causing further habitat degradation, and a
appears short for a Roloway, raising the possibility community-based conservation program initiated
that surviving individuals in this portion of the in 2006 has helped reduce poaching in this forest
interfluvial region may in fact be hybrids. In any (Koné 2015). Thanks to continued efforts of
case, no sightings of Roloway Monkeys have communities supported by the Centre Suisse
been made in the Dassioko Sud or Port Gauthier de Recherches Scientifiques (CSRS) and various
forest reserves since 2012. These reserves are partners including the Association Française
described as coastal evergreen forests, and both des Parc Zoologique (AFdPZ), WAPCA, Parcs de
are heavily degraded due to a large influx of Noé, the Critical Ecosystem Partnership Fund
farmers and hunters from the north of the country (CEPF), Rainforest Trust, and Re:wild, the Tanoé-
(Bitty et al. 2013). Gonedelé Bi and colleagues, in Ehy forest was recently officially designated as a
cooperation with Société de Développement des community reserve, the first of its kind in Côte
Forêts (SODEFOR) and local communities, have d’Ivoire. Hunting and chainsaw milling still
organized regular surveys aimed at removing occur in that forest, posing serious threats to
illegal farmers and hunters from both reserves. the survival of its primate populations. Water
However, surveys made in August 2015 revealed pollution linked to gold mining at the periphery
that a logging company (Société Industrielle de of the Tanoé-Ehy Forest represents an emerging
Bois et Débités) had begun clearing a portion of threat. In 2021, the CSRS and several partners
the Port Gauthier Reserve. A recent recce survey succeeded in stopping the launch of a gold
conducted by Lezou in September 2021 indicated mining project at the north-western periphery of
that the situation has become more dire. During the Tanoé-Ehy forest (Koné et al. 2021).
this survey, no primates were observed, poaching
signs inside the reserve were high (1.46 signs per As the potential last refuge for Roloway Monkeys
km), and traditional hunters (Dozo) were acting and one of the last refuges for White-naped
as a military force inside the reserve (Lezou 2021). Mangabeys and White-thighed Black-and-white
Colobus, the protection of the Tanoé-Ehy Forest
Thus, the only forest in Côte d’Ivoire where in Côte d’Ivoire and the adjacent Ankasa-Tano
Roloway Monkeys are confirmed to still exist Community Rain Forest in Ghana should be the
is the Tanoé-Ehy forest adjacent to the Ehy highest conservation priority. By any measure,
Lagoon, and immediately across the Tano River the Roloway Monkey must be considered one
from the Ankasa-Tano Community Rainforest of the most Critically Endangered monkeys in
in Ghana. This wet forest also harbors one of Africa and is evidently on the verge of extinction
the few remaining populations of White-naped (Oates 2011). In addition, the captive population
Mangabeys (Cercocebus lunulatus) and White- is now so small that extinction in captivity is also
thighed Black-and-white Colobus (Colobus a strong possibility (Lefaux and Montjardet 2016).

45
 AFRICA

Benefoh, D.T., Villamorvan Noordwijk, M., Borgemeister, and Actions for the Community-based Conservation
C., Asante, W.A. and Asubonteng, K.O. (2018). Assessing of Critically Endangered Primates and their habitat.
land-use typologies and change intensities in a 2018–2020 Activity report. Abidjan: Centre Suisse de
structurally complex Ghanaian cocoa landscape. Appl. Recherches Scientifiques (CSRS).
Geogr. 99: 109–119. Kyere-Boateng, R., Marek, M.V., Huba, M. and Kluvankova, T.
Bitty, E.A., Gonedelé Bi, S. and McGraw, W.S. (2013). (2021). Perceived injustices in forest policy interventions
Accelerating deforestation and hunting in protected are causes of forest resources degradation and loss in
reserves jeopardize primates in southern Côte d’Ivoire. Ghana: a review. Open J. For. 11: 171–191.
Am. J. Phys. Anthropol. Suppl. 56: 81–82. Lefaux, B. and Montjardet, L. (2016). 7th European Roloway
Buzzard, P.J. and Parker, A.J.A. (2012). Surveys from the Subri Monkey Studbook (Cercopithecus roloway). Edited by
River Forest Reserve, Ghana. Afr. Primates. 7: 175–183. Mulhouse Zoological and Botanical Park, Mulhouse,
Dempsey, A. (2014). Save Our Forest, Save Our Future: A France.
Survey of the Tanoe-Ankasa Community-owned Forest Lezou, D.P.A. (2021). Inventaire des primates et des menaces
for Primate Presence and Illegal Activity, Introducing dans la forêt classée de Dassioko sud, Côte d’Ivoire.
SMART Technology. Master’s thesis, Oxford Brookes Master’s dissertation, Université Félix Houphouët Boigny,
University, Oxford, England. Abidjan, Côte d’Ivoire.
Gatti, S. (2010). Status of Primate Populations in Protected McGraw, W.S. (1998). Surveys of endangered primates in the
Areas Targeted Under the Community Forest Biodiversity forest reserves of eastern Côte d’Ivoire. Afr. Primates. 3:
Project. CFBP Report, West African Primate Conservation 22–25.
and Ghana Wildlife Division/Forestry Commission, McGraw, W.S. (2005). Update on the search for Miss Waldron’s
Accra, Ghana. Available online <http://www.primate-sg. red colobus monkey (Procolobus badius waldroni). Int. J.
org/ghana-unpublished-reports/>. Primatol. 26: 605–619.
Gonedelé Bi, S., Koné, I., Béné, J.-C.K., Bitty, A.E., Akpatou, Oates, J.F. (2006). Primate Conservation in the Forests of
B.K., Goné Bi, Z., Ouattara, K. and Koffi, D.A. (2008). Western Ghana: Field Survey Results, 2005–2006. Ghana:
Tanoé forest, south-eastern Côte d’Ivoire identified Wildlife Division, Forestry Commission.
as a high priority site for the conservation of Critically Oates, J.F. (2011). Primates of West Africa: A Field Guide
Endangered primates in West Africa. Trop. Conserv. Sci. and Natural History. Arlington, VA: Conservation
1: 265–278. International.
Gonedelé Bi, S., Koné, I., Bitty, A.E., Béné, J.-C.K., Akpatou, Osei, D., Horwich, R.H. and Pittman, J.M. (2015). First
B. and Zinner, D. (2012). Distribution and conservation sightings of the roloway monkey (Cercopithecus diana
status of catarrhine primates in Côte d’Ivoire (West roloway) in Ghana in ten years and the status of other
Africa). Folia Primatol. 83: 11–23. endangered primates in southwestern Ghana. Afr.
Gonedelé Bi, S., Béné, J.-C.K., Bitty, A.E., N’Guessan, A., Primates 10: 25–40.
Koffi, A.D., Akpatou, B. and Koné, I. (2013). Roloway O’Sullivan, R., Roth, M., Antwi, Y.A., Ramirez, P. and
guenon (Cercopithecus diana roloway) and white-naped Sommerville, M. (2018). Land and Tree Tenure Innovations
mangabey (Cercocebus atys lunulatus) prefer mangrove for Financing Smallholder Cocoa Rehabilitation in Ghana.
habitats in Tanoé Forest, southeastern Ivory Coast. J. Presentation at the 2018 World Bank Conference on Land
Ecosystem. Ecography. 3: 126. and Poverty, Washington DC, 19–23 March 2018.
Gonedelé Bi, S., Bitty, E.A. and McGraw, W.S. (2014). Trench, P. (2000). The many faces of community wildlife
Conservation of threatened primates in Dassioko Sud management in Central and West Africa. In: J. Abbott,
and Port Gauthier forest reserves: use of field patrols to F.G. Ananze, N. Barning, P. Burnham, E. de Merode,
assess primate abundance and illegal human activities. A. Dunn, E. Fuchi, E. Hakizumwami, C. Hesse, R.
Am. J. Phys. Anthropol. Suppl. 58: 127. (Abstract) Mwinyihali, M. Mahaman Sani, D. Thomas, P. Trench, and
Koné, I. (2015). Community empowerment for the R. Tshombe (eds.), Promoting Partnerships: Managing
conservation of Critically Endangered primates and Wildlife Resources in Central and West Africa, pp. 9–49.
their habitat in south-eastern Côte d’Ivoire. In: The IUCN London, UK: International Institute for Environment and
Programme for African Protected Areas & Conservation Development (IIED) Publications.
(ed.), Community Empowerment for the Conservation WAPCA (2014). Annual Report. Accra, Ghana: West African
of Critically Endangered Primates and Their Habitat Primate Conservation Action (WAPCA).
in South-eastern Côte d’Ivoire. Twenty-two Stories of Wiafe, E.D. (2013). Status of the Critically Endangered
Conservation in Africa: Key Elements for Effective and roloway monkey (Cercopithecus diana roloway) in the
Well-Governed Protected Areas in Sub-Saharan Africa, Dadieso Forest Reserve, Ghana. Afr. Primates 8: 9–15.
pp. 36–39. IUCN Programme for African Protected Areas Wiafe, E.D. (2021). Distribution and population density
& Conservation (PAPACO). of endangered primate species in Ghana’s forest
Koné, I. and Akpatou, B.K. (2005). Recherche en Côte reserves. Forestist 71(3): 238–247. doi.10.5152/
d’Ivoire de trois singes gravement menacés d’extinction. forestist.2021.20040
CEPA Magazine 12: 11–13.
Koné, I., Oates, J.F., Dempsey, A., Gonedelé Bi, S.,
McGraw, S. and Wiafe, E. (2019). Cercopithecus roloway.
The IUCN Red List of Threatened Species 2019:
e.T4232A92384429.
Koné, I., Koffi, D.A., and Bogui, E.B. (2021). The Tanoé-Ehy
Forest Conservation Programme, Côte d’Ivoire: Research

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© Rachel Ashegbofe Ikemeh

48
 NIGER DELTA RED COLOBUS
Piliocolobus epieni (Grubb and Powell, 1999)

Nigeria
(2008, 2010, 2016, 2018, 2022)

Rachel Ashegbofe Ikemeh, John F. Oates and Tukuru Seibokuro

The Niger Delta Red Colobus (Piliocolobus in the mid-1990s, this red colobus was locally
epieni) is endemic to the marsh forests in the common, especially in forests near the town of
central part of the Niger Delta of Nigeria (Oates Gbanraun, but it was beginning to come under
2011). Its species name is derived from its name intense pressure from degradation of its habitat
in the language of the Ijaw people who inhabit and commercial hunting. Important colobus
the limited area of about 1,500 km2 in Bayelsa food trees, especially Fleroya ledermannii, were
State where it occurs. Piliocolobus epieni being felled at a high rate by artisanal loggers,
only became known to science in 1993, in the and the logs were floated out of the delta on rafts
course of a biodiversity survey co-ordinated to processing centers in Lagos and elsewhere. In
by C. Bruce Powell (Blench 2007). Studies of addition, large canals dug as part of oil extraction
vocalizations and mitochondrial DNA suggest activities, as well as smaller canals dug by loggers
that this population is not closely related to its into the interior swamps, were changing local
closest relatives geographically, the Bioko Red hydrology (Werre and Powell 1997, Grubb and
Colobus (Piliocolobus pennantii) and Preuss’s Powell 1999). The Ijaw people are traditionally
Red Colobus (Piliocolobus preussi) of eastern fishermen, but outside influences introduced by
Nigeria and western Cameroon, leading Ting the oil industry have encouraged commercial
(2008) to treat this monkey not as a subspecies of bushmeat hunting and logging throughout the
P. pennantii (see Groves 2001, Grubb et al. 2003) Niger Delta.
but as a distinct species. Groves (2007) regarded
almost all the different red colobus monkeys, The most recent range-wide assessment of P.
including epieni, pennantii and preussi, as epieni, conducted in 2013, suggests that as a
separate species in the genus Piliocolobus – a result of habitat destruction and hunting the
taxonomy that we follow here. The IUCN Red population has declined significantly since the
List of Threatened Species has regarded P. epieni 1990s, and that it may now be around 90% lower
as Critically Endangered since 2008 (Ikemeh et al. than estimated 25 years ago (Ikemeh 2015). In
2019). the 2013 survey, the presence of P. epieni was
confirmed only in four forest areas, and it was
The marsh forests where the Niger Delta Red considered extirpated from 11 other forests
Colobus is found have a high water table all year where it had been reported in the 1990s by Werre
round but do not suffer deep flooding or tidal (2000). Cumulative survey data indicate that the
effects. The most intensive ecological study of current number of individuals surviving in the
this monkey, by Lodjewijk Werre in 1994–1996 wild may be only a few hundred (Ikemeh 2015).
suggested that the clumped distribution of food Insecurity in the region and the consequences
species in the marsh forest is a key factor restricting of poor governance, amongst other factors,
P. epieni to its limited range, which is demarcated have exacerbated the major threats of habitat
by the Forcados River and Bomadi Creek in the degradation and commercial hunting. Because
northwest, the Sagbama, Osiama and Apoi red colobus monkeys are known to be sensitive
creeks in the east, and the mangrove belt to the to habitat disturbance and hunting in other parts
south (Werre 2000). At the time of its discovery of Africa (Struhsaker 2005), it is feared that the

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AFRICA

Niger Delta Red Colobus, with its very restricted

range, is at high risk of extinction.

The red colobus monkeys are probably more

threatened than any other taxonomic group of
primates in Africa (Oates 1996, Struhsaker 2005).
Piliocolobus preussi (western Cameroon and
eastern Nigeria), Piliocolobus pennantii (Bioko
Island, Equatorial Guinea) and Piliocolobus
rufomitratus (Tana River, Kenya) are all regarded
as Critically Endangered from different
combinations of habitat loss and hunting, while
Piliocolobus waldroni (eastern Côte d’Ivoire and
western Ghana) may already be extinct (Oates
2011).

The Niger Delta Red Colobus shares its marsh

forest habitat with two other threatened
primates; the Nigerian White-throated Guenon
(Cercopithecus erythrogaster pococki) and the
Red-capped Mangabey (Cercocebus torquatus)
(Ikemeh 2015). Also found in these forests are the
Putty-nosed Monkey (Cercopithecus nictitans),
the Mona Monkey (Cercopithecus mona) and
the Olive Colobus (Procolobus verus) (Efenakpo
et al. 2018). However, political instability in the
delta, related in the most part to disputes over
the allocation of oil revenues, has hindered
progress in biodiversity conservation during the
last decade (Ikemeh 2015).

The two most important remaining areas for P.

epieni conservation have been identified as the
Apoi Creek Forests, flanked by the communities
of Gbanraun, Apoi and Kokologbene, and forests
near Kolotoro (Ikemeh 2015). However, in the
absence of conservation intervention, the Otolo-
Kolotoro-Ongoloba area has been devastated
by excessive logging, driving the population
in that area to the edge of extinction. In 2021,
a community-based conservation area was
established covering 1,109 ha in Apoi Creek
Forest. This area is under the customary authority
of the Apoi community, and protects 150–300
Niger Delta Red Colobus Monkeys, possibly the
most significant population anywhere. The effort
to create a community conservancy has been
coordinated by a local conservation group – the
Southwest Niger Delta Forest Project. Further
priority actions needed to secure the survival of this
species include establishing appropriate laws at
the state and national level to protect the species

50
and its habitat and conducting new surveys to
identify sites where other viable populations still
exist and where additional protected areas might
be created. Finally, efforts need to continue to
increase local level awareness on the ecological
importance of the species and its habitat.

Blench, R. (2007). Mammals of the Niger Delta, Nigeria.

Port Harcourt, Nigeria: Kay Williamson Education
Foundation.
Efenakpo, O., Ijeomah, H. and Ayodele, A.I. (2018). Threat
and Conservation of Wildlife Resources in Niger Delta
Region of Nigeria. In: Proceedings of 2nd Wildlife Society
of Nigeria conference, Akure, Nigeria (pp. 259–267).
Akure, Nigeria.
Groves, C.P. (2001). Primate Taxonomy. Washington, DC:
Smithsonian Institution Press.
Groves, C.P. (2007). The taxonomic diversity of the Colobinae
of Africa. J. Anthropol. Sci. 85: 7–34.
Grubb, P. and Powell, C.B. (1999). Discovery of red colobus
monkeys (Procolobus badius) in the Niger delta with
the description of a new and geographically isolated
subspecies. J. Zool., Lond. 248: 67–73.
Grubb, P., Butynski, T.M., Oates, J.F., Bearder, S.K., Disotell,
T.R., Groves, C.P. and Struhsaker, T.T. (2003). Assessment
of the diversity of African primates. Int. J. Primatol. 24(6):
1301–1357.
Ikemeh, R.A. (2015). Assessing the population status of
the Critically Endangered Niger Delta Red Colobus
(Piliocolobus epieni). Primate Conserv. (29): 1–10.
Ikemeh, R.A., Oates, J.F. and Inaoyom, I. (2019). Piliocolobus
epieni. The IUCN Red List of Threatened Species 2019:
e.T41024A92656391.
Oates J.F. (1996). African Primates: Status Survey and
Conservation Action Plan. Revised Edition. Washington,
D.C.: IUCN/SSC Primate Specialist Group.
Oates, J.F. (2011). Primates of West Africa: A Field Guide
and Natural History. Arlington, VA: Conservation
International.
Struhsaker, T.T. Conservation of red colobus and their
habitats. Int. J. Primatol. 26: 525–538.
Ting, N. (2008). Molecular Systematics of Red Colobus
Monkeys (Procolobus [Piliocolobus]): Understanding
the Evolution of an Endangered Primate. Doctoral
dissertation, City University of New York, New York, NY.
Werre, J.L.R. (2000). Ecology and Behavior of the Niger
Delta Red Colobus Monkey (Procolobus badius epieni).
Doctoral dissertation, City University of New York, New
York, NY.
Werre, J.L.R. and Powell, C.B. (1997). The Niger Delta
colobus – discovered in 1993 and now in danger of
extinction. Oryx 31: 7–9.

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© Ekwoge Abwe

52
 NIGERIA-CAMEROON
CHIMPANZEE
Pan troglodytes ellioti (Matschie,1914)

Cameroon, Nigeria
(2022)

Ekwoge Abwe, Osiris Doumbé, Paul Dutton, Chefor Fotang, Mary K. Gonder, Rachel
Ashegbofe Ikemeh, Inaoyom Imong, Serge A. Kamgang, Bethan J. Morgan and
Amadou Ngouh

The Nigeria-Cameroon Chimpanzee (Pan Djerem National Park (at least 500 individuals;
troglodytes ellioti), until recently known as P. t. Kamgang et al. 2020). Recent genetic studies
vellerosus, ranges from Cameroon, west of the especially in the mountainous region along the
Sanaga River, to Nigeria (Gonder et al. 2006, Nigeria-Cameroon border and south of the
Oates et al. 2009). Pan t. ellioti has two distinct Adamawa Plateau in Central Cameroon estimate
genetic groups, one associated with lowland and the current effective population size of P. t. ellioti
mountainous rainforest in west Cameroon and in Cameroon at 3,000–4,500 individuals (Mitchell
east Nigeria, and the second with the forest- et al. 2015b).
woodland-savannah mosaic in central Cameroon
(Mitchell et al. 2015a). It inhabits primary and The Nigeria-Cameroon Chimpanzee is classified
secondary moist lowland forest, montane and as Endangered on the IUCN Red List of Threatened
submontane forest, dry forest, gallery forest in Species based on an inferred population size
savanna woodland, and farmland, and spans an reduction of 50% over a three-generation period
altitudinal range from sea level to 2000 m (Oates from the mid-1980s to 2060 (Oates et al. 2016).
2011, Sesink Clee et al. 2015, Abwe et al. 2020). Within its larger range, P. t. ellioti is most seriously
threatened in two subregions: southwestern
Pan t. ellioti has the smallest geographic Nigeria and northwestern Cameroon. In each of
range and is the least numerous subspecies these subregions, total chimpanzee population
of chimpanzee with a total population that is numbers are very small (probably less than 250),
almost certainly less than 9,000 individuals and suitable habitat is highly fragmented, and hunting
probably less than 6,000 remaining in the wild pressure is intense, and population declines
(Morgan et al. 2011). One of the largest, and exceeding 80% are likely in the 1985–2060 period
probably most secure, subpopulations of P. (Greengrass 2009, Ikemeh 2013, Fotang et al.
t. ellioti is in Gashaka-Gumti National Park in 2021a). The reduction in P. t. ellioti populations
Nigeria, which has an estimated subpopulation is due to increasing anthropogenic pressure
of 900–1,000 individuals (Ogunjemite and exacerbated by human population growth in
Ashimi 2010, Adanu et al. 2011). Other major Cameroon and Nigeria, and linked with poaching
subpopulations are found in Cameroon in the (for bushmeat, traditional medicine, and the pet
Banyang-Mbo Wildlife Sanctuary (estimated at trade) and habitat loss from farming, logging,
500–900 or 800–1,450 individuals, depending fire, and commercial plantations (Morgan et al.
on the nest-decay parameters used; Greengrass 2011). Analysis of the viability of this subspecies
and Maisels 2007), in the Ebo forest (estimated in Cameroon and Nigeria through extinction-
at 626–1,480 individuals; M.S. Ndimbe and B.J. risk modeling has shown that the species could
Morgan pers. comm. 2015), and in Mbam and become extinct in 2035 if appropriate conservation

53
AFRICA

measures are not initiated and implemented

(Hughes et al. 2011). Climate change is expected
to further shrink P. t. ellioti habitat in the forest-
woodland-savannah mosaic over the coming
century (Sesink Clee et al. 2015). Since 2018,
political instability, especially in Cameroon’s
Northwest and Southwest Regions and along the
entire boundary with Nigeria, has stalled research
and conservation efforts in the central area of the
P. t. ellioti range. The proliferation of arms and
ammunition and local communities taking refuge
in the forest could jeopardize not only fragile
chimpanzee communities (Mbembe forest, Kom-
Wum Forest Reserve) but also well-established
populations, including in Korup National Park
and Banyang-Mbo Wildlife Sanctuary. The
species has already disappeared from some
sites such as the Bafut-Ngemba Forest Reserve,
Nkwende Hills (Bobo et al. 2013), South Bakundu
Forest Reserve (Eno-Nku 2004) in Cameroon and
the Ala, Oba Hills and Ogbesse Forest Reserves
in Nigeria (Greengrass 2006).

Pan t. ellioti occurs in several important

protected areas, including the Gashaka-Gumti
National Park, Cross River National Park and
Afi Mountain Wildlife Sanctuary in Nigeria,
and Mbam and Djerem National Park, Korup
National Park, Takamanda National Park, Mount
Cameroon National Park, and Banyang-Mbo
Wildlife Sanctuary in Cameroon (Morgan et al.
2011). More recently, the presence of a viable
population of Nigeria-Cameroon chimpanzee
was confirmed in Mpem and Djim National Park, in
the forest-woodland-savannah mosaic of central
Cameroon (O. Doumbé and A. Ngouh unpubl.
data 2021). Management effectiveness varies
across protected areas; poaching for bushmeat
occurs, in most of them, sometimes at high levels.
Biological surveys, including non-invasive sample
collection efforts for genetic studies throughout
much of the subspecies’ range in Cameroon,
have shown that many chimpanzees still survive
outside protected areas (Mitchell et al. 2015b).
One is the Yabassi Key Biodiversity Area which
encompasses the Ebo forest, where chimpanzee
and gorilla conservation efforts have been
underway since 2005 (Abwe and Morgan 2008).
Pan t. ellioti also survives in community-managed
forests such as the Mbe Mountains in Nigeria.

54
Like other chimpanzees, P. t. ellioti is omnivorous. Nigeria-Cameroon chimpanzee (Pan troglodytes ellioti).
Fruits (including figs) make up a large proportion Int. J. Primatol. 41(1): 81–104.
Adanu, J., Sommer, V. and Fowler, A. (2011). Hunters, fire,
of the diet, but leaves, bark, stems and animals cattle. Conservation challenges in eastern Nigeria, with
are also eaten, especially when fruits are scarce special reference to chimpanzees. In: V. Sommer and
(Dutton and Chapman 2015, Abwe et al. 2020). C. Ross (eds.), Primates of Gashaka: Socioecology and
Tools made from plant parts are used to extract Conservation in Nigeria’s Biodiversity Hotspot, pp. 55–
100. New York: Springer.
honey and also ants and termites from their nests Bobo, K., Ntumwel, B.C. and Aghomo, F.F.M. (2013). Local
(Fowler and Sommer 2007, Abwe and Morgan perception on wildlife uses and related loss of cultural
2008), stone and wooden hammers are used to values around the Nkwende Hills Forest Reserve, south-
crack nuts (Morgan and Abwe 2006), and monkeys west Cameroon. Life Sciences Leaflets. 5: 1–8.
and other mammals are probably captured for Dutton, P. and Chapman, H. (2015). Dietary preferences of
a submontane population of the rare Nigeria-Cameroon
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(Morgan et al. 2011). Conservation outreach Primatol. 27(4): 1103–1127.
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© Aug Aconk

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 JAVAN SLOW LORIS
Nycticebus javanicus É. Geoffroy Saint-Hilaire, 1812

Indonesia
(2008–2018, 2022)

K. Anna I. Nekaris and Vincent Nijman

Habitat loss and habitat degradation throughout Javan Slow Lorises are listed as Critically
Southeast Asia threaten all nine species of Endangered on the IUCN Red List of Threatened
slow and pygmy loris with extinction – Pygmy Species, thus here we use this species as the
Xanthonycticebus pygmaeus, Greater Nycticebus flagship for slow and pygmy loris conservation
coucang, Bengal N. bengalensis, Philippine N. (Nekaris et al. 2013b). Since being re-recognized
menagensis, Bornean N. borneanus, Kayan N. as a species by IUCN in 2006, work on the Javan
kayan, Sody’s N. bancanus, Sumatran N. hilleri, Slow Loris has increased and provides a sound
and Javan N. javanicus (Munds et al. 2013, Pozzi example of understanding and mitigating the
et al. 2014, Rowe and Myers 2016, Nekaris and threats to a highly threatened species.
Nijman 2022). Slow and pygmy lorises exhibit
numerous unique traits including slow life history, Capture to meet the demand for pets is the most
locomotion and digestion, and the ability to severe threat to the survival of Javan Slow Lorises.
enter torpor and hibernate, and they are the only Despite being legally protected in Indonesia
venomous primates (Nekaris 2014). Still, wild slow since 1973, with their striking coloration and their
lorises have seldom been studied for more than presence on Java, Indonesia’s commercial center,
a year (Cambodia X. pygmaeus, Starr et al. 2011; it is no wonder that Indonesian pet traders in the
Malaysia N. coucang, Wiens et al. 2006; India, N. 1990s targeted Javan Slow Lorises above other
bengalensis, Das et al. 2014), with only N. javanicus endemic slow loris species. Analysis of slow
being the focus of a long-term study (Nekaris lorises in the wildlife markets of Java and Bali
2016). Many researchers and conservationists shows that the overall number openly offered
have only ever seen a slow or pygmy loris in the for sale remains fairly constant, but the species
illegal wildlife trade, either dried on bamboo composition changed, with fewer Javan Slow
sticks in preparation for traditional medicine, Lorises counted (Nijman et al. 2017). Traders
paraded as a photo prop on a tourist beach, or claim that Javan Slow Lorises are increasingly
sold as a pet (Schulze and Groves 2004, Das et al. difficult to obtain and, as with other wildlife,
2009, Nijman et al. 2015, Osterberg and Nekaris trade has moved from animal markets only, to
2015). The extreme popularity of viral slow loris both animal markets as well as online trade.
internet videos is a double-edged sword, to some COVID-19 thus far has had limited effect on the
extent making the public aware of their decline, wildlife markets and trade in Javan Slow Lorises;
but also causing the public to think that they during lockdowns and times of regional travel
are not threatened (Nekaris et al. 2013a). The restrictions the number of visitors and vendors
level of international trade was sufficiently large may have dropped temporarily, but in general
for the genus Nycticebus (now Nycticebus and wildlife markets have remained open.
Xanthonycticebus) to be transferred to CITES
Appendix I in 2007 (Nekaris and Nijman 2007, Successful prosecution of lawbreakers buying
2022), meaning that all commercial international or selling slow lorises in Indonesia is a very
trade is prohibited. rare occurrence and despite hundreds of slow
lorises having been confiscated from traders
over the last decade, we are only aware of a

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handful of successful prosecutions (Nekaris and

Nijman 2018). Wildlife traders in Indonesia have
increasingly turned to social media to advertise
their illegal stock, including Javan Slow Lorises.
The huge rise in Facebook, Instagram, and
WhatsApp means that many are sold via social
media without ever being seen in a wildlife
market. An ongoing online monitoring program
by the Little Fireface Project suggests that dozens
of live Javan Slow Lorises are offered for sale in
online forums. This monitoring revealed that slow
lorises in Indonesia are also traded for medicinal
purposes, mainly in the form of slow loris oil, and
Javan Slow Lorises are one of three species most
affected by this trade (Nekaris et al. 2020).

Slow lorises are the only venomous primates and,

amongst other functions, use their venom for
interspecific competition (Nekaris et al. 2020). To
avoid being bitten by the venomous slow lorises,
traders habitually cut or pull out an animal’s lower
front teeth prior to selling them (Nekaris et al.
2013c). Traders may also cut teeth prior to packing
slow lorises tightly into crates, as they often
damage each other with their venomous bites
during transport. Fuller et al. (2018) showed that
following confiscations and during rehabilitation,
nearly 30% die in the first 6 months, with
morbidity from wounds (mainly bites) being the
main cause. Other causes of death due to dental
removal include dental abscess or pneumonia
(Nekaris and Starr 2015). Those that do survive
are no longer able to eat their preferred food
(gum) (Das et al. 2014) or engage in the important
behavior of social grooming with the toothcomb,
meaning that any confiscated animals are unlikely
to survive if released to the wild.

Reintroduction itself is a threat to the Javan Slow

Loris. In the major markets in Java, at least four
other slow loris species from Borneo and Sumatra
are traded alongside the Javan Slow Loris, and in
the markets in Sumatra at least three species are
regularly traded, including Javan Slow Lorises.
Likewise, traders based in Java and Sumatra
offer several species for sale, including ones that
do not occur naturally on the island where the
traders reside. The similar appearance of slow
lorises, to the untrained eye, results in release
of slow loris species from Sumatra and Borneo
into Java, and Javan Slow Lorises in Sumatra
and Borneo, causing potential for hybridization

62
or even displacement of native species by dependent on local people for their protection,
introduced ones (Nekaris and Starr 2015). The feeding on human-planted tree species and
ability of slow lorises to persist in human habitat residing in farmland. Thus, a major conservation
if left undisturbed means that well-meaning program combining empowerment activities,
people may translocate animals to habitat that conservation education, and village events has
is unknown to the animals, exacerbating these been launched, and it is hoped that it can be
problems (Kumar et al. 2014). used as a model for other key slow loris sites in
Indonesia (Nekaris and Starr 2015). One such
Moore et al. (2014) assessed the success of Javan program involved building bridges for slow
Slow Loris reintroductions, finding a death rate of lorises to connect habitat. Most of these bridges
up to 90%. Illness, hypothermia and exhaustion are made of water hose pipes that also help to
were all implicated in the death of slow lorises. irrigate the land of farmers. This program was
Reintroductions were started before the basics accompanied by an intensive education program
were known about the Javan Slow Loris’ behavior, for children and their parents, making people
ecology, or distribution. No habitat suitability aware of the lorises who live on their land (Birot
assessment could be made, since details were et al. 2020, Nekaris et al. 2021). This program
lacking on the type of habitat the species has been reproduced by other loris conservation
preferred and what it avoided. It has recently projects in Bangladesh and in Thailand, where
been reported by rescue centers that the success artificial bridges have also been used successfully
rates of Javan Slow Loris reintroductions are by N. bengalensis. Because many slow loris
improving, but unfortunately no published data species rely on non-protected farmland, helping
are available to verify these claims. Newspaper local people to understand their important
reports show that up to 30 slow lorises are role in pollination and pest control can be vital
released in one site at one time, but the highly for populations to persist. With this in mind,
territorial and venomous nature of slow lorises extending on the building bridges program, the
means that such releases are destined to have a Little Fireface Project also developed an eco-
high failure rate. A related study of pygmy lorises friendly coffee program, with cooperatives of
in Vietnam found that the season of release farmers achieving Wildlife Friendly Certification.
and age should be considered to increase the The certification involved a total hunting ban in
likelihood of survival (Kenyon et al. 2014). the area, as well as significant reforestation of
endemic tree species (Campera et al. 2021).
To obtain vital information on the Javan Slow
Lorises, in 2011 the Little Fireface Project For a long time, slow lorises were thought to be
instigated a study of the species’ behavioral common throughout Indonesia, and the presence
ecology in Garut District of West Java, Indonesia of animals in trade was believed to be an indicator
(Nekaris 2016). This multi-disciplinary project of their abundance. We are only beginning to
has obtained data on home range size, social unravel the complexity of their taxonomy and
organization, infant dispersal, and feeding distribution, leading to a bleak overall picture.
ecology. It was found that both sexes disperse While Java has an impressive and comprehensive
from their natal range at about 20 months old, protected area network, encompassing over 120
dispersal distances are 1–3 km from the natal terrestrial conservation areas and covering 5,000
range, home range sizes are large relative to the km², enforcement of environmental laws and
size of the animal (5–10 ha), the species goes active protection in most of these parks is lacking
into torpor, and the diet comprises mainly gum, and, as indicated above, many of the Javan Slow
supplemented with nectar and insects (Cabana Lorises are found outside the protected area
et al. 2017, 2019). Several initiatives have been network. Besides curbing the illegal trade, it is
put into place to conserve slow lorises in the area paramount that these conservation areas, and
and in other parts of Java. National workshops indeed all other remaining forest areas on the
have been held for law enforcement officers and island, are effectively protected.
rescue center employees to provide essential
data for a national slow loris action plan. At
the local level, slow lorises are often totally

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Birot, H., Campera, M., Imron, M.A. and Nekaris, K.A.I. Imron, M.A. (2020). Slow lorises use venom as a weapon in
(2020). Artificial canopy bridges improve connectivity in intraspecific competition. Curr. Biol. 30(20): 1252–1253.
fragmented landscapes: the case of Javan slow lorises Nekaris, K.A.I., Handby, V. and Campera, M. (2021). Impact
in an agroforest environment. Am. J. Primatol. 82(4): of weather conditions, seasonality and moonlight on the
23076. use of artificial canopy bridges by nocturnal arboreal
Cabana, F., Dierenfeld, E., Wirdateti, W., Donati, G. and mammals. Biodiv. Conserv. 30(12): 3633–3645.
Nekaris, K.A.I. (2017). The seasonal feeding ecology of Nekaris, K.A.I., Moore, R.S., Rode, E.J. and Fry, B.G. (2013c).
the Javan slow loris (Nycticebus javanicus). Am. J. Phys. Mad, bad, and dangerous to know: the only venomous
Anthropol. 162: 768–781. primates, slow lorises. J. Venom. Anim. Toxins Incl. Trop.
Cabana, F., Clayton, J.B., Nekaris, K.A.I., Wirdateti, W., Dis. 19: 21.
Knights, D. and Seedorf, H. (2019). Nutrient-based diet Nekaris, K.A.I. and Nijman, V. (2007). CITES proposal
modifications impact on the gut microbiome of the Javan highlights rarity of Asian nocturnal primates (Lorisidae:
slow loris (Nycticebus javanicus). Sci. Rep. 9(1): 1–11. Nycticebus). Folia Primatol. 78: 211–214.
Campera, M., Budiadi, B., Adinda, E., Ahmad, N., Balestri, Nekaris, K.A.I. and Nijman, V. (2018). Successful prosecution
M., Hedger, K., Imron, M.A., Manson, S., Nijman, V. of slow loris traders in Indonesia. Oryx 52(3): 411–411.
and Nekaris, K.A.I. (2021). Fostering a wildlife-friendly Nekaris, K.A.I. and Nijman, V. (2022). A new genus name for
program for sustainable coffee farming: the case of pygmy lorises, Xanthonycticebus gen. nov. (Mammalia,
small-holder farmers in Indonesia. Land 10(2): 121. Primates). Zoosyst. Evol. 98: 87–92.
Das, N., Biswas, J., Das, J., Ray, P.C., Sangma, A. and Nekaris, K.A.I., Shekelle, M., Wirdateti, W., Rode, E.J. and
Bhattacharjee, P.C. (2009). Status of Bengal slow loris Nijman, V. (2013b). Nycticebus javanicus. The IUCN Red
Nycticebus bengalensis (Primates: Lorisidae) in Gibbon List of Threatened Species 2013. doi.org/10.2305/IUCN.
Wildlife Sanctuary, Assam, India. J. Threat. Taxa. 1: 558– UK.2013-2.RLTS.T39761A17971158.en
561. Nekaris, K.A.I. and Starr, C.R. (2015). Conservation and
Das, N., Nekaris, K.A.I. and Bhattacharjee, P.C. (2014). ecology of the neglected slow loris: priorities and
Medicinal plant exudativory by the Bengal slow loris prospects. Endang. Species Res. 28: 87–95.
Nycticebus bengalensis. Endang. Species Res. 23: 149– Nijman, V., Shepherd, C.R. and Nekaris, K.A.I. (2015). Trade
157. in Bengal slow lorises in Mong La, Myanmar, on the China
Fuller, G., Eggen, W.F., Wirdateti, W. and Nekaris, K.A.I. border. Primate Conserv. (28): 139-142.
(2018). Welfare impacts of the illegal wildlife trade in a Nijman, V., Spaan, D., Rode-Margono, E.J. and Nekaris,
cohort of confiscated greater slow lorises, Nycticebus K.A.I. (2017). Changes in the primate trade in Indonesian
coucang. J. Appl. Anim. Welf. Sci. 21(3): 224–238. wildlife markets over a 25-year period: Fewer apes and
Kenyon, M., Streicher, U., Loung, H., Tran, T., Tran, M., Vo, langurs, more macaques and slow lorises. Am. J. Primatol.
B. and Cronin, A. (2014). Survival of reintroduced pygmy 79(11): 22517.
slow loris Nycticebus pygmaeus in South Vietnam. Osterberg, P. and Nekaris, K.A.I. (2015). The conservation
Endang. Species Res. 25: 185–195. implications of the use of photo prop animals for tourists
Kumar, A., Sarma, K., Panvor, J., Mazumdar, K., Devi, A., in Thailand: a slow loris case study (Nycticebus spp.).
Krishna, M. and Ray, P.C. (2014). Threats to the Bengal TRAFFIC Bulletin 27: 13–18.
slow loris Nycticebus bengalensis in and around Pozzi, L., Hodgson, J.A., Burrell, A.S., Sterner, K.N., Raaum,
Itanagar Wildlife Sanctuary, Arunachal Pradesh, India: R.L. and Disotell, T.R. (2014). Primate phylogenetic
impediments to conservation. Endang. Species Res. 23: relationships and divergence dates inferred from
99–106. complete mitochondrial genomes. Mol. Phylogenet.
Moore, R., Wihermanto, S. and Nekaris, K.A.I. (2014). Evol. 75: 165–183.
Compassionate conservation, rehabilitation and Rowe, N. and Myers, M. (2016). All the World’s Primates.
translocation of Indonesian slow lorises. Endang. Species Charlestown, RI: Pogonias Press.
Res. 26: 93–102. Schulze, H. and Groves, C.P. (2004). Asian lorises: taxonomic
Munds, R.A., Nekaris, K.A.I. and Ford, S.M. (2013). Taxonomy problems caused by illegal trade. In: T. Nadler, U. Streicher
of the Bornean slow loris, with new species Nycticebus and Ha Thang Long (eds.), Conservation of Primates
kayan (Primates, Lorisidae). Am. J. Primatol. 75: 46–56. in Vietnam, pp. 33–36. Frankfurt, Germany: Frankfurt
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evolution of Asian lorises. Evol. Anthropol: 23: 177– Starr, C., Nekaris, K.A.I., Streicher, U. and Leung, L.K.P.
187. (2011). Field surveys of the vulnerable pygmy slow
Nekaris, K.A.I. (2016). The Little Fireface Project: community loris Nycticebus pygmaeus using local knowledge in
conservation of Asia’s slow lorises via ecology, Mondulkiri Province, Cambodia. Oryx 45: 135–142.
education, and empowerment. In: M.T. Waller (ed.), Wiens, F., Zitzmann, A. and Hussein, N.A. (2006). Fast food
Ethnoprimatology, pp. 259–272. New York: Springer. for slow lorises: is low metabolism related to secondary
Nekaris, K.A.I., Campbell, N., Coggins, T.G., Rode, E.J. and compounds in high-energy plant diet? J. Mammal. 87:
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Margono, E.J., Fry, B.G., Weldon, A., Wirdateti, W. and

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© Myron Shekelle

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 SANGIHE TARSIER
Tarsius sangirensis Meyer, 1897

Indonesia
(2022)

Myron Shekelle

Sometime ago in the distant past, a small family highest value conservation habitats are on the
of tarsiers found itself trapped on a floating southern end of Sangihe (Whitten, 2006), in the
island of vegetation, tossed about on the Pacific gold mine concession.
Ocean (Shekelle et al. 2013). Drifting northward
from their home on Sulawesi, they travelled for Sangihe Island is famous to bird conservationists
days, up to 200 km from where their journey for its highly threatened avifauna. The Cerulean
began, their life raft touched on the tiny island Flycatcher (Eutrichomyias rowleyi) graced the
of Sangihe. Sangihe Island is an active volcano first cover of the journal, Conservation Biology,
that pushed upward from the ocean floor several accompanying an article concerned that it
thousands of meters below the surface. Thus, had gone extinct (see Whitten 2006). When its
like the more famous island chains of Hawaii rediscovery in the wild was announced in 2006, its
and Galapagos, all life found on Sangihe Island conservation hinged on a 940-ha patch of mixed
arrived as wandering vagabonds, just as these primary and secondary forest that had been zoned
tarsiers did. For a million years or more, these by the community as a water catchment, a level of
tarsiers lived in isolation, evolving into the species protection so limited that it did not even prohibit
Tarsius sangirensis. hunting (Whitten 2006). In an amazing contrast
to the near disregard of this forest by anyone
Isolated no more, Sangihe Island has the bustling other than the local community, it was known to
port city of Tahuna, home to 33,000 people, shelter three species of Critically Endangered
an airport, and many of the conveniences birds – no other place in Indonesia has more than
of industrialized life. The island, of 547 km² one Critically Endangered bird species (Whitten
(Shekelle and Salim 2009), is about one-fifth the 2006). In addition to the Cerulean Flycatcher, the
size of Luxembourg. With a human population Sangihe Shrike-Thrush (Coracornis sanghirensis)
of 140,000 (Badan Pusat Statistik, 2022) crowded and the Sangihe White-eye (Zosterops nehrkorni)
into 255 people per km², the island would rank are also Sangihe Island endemics (Whitten 2006).
tenth highest in terms of population density – All rely on that 940-ha patch of forest, a little less
just ahead of Luxembourg – were it a European than three times the size of New York’s Central
nation. Looked at another way, Sangihe Island Park, now within the gold mine concession.
is slightly smaller than the Isle of Man, but its
population is about 175% larger. We can add to this list a small primate, Tarsius
sangirensis. It is listed as Endangered on the
The northern half of Sangihe Island is dominated IUCN Red List of Threatened Species (Shekelle
by the volcano, Mt. Awu, ranked as the fourth 2020). The presence of the gold mine across the
most deadly volcano in Indonesia’s “Ring of southern half of the island, and the loss of the
Fire” (Bani et al. 2020). In 2021, a concession local community’s ability to manage their own
for a gold mine was ceded on the southern half forests, has greatly changed the threat status
of the island. The BBC named Sangihe Island, to the Sangihe Island Tarsier, and to all of the
“Indonesia’s Gold Island”, an island made of gold endemic biodiversity of the Island. Shekelle and
(BBC, 2021). Unfortunately, as far as is known, the Salim (2009) found that the Sangihe Island Tarsier

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was at risk from a small extent of occurrence and

area of occupancy, small population size, high
risk of volcanism, high human population density,
fragmented populations (many of which are in
marginal habitat), and lack of conservation areas.
There are no ex situ conservation options for any
tarsier species, should their extinction in the wild
become imminent.

Local people have protested the gold mine, and

one anti-mine government official died under
conditions that some find suspicious. They have
created a petition on Change.Org that has nearly
150,000 signatures. If it is determined that the
gold mining company, PT. Tambang Mas Sangihe
(TMS), the Indonesian company registered by
BARU Gold Corporation, has a legal right to
mine gold on Sangihe Island, then that must be
respected. Listing Tarsius sangirensis among the
World’s 25 Most Endangered Primates tells the
gold mine that the eyes of the world are focused
on its management of the biodiversity that it has
been entrusted by Indonesian mining laws to
preserve.

BBC. (2021). The Battle for Indonesia’s Gold Island. BBC

News Video. Available online: <https://www.bbc.co.uk/
news/av/world-asia-57902815>.
Badan Pusat Statistik. (2022). Available online: <https://www.
bps.go.id/>.
Bani, P., Kristianto, Kunrat, S. and Syahbana D.K. (2020).
Insights into the recurrent energetic eruptions that drive
Awu, among the deadliest volcanoes on Earth. Nat.
Hazards Earth Syst. Sci. 20: 2119–2312.
Shekelle, M. (2020). Tarsius sangirensiss. The IUCN Red List of
Threatened Species 2020: e.T21493A17977351.
Shekelle, M. and Salim, A. (2009). An acute conservation
threat to two tarsier species in the Sangihe Island chain,
North Sulawesi, Indonesia. Oryx 43: 419–426.
Shekelle, M., Gursky-Doyen, S. and Richardson, M.C.
(2013). Family: Tarsiidae (Tarsiers). In: R.A. Mittermeier,
A.B. Rylands and D.E. Wilson (eds.), Handbook of the
Mammals of the World. Volume 3. Primates, pp. 236–261.
Barcelona: Lynx Edicions.
Whitten, T. (2006). Cerulean paradise-flycatcher not extinct:
subject of first cover lives. Conserv. Biol. 20: 918–920.

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© Rasanayagam Rudran

70
 PURPLE-FACED LANGUR
Semnopithecus vetulus (Erxleben, 1777)

Sri Lanka
S. v. nestor (2004–2022)
S. v. vetulus, S. v. monticola, S. v. philbricki (2018–2022)

Rasanayagam Rudran

Extensive deforestation occurred following subspecies, S. v. nestor, is Critically Endangered

the cessation of Sri Lanka’s 26-year civil war in (Rudran et al. 2020a). The range of the Western
2009, escalating human-primate conflicts and Purple-faced Langur (Semnopithecus vetulus
undermining the long-term survival of three Sri nestor) includes the most densely populated
Lankan primates, Semnopithecus vetulus, S. region around Colombo, the country’s capital.
priam thersites and Macaca sinica that are not Urbanization poses a serious threat, therefore,
only endemic, but also threatened with extinction. to the long-term survival of this Critically
As public outcry and political pressure mounted Endangered and endemic subspecies (Rudran
to resolve these conflicts, several government et al. 2009, 2020a). A survey conducted in 2007
institutions and non-governmental organizations, (Rudran 2007) indicated that 81% of S. v. nestor’s
led by SPEARS Foundation, helped the country’s historical range (Hill 1934, Phillips 1981) had been
Department of Wildlife Conservation (DWC) to deforested and converted to human altered
develop an action plan for people to conserve landscapes. Due to this habitat reduction, much
and coexist with all species of monkeys. The plan of S. v. nestor’s current population subsists mainly
was submitted to the country’s government in on fruit from home gardens (Dela 2007, Rudran
March 2016 for cabinet approval. 2007). Nutritional consequences of feeding on a
low diversity diet of cultivated fruits are unclear
While awaiting approval, SPEARS Foundation but considered detrimental to the folivorous S. v.
used funds from foreign donors to implement nestor (Rudran 2015).
some key elements of the plan. One was to
develop Community Conservation Areas (CCAs), Besides depleting natural food sources,
which, when established, would be administered deforestation causes habitat fragmentation,
and managed sustainably by local communities forcing animals to travel on the ground and along
under DWC supervision. To find suitable sites power lines to move between fragments. These
for CCAs, the SPEARS team analyzed complaints movements increase mortality by exposing
of human-monkey conflicts received by the DWC them to attacks by dogs, speeding vehicles, and
between 2007 and 2015. The analysis indicated electrocution (Parker et al. 2008). In some parts
that conflicts occurred throughout the country, of its range, S. v. nestor is occasionally shot and
but their frequency varied between localities killed as a pest while feeding in home gardens
(Cabral et al. 2018). Thirteen field surveys were (Dela 2004). Such human-induced fatalities
conducted, therefore, from 2016 to 2018, to reduce group sizes and appear to lead to local
locate sites best suited for the establishment of extinctions in S. v. nestor’s range (Rudran 2007).
CCAs. Information from these surveys and other
relevant data on all four Purple-faced Langur The Highland Purple-faced Langur (S. v.
subspecies are presented below. monticola), also known as the Bear Monkey, was
studied for two years at Horton Plains by Rudran
Three of the four subspecies of Semnopithecus (1973a, 1973b) nearly fifty years ago. When the
vetulus are Endangered. The fourth, the western area was surveyed again in 2016, Rudran noted

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appreciable changes to the vegetation. Many The Northern Purple-faced Langur (S. v.
species previously recorded as important food philbricki) was investigated for two years in the
plants of the Bear Monkey were dead or dying. late 1960s (Rudran 1973a, 1973b) when conflicts
This appeared to be primarily due to debarking with humans were not a serious issue. In the late
of the adult trees and consumption of saplings 1970s, however, the impact of the Accelerated
by the sambar (Cervus unicolor) population, Mahaweli Development Program (AMDP) on
which had increased in numbers because of the wildlife in S. v. philbricki’s range became a
invasive soft grass introduced to Horton Plains serious concern. To mitigate this concern, an
with the fertilizer used by a now defunct potato Environmental Impact Assessment (EIA) of the
farm (Adikaram et al. 1999). The death and lack AMDP recommended the establishment of four
of regeneration of food plants appear to have new national parks around the development area
undermined Bear Monkey survival. A census (Tippetts-Abbett-McCarthy-Stratton 1980). While
was not conducted in 2016, but early morning these areas provided protection to S. v. philbricki,
loud calls of harem males were considerably less serious threats such as habitat fragmentation and
frequent than before, indicating a population hunting for food, medicinal purposes and rituals
decline. The area was surveyed again in 2017 still remained in other areas (Wickremasinghe et
to collect data on crop damage and human al. 2016). Similar findings have also been reported
attitudes towards monkeys. by other investigators (Nahallage and Huffman
2013). Two surveys conducted in 2018 by SPEARS
Three surveys were conducted in the range Foundation staff found that populations of the
of the Southern Purple-faced Langur (S. v. highly arboreal S. v. philbricki were fewer than
vetulus) in 2017. In addition, a long-term study that of the other two subspecies in the area due
of S. v. vetulus (Roscoe et al. 2013) reported to the fragmentation of their habitat.
several threats to the future survival of this
subspecies. These threats were the same as In the last couple of years, the COVID-19
those experienced by S. v. nestor. Additionally, a pandemic has hampered the SPEARS team’s
major highway constructed through S. v. vetulus’ field activities, though the pandemic did provide
range is expected to create a permanent barrier the team with opportunities to publish data that
to gene flow between the populations found it had on file, to produce video documentaries
along the coast and the interior of the country. about their efforts, and to develop a device

72
designed to reduce financial losses due to crop expected to travel extensively to assess the
damage by monkeys. damage before providing compensation. The
SPEARS Team, therefore, recommended that
In regards to publications published during local community organizations be tasked with
COVID-19, these were based on data that assessing the damage (Rudran et al. 2021). The
had been collected prior to the onset of recommendation also stressed that requests for
pandemic. One of the articles discussed an compensation payments should be submitted
ethno-primatological approach to conserving to wildlife authorities only with convincing and
the Critically Endangered Western Purple-faced verifiable supporting evidence. This arrangement
Langur (Rudran et al. 2020). This paper was the was recommended to help prevent bogus
outcome of a 14-year effort during which time claimsfor crop damage by monkeys.
the SPEARS team implemented annual outreach
activities with the aim of building support in The SPEARS team’s articles were published
rural communities, so that they would become in scientific journals not easily accessible to
staunch allies in promoting the conservation of people living in Sri Lanka. To stimulate local
the species. These efforts were managed by a interest about Sri Lanka’s monkeys and its
non-governmental organization, because the Sri natural habitats, the SPEARS team used video
Lankan government’s wildlife agency did not have footage taken before the pandemic to produce
adequate staff and financial resources to deal with nine presentations in English and the two local
critically important wildlife conservation issues. languages. Each presentation lasted anywhere
Donor agencies are urged, therefore, to provide from two to nearly 30 minutes, and introduced
long-term financial support to the country’s local people to the objectives of the SPEARS
non-governmental organizations that have team, the diversity of Sri Lankan monkeys, and
demonstrated their unwavering commitment to causes of human-monkey conflicts. Some
promote wildlife conservation. Such support may presentations also provided information on
also be useful in other countries that have rural interesting local species and important natural
communities and wildlife protection agencies habitats, such as the Malabar Pied Hornbill and
with characteristics similar to those in Sri Lanka. the country’s vanishing wetlands. These popular
video presentations were shown at several public
As another example, the SPEARS team meetings held by the SPEARS team before the
published research about the use of an pandemic took hold in Sri Lanka.
ethnoprimatological approach to mitigate Sri
Lanka’s human-monkey conflicts (Rudran et al. Another pre-pandemic project launched by the
2021). This problem had skyrocketed after 2009, SPEARS Team involved the development of a
when the end of Sri Lanka’s 26-year ethnic war motion sensing device designed to emit loud
precipitated extensive deforestation to expand alarm signals when monkeys enter home gardens
the country’s agricultural base and economy. To and croplands. When the alarm is activated,
assess the intensity of this country-wide problem, the homeowner was expected to use his/her
the SPEARS team conducted field surveys, cell phone to alert the crop protection brigade
for five days each month from January 2016 to (Rudran et al. 2021) composed of unemployed
June 2018, in 11 of Sri Lanka’s 25 administrative and underemployed youngsters, who have
districts. During these 30 months, the survey their own modes of conveyance and expertise
team held semi-structured interviews with more in using catapults and other non-lethal devices
than 1,600 adults, which were used to craft to chase away monkeys. When the monkeys
recommendations, including a recommendation are evicted, homeowners whose crops were
to establish a private insurance scheme or saved would compensate the crop protection
a government-sponsored compensation brigade. In this manner the crop protection
program to deal with humanmonkey conflicts program will not only help minimize financial
(Rudran et al. 2021). In addition, and since losses incurred by homeowners but also provide
human-monkey conflicts occurred throughout gainful employment to local youngsters. This
the country, officers of the understaffed and program is expected to become functional after
underfunded wildlife agency could not be

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the pandemic dies out and does not pose any Roscoe, C.J., De Silva, M.A., Hapuarachchi, N.C. and Rohan
further threats to human health. Krishantha, P.A. (2013). A new color morph of the southern
purple-faced langur (Semnopithecus vetulus vetulus)
from the rainforests of southwestern Sri Lanka. Primate
In conclusion, although Sri Lanka’s monkeys Conserv. (26): 115–124.
face a perilous future (Rudran 2013), there is Rudran, R. (1973a). The reproductive cycles of two subspecies
hope that they can be conserved. One reason of purple-faced langurs (Presbytis senex) with relation to
for hope is that most Sri Lankans follow the environmental factors. Folia Primatol. 19: 41–60.
Rudran, R. (1973b). Adult male replacement in one-male
Buddhist doctrine of compassion towards all troops of purple-faced langurs (Presbytis senex) and its
living things. Promoting this doctrine and effect on population structure. Folia Primatol. 19: 166–
Buddha’s own reverence of the forest presents, 192.
therefore, opportunities to deter deforestation Rudran, R. (2007). A survey of Sri Lanka’s endangered and
in a country steeped in cultural traditions but endemic western purple-faced langur (Trachypithecus
vetulus nestor). Primate Conserv. (22): 139–144.
ignorant of the detrimental effects of habitat Rudran, R. (2013). Purple-faced langur, Trachypithecus
destruction. Another reason for optimism stems vetulus. In: J.C. Daniel and N. Manjrekar (eds.), Mammals
from a decision by successive governments to of South Asia, pp. 317–331. Hyderabad, India: Universities
increase Sri Lanka’s forest cover from 27% to Press.
Rudran, R. (2015). Western purple-faced langur
36% using native plants, to achieve the country’s Semnopithecus vetulus nestor (Bennett, 1833). In: C.
economic development goals (Yatawara 2011). Schwitzer et al. (eds.), Primates in Peril: The World’s
The political will to increase forest cover augurs 25 Most Endangered Primates 2014–2016, pp. 63–66.
well for the future protection of wildlife. It Arlington, USA: IUCN SSC Primate Specialist Group (PSG),
is important that the Sri Lankan government International Primatological Society (IPS), Conservation
International (CI) and Bristol Zoological Society.
approves the 2016 action plan in order to ensure Rudran, R., Weerakoon, K. and Wanasinghe, A. 2009. Western
a steady flow of financial support to conserve Sri purple-faced langur Trachypithecus (Semnopithecus)
Lanka’s monkeys. vetulus nestor (Bennett, 1833). In: R.A. Mittermeier et
al. (eds.), The World’s 25 Most Endangered Primates
2008–2010, pp. 24–25. Arlington, USA: IUCN SSC Primate
Specialist Group (PSG), International Primatological
Society (IPS), Conservation International (CI) and Bristol
Adikaram, N.K.B., Ranawana, K.B., Dissanayake, C.B. Zoological Society.
and Ranasinghe, P.N. (1999). Forest dieback in Horton Rudran, R., Dittus, W., Gamage, S.N. and Nekaris, K.A.I.
Plains National Park. Project Terminal Report. Global (2020a). Semnopithecus vetulus. The IUCN Red List of
Environmental Facility Project of the Department of Threatened Species 2020: e.T22042A17959452.
Wildlife Conservation. Rudran, R., Cabral de Mel, S.J., Salindra, H.G., Dayananda,
Cabral, S.J., Sumanapala, A.P., Weerakoon, D.K., K., Ratnayake, R.M.R.R., Weerakkody, S. and de Mel, R.K.
Kotagama, S.W. and Rudran, R. (2018). Have habitat loss (2020b). An ethnoprimatological approach to conserving
and fragmentation affected the social organization of Sri Lanka’s critically endangered western purple-faced
the western purple-faced langur (Semnopithecus vetulus langur. Primate Conserv. (34): 227–233.
nestor)? Primate Conserv. (32): 159–166. Rudran, R., Cabral de Mel, S.J., Sumanapala, A., de Mel, R.K.
Dela, J.D.S. (2004). Protecting the endemic purple-faced and Mahindarathna K.K.T.I. (2021). An ethnoprimatological
langur. Loris 23: 14–22. approach to mitigating Sri Lanka’s human-monkey
Dela, J.D.S. (2007). Seasonal food use strategies of conflicts. Primate Conserv. (35): 189–198.
Semnopithecus vetulus nestor at Panadura and Tippetts-Abbett-McCarthy-Stratton (TAMS). (1980).
Piliyandala, Sri Lanka. Int. J. Primatol. 28: 607–626. Environmental Impact Assessment, Accelerated
Hill, W.C.O. (1934). A monograph on the purple-faced leaf Mahaweli Development Program. Volume II - Terrestrial
monkeys (Pithecus vetulus). Ceylon J. Sci. 29(1): 23–88. Environment. Ministry of Mahaweli Development,
Nahallage, C.A.D. and Huffman, M.A. (2013). Macaque- Democratic Socialist Republic of Sri Lanka.
human interactions in the past and present day Sri Lanka. Wickremasinghe, S., Unantenna, J.R.K.W. and Dissanayake,
In: S. Radhakrishna. M.A. Huffman and A. Sinha (eds.), D.S.B. (2016). Distribution of Trachypithecus vetulus
The Macaque Connection: Cooperation and Conflict philbricki in selected forest patches in the Anuradhapura
between Humans and Macaques, pp. 135–147. New District, Sri Lanka. 5th Asian Primate Symposium 18–22
York, NY: Springer. October 2016. Sri Jayawardeepura University, Sri Lanka.
Parker, L., Nijman, V. and Nekaris, K.A.I. (2008). When there Yatawara, S.B. (2011). Let’s preserve trees, before the last
is no forest left: fragmentation, local extinction, and small tree vanishes. Available online: <www.sundayobserver.
population sizes in the Sri Lankan western purple-faced lk/2011/11/06/fea05.asp>. Accessed 27 February 2014.
langur. Endang. Species. Res. 5: 29–36.
Phillips, W.W.A. (1981). Manual of the Mammals of Sri
Lanka. Part II. 2nd edition. Colombo, Sri Lanka: Wildlife
and Nature Protection Society.

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© Neahga Leonard

76
 GOLDEN-HEADED LANGUR
or Cat Ba Langur
Trachypithecus poliocephalus (Pousargues, 1898)

Vietnam
(2000–2022)

Neahga Leonard, Richard J. Passaro, Daniela Schrudde, Roswitha Stenke,

Phan Duy Thuc and Martina Raffel

The Cat Ba Langur (also known as the Golden- as langur numbers recover interest in poaching
headed Langur), Trachypithecus poliocephalus, by people from adjacent regions may also revive
is assessed as Critically Endangered on the IUCN (Leonard et al. 2016).
Red List of Threatened Species and is probably
the most endangered of the Asian colobines Although the growth of the population is
(Rawson et al. 2020). It occurs only on Cat Ba encouraging, the overall status of the species
Island in the Gulf of Tonkin, off the north-eastern remains critical, and the total population is
Vietnamese shore (Stenke and Chu 2004). The Cat worryingly small. Habitat fragmentation and
Ba Archipelago is adjacent to the world-famous hunting has divided the remaining population
Ha Long Bay, a spectacular karst formation that into several isolated sub-populations, some of
was invaded by the sea following the last major which are non-reproducing social units. A surplus
glaciation. The favored habitat of the Cat Ba of young males is a cause for concern as take-
Langur is tropical moist forest on limestone karst over attempts can lead to infanticide, inadvertent
hills, a habitat preference it shares with the other infant deaths, and group fragmentation, all
six to seven taxa of the T. francoisi group. of which have been recorded since 2018 (N.
Leonard unpubl. data). Reproduction appears to
While there are no systematic and reliable only take place in groups above a threshold size,
data available on the historic density of the making group fragmentation a cause of concern
langur population on Cat Ba Island, reports by for population growth.
indigenous people suggest the entire island
of Cat Ba (140 km²) and some smaller offshore The total reproductive output of Trachypithecus
islands were previously densely populated by poliocephalus has been low due to the small
langurs. Hunting has been identified as the sole population and the long inter-birth cycle but
cause for the dramatic and rapid population records indicate that the birth rate is increasing,
decline from an estimated 2,400–2,700 in the with 55% of the total births recorded between
1960s to approximately 50 individuals by 2000 2000 and 2022 having taken place from 2017
(Nadler and Ha 2000). The langurs were poached to 2022. Births occur throughout the year, with
mainly for trade in traditional medicines and a peak in February–April, just prior to the rainy
for sport. Folllowing the implementation of season (Leonard et al. 2016), with a corresponding
strict protection measures in 2000, the langur peak in conception in August-October at the
population on Cat Ba Island stabilized (Nadler conclusion of the rainy season. The portion of
et al. 2003) and since 2003 has been on the reproductively active females giving birth each
increase (Leonard et al. 2016). In the latter half year varies, but on average approximately 30%
of 2015, however, numbers fell from the mid- of the active females do so (N. Leonard unpubl.
to high 60s to the low 50s and have since been data).
slowly recovering. This has raised concerns that

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In 2012, after many years of planning and

preparation, two females were successfully
translocated from a small offshore islet, where
they had become stranded, to the relative
safety of the strictly protected core zone of Cat
Ba National Park. There they assimilated into
existing groups containing males, thus giving
them the opportunity to reproduce for the first
time. It is hoped that continued protection
efforts and additional population management
interventions such as these will enhance the
rebound of this species.

The Cat Ba Archipelago and adjacent Ha Long

Bay are nationally and internationally recognized
for their importance to biodiversity conservation.
Cat Ba National Park was established in 1986.
It presently covers more than half of the main
island. Ha Long Bay was established as a
World Heritage site in 1994, and the combined
archipelago includes about 1,500–2,000 large
and small islands, cliffs and rocks. In 2004, the
Cat Ba Archipelago was designated a UNESCO
Man and the Biosphere Reserve. Despite the
conservation designations and laws to protect
the region, nature and wildlife protection on Cat
Ba Island is deficient. Environmental awareness
and commitment among the local communities
is slowly increasing, and hunting/trapping of
all animals on the island illegal. Unfortunately,
efforts to effectively conserve the langurs and
their habitat continue to face major obstacles
from increasing tourism development, increasing
human population and severe deficiencies in law
enforcement (Stenke 2005, Leonard 2018). As
is common elsewhere in the region, poaching
by local people is driven by livelihood issues,
brought about by low incomes and lack of
employment opportunities. Immense local and
regional demand for wildlife and animal parts for
food and dubious traditional medicines provide a
market for poached animals and plants. Although
langur hunting ostensibly stopped years ago, the
2015 decline in numbers raises doubts as to the
permanence of the hunting cessation. Regardless,
hunters continue to poach other animals and
plants in langur areas, thus jeopardizing langur
habitat. Strict enforcement of the established
protections is therefore necessary for the survival
of all species on Cat Ba Island that are targeted
by the illegal Asian wildlife trade.

78
A conservation program for the Cat Ba Langur
is supported by Zoo Leipzig, Zoological Society
for the Conservation of Species and Populations
(ZGAP), and the Allwetterzoo Münster in
Germany. The project was initiated on Cat
Ba Island in November 2000 by Allwetterzoo
Münster and ZGAP. The aim of the Cat Ba Langur
Conservation Program is to provide protection
for the langurs and their habitat, to conduct
research that will help inform future population
management decisions, and to help contribute
to the conservation of the overall biodiversity of
the Cat Ba Archipelago, all in collaboration with
Vietnamese authorities.

Leonard, N. (2018). Is Primate tourism a Practical Protection

Tool? The Case of Cat Ba Island, Vietnam. Presentation,
XXVIIth Congress of the International Primatological
Society, Nairobi, Kenya, 19–25 August 2018.
Leonard, N., Mai, L.S., Raffel, M., Adler, J., Passaro, R.,
Schrudde, D. and Stenke, R. (2016). Current status of the
Cat Ba langur population: numbers, population trend,
and birth rates. Presentation, Joint Meeting of XXVIth
Congress of the International Primatological Society and
the American Society of Primatologists, Chicago, 21–27
August 2016.
Nadler, T. and Ha, T.L. (2000). The Cat Ba Langur: Past, Present
and Future. The Definitive Report on Trachypithecus
poliocephalus – the World’s Rarest Primate. Hanoi,
Vietnam: Frankfurt Zoological Society.
Nadler, T., Momberg, F., Nguyen, X.D. and Lormee, N.
(2003). Vietnam Primate Conservation Status Review
2002. Part 2: Leaf Monkeys. Hanoi, Vietnam: Fauna and
Flora International and Frankfurt Zoological Society.
Rawson, B.M., Leonard, N., Covert, H. and Nadler, T. (2020).
Trachypithecus poliocephalus. The IUCN Red List of
Threatened Species 2020: e.T39871A17959804.
Stenke, R. (2005). Conservation of the Golden-headed
Langur (Trachypithecus poliocephalus poliocephalus)
on Cat Ba Island, Vietnam: Status of the Golden-
headed Langur and of Human Impact in Langur Areas.
Report, People’s Committee of Hai Phong (Provincial
Government), Cat Ba, Hai Phong, Vietnam.
Stenke, R. and Chu, C.X. (2004). The golden-headed langur
(Trachypithecus poliocephalus poliocephalus) on Cat Ba
Island – status, threat factors, and recovery options. In: T.
Nadler, U. Streicher and T.L. Ha (eds.), Conservation of
Primates in Vietnam, pp. 72–77. Hanoi, Vietnam: Frankfurt
Zoological Society.

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© Andie Ang

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 RAFFLES’ BANDED LANGUR
Presbytis femoralis (Martin, 1838)

Malaysia, Singapore
(2022)

Andie Ang, Sabrina Jabbar, Zan Hui Lee and Nadine Ruppert

Presbytis femoralis was formerly regarded as a Presbytis femoralis eats young leaves, fruits,
species with three subspecies: P. f. femoralis, P. f. seeds, and flowers. A total of 61 plant species
percura and P. f. robinsoni (Groves 2001, Roos et from 34 families were identified in the diet of P.
al. 2014). Based on multiple species delimitation femoralis in Singapore (Srivathsan et al. 2016,
algorithms applied to a dataset covering 40 Ang and Jabbar 2022). In Malaysia, 27 plant
species and 43 subspecies of Asian colobines, species from 17 families in Kampung Johor Lama
including the complete mitochondrial genomes (Najmuddin et al. 2019a) and no less than 38
of the three subspecies of P. femoralis, all three species from 20 families in Gunung Lambak (Z.
were resurrected to species (Ang et al. 2020). H. Lee unpubl. report) were documented as food
for the langurs.
The Raffles’ Banded Langur (P. femoralis) is
found only in Singapore and southern Peninsular The species is classified as Critically Endangered
Malaysia (in Johor state and a small part of on the IUCN Red List of Threatened Species
Pahang state). The populations in Singapore (Ang et al. 2021a) and is listed in Appendix II
and Malaysia are isolated from one another by of CITES. It is protected in Peninsular Malaysia
the Strait of Johor. In Singapore, this primate is under Schedule 2 of the Wildlife Conservation
found mainly in the Central Catchment Nature (Amendment of Schedule) Order 2012 under
Reserve (CCNR), the largest in the country. The the Presbytis spp. group. In Singapore, it is
CCNR comprises 2,880 hectares of lowland protected under the Protected Wildlife Species
primary and secondary forest, and freshwater Rules 2020 of the Wildlife Act (Chapter 351) as
swamp forest. In Malaysia, most populations Presbytis femoralis femoralis.
occur in the state of Johor, i.e., in Endau Rompin
National Park, Gunung Arong, Gunung Belumut, Deforestation and habitat conversion continue to
Gunung Lambak, Gunung Panti, Gunung Pulai, be the major threats to this species. It is particularly
Kampung Johor Lama, which are isolated from affected by forest clearance and disturbance
each other, with just one known population in the from urban development in Singapore and from
state of Pahang (Rompin State Park). oil palm plantations in Malaysia. As a result,
known populations are distributed in fragmented
As of 2021/22, there are 70 individuals in habitats, and fragmentation is thus recognized as
the Singapore population with a sex ratio an additional stressor. Presbytis femoralis shows
(male:female:unknown) of 26:24:20 (Ang and low genetic variability (Ang et al. 2012, Srivathsan
Jabbar 2022). There are no reliable population et al. 2016) in Singapore, and there is still a lack
estimates available for the conspecifics in of data on this species from Malaysia, where
Malaysia, but it is believed that only a few hundred population numbers and the distribution are not
individuals remain (Abdul-Latiff et al. 2019, Ang up to date.
and Jabbar 2022). The overall population of
P. femoralis, therefore, could be less than 250 Casualties in both countries have been recorded
mature individuals in the wild. as individuals attempt to travel between
fragmented habitats using roads and electric

81
ASIA

cables (Ang and Jabbar 2020; N. Ruppert, pers.

obs.). Predation events are rare, although the
langurs have been seen to be killed by eagles
(Fam and Nijman 2011) and dogs (Yang and
Lua 1988, Najmuddin et al. 2019b). In Malaysia,
additional threats come from human-wildlife
interactions when the langurs enter suburban
and residential areas to eat garden fruits (Z.H.
Lee pers. obs.).

In 2016, an IUCN Species Action Plan for the

conservation of P. femoralis was developed, and
the Raffles’ Banded Langur Working Group was
formed with representatives from government
agencies, non-governmental organizations,
universities, and experts from both countries.
The goals of the action plan are threefold: (i)
to recover and protect P. femoralis in the wild;
(ii) to gather key data through ongoing studies;
and (iii) to secure the necessary resources and
commitments for the long-term conservation of
the species (Ang et al. 2016).

In Singapore, two experimental rope bridges were

installed by the National Parks Board to facilitate
safe crossing of the langurs over roads (Ow et al.
2022). In Malaysia, a rope structure was installed
by the Malaysian Nature Society (Johor) in Gunung
Panti for primates, including P. femoralis (Chong
2020). In 2016, a citizen science program was set
up in Singapore where volunteers are trained to
collect data on the langurs in the CCNR (Ang et al.
2021b). In Malaysia, the Universiti Sains Malaysia,
in collaboration with Malaysian Primatological
Society, WWF (Malaysia, and Singapore) and
Malaysian Nature Society (Johor), is conducting
research on the behavior and ecology of the
species and planning for public outreach and
stakeholder engagement programs. Studies
on the behavior and potential for nature-based
tourism activities of this species have also been
conducted by the Universiti Tun Hussein Onn
Malaysia in Johor in recent years.

In the long term, conservation translocation of

individuals of P. femoralis between Singapore
and Malaysia might need to be considered to
restore and maintain the genetic diversity of the
populations and increase their genetic adaptive
potential. The recognition of P. femoralis as

82
one of the Top 25 Most Endangered Primates Ow, S., Chan, S., Toh, Y.H., Chan, S.H., Lakshminarayanan, J.,
can expedite urgent communication and Jabbar, S., Ang, A. and Loo, A. (2022). Bridging the gap:
assessing the effectiveness of rope bridges for wildlife
collaboration between stakeholders in Singapore in Singapore. Folia Primatol. doi.org/10.1163/14219980-
and Malaysia to foster more effective research on 20211110
and conservation actions to protect this species, Roos, C., Boonratana, R., Supriatna, J., Fellowes, J.R.,
its habitat, and the shared natural heritage of the Groves, C.P., Nash, S.D., Rylands, A.B. and Mittermeier,
R.A. (2014). An updated taxonomy and conservation status
two countries. review of Asian primates. Asian Primate J. 4(1): 2–38.
Srivathsan, A., Ang, A., Vogler, A.P. and Meier, R. (2016). Fecal
Abdul-Latiff, M.A.B., Baharuddin, H., Abdul-Patah, P. metagenomics for the simultaneous assessment of diet,
and Md-Zain, B.M. (2019). Is Malaysia’s banded langur, parasites, and population genetics of an understudied
Presbytis femoralis femoralis, actually Presbytis neglectus primate. Front. Zool. 13: 17.
neglectus? Taxonomic revision with new insights on Yang, C.M. and Lua, H.K. (1988). A report of a banded
the radiation history of the Presbytis species group in leaf monkey found dying near the Bukit Timah Nature
Southeast Asia. 60(1): 63–79. Reserve. Pangolin 1: 23.
Ang, A. and Jabbar, S. (2020). Use of forest in state land
along Upper Thomson Road by Raffles’ banded langurs
Presbytis femoralis (Martin, 1838) in Singapore. Nat.
Singap. 13: 7–10.
Ang, A. and Jabbar, S. (2022). Raffles’ Banded Langur: The
Elusive Primate of Singapore and Malaysia. Singapore:
World Scientific.
Ang, A., Srivathsan, A., Md-Zain, B.D., Ismail, M.R.B. and
Meier, R. (2012). Low genetic variability in the recovering
urban banded leaf monkey population of Singapore.
Raffles Bull. Zool. 60(2): 589–594.
Ang, A., D’Rozario, V., Jayasri, S.L., Lees, C.M., Li, T.J. and
Luz, S. (2016). Species Action Plan for the Conservation of
Raffles’ Banded Langur (Presbytis femoralis femoralis) in
Malaysia and Singapore. Report to IUCN SSC Conservation
Breeding Specialist Group, Minnesota, USA.
Ang, A., Roesma, D.I., Nijman, V., Meier, R., Srivathsan, A.
and Rizaldi. (2020). Faecal DNA to the rescue: shotgun
sequencing of non-invasive samples reveals two
subspecies of Southeast Asian primates to be Critically
Endangered species. Sci. Rep. 10: 9396.
Ang, A., Boonratana, R. and Nijman, V. (2021a). Presbytis
femoralis. The IUCN Red List of Threatened Species 2021:
e.T39801A205832720.
Ang, A., Jabbar, S., D’Rozario, V. and Lakshminarayanan, J.
(2021b). Citizen science program for critically endangered
primates: a case study from Singapore. Primate Conserv.
(35): 179–188.
Chong, C.H.A. (2020). Aerial bridges for primates at Panti
Bird Sanctuary. Panti Scientific Expedition (Part 1).
Malaysian Naturalist 74(1): 83–85.
Fam, S.D. and Nijman, V. (2011). Spizaetus hawk-eagles as
predators of arboreal colobines. Primates 52(2): 105–110.
Groves, C.P. (2001). Primate Taxonomy. Smithsonian
Institution Press, Washington, DC.
Najmuddin, M.F., Haris, H., Shahrool-Anuar, R., Md-Zain,
B.M. and Abdul-Latiff, M.A.B. (2019a). Primate tourism:
plant selection by Schlegel’s banded langur Presbytis
neglectus in Johor. IOP Conference Series: Earth and
Environmental Science. 269: 012036.
Najmuddin, M.F., Haris, H., Norazlimi, N., Md-Zain, B.M.,
Mohd-Ridwan, A., Shahrool-Anuar, R., Husna, H.A. and
Abdul-Latiff, M.A.B. (2019b). Predation of domestic dogs
(Canis lupus familiaris) on Schlegel’s banded langur
(Presbytis neglectus) and crested hawk-eagle (Nisaetus
cirrhatus) on dusky leaf monkey (Trachypithecus obscurus)
in Malaysia. J. Sustain. Sci. Manag. 14(6): 39–50.

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© Kuan-Liang Ding

84
 GUIZHOU SNUB-NOSED MONKEY
or Gray Snub-nosed Monkey
Rhinopithecus brelichi Thomas, 1903

China
(2002, 2022)

Jiang Zhou, Baoping Ren and Paul A. Garber

The Guizhou or Gray Snub-nosed Monkey As is the case for all five species of extant snub-
(Rhinopithecus brelichi) is endemic to nosed monkeys, the social organization of
southwestern China. The last remaining wild Rhinopithecus brelichi is best described as a
population of this species inhabits evergreen and large multilevel or modular society composed
deciduous broadleaf mixed forest at an altitude of several adult males, multi-adult females plus
of 1400–2100 m in the Fanjingshan National offspring units (OMUs) that feed, forage, rest,
Nature Reserve, Guizhou Province (108˚45’55”– and travel together throughout the year. These
108°48’30”E, 27˚49’50”–28˚1’30”N) (Guo et al. OMUs collectively form a breeding band that is
2020). The reserve was established in 1986 and followed by an all-male unit (AMU) composed
encompasses an area of 419 km². It is surrounded of juvenile, subadult, and adult males (Qi et al.
by local villages and small forest fragments (Xiang 2014, Tan and Bleisch 2016).
et al. 2009) and is separated from nearby lowland
forested areas by a road built in 2010. Outside Over the past several decades, deforestation and
Guizhou province, there are also five Guizhou habitat conversion for agriculture, mining, the
Snub-nosed monkeys housed at the Beijing Zoo. collection of firewood and medicinal plants, and
infrastructure development have significantly
The Guizhou Snub-nosed Monkey is listed as altered and severely fragmented the remaining
Endangered on the IUCN Red List of Threatened habitat available to the Guizhou Snub-nosed
Species (Long et al. 2020), a Class I key protected Monkey (Xiang et al. 2009, Guo et al. 2020).
species under the Chinese Wild Animal Observations of this species in the 1960s and
Protection Law, and as Critically Endangered on 1980s often reported individuals exploiting
the Red List of China’s Vertebrates (Xiang et al. forests at an elevation of 500–800 m (Zhou and
2009, Jiang et al. 2016). It was initially regarded Deng 2019). Today, individuals are principally
as a subspecies of the Golden Snub-nosed found in a narrow forest band between 1400
Monkey (Rhinopithecus roxellana, Quan and Xie and 2100 m. Across their range, trees of the
1981). Based on genomic data, however, these genera Cyclobalanopsis, Betula, Acer, Fagus,
two lineages appear to have diverged some 1.6 and Lithocarpus are the most common (Guo et
mya (Zhou et al. 2014). Both R. brelichi and R. al. 2018). Guizhou Snub-nosed Monkeys have
roxellana are considered ‘northern’ species of been reported to feed on 104 plant species (Guo
snub-nosed monkeys (Zhou et al. 2014). A recent et al. 2018).
study of the mitochondrial DNA D-loop based
on over 300 R. brelichi fecal samples identified A recent study found that only 16.6% (69.6 km²) of
11 haplotypes (Wang and Zhou 2021). Haplotype the Fanjingshan National Nature Reserve contains
diversity in R. brelichi was lower than that reported habitat suitable for the monkeys (Guo et al. 2020).
in R. roxellana and R. bieti. These latter two snub- And, although the hunting of Guizhou Snub-
nosed monkey species have larger populations nosed Monkeys has been largely eliminated, the
(Li et al. 2018). construction of roads, shops, accommodation,

85
ASIA

and facilities for the more than 1.4 million tourists

that have visited the reserve since 2018 (Guizhou
Provincial Government 2021), has impacted
Snub-nosed Monkey patterns of ranging and
distribution. For example, the construction of
an aerial tram in the reserve in 2009, divided
the remaining area of suitable habitat into two
isolated regions: a northern region of 41.1 km²
and a southern region of 28.5 km² (Guo et al.
2020). Extensive field surveys indicate there are
no Snub-nosed Monkeys present in the southern
region, and a fine-grained analysis of habitat
disturbance and forest fragmentation in the
northern region found that the area of remaining
suitable habitat totals only 27.8 km² (Guo et al.
2020).

Population estimates for the Guizhou Snub-

nosed Monkey have varied over the past 35
years, from a low of 90 individuals (Quan and
Xie 1981) to a high of 750 (Xiang et al. 2009). A
recent study based on direct field observations
to calculate the population density of Guizhou
Snub-nosed Monkeys in the reserve found that
the remaining population totals only between
125 and 336 individuals and appears to represent
a single multilevel society.

Guo et al. (2020) outline a set of conservation

recommendations designed to expand existing
areas of suitable habitat for this last remaining
wild population of Guizhou Snub-nosed
Monkeys. These include: 1) implementing an
aggressive program of the targeted reforesting
of fragmented habitat in the northern part of the
reserve in order to expand the ability of these
monkeys to reach small disconnected patches
of forest that contain feeding trees; 2) hiring
local villagers to plant deciduous and evergreen
broadleaf mixed forest trees throughout the
reserve; and 3) given the small size of the Guizhou
Snub-nosed Monkey population, moving its
conservation status to Critically Endangered
on the IUCN Red List. We also suggest a
moratorium on expanding tourist infrastructure
in the Fanjingshan National Nature Reserve and
limiting the number of tourists visiting the reserve
each year. In addition, nearby the Fanjingshan
National Nature Reserve there is another nature
reserve, the Yangxi Provincial Nature Reserve
(218 km²), which is largely unexplored. It remains

86
uncertain whether there are any Guizhou Snub-
nosed Monkeys present there.

Guizhou Provincial Government (2021). Available online:

<https://www.sohu.com/a/392098644_114988>.
Guo, Y., Zhou, J., Xie, J.H., Garber, P.A., Michael, B., Ren,
B.P., Ni, D.Y. and Zhou, J. (2018). Altitudinal ranging of
the Guizhou golden monkey (Rhinopithecus brelichi):
patterns of habitat selection and habitat use. Glob. Ecol.
Conserv. 16. doi.10.1016/j.gecco.2018.e00473
Guo, Y. et al. (2016). Red List of China’s Vertebrates. Biodiv.
Sci. 24(5): 500–551.
Li, B.G. et al. (2018). The primate extinction crisis in China:
immediate challenges and a way forward. Biodiv..
Conserv. 27: 3301–3327.
Long, Y., Bleisch, W.V., Richardson, M. and Baoguo, L. 2020.
Rhinopithecus brelichi. The IUCN Red List of Threatened
Species 2020: e.T19595A185220311.
Qi, X.G., Garber, P.A., Huang, Z.P., Huang, K. and Li, B.G.
(2014). Satellite telemetry and social modeling offer new
insights into the origin of primate multilevel societies.
Nature Commun. 5: 5296.
Quan, G.Q. and Xie, J.H. (1981). Notes on Rhinopithecus
roxellanae brelichi Thomas. Acta Theriologica Sinica. 1:
113–116.
Wang, S.W. and Zhou, J. (2021). Limited heredity diversity of
the critically endangered Guizhou golden monkeys. Res.
Ecol. 3(2): 53–55.
Tan, C.L. and Bleisch, W. (2016). Guizhou snub-nosed monkey.
In: N. Rowe and M. Myers (eds.), All the World’s Primates,
pp. 624–625. Charlestown, RI: Pogonias Press.
Xiang, Z.F., Nie, S.G., Lei, X.P., Chang, Z.F., Wei, F.W. and
Li, M. (2009). Current status and conservation of the gray
snub-nosed monkey Rhinopithecus brelichi (Colobinae)
in Guizhou, China. Biol. Conserv. 142: 469e476.
Zhou, J. and Deng, H. (2019). Two Endemic primate species
in China: Hainan gibbon and Guizhou snub-Nosed
monkey. In: E.C. Carmona, C.M. Musarella and A.C.
Ortiz (eds.), Endemic Species. Intech Open. doi.10.5772/
intechopen.85933
Zhou, X. et al. (2014). Whole-genome sequencing of the
snub-nosed monkey provides insights into folivory and
evolutionary history. Nat. Genet. doi.10.1038/ng.3137

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© Fan Pengfei

88
 GAOLIGONG HOOLOCK GIBBON
or Skywalker Hoolock Gibbon
Hoolock tianxing Fan et al., 2017

China, Myanmar
(2018, 2022)

Pengfei Fan, Hanlan Fei, Lu Zhang and Susan M. Cheyne

Hoolock gibbons were first described scientifically estimated that the total population was less than
by Harlan (1834) under the name Simia hoolock. 150 individuals, made up of 26 family groups and
They were subsequently transferred to the 11 solitary individuals across 17 subpopulations
genus Hylobates, and then assigned to their (Zhang et al. 2020). The largest subpopulation
own distinct subgenus (later elevated to genus), has seven groups, and five of the subpopulations
first Bunopithecus (later restricted to an extinct only have one group remaining. Although the
Quaternary gibbon from China) (Prouty et al. population remained relatively stable from 2009
1983; Groves 2001) and then Hoolock (Mootnick to 2017, it is isolated from other populations
and Groves 2005). Taxonomic variation between by distance, villages and roads, and has a low
different hoolock populations was first recognized birth rate. For example, the reproductive rates
by Groves (1967), who identified a major east- of three mature females were tracked between
west morphological division and described 2008 and 2020. One of these females produced
Hylobates hoolock leuconedys to distinguish two offspring in this time (November 2008 and
eastern hoolock populations from those in the December 2012), whilst the others produced just
west, geographically isolated by the Chindwin one (2008 and 2012). Hoolock tianxing is listed as
River. Both subspecies were latterly elevated Endangered on the IUCN Red List of Threatened
to full species: the western (Hoolock hoolock) Species (Fan et al. 2020).
and eastern hoolock (H. leuconedys) gibbons.
Fan et al. (2017) assessed the morphological Agricultural encroachment, commercial logging,
and genetic characteristics of wild gibbons and habitat fragmentation and isolation, and hunting
museum specimens to evaluate the taxonomic (for bushmeat and pet trade) are major threats to
status of the hoolock population in China. The H. tianxing. The population is also threatened
results suggested that hoolocks distributed to by stochastic loss, in which subpopulations
the east of the Irrawaddy and Nmai Hka rivers, are reduced to 1-2 groups with no opportunity
which were previously assigned to H. leuconedys, for dispersal or gene-flow. Population linking,
are morphologically and genetically distinct protection and habitat restoration are urgently
from those to the west of the rivers, resulting in needed, and the translocation of non-viable
them now being recognized as a new species: sub-populations may also be required. There
the Gaoligong Hoolock Gibbon or Skywalker is also a hoolock population in Myanmar. While
Hoolock Gibbon, Hoolock tianxing Fan et al. unstudied, it is likely that the population faces
2017. similar difficulties to those in China, (i.e., habitat
loss and poaching) but there is comparatively
Hoolock tianxing was once widely distributed less conservation action and law enforcement in
around the west bank of the Salween River, west Myanmar. As the population in China decreases,
of Yunnan, China, but >90% of its habitat was lost the importance of the Myanmar population
by 1994 (Fan et al. 2017). In 2009, the population increases. Therefore, although demand for
was estimated to be <200 individuals (Fan et conservation intervention in Myanmar is high, a
al. 2011). In 2017, a second population survey

89
ASIA

careful approach is advised to safely navigate the

recent political unrest.

The following actions are needed: (1) Raise

awareness of this species, especially in China
through targeted campaigns; (2) Determine
population status in Myanmar through
population surveys; (3) Address threats at a local
scale through an ethnographic approach; and (4)
Investigate possibilities for connecting populated
forest fragments and/or translocation of isolated
groups/individuals.

Fan, P.-F., Xiao, W., Huo, S., Ai, H.S., Wang, T.C. and Lin, R.T.
(2011). Distribution and conservation status of Hoolock
leuconedys in China. Oryx 45 (1): 129–134.
Fan, P.-F., He, K., Chen, X., Ortiz, A., Zhang, B., Zhao, C.,
Li, Y.-Q., Zhang, H.-B., Kimock, C., Wang, W.-Z., Groves,
C., Turvey, S.T., Roos, C., Helgen, K.M. and Jiang, X.-L.
(2017). Description of a new species of hoolock gibbon
(Primates: Hylobatidae) based on integrative taxonomy.
Am. J. Primatol. 79: e22631. 15pp.
Fan, P.-F., Turvey, S.T. and Bryant, J.V. 2020. Hoolock tianxing
(amended version of 2019 assessment). The IUCN Red List
of Threatened Species 2020: e.T118355648A166597159.
Groves, C.P. (1967). Geographic variation in the hoolock or
white-browed gibbon (Hylobates hoolock Harlan, 1834).
Folia Primatol. 7: 276–283.
Groves C.P. (2001). Primate Taxonomy. Washington, DC:
Smithsonian Institution Press.
Harlan, R. (1834). Description of a species of Orang, from
the north-eastern province of British East India, lately the
Kingdom of Assam. Trans. Am. Phil. Soc. 4: 52–59.
Mootnick, A. and Groves, C.P. (2005). A new generic name
for the hoolock gibbon (Hylobatidae). Int. J. Primatol.
26(4): 971–976.
Prouty, L.A., Buchanan, P.D., Pollitzer, W.S.and Mootnick A.R.
(1983). Bunopithecus: A genus-level taxon for the hoolock
gibbon (Hylobates hoolock). Am. J. Primatol. 5: 83–87.
Zhang, L., Guan, Z., Fei, H., Yan, L., Turvey, S.T. and Fan, P.
(2020). Influence of traditional knowledge on conservation
of the skywalker hoolock gibbon (Hoolock tianxing)
outside nature reserves. Biol. Conserv. 241: 108276.

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© Serge Wich

92
 TAPANULI ORANGUTAN
Pongo tapanuliensis Nurcahyo, Meijaard, Nowak, Fredriksson
& Groves in Nater et al., 2017

Indonesia
(2018, 2022)

Serge Wich, Erik Meijaard, Didik Prasetyo, Wanda Kuswanda,

Panut Hadisiswoyo and Jatna Supriatna

The Tapanuli Orangutan, Pongo tapanuliensis, to have experienced a significant population

was only formally described in 2017 when it was reduction in the past 150 years (Nowak et al. 2017,
shown that an isolated orangutan population Meijaard et al. 2021). With a population estimate
in the Batang Toru region (which used to be of 767 individuals (95% confidence intervals: 231–
considered the southernmost range of extant 1,597 individuals; Wich et al. 2019), the Tapanuli
Sumatran orangutans, Pongo abelii), south Orangutan is the least numerous of all great ape
of Lake Toba, is distinct from other Sumatran species. Its distribution is separated by around
and Bornean populations (Nater et al. 2017). 100 km from the closest population of the
Through a comparison of cranio-mandibular Sumatran Orangutan to the north. A combination
and dental characters from an orangutan of small population size and geographic isolation
killed during human-orangutan conflict to a is something of particularly high conservation
comparative sample of adult male orangutans concern. This may lead to inbreeding depression
of similar developmental stage, Nater et al. (Hedrick and Kalinowski 2000) and threaten
(2017) found consistent differences between population persistence (Allendorf et al. 2013). For
the Batang Toru individual and other extant example, in the conflict area of South Tapanuli,
Ponginae. Comparisons of adult male long calls only 155 individuals were found (95% confidence
from two Tapanuli males with those of a large interval: 121–187 individuals; Kuswanda et al.
sample of Bornean and Sumatran males also 2020). Nater et al. (2017) recorded extensive
revealed a unique mix of long call characteristics. runs of homozygosity in the genomes of two
Model-based approaches based on the Tapanuli Orangutan individuals, pointing to the
analyses of 37 orangutan genomes supported occurrence of recent inbreeding.
the morphological results, revealing that the
deepest split in the evolutionary history of extant The only known population of Tapanuli
orangutans occurred around 3.38 million years Orangutans is in the uplands of the Batang
ago between the Batang Toru population and Toru Ecosystem, an area of roughly 1,500 km²
those to the north of Lake Toba. In comparison, consisting of three forest blocks, of which 1,023
the Bornean Orangutan and Sumatran Orangutan km² is suitable orangutan habitat (Wich et al.
separated much later at about 674 ka. The 2016, 2019, Rahman et al. 2019, Kuswanda et al.
analyses show that there was some gene flow 2021a). Most of this is ecologically suboptimal
between the Sumatran and Tapanuli Orangutan upland forest (>500 m asl, up to 1800 m asl),
species until 10–20 ka. Combined, these analyses covering the upper watersheds of nine river
supported a new classification of orangutans into systems and providing fresh water for over
three extant species. 100,000 people across Tapanuli, which covers
26 sub-districts and 187 villages (Putro et al.
Due to high levels of habitat conversion and 2019). Forest loss data indicate that orangutan
fragmentation, along with illegal hunting and habitat below 500 m asl was reduced by 60%
poaching, the Tapanuli Orangutan is estimated between 1985 and 2007 for both the Tapanuli

93
ASIA

and the Sumatran Orangutan (Wich et al. 2008,

2011). A recent analysis indicated that the current
Tapanuli Orangutan distribution is approximately
2.5% and 5.0% of what it was in the 1890s and
1940s, respectively (Meijaard et al. 2021). It is
thought that more Tapanuli Orangutan habitat
will be lost as significant areas of forest within its
range remain under considerable threat (Wich
et al. 2016, 2019, Sloan et al. 2018) from habitat
conversion for small-scale agriculture, crop-
conflict and related killing, mining exploration and
exploitation, a hydroelectric scheme, geothermal
development and agricultural plantations. The
habitat of the Tapanuli Orangutan consists of
7% conservation forest, 64% protection forest,
4% production forest and the remaining 25% in
other use areas/cultivated land (Putro et al. 2019).
Orangutan populations are easy to find on land
managed by farmers and this poses a high threat
to orangutans.

Across the species’ range, the protected areas

are not immune from the above threats (Wich
et al. 2008, 2011, 2016) and orangutans in these
areas are also hunted (Wich et al. 2012). Due
to their slow life history, with a generation time
of at least 25 years, and an interbirth interval of
8-9 years, orangutans on Sumatra are unable
to sustain substantial and continual loss of
individuals (Wich et al. 2004, 2009, Marshall et
al. 2009, Kuswanda et al. 2021b). Recent studies
indicate that the eastern subpopulation may be
growing (Kuswanda et al. 2021b). Nevertheless,
annual removal rates (killing, translocations,
rescues) appear to exceed those needed for
maintaining viable populations (Meijaard et al.
2021), while disease risk poses another significant
threat (Sherman et al. 2021).

The Tapanuli Orangutan was more widespread

until quite recently as indicated by the Meijaard
et al. (2021) study, with sightings further south in
the lowland peat swamp forests in the Lumut area
(Wich et al. 2003) and several nests encountered
during a rapid survey in 2010 (G. Fredriksson pers.
obs.). The forests in the Lumut area have, however,
been almost completely converted to oil-palm
plantations in recent years. Observations were
also made of a male orangutan in the Adiankoting
subdistrict in North Tapanuli, north of the Batang
Toru West forest block, during a human conflict
situation where the orangutan was shot at with

94
an air rifle when it was found foraging on Durian (Laurance et al. 2020, Prasetyo et al. 2021).
fruits (G. Fredriksson pers. obs.). The persistence Opinions about impacts differ significantly, even
of viable subpopulations in these areas is not among orangutan conservation scientists and
known. practitioners. On the one hand, opponents of
the dam argue that the dam could impact roughly
Tapanuli Orangutans have been observed 100 km² of Tapanuli Orangutan habitat, or nearly
feeding on a number of tree species that have 10% of the entire species population (Sloan et al.
not previously been recorded in orangutan diets. 2018). The dam would jeopardize the chances
These unique species include Gymnostoma of maintaining and restoring habitat corridors
sumatranum from the Casuarinaceae family, between the western and eastern Tapanuli
and Dacrycarpus imbricatus, Dacrydium Orangutan ranges and a strict nature reserve with
beccarii, Dacrydium comosum, and Podocarpus a small population of Tapanuli Orangutans (Wich
neriifolius from the Podocarpaceae family. At the et al. 2019). If the connectivity between these
Sumatran Orangutan Conservation Programme’s populations is not restored, it is argued that the
(SOCP) long-term monitoring station in the long-term survival of the Tapanuli Orangutan will
Batang Toru Ecosystem, 21.9% of all feeding be severely threatened (Wich et al. 2019).
observations recorded between 2011 and
2015 were represented by five conifer species Proponents of the dam, on the other hand, have
(Araucariaceae and Podocarpaceae) and one argued that the dam is a legitimate development
non-conifer evergreen species (Casuarinaceae). program by the Indonesian government needed
Seeds of Agathis borneensis from the to meet Sumatra’s clean energy needs. Most of
Araucariaceae family have been considered the infrastructure development is carried out on
a ‘fallback’ fruit, frequently consumed when rubber plantations and community agricultural
few other fruits are available (Nater et al. 2017; lands that are no longer habitat for orangutans.
SOCP unpubl. data). In agroforestry, orangutans The hydro-dam company has also implemented
have also been identified consuming Hevea various conservation programs, such as the
brasiliensis and Coffea arabica fruits (Kuswanda development of arboreal bridges and vegetation
et al. 2021c). Thus, a significant proportion of corridors, re-vegetation of degraded areas, and
the dietary profile of Tapanuli Orangutans is the establishment of a special conservation
markedly different from that of previously studied program for the species. They believe that
orangutan populations. This may indicate that these measures will reduce the total impact of
this is a refugee species that has been pushed the hydroelectric project and that, in fact, the
out of more optimal lowland habitat by past hydrocompany can make a significant positive
hunting (Meijaard et al. 2021). contribution to the likelihood of survival of the
species (Prasetyo et al. 2021).
Due to the extremely rugged terrain, external
threats have been primarily limited to illegal In order to safeguard the future of the most
clearing of protected and production forests, endangered great ape species, all possible
hunting and killing during crop conflict, and efforts must now be made to prevent any further
trade in young orangutans (Wich et al. 2012, degradation of Tapanuli Orangutan habitat,
2016, Kuswanda et al. 2021b). In the southwest and to reconnect its three habitat fragments to
corner of the Batang Toru Ecosystem, a large restore genetic exchange. As it currently stands,
gold and silver mine has converted key lowland two of the three habitat fragments do not have
habitat of the Tapanuli Orangutan and retains viable populations, leaving only one viable but
controversial mining permits overlapping parts highly threatened population to safeguard the
of the remaining Tapanuli Orangutan range. future of the species. Lastly, field management
Recent expansion of the mine has led to further activities need to be established to prevent
deforestation in the area where the Tapanuli further hunting and encroachment, with clear
Orangutan occurs. and enforced boundary demarcation, and active
human-orangutan conflict mitigation efforts
A hydro-electric development has been underway must be put in place. The Tapanuli Selatan
in the Tapanuli Orangutan’s range since 2017 government is urged to legalize the animal (such

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1
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tapanuliensis) within the renewable energy development
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Selatan district regulation number 63, 2020. 493.
Putro, H.R., Rinaldi, D., Rahman, D.A. and Kuswanda, W.
(2019). The Ecology of Tapanuli Orangutan. Bogor,
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Hedrick, P.W. and Kalinowski, S.T. (2000). Inbreeding
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go.id)>.

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NEOTROPICS

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© Sarisha Trindade

100
 BUFFY-HEADED MARMOSET
Callithrix flaviceps (Thomas, 1903)

Brazil
(2022)

Fabiano R. de Melo, Orlando Vital, Sarisha Trindade do Carmo, Rodrigo Salles de Carvalho,
Mônica M. Valença-Montenegro, Sérgio Lucena Mendes and Leandro Jerusalinsky

The Buffy-headed Marmoset, Callithrix flaviceps, ecological and behavioral studies of groups of C.
is a small primate endemic to the Atlantic Forest flaviceps and C. aurita.
of Southeast Brazil. It occurs in the Serra da
Mantiqueira, southern Espírito Santo, south of As with the other marmosets, C. flaviceps eats
the Rio Doce at least to the state boundary with fruits, flowers, nectar, plant exudates (gums,
Rio de Janeiro. In the past, its range possibly saps, latex), fungi and animal prey (including
reached the north of the state of Rio de Janeiro, frogs, snails, lizards, spiders, and insects) (Ferrari
in the municipalities of Natividade, Porciúncula 1988, 1991a, Corrêa et al. 2000, Hilário and Ferrari
and the north of Bom Jesus do Itabapoana, 2010a). Marmosets also have morphological
when this area was covered with native forests and behavioral adaptations for gouging tree
(Melo et al. 2021). It extends west into eastern trunks, branches, and vines of certain species to
Minas Gerais in scattered localities in the highly stimulate the flow of gum, which they eat, forming
fragmented forests of the Rio Manhuaçu basin as a notable component of their diet (Coimbra-Filho
far as Manhuaçu (40°02’W), as noted by Coimbra- 1972, Rylands 1984). Nevertheless, C. flaviceps
Filho (1986a), Coimbra-Filho et al. (1981) and rarely gouges trees in order to stimulate exudate
Mendes and Melo (2007), reaching an area of production and feeds primarily on exudates
approximately 30,000 km² – among the smallest already available (Ferrari 1991b, Hilário and
ranges (Extent of Occurrence) of any platyrrhine Ferrari 2010a).
(Ferrari and Mendes 1991).
Buffy-headed Marmosets live in extended family
In the past, the eastern Brazilian marmosets groups of 3 to 20 individuals (Alves 1986, Ferrari
of the “jacchus group” were considered to 1991b, Ferrari and Digby 1996, Guimarães 1998,
be subspecies of Callithrix jacchus, following Hilário and Ferrari 2010a), and groups present
Hershkovitz (1977). All are now considered weak territoriality and occupy large home ranges
to be full species (Vivo 1991, Mendes 1997a, (33.9 to 138.3 ha) compared to other congeneric
1997b, Marroig et al. 2004, Coimbra-Filho et al. species (Ferrari 1991, Guimarães 1998, Hilário
2006). Coimbra-Filho (1986a, 1986b, 1990) and 2009). Generally, only a single dominant female
Coimbra-Filho et al. (1993) argued that Callithrix breeds, although up to four females have been
flaviceps should be considered a subspecies of seen to reproduce simultaneously in one large
C. aurita. The similarities between C. flaviceps group (Ferrari 1991, Guimarães 1998, Hilário and
and C. aurita in pelage patterns such as the ear Ferrari 2010b). Births often occur twice a year,
tufts and the skull-like facial mask, ecological however, the reproductive peak in the dry season
adaptations, ontogeny (the infants are extremely is less pronounced (Ferrari 1991), with some
similar), vocalizations and clinal variation in populations even avoiding reproduction in this
overall pelage color, are the basis for this opinion season (Hilário and Ferrari 2010a).
(Coimbra-Filho 1986a, 1986b, Coimbra-Filho et
al. 1993). Ferrari et al. (1996), however, rejected
the subspecies status of C. flaviceps based on

101
NEOTROPICS

Callithrix flaviceps and C. aurita are the

southernmost marmosets in terms of the
natural range of the genus. Callithrix jacchus, C.
penicillata and C. geoffroyi have been introduced
further south in the Brazilian states of São Paulo,
Paraná, Santa Catarina, Rio Grande do Sul and in
Argentina (Santos et al. 2005). They occur in the
montane Atlantic coastal forest and forests of the
inland plateau, at altitudes up to 1,200 m where
dry season temperatures can fall close to freezing
(Ferrari et al. 1996). They show some level of
tolerance and flexibility to habitat disturbance,
being sometimes rare in old growth forest with
sparse understories (Ferrari 1988, Ferrari and
Mendes 1991). There is a natural hybrid zone with
C. aurita, in the region of Carangola, Tombos and
Caiana municipalities, in south-eastern Minas
Gerais (Coimbra-Filho et al. 1993, Cosenza 1993,
Mendes 1997a, Cosenza and Melo 1998).

Callithrix flaviceps has a restricted range in an

area where the forest is extremely fragmented
due to expanding agriculture, cattle ranching,
tree plantations (Eucalyptus), urbanization,
mining and dam construction (Coimbra-Filho
1986a, 1990, Ferrari and Mendes 1991, Mendes
and Melo 2007). Mendes and Melo (2007)
surveyed forest fragments in the Zona da Mata
of the state of Minas Gerais, and recorded the
presence of introduced populations of Callithrix
jacchus, C. penicillata, and C. geoffroyi, which
they believe are displacing C. flaviceps. They
recommended awareness campaigns to reduce
the habit of releasing exotic marmosets in the
forests surrounding the town of Manhuaçú. Since
this study, the presence of invasive congenerics,
resulting in mixed groups and hybrid specimens,
has unfortunately become common and results
in ecological competition and genetic erosion for
the native and threatened Callithrix flaviceps (see
Malukiewicz et al. 2021). A recent survey in the
Caratinga Biological Station/Reserva Particular
do Patrimônio Natural (RPPN) Feliciano Miguel
Abdala indicated a 90% population decline since
2005, primarily due to a yellow fever outbreak
(Possamai et al. 2019). Other subpopulations
were highly impacted by this epidemic in Espírito
Santo (S.L. Mendes, pers. obs.), and this may be
the case throughout the species’ range.

This species has been estimated to have a mean

density of 7.1 individuals/km² (range: 3.4 to 18

102
individuals/km²) in the Augusto Ruschi Biological The Mountain Marmosets Conservation Center
Reserve, Espírito Santo (Pinto et al. 1993), while of the Federal University of Viçosa (CCSS-UFV)
Hilário (2009) recorded a density of 15.4 individuals/ was recently created with the specific goal
km². At the Caratinga Biological Station/RPPN to breed and maintain captive groups of C.
Feliciano Miguel Abdala, Minas Gerais, the aurita and C. flaviceps, and to set up an ex situ
recorded densities were 40 individuals/km² management program to preserve the species’
(Ferrari 1988) and 13 individuals/km2 (Almeida- genetic diversity and support releases in key
Silva et al. 2005). A recent study of occupancy areas. A group of researchers and managers from
in the northwest boundary of the species’ range several institutions (universities, NGOs, zoos,
(in Atlantic Forest patches in private reserves and public agencies) has established a series of tools
their surroundings in the central region of Minas and methodologies to increase our knowledge
Gerais), detected healthy, pure-breeding groups of the species, including developing a Callithrix
in 25 of 145 sites surveyed, ranging from 2–12 occurrence database to identify priority areas for
individuals, and two hybrid groups (Carmo 2022). surveying, conserving, and managing populations
Using a conservative approach (with the lowest of these species. These initiatives are part of the
density reported for the species), we estimate a Mountain Marmosets Conservation Program
total population of 4,440, consisting of less than (MMCP), which is developing an operational
2,500 mature individuals (Brazil, ICMBio 2018). agenda for the effective implementation of
Considering the highly fragmented landscape in the conservation strategies established by the
which C. flaviceps occurs, it is also unlikely that National Action Plan for the Conservation of the
any subpopulation has more than 250 mature Atlantic Forest Primates and the Maned Sloth
individuals (Melo et al. 2019). (Brazil, ICMBio, 2018).

A population reduction of at least 80% over 18

years (three generations) has been inferred based
on cumulative and synergistic impacts of the Almeida-Silva, B., Cunha, A.A., Boubli, J.P., Mendes, S.L.
historical habitat loss in the species’ restricted and Strier, K.B. (2005). Population density and vertical
stratification of four primate species at the Estação
range, the diminished and severally fragmented
Biológica de Caratinga/RPPN-FMA, Minas Gerais, Brazil.
habitat remnants, ecological competition Neotrop. Primates 13: 25–30.
and genetic erosion caused by the invasion Alves, M.C. (1986). Observações sobre o Callithrix flaviceps
of congenerics, and the recent yellow fever (Thomas, 1903) na Estação Ecológica de Caratinga –
outbreak. Thus, Callithrix flaviceps was classified EBC/FBCN, MG (Callitrichidae, Primates). In: M.T. de
Mello (ed.), A Primatologia no Brasil – 2, pp. 205–206.
as Critically Endangered on the IUCN Red List of Sociedade Brasileira de Primatologia, Brasília.
Threatened Species (Melo et al. 2021), as it was Brazil, ICMBio (2018). Aprova o Plano de Ação Nacional
for the national Brazilian assessment (ICMBio in para a Conservação dos Primatas da Mata Atlântica e
prep.). da Preguiça-de-Coleira. Portaria n° 702, de 07 de agosto
de 2018 – Instituto Chico Mendes de Conservação da
Biodiversidade. Diário Oficial da União – Seção 1 153:
The larger protected areas where the species
57.
occurs are the Caparaó National Park, the Carmo, S.T. (2022). Probabilidade de Ocupação e Detecção
Augusto Ruschi Biological Reserve, the Rio Doce de Sagui-da-Serra (Callithrix flaviceps Thomas, 1903)
State Park, and the Pedra Azul State Reserve. (Primates: Callitrichidae) na RPPN Fazenda Macedônia
It is also present in a series of privately owned e em Fragmentos de Mata Atlântica no Seu Entorno,
Ipaba, MG. Masters dissertation, Universidade Federal
reserves, including the Caratinga Biological de Viçosa, MG, Brazil.
Station/RPPN Feliciano Miguel Abdala, the RPPN Coimbra-Filho, A.F. (1972). Aspectos inéditos do
Mata do Sossego, and the Montes Verdes Forest comportamento de sagüis do gênero Callithrix
Reserve. It is not clear, however, whether (and to (Callithricidae, Primates). Revista Brasil. Biol. 32: 505–
512.
what extent) these areas are free from the invasive
Coimbra-Filho, A.F. (1986a). Sagui-da-serra Callithrix
and hybrid forms. This highlights the crucial flaviceps (Thomas, 1903). FBCN/Inf., Rio de Janeiro 10(1):
need to develop effective methods to control 3.
the invasive congenerics and the resulting hybrid Coimbra-Filho, A.F. (1986b). Sagui-da-serra-escuro Callithrix
populations, as well as implementing recovery aurita (E. Geoffroy, 1812). FBCN/Inf., Rio de Janeiro 10(2):
3.
strategies for native populations of C. flaviceps.

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and Callithrix flaviceps) – how different, how similar? In: Santos, C.V., Luz, K.P. and Sant’anna, F.S. (2005). As três
M.A. Norconk, A.L. Rosenberger and P.A. Garber (eds.), espécies de primatas do gênero Callithrix (C. jacchus, C.
Adaptive Radiations of Neotropical Primates, pp. 157– penicillata e C. geoffroyi) introduzidos na Ilha de Santa
171. New York: Plenum Press. Catarina – SC: importância de pesquisa na implantação
Guimarães, A. (1998). Ecologia, Comportamento do manejo. In: Resumos: XI Congresso Brasileiro de
Reprodutivo e Marcação de Cheiro em Um Grupo de Primatologia, Porto Alegre, RS, Brasil. 13 a 18 de fevereiro
Callithrix flaviceps (Callitrichidae, Primates), na Estação de 2005. [Abstract]
Biológica de Caratinga, Minas Gerais. Masters thesis, Vivo, M. de. 1991. Taxonomia de Callithrix Erxleben, 1777
Universidade Federal de Minas Gerais, Belo Horizonte, (Callitrichidae, Primates). Belo Horizonte: Fundação
MG, Brazil. Biodiversitas.
Hershkovitz, P. (1977). Living New World monkeys
(Platyrrhini), With an Introduction to Primates. Chicago,
IL: University of Chicago Press.
Hilário, R.R. (2009). Padrão de Atividades, Dieta e Uso do
Habitat por Callithrix flaviceps na Reserva Biológica
Augusto Ruschi, Santa Teresa, ES. Masters thesis,
Universidade Federal de Minas Gerais, Belo Horizonte,
MG, Brazil.

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105
 NEOTROPICS

© Fabiano R. de Melo

106
 KA’APOR CAPUCHIN
Cebus kaapori Queiroz, 1992

Brazil
(2012, 2014, 2016, 2022)

Gerson Buss, Tatiane dos Santos Cardoso, Helder Queiroz, Ana Cristina Mendes de
Oliveira, Fabiano R. de Melo and Leandro Jerusalinsky

The Ka’apor Capuchin (Cebus kaapori), first and habitat degradation in the entire Brazilian
described just over 30 years ago, is endemic Amazon (Carvalho et al. 1999). More than 70%
to the eastern edge of the Brazilian Amazon, of the forest has been destroyed, converted
in the so-called Centro de Endemismo Belém to farmland and pasture (Almeida and Vieira
(Belém Endemism Center), in the north-east of 2010). Deforestation continues, and most of the
the state of Pará and the north-west of the state remaining forests are isolated and degraded
of Maranhão (Queiroz 1992). Its range extends patches in farmland where this species is also
from the east of the lower Rio Tocantins to the hunted. Habitat loss across the species’ range
Rio Grajaú, where it enters the Zona dos Cocais from 1985 to 2020 was estimated at 32.8% (Butti
(Queiroz 1992, Ferrari and Queiroz 1994, Ferrari et al. in prep.). A species distribution model
and Souza 1994, Silva and Cerqueira 1998, indicates that C. kaapori could lose all of its forests
Carvalho et al. 1999, Cunha et al. 2007). The because of climate change and deforestation
Extent of Occurrence is 206,081 km² (Butti et al. over the next 30 years (da Silva et al. 2022).
in prep.). This species is usually observed in tall
lowland terra firma forest, generally below 300 m Cebus kaapori occurs in two protected areas:
above sea level, and has not been recorded in the Lago de Tucuruí Environmental Protection
seasonally inundated or secondary forest (Silva Area of 5,687 km² that allows for sustainable use;
Jr. et al. 2009, 2010, Rylands and Mittermeier and the Gurupi Biological Reserve of 2,712 km²
2013). It is generally seen in small groups of up that is under strict protection. In 2015, Gurupi
to ten individuals, sometimes accompanying and contiguous indigenous lands were seriously
the Endangered Black Saki (Chiropotes satanas) affected by forest fires. The Chico Mendes
(Ferrari and Lopes 1996, Carvalho et al. 1999). Institute for Biodiversity Conservation (ICMBio)
Besides surveys and abundance studies, recent estimated that 1,330 km² of the Gurupi reserve
research has provided additional ecological was impacted (Buss et al. 2017), reducing the
information about the species (Oliveira et al. available habitat for C. kaapori.
2014).
Lopes (1993) recorded three groups over 480 km
Due to the threats of habitat loss and hunting, surveyed in the Gurupi Biological Reserve (0.06
and a drastic population reduction, C. kaapori groups/10 km). Carvalho et al. (1999) registered
is classified as Critically Endangered on the a relative abundance of 0.99 groups/10 km in the
IUCN Red List of Threatened Species (Fialho Fazenda Cauaxi, in Paragominas. More recently,
et al. 2021), likewise in the national assessment Buss et al. (2017) found 0.25 groups/10 km in a
of Brazil (Brazil, MMA 2014, Fialho et al. 2018). survey of 320 km, in the Gurupi reserve. These
Lopes and Ferrari (1993) and Ferrari and Queiroz results indicate that the Ka’apor Capuchin is
(1994) concluded that C. kaapori is one of the naturally rare; it is hunted and is susceptible to
most threatened of all Amazonian primates. The any, even light, disturbance, or degradation of
Ka’apor Capuchin inhabits a densely populated its habitat. Selective logging of fruit trees that
region with the highest level of deforestation form a significant part of the species’ diet is a

107
NEOTROPICS

considerable threat (Lopes 1993). Its rarity may

be related to competition with the sympatric
Black-capped Capuchin (Sapajus apella) and
naturally low densities may reflect the need for
large home ranges.

A Population Viability Analysis using Vortex

software indicated only three viable populations
over the next 100 years (Campos 2009): a complex
of Indigenous Territories in the state of Maranhão
(Caru, Awá, Alto Turiaçu, Araribóia); Alto Rio
Guama, an Indigenous Territory in the state of
Pará; and the Gurupi Biological Reserve along
the border between the two states.

Ka’apor Capuchins are maintained in only a few

zoological institutions, such as São Paulo Zoo
(Fundação Parque Zoológico de São Paulo), Rio
de Janeiro Zoo (BioParque do Rio de Janeiro) and
the National Primate Center (Centro Nacional de
Primatas [CENP], linked to the Brazilian Ministry
of Health) (Mônica M. Valença-Montenegro -
ICMBio/CPB, pers. comm.). Guajá Indians often
keep them as pets (Queiroz 1992).

A study on the ecology of C. kaapori in the Gurupi

Biological Reserve, aiming to generate the first
systematic information on home range, habitat
use, diet and activity pattern of the species in
continuous forest, is being carried out by Tatiane
S. Cardoso. Preliminary data indicate large home
ranges of approximately 300 ha for a group with
12 individuals, among the largest estimated for
the genus Cebus (Cardoso 2021).

Strategies to promote the conservation of C.

kaapori are included in the Brazilian National
Action Plan for the Conservation of the
Amazonian Primates (Brazil, MMA 2017). The
Gurupi Biological Reserve is one of the study
areas of the project “Primates in Protected Areas
of the Brazilian Amazon: assessing the impacts
of forest fires over endangered primates,”
coordinated by the National Center for Research
and Conservation of Brazilian Primates (ICMBio/
CPB) and supported by Re:wild. This project
aims to evaluate the occurrence and abundance,
conservation status and impacts of fires on
endangered primate populations in protected
areas along the Brazilian Amazon’s Arc of
Deforestation, to provide information to support
further protection and management actions.

108
Almeida, A.S. and Vieira, I.C.G. (2010). Centro de Endemismo Fialho, M.S., Jerusalinsky, L., Moura, E.F., Ravetta, A.L.,
Belém: status da vegetação remanescente e desafios Laroque, P.O., Queiroz, H.L., Boubli, J.P. and Lynch Alfaro,
para a conservação da biodiversidade e restauração J.W. (2021). Cebus kaapori (amended version of 2020
ecológica. REU Sorocaba 36(3): 95–111. assessment). The IUCN Red List of Threatened Species
Brazil, MMA. (2014). Reconhecer como espécies da fauna 2021: e.T40019A191704766.
brasileira ameaçadas de extinção aquelas constantes da Lopes, M.A. (1993). Conservação do Cuxiú-preto,
“Lista Nacional Oficial de Espécies da Fauna Ameaçadas Chiropotes satanas satanas (Cebidae: Primates) e de
de Extinção.” Portaria 444, de 17 de dezembro de 2014. Outros Mamíferos na Amazônia Oriental. Master’s thesis,
Ministério do Meio Ambiente. Diário Oficial da União, Universidade Federal do Pará, Belém.
Seção 1, número 245: 121–126. Lopes, M.A. and Ferrari, S.F. (1993). Primate conservation in
Brazil, MMA. (2017). Aprova o Plano de Ação Nacional para eastern Brazilian Amazonia. Neotrop. Primates 1: 8–9.
a Conservação dos Primatas Amazônicos – PAN Primatas Oliveira, S.G., Alfaro, J.W. and Veiga, L.M. (2014). Activity
Amazônicos. Portaria 792, de 01 de dezembro de 2017. budget, diet, and habitat use in the critically endangered
Ministério do Meio Ambiente. Diário Oficial da União, Ka’apor capuchin monkey (Cebus kaapori) in Pará State,
Seção 1, número 232: 24–61. Brazil: a preliminary comparison to other capuchin
Buss, G., Fialho, M.S., Jerusalinsky, L., Azevedo, R.B., Alves, monkeys. Am. J. Primatol. 76(10): 919–931.
S.L., Vidal, M.D. and Mendonça, E.N. (2017). Abundância Queiroz, H.L. (1992). A new species of capuchin monkey,
e densidade de primatas na Reserva Biológica do Gurupi, genus Cebus Erxleben 1977 (Cebidae, Primates), from
Maranhão, Brasil. Biodiv. Brasil. 7(2): 47–57. eastern Brazilian Amazonia. Goeldiana Zool. 15: 1–13.
Butti, M., Pacca, L., Santos, P.M., Alonso, A.C., Buss, G., Rylands, A.B. and Mittermeier, R.A. (2013). Ka’apor capuchin
Ludwig, G., Jerusalinsky, L. and Martins, A.B. In prep. Cebus kaapori. In: R.A. Mittermeier, A.B. Rylands and D.E.
Habitat loss estimation to assess terrestrial mammal Wilson (eds.), Handbook of the Mammals of the World.
species extinction risk: an open data framework. Volume 3. Primates, p. 410. Barcelona: Lynx Edicions.
Campos, I.B. (2009). Identificando Vacíos Clave de Silva Jr., J.S. and Cerqueira, R. (1998). New data and a
Información y Posibles Acciones Conservacionistas a historical sketch on the geographical distribution of
través de un Análisis de Viabilidade Poblacional para the Ka’apor capuchin, Cebus kaapori Queiroz, 1992.
Cebus kaapori Queiroz (1992) (Cebidae-Primate), un Neotrop. Primates 6: 118–121.
Primate del Este Amazónico en Peligro Crítico de Silva Jr., J.S., Lima, E.M., Camargo, C.C. and Ramos, R.M.
Extinción. Master’s thesis, Universitat de Barcelona, (2009). Mamíferos de médio e grande porte. In: M.A.
Barcelona, Spain. Monteiro, M.C.N. Coelho and E.J.S. Barbosa (eds.),
Cardoso, T.S. (2021). Ecologia Comportamental de Cebus Atlas Sócioambiental: Municípios de Tomé-Açu, Aurora
kaapori na Reserva Biológica do Gurupi, Maranhão. do Pará, Ipixuna do Pará, Paragominas e Ulianópolis,
Relatório Técnico. Programa de Capacitação Institucional pp. 125–300. Belém, Pará: Núcleo de Altos Estudos
(PCI/CNPq). Belém, PA: Museu Paraense Emílio Goeldi. Amazônicos (NAEA).
Carvalho, O., Jr, de Pinto, A.C.B. and Galetti, M. (1999). Silva Jr., J.S., Ohana J.A.B., Silva, C.R., Cardoso, E.M., Avelar,
New observations on Cebus kaapori in eastern Brazilian A.A., Silva, V.F. and Silva, L.S. (2010). Mamíferos terrestres
Amazonia. Neotrop. Primates 7: 41–43. de médio e grande porte no litoral da Amazônia
Cunha, F.A., Lopes, M.A., Dantas, S. de M., Carmo, N.A.S. Brasileira. In: L.M. Pessoa, W.C. Tavares and S. Siciliano
do and Silva S. do S.B. da. (2007). Registro de ocorrência (eds.), Mamíferos de Restingas e Manguezais, pp. 19–44.
de Cebus kaapori (Cebidae: Primates) na APA Lago de Rio de Janeiro: Sociedade Brasileira de Mastozoologia.
Tucuruí. Neotrop. Primates. 14: 84–85. Museu Nacional do Rio de Janeiro.
da Silva, L.B., Oliveira, G.L., Frederico, R.G., Loyola, R.,
Zacarias, D., Ribeiro, B.R. and Mendes-Oliveira, A.C.
(2022). How future climate change and deforestation can
drastically affect the species of monkey endemic to the
eastern Amazon, and priorities for conservation. Biodiv.
Conserv. 18pp. doi.org/10.1007/s10531-022-02373-1
Ferrari, S.F. and Lopes, M.A. (1996). Primate populations in
eastern Amazonia. In: M.A. Norconk, A.L. Rosenberger
and P.A. Garber (eds.), Adaptive Radiations of Neotropical
Primates, pp. 53–67. New York: Plenum Press.
Ferrari, S.F. and Queiroz, H.L. (1994). Two new Brazilian
primates discovered, endangered. Oryx 28: 31–36.
Ferrari, S.F. and de Souza Jr., A.P. (1994). More untufted
capuchins in southeastern Amazonia? Neotrop. Primates
21: 9–10.
Fialho, M.S., Moura, E.F., Ravetta, A.L., Laroque, P.O. and
Queiroz, H.L. (2018). Avaliação do Risco de Extinção de
Cebus kaapori Queiroz, 1992. In: Instituto Chico Mendes
de Conservação da Biodiversidade (ed.), Livro Vermelho
da Fauna Brasileira Ameaçada de Extinção: Volume II –
Mamíferos, pp. 253–259. Brasília: MMA, ICMBio.

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© Alejandra Niño/Proyecto Washu

110
 ECUADORIAN WHITE-FRONTED
CAPUCHIN
Cebus aequatorialis J.A. Allen, 1914

Ecuador, Peru
(2018, 2022)

Stella de la Torre, Fanny Cornejo, María Fernanda Solórzano, Laura Cervera, Diego G. Tirira,
Felipe Alfonso-Cortes, Nathalia Fuentes, Esteban Rivera and Mónica Alzamora

The Ecuadorian White-fronted Capuchin, The reproductive biology of C. aequatorialis is

Cebus aequatorialis, is classified as Critically unknown. In other species of the genus, sexual
Endangered on the IUCN Red List of Threatened maturity in females occurs when they are 4–7
Species (Tirira 2011, 2021a, Moscoso et al. 2021). years old and one year later in males. However,
It is found in western Ecuador and northwestern both sexes only reach adult body size at about
Peru from sea level to 2420 m (Tirira 2017, Brito 15 years old (Rylands et al. 2013). Considering
et al. 2018, Moscoso et al. 2021). In the north of that successful reproduction usually occurs only
its range, most records now are south of the Río when animals have attained adult body size, the
Guayllabamba, and its southernmost distribution generation time is estimated to be about 15–16
appears to be the Cerros de Amotape National years (Moscoso et al. 2021).
Park in Peru (Tirira 2017, Cervera et al. 2018a,
2018b, Moscoso et al. 2021). The species is The main threats to C. aequatorialis are forest loss
included in CITES Appendix II and it is illegal to and fragmentation, which have been particularly
hunt or trade C. aequatorialis in both Ecuador severe in western Ecuador. About 70% of the
(Tirira 2011) and Peru (SERFOR 2015). original forest cover in this region has been
converted, mainly for agriculture and ranching
The species inhabits tropical and subtropical (Ministerio del Ambiente 2012, Sierra 2013,
forests of the Chocó and Tumbes eco-regions Gonzalez-Jaramillo 2016, Cervera et al. 2018a).
(Albuja 2002, Albuja and Arcos 2007). It occurs in It is estimated that the species distribution has
primary and secondary forest as well as orchards been reduced to less than 1% of its original
and occasionally goes to the ground (Campos range in the last few decades (Albuja and Arcos
and Jack 2013, Tirira 2017). It feeds mainly on 2007), and different modelling methods have
mature fruits, complementing its diet with animal indicated that only 5,000 km² (Campos and
prey (insects, eggs, and small vertebrates). In Jack 2013) or 8,600 km² (Albuja et al. 2018) of
Ecuador, about 30 different plant species are suitable habitat remains. A recent assessment by
known to be part of its diet (Albuja et al. 2018). Tirira (2021b) estimated that the remaining area
Groups vary in size from 5 to 20 individuals. The of native forests for the species in Ecuador is
adult sex ratio appears to be 1:1 or mildly skewed 10,701 km², representing an 83% reduction of its
towards females 0.8:1 (Jack and Campos 2012, historical area of occurrence and a 75% reduction
Rylands et al. 2013). Group home range size of its suitable habitat (suitable habitat within the
appears to be large, at about 500 ha (Jack and area of occurrence). The author found that only
Campos 2012, Rylands et al. 2013, Cervera et al. 39% of the remaining area is under protection
2018b), but more studies are needed to confirm (national and private protected areas) and, based
this estimation. on current deforestation trends, estimated an
additional 7% loss of suitable habitat by 2050.

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NEOTROPICS

To date, however, the presence of the species

in most of this estimated area has not been
confirmed.

Cebus aequatorialis is considered a pest in

plantations of corn as well as bananas, plantain,
and cacao, and hence is persecuted, poisoned,
and hunted. In some areas of mangrove, local
people see it as a competitor in crab-hunting
and persecute it. Captive animals have been
observed in local villages in western Ecuador
(Moscoso et al. 2021). The species is also used
as a tourist attraction in some areas, such as the
western slopes of the Cotopaxi province, where
capuchins have been habituated to approach the
vicinities of road restaurants to obtain food. The
owners of the restaurants give visitors bananas
and other fruits to attract the capuchins (F.
Alfonso-Cortes and N. Fuentes, pers. obs.).

In Ecuador, C. aequatorialis has mostly been

reported to occur in public and private protected
areas (see below), which are the only sites with
sufficient forest to support the species. In
disturbed areas, i.e., most localities in Ecuador, the
species is elusive, tending to flee upon sighting.
In a census of four species of western Ecuadorian
primates carried out from October 2016 – March
2017, in 83 localities of 13 provinces, only 13
out of 260 records (5%) were of C. aequatorialis
(Cervera et al. 2018a). Surveys from previous
years evidenced a relatively wide variability
in local abundance (Cervera et al. 2018b). In
central-western Ecuador, Jack and Campos (2012)
estimated densities of 2–22 ind/km² (mean 2.4
ind/km²). In central and southwestern Ecuador,
Albuja and Arcos (2007) estimated densities of
3.5 and 3.9 ind/km².

Information about the species’ demography and

distribution in western Ecuador has been provided
by Albuja and Arcos (2007), Jack and Campos
(2012), Campos and Jack (2013) and Cervera et
al. (2018a). According to a recent assessment
carried out by Tirira (2021b), the habitat of this
species is severely fragmented. The largest
forest fragment, located between Cordillera
Chongón-Colonche and Machalilla National Park,
corresponds to 21% of the suitable habitat; the
average area for fragments larger than 1 km² (n =
730) is about 13 km². It is predicted that about 8%

112
of the species suitable habitat would be lost by data from this study, they estimated a density of
2050 due to climate change (Tirira 2021b). 1.36 ind/km². The species has been recorded by
park rangers in a conservation unit adjacent to
A recent survey determined two conservation the Cerros de Amotape National Park called the
priority areas for the species, covering a total Angostura Faical Regional Conservation Area.
of 129 km² of forest in the Azuay province, in
southwestern Ecuador. Connectivity between the Given the degree of fragmentation across the
two areas is limited, however, and the possibility species’ range, targeted efforts are required to
of mining activities taking place in these areas better understand its relationship with humans
creates a complex scenario for the species’ viability and how the disturbed landscape affects its
(Tirira and Gallo-Viracocha 2021). In primate demography, ecology, and behavior. The
censuses carried out by Proyecto Washu (PW) in a Peruvian population in the protected areas can
600-ha forest fragment in the canton Flavio Alfaro be used as a baseline for comparisons. This
of the Manabí province in 2019, C. aequatorialis information is imperative for conservation
was encountered three times (Duch-Latorre et al. planning, so that actions such as the creation
2019). In 2020, PW surveyed 41 forest fragments of habitat corridors, conservation education,
(ranging in size from 10 to 100 ha) in northern and and resolution of human-wildlife conflicts can
eastern Manabí and recorded the presence of the be undertaken. Improving forest connectivity
species in only four fragments (9.7%), with a total along Ecuador and Peru’s border is imperative to
of six encounters. The number of individuals maintain the species in both countries (Hurtado
in the sightings ranged from 5 to 12 individuals et al. 2016). In 2019, a bi-national collaborative
(Rivera et al. 2020). In November 2021, PW and project led by the Universidad San Francisco de
Naturaleza y Conservación Internacional (NCI) Quito, in Ecuador, and Fundación Yunkawasi,
carried out a primate survey in the Cazaderos in Perú, was initiated to address these research
Private Reserve, in Loja province; but the species and management priorities. Although several
was not found (Vega et al. 2022). Other, smaller, project activities that were planned for 2020
short-term studies have provided information and 2021 were not carried out because of the
about local abundance and conservation threats COVID-19 pandemic, local field assistants were
(Moscoso-Silva 2013, Solórzano 2014, Cervera et able to intermittently monitor two groups of the
al. 2015), but the species remains poorly studied species in the fragmented Ecuadorian forests
across most of its potential range. of La Libertad parish, in the El Oro province,
close to the Peruvian border. Data from these
In Peru, studies on C. aequatorialis are scarce. monitoring efforts are being analyzed and will be
It is known to occur only in government published in 2022 (de la Torre et al. in prep.). The
protected areas that provide a certain degree activities of this bi-national project are expected
of protection. However, there is very little to resume in 2022.
information on its status in these protected areas
and limited capacity to monitor them. In these The species has been reported to occur in various
areas, Hurtado et al. (2016) reported a group public and private protected areas. In Ecuador
size of 3–12 individuals and an encounter rate (Cervera et al. 2018b) these are: Chocó Andino
of 0.3 ind/km (based on 7 sightings during 112 de Pichincha, Parque Nacional Machalilla,
km of transects). Previously, group sizes of 3–5 Reserva Ecológica Los Ilinizas, Reserva Ecológica
individuals were reported in 1980 (Saavedra Mache-Chindul, Reserva Ecológica Manglares
and Velarde 1980) and 1994 (Encarnación and Churute, Refugio de Vida Silvestre Manglares
Cook 1998). Improving forest connectivity along Estuario Río Muisne, Refugio de Vida Silvestre
Ecuador and Peru’s border is imperative to Marino y Costera Pacoche, Área importante para
maintain the species in both countries (Hurtado las Aves Tito Santos, Bosque Protector Puyango,
et al. 2016). Between October 2019 and April Bosque Protector Bellavista, Bosque Protector
2020, Fundación Yunkawasi observed and Buenaventura, Bosque Protector Cambugán,
followed a group of 10 individuals and a single Bosque Protector Cerro de Hayas, Bosque
male in the sector known as Campo Verde inside Protector Cerro Blanco, Bosque Protector
the Cerros de Amotape National Park. With the Jama-Coaque, Bosque Protector Yaku Sinchi,

113
 NEOTROPICS

Bosque Protector Jauneche, Bosque Protector Forest Fragment Rio de Oro, Ecuador. Technical report.
La Hesperia, Bosque Protector La Otonga, Washington, DC: Primate Action Fund, Re:wild.
Ecuador, Ministerio del Ambiente. (2012). Línea Base
Bosque Protector Lalo Loor, Bosque Protector de Deforestación del Ecuador Continental. Quito,
Maquipucuna, Bosque Protector Mashpi, Bosque Ecuador.
Protector Mindo- Nambillo, Bosque Protector Encarnación, F. and Cook, G. (1998). Primates of the tropical
Río Guajalito. In Peru, it is reported from Cerros forest of the Pacific Coast of Peru: the Tumbes Reserved
Zone. Primate Conserv. (18): 15–20.
de Amotape National Park, Angostura Faical Gonzalez-Jaramillo, V., Fries, A., Rollenbeck, R., Paladines,
Regional Conservation Area, and Tumbes J., Onate-Valdivieso, F. and Bendix, J. (2016). Assessment
National Reserve. Finally, it is also reported from of deforestation during the last decades in Ecuador using
Bosques de Paz, a bi-national reserve of Ecuador NOAA-AVHRR satellite data. Erdkunde. 70(3): 217–235.
and Peru. Jack, K.M. and Campos, F.A. (2012). Distribution, abundance,
and spatial ecology of the Critically Endangered
Ecuadorian Capuchin (Cebus albifrons aequatorialis).
Trop. Conserv. Sci. 5(2): 173–191.
Hurtado, C.M., Serrano-Villavicencio, J.S. and Pacheco, V.
Albuja, L. (2002). Mamíferos del Ecuador. In: G. Ceballos (2016). Population density and primate conservation in
and J.A. Simonetti (eds.), Diversidad y Conservación the Noreste Biosphere Reserve, Tumbes, Peru. Revista
de Los Mamíferos Neotropicales, pp. 271–327. Mexico Peru. Biol. 23(2): 151–158.
City: CONABIO / Universidad Nacional Autónoma de Moscoso-Silva, F.J. (2013). Estado de Conservación del
México. Hábitat del Mono Machín (Cebus albifrons aequatorialis)
Albuja, L. and Arcos, R. (2007). Evaluación de las poblaciones en la Localidad de Las Tolas, Noroccidente del Distrito
de Cebus albifrons cf. aequatorialis en los bosques Metropolitano de Quito, Provincia de Pichincha.
suroccidentales ecuatorianos. Revista Politécnica. 27(4), Bachelor’s dissertation, Pontifica Universidad Católica del
Biología 7: 58–67. Ecuador, Quito, Ecuador.
Albuja, L., Moreno-Cárdenas, P. and Solórzano, M.F. (2018). Moscoso, P., de la Torre, S., Cornejo, F.M., Mittermeier, R.A.,
Aspectos taxonómicos y ecológicos del capuchino Lynch, J.W. and Heymann, E.W. (2021). Cebus aequatorialis
ecuatoriano, Cebus albifrons aequatorialis (Primates: (amended version of 2020 assessment). The IUCN Red
Cebidae) en el Ecuador. In: B. Urbani, M. Kowalewski, List of Threatened Species 2021: e.T4081A191702052.
R.G.T. Cunha, S. de la Torre and L. Cortés-Ortiz (eds.), Peru, SERFOR. (2015). Ley Forestal y de Fauna Silvestre
La Primatología en Latinoamérica 2 – A Primatología Ley Nº 29763 y sus Reglamentos. Depósito Legal en
na America Latina 2. Tomo II Costa Rica-Venezuela, pp. la Biblioteca Nacional del Perú Nº 2015-17751. Lima,
411–426. Caracas, Venezuela: Ediciones IVIC, Instituto Peru: Servicio Nacional Forestal y de Fauna Silvestre
Venezolano de Investigaciones Científicas. (SERFOR).
Brito, J., Garzón-Santomaro, C., Mena-Valenzuela, P., Rivera, E., Estay-Araya, S., Alfonso-Cortes, F. and Fuentes, N.
González-Romero, D. and Mena-Jaén, J. (2018). (2020). Monitoreo de Primates en la Línea de Transmisión
Mamíferos de la Provincia de El Oro: Una Guía de La Concordia – Pedernales 230kV. Unpublished report.
Identificación de Especies de Mamíferos Páramo al Mar. Quito, Ecuador: Corporación Eléctrica Nacional.
Serie de Publicaciones Misceláneas 8. Quito, Ecuador: Rylands, A.B., Mittermeier, R.A., Bezerra, B.M., Paim, F.P. and
Gobierno Autónomo Descentralizado de la provincia de Queiroz, H.L. (2013). Family Cebidae (squirrel monkeys
El Oro e Instituto Nacional de Biodiversidad. and capuchins). In: R.A. Mittermeier, A.B. Rylands and
Campos, F.A. and Jack, K.M. (2013). A potential distribution D.E. Wilson (eds.), Handbook of the Mammals of the
model and conservation plan for the Critically Endangered World. Volume 3: Primates, pp. 348–413. Barcelona: Lynx
Ecuadorian Capuchin, Cebus albifrons aequatorialis. Int. Edicions.
J. Primatol. 34(5): 899–916. Saavedra, C.J. and Velarde, D. (1980). Evaluación Preliminar
Cervera, L., Lizcano, D.J., Tirira, D.G. and Donati, G. (2015). de la Distribución y el Estatus del Coto de Tumbes
Surveying two endangered primate species (Alouatta (Alouatta palliata) en el Bosque Nacional de Tumbes.
palliata aequatorialis and Cebus aequatorialis) in the Unpublished report. Lima: Dirección General Forestal y
Pacoche Marine and Coastal Wildlife Refuge, West de Fauna. Ministerio de Agricultura.
Ecuador. Int. J. Primatol. 36(5): 933–947. Sierra, R. (2013). Patrones y Factores de Deforestación en
Cervera, L. et al. (2018a). Working together towards one el Ecuador Continental, 1990–2010, y un Acercamiento a
goal: results of the first primate census in western los Próximos 10 Años. Quito: Conservación Internacional
Ecuador. Primate Conserv. (32): 49–56. Ecuador y Forest Trends.
Cervera, L. et al. (eds.), Estado de Conservación de los Solórzano, M.F. (2014). Estado de Conservación de
Primates del Ecuador, pp. 54–63. Publicación Especial las Poblaciones del Capuchino Ecuatoriano (Cebus
sobre los mamíferos del Ecuador 12. Quito, Ecuador: aequatorialis), al Noroccidente del Distrito Metropolitano
Grupo de Estudio de Primates del Ecuador and de Quito (DMQ), Localidad de Las Tolas, Provincia de
Asociación Ecuatoriana de Mastozoología. Pichincha. Bachelor’s thesis, Universidad Central del
Duch-Latorre, I., Alfonso-Cortes, F., de la Torre, S. and Ecuador, Quito.
Strier, K. (2019). First Assessment of the Brown-headed Tirira, D.G. (ed.). (2011). Libro Rojo de los Mamíferos del
Spider Monkey, Ateles fusciceps fusciceps, in the Ecuador. 2nd edition. Publicación especial sobre los
mamíferos del Ecuador 8. Quito: Fundación Mamíferos

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 y Conservación, Pontificia Universidad Católica del
Ecuador and Ministerio del Ambiente del Ecuador.
Tirira, D.G. (2017). A Field Guide to the Mammals of
Ecuador. Publicación Especial sobre los mamíferos del
Ecuador 10. Quito, Ecuador: Asociación Ecuatoriana de
Mastozoología and Editorial Murciélago Blanco.
Tirira, D.G. (ed.) (2021a). Lista Roja de los mamíferos del
Ecuador. In: Libro Rojo de los Mamíferos del Ecuador.
3rd edition. Publicación Especial sobre los mamíferos del
Ecuador 13. Quito, Ecuador: Asociación Ecuatoriana de
Mastozoología, Fundación Mamíferos y Conservación,
Pontificia Universidad Católica del Ecuador and
Ministerio del Ambiente, Agua y Transición Ecológica del
Ecuador.
Tirira, D.G. (2021b). Primates del Ecuador: Aportes
al Conocimiento de su Diversidad, Distribución y
Conservación. Doctoral dissertation, Universidad de
Salamanca, Salamanca, Spain.
Tirira, D.G. and Gallo-Viracocha, F. (2021). Áreas prioritarias
para la conservación y vulnerabilidad al cambio climático
de Alouatta palliata aequatorialis (Atelidae) y Cebus
aequatorialis (Cebidae) en la provincia de Azuay, Ecuador.
Mammal. Aequatorialis 3: 37–57.
Vega, S., Alfonso-Cortes, F., Rivera, E., Fuentes, N., Nuñez,
D. and Sanmartín, D. (2022). Primer Censo de Primates
en el Bosque Seco de la Reserva Cazaderos, Provincia
de Loja. Unpublished report. Loja, Ecuador: Naturaleza y
Cultura Internacional.

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© Francielly Reis

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 GROVES’ TITI MONKEY
Plecturocebus grovesi Boubli et al. 2019

Brazil
(2022)

Jean Philippe Boubli, Gustavo Rodrigues Canale, Thiago B.F. Semedo,

Fabiano R. de Melo and Leandro Jerusalinsky

Groves’ Titi Monkey, Plecturocebus grovesi, the next 24 years, which will amount to 50%
occurs in the southern Brazilian Amazon and its under an optimistic “governance” scenario or to
ecotonal zone with the Brazilian Cerrado. It is a 86% under a “business-as-usual” scenario. The
member of the subfamily Callicebinae, the most latter is most likely, given the current process of
diverse group of Amazonian primates. protected area downgrading, downsizing, and
degazettement (PADDD) in the Brazilian Amazon
Titi monkeys of the genus Plecturocebus are and the planned complex of hydroelectric dams
small (about 1 kg). They are monogamous – the for this region (Bernard et al. 2014, Ferreira et
male provides much of the infant care (Spence- al. 2014, Pack et al. 2016, Fernandes et al. 2017,
Aizenberg et al. 2016), they duet and, when UHE Teles-Pires 2018). Based on these studies,
the pair are together, they tend to intertwine and other evidence, Groves’ Titi was classified
their tails. The Amazonian species use all forest as Critically Endangered on the IUCN Red List
strata, but forage and travel mostly in the dense of Threatened Species (Boubli et al. 2020). This
understory (0.5 to 10 m). They sometimes go to was substantiated by evidence that a future
the ground at times of fruit scarcity (Souza-Alves population reduction of at least 80% is suspected
et al. 2019). The small groups – an adult pair and to take place over the next three generations (c.
one or two offspring – have small home ranges, 24 years) (Veiga et al. 2011, Defler and García
generally less than 10 ha. Small fruits (whole 2012, Boubli et al. 2019, 2020). The species
fruits, pulp, aril, and seeds) and insects make was assessed as Endangered in the Brazilian
up the bulk of their diet (Bicca-Marques and national assessment, using the same criterion,
Heymann 2013). Plecturocebus grovesi has yet but adopting the “governance” scenario as more
to studied in the wild. probable, thus projecting a population reduction
of at least 50% in the next three generations
The range of Groves’ Titi is delimited by the (Brazil, ICMBio in prep.).
rios Juruena and Arinos in the west and the Rio
Teles-Pires in the east. To the north, the range Habitat destruction and fragmentation is
reaches the Juruena National Park, while the widespread in the species’ range. Its distribution
southernmost record is approximately 10°S. It is entirely within the Amazon’s arc of deforestation,
is probable that the Amazon forest – Cerrado the fast-advancing agribusiness frontier of Brazil.
ecotone – marks the limit to its range in the south This region contains the largest tropical ecotone
(about 13°S). More surveys, however, are needed in the world, where the megadiverse Amazon
to confirm this (Boubli et al. 2019). forests meet tropical savanna, the Brazilian
Cerrado, the latter one of the most threatened
Boubli et al. (2019) estimated that, to date, 42% biodiversity hotspots. Habitat restoration in
of its forest habitat has been lost (excluding the southwestern Amazon might be extremely
savannas), corresponding to 39% of the species’ difficult and costly in the following decades,
total distribution (forest and savannas). In due to the savannization of forests associated
addition, they forecast further habitat loss over with longer droughts and higher temperatures.

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NEOTROPICS

Members of the Mato Grosso state legislature

are trying to pass the ordinance law PL-337/2022
which will withdraw Mato Grosso state from
the Legal Amazon umbrella; all Brazilian States
within the Legal Amazon status are obliged by
law to preserve 80% of their forest cover. If this
new ordinance is approved, deforestation will
quickly escalate to a level that will put at risk
many southern Amazonian species including
the already Critically Endangered Groves’ Titi
Monkey.

Forest loss must be mitigated or avoided by the

creation of private and governmental reserves,
the enforcement of the law to protect forest set-
asides (‘Legal Reserves’), and the replacement of
large areas of chemical-dependent monocultures
of commodity crops by more sustainable models
of land use, such as agroforests and agroecological
food production. Forest degradation must
also be avoided by the prevention of fires and
logging, which has recently been stimulated by
land-grabbers incited by the lack of enforcement.
The geographic range of Groves’ Titi Monkeys
encompasses nine protected areas in Mato
Grosso, most of them located in the north
portion of the species distribution, but with only
one record to date for Groves’ Titi Monkeys in
one of them. These protected areas range from
indigenous lands (Kaiaby, Batelhão, Apiaká-
Kayabi and Apiaká do Pontal e Isolados), to
national parks (Parque Nacional do Juruena),
state reserves (Reserva Ecológica Apiacás) and
privately-owned, sustainable-use reserves (RPPN
Reserva Ecológica América Amazônia, RPPN
Reserva Ecológica Verde Amazônia). There are
no protected areas in the central and southern
portions of the species distribution.

The inclusion of Plecturocebus grovesi in the

National Action Plan for the Conservation of the
Amazonian Primates (Brazil/ICMBio 2017) – the
Brazilian public policy to establish conservation
strategies for threatened species – will take place
the moment the species is formally recognized as
under extinction risk and included in the National
Official List of Threatened Species (expected to
occur in 2022 or 2023). We suggest that Groves’
Titi Monkey be adopted as the flagship species
for Mato Grosso – the species is endemic to this
state, it is charismatic, and it is now listed here as
one of the World’s 25 Most Endangered Primates.

118
We can use the species to raise local awareness
in order to mitigate threats to the forests where
the titi still struggles to survive in the Brazilian
epicenter of agribusiness.

Bernard, E., Penna, L.A.O. and Araujo, E. (2014).

Downgrading, downsizing, degazettement, and
reclassification of protected areas in Brazil. Conserv. Biol.
28: 939–950.
Bicca-Marques, J.C. and Heymann, E.W. (2013). Ecology
and behavior of titi monkeys (genus Callicebus). In: L.M.
Veiga, A.A. Barnett, S.F. Ferrari and M.A. Norconk (eds.),
Evolutionary Biology and Conservation of Titis, Sakis and
Uacaris, pp. 196–207. Cambridge: Cambridge University
Press.
Boubli, J.P., Byrne, H., da Silva, M.N.F., Silva-Júnior, J.,
Costa Araújo, R., Bertuol, F., Gonçalves, J., de Melo, F.R.,
Rylands, A.B., Mittermeier, R.A., Silva, F.E., Nash, S.D.,
Canale, G., Alencar, R. de M., Rossi, R.V., Carneiro, J.,
Sampaio, I., Farias, I.P., Schneider, H. and Hrbek, T. (2019).
On a new species of titi monkey (Primates: Plecturocebus
Byrne et al., 2016), from Alta Floresta, southern Amazon,
Brazil. Mol. Phylogenet. Evol. 132: 117–137.
Boubli, J., de Melo, F.R. and Rylands, A.B. (2020).
Plecturocebus grovesi. The IUCN Red List of Threatened
Species 2020: e.T172272064A172272430.
Brazil, MMA. (2017). Aprova o Plano de Ação Nacional para
a Conservação dos Primatas Amazônicos – PAN Primatas
Amazônicos. Portaria 792, de 01 de dezembro de 2017.
Ministério do Meio Ambiente. Diário Oficial da União,
Seção 1, número 232: 24–61.
Defler, T.R. and Garcia, J. (2012). Plecturocebus caquetensis.
The IUCN Red List of Threatened Species 2012:
e.T14699281A14699284.
Fernandes, G.W. et al. (2017). Dismantling Brazil’s science
threatens global biodiversity heritage. Perspect. Ecol.
Conserv. 15(3): 239–243.
Ferreira, J. et al. (2014). Brazil’s environmental leadership at
risk. Science. 346: 706–707.
Pack, S.M., Ferreira, M.N., Krithivasan, R., Murrow, J.,
Bernard, E. and Mascia, M.B. (2016). Protected Area
Downgrading, Downsizing, and Degazettement (PADDD)
in the Amazon. Biol. Conserv. 197, 32–39.
Souza-Alves, J.P. et al. (2019). Terrestrial behavior in titi
monkeys (Callicebus, Cheracebus, and Plecturocebus):
potential correlates, patterns, and differences between
genera. Int. J. Primatol. 40(4): 553–572.
Spence-Aizenberg, A., Di Fiore, A. and Fernandez-Duque, E.
(2016). Social monogamy, male-female relationships, and
biparental care in wild titi monkeys (Callicebus discolor).
Primates 57(1): 103–112.
UHE Teles-Pires. (2018). Available online: <http://www.
uhetelespires.com.br/site/?page_id=27#body>.
Accessed 1 July 2018).
Veiga, L.M., Boveda-Penalba, A., Vermeer, J., Tello-
Alvarado, J.C. and Cornejo, F. (2011). Plecturocebus
oenanthe. The IUCN Red List of Threatened Species
2011: e.T3553A9939083.

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© Gerson Buss

120
 BROWN HOWLER MONKEY
Alouatta guariba (Humboldt, 1812)

Argentina, Brazil
(2012, 2014, 2016, 2018, 2022)

Luciana I. Oklander, Gerson Buss, Júlio César Bicca-Marques, Zelinda B. Hirano,

Óscar M. Chaves, Márcia M. Assis Jardim, Mônica M. Valença-Montenegro
Sérgio L. Mendes, Leonardo G. Neves, Martin Kowalewski, Fabiano R. de Melo,
Anthony B. Rylands and Leandro Jerusalinsky

Alouatta guariba is endemic to the Atlantic uni-female, and multimale-multifemale groups

Forest in eastern Brazil and northeastern have been reported (Glander 2013). The size of
Argentina. In the south, its range is limited by the an adult male is 50–60 cm (head-body) and 52–67
Camaquã river basin in the state of Rio Grande cm (tail), while an adult female is 44–54 cm (head-
do Sul (Printes et al. 2001) and, in the past, to body) and 48–57 cm (tail). Adult males weigh
the north by the Rio Paraguaçu in the state of 5.3–7.2 kg and adult females weigh 4.1–5 kg
Bahia (Gregorin 2006). The western boundary (Glander 2013). Longevity is estimated at 15–20
is marked by the limits of the Atlantic Forest. years (Strier 2004). Females reach sexual maturity
In Argentina, the species only occurs in the at between three and six years and males at five
province of Misiones (Agostini et al. 2014), while years (Strier et al. 2001), the gestation period is
in Brazil it is present in the states of Rio Grande approximately 190 days (Steinmetz 2000, Jardim,
do Sul, Santa Catarina, Paraná, São Paulo, Rio de 2005). Females have singletons, with an interbirth
Janeiro, Minas Gerais, Espírito Santo, and Bahia interval of 9–22 months (Strier et al. 2001).
(Bicca-Marques et al. 2018, Neves et al. 2018).
Although somewhat uncertain, there are currently As for all the Atlantic Forest primates, the Brown
two subspecies recognized: the Southern Brown Howler has suffered extensive habitat loss since
Howler, A. guariba clamitans Cabrera, 1940, and the European arrival in Brazil more than five
the Northern Brown Howler, Alouatta g. guariba, centuries ago. The principal economic cycles
north of the rios Jequitinhonha or Doce (Rylands during colonization broadly devastated the forest
et al. 2000, Glander 2013). cover in this region, which today concentrates
around 70% of the Brazilian population (about
The Brown Howler is a folivore-frugivore, 150 million people), as well as the principal capital
including more or less fruit in its diet according to cities, with corresponding industrial activity and
seasonal availability (Neville et al. 1988, Chaves urbanization (Scarano and Ceotto 2015). In Brazil,
and Bicca-Marques 2013, 2016, Chaves et al. the Atlantic Forest has been reduced to 11.7%
2018). As such, Brown Howlers are important of its original coverage (Ribeiro et al. 2009). The
seed dispersers for numerous plant species remaining forest is immensely fragmented into
(Chaves et al. 2018). Home range size varies hundreds of thousands of patches, the great
between study sites but averages 13 ha (Fortes majority of which are 50 ha or less (Ribeiro et al.
et al. 2015). Ranges of 15 groups studied varied 2009), hence unsuitable to support viable Brown
from 1.8 to 33 ha (Miranda and Passos 2011). Day Howler Monkey populations in the long term.
range varies from 50 m to 1,677 m (Fortes et al. Being one of the largest primates in the Atlantic
2015). Groups average 4 to 6 individuals but Forest, the species has been extensively hunted,
can be as large as 13 (Jardim 2005, Miranda and and also suffers to some extent from the pet
Passos 2005, Ingberman et al. 2009). Uni-male, trade.

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NEOTROPICS

Disease epidemics are an additional and very

serious threat. Howlers are highly susceptible to
yellow fever, and two recent outbreaks (2008/2009,
2016/2021), have severely affected their numbers
throughout the Atlantic Forest (Holzmann et al.
2010, Almeida et al. 2012, Bicca-Marques et al.
2017, Silva et al. 2020, Andrade et al. 2021). Due to
misinformation and the dissemination of the fear
that humans could be infected directly through
contact or proximity with monkeys, howlers were
persecuted, with many injured and killed during
the outbreaks (Bicca-Marques and Freitas 2010,
Bicca-Marques et al. 2017, Bicca-Marques 2018).
In the next few decades, pathogen exposure
could act synergistically with other threats such
as habitat loss, putting populations at high
extinction risk (Bicca-Marques et al. 2020).

In the south, Brown Howlers (Alouatta guariba

clamitans) occur in lowland forests along
Brazil’s coast as well as in higher elevation sub-
montane and montane forests and seasonal
semi-deciduous forests inland (Bicca-Marques
et al. 2018). In southern Brazil and northeast
Argentina, they also occupy a transition of mixed
Upper Paraná Atlantic Forest and Araucaria Moist
Forest (Miranda and Passos 2005, Agostini et al.
2014). Aguiar et al. (2007) recorded the species in
periodically flooded and semi-deciduous forests
in the Rio Paraná floodplains.

The primary threats are widespread forest loss and

fragmentation throughout the subspecies’ range
due to logging, agriculture, and cattle-ranching
(Bicca-Marques 2018). The most common causes
of death and injury of Brown Howlers in urban and
peri-urban (and even, rural) regions of southern
Brazil are electrocution on power lines (37%), dog
attacks (34%), vehicle collisions (17%), and human
mistreatments (12%) (Chaves et al. 2022).

The design and implementation of conservation

strategies for the Southern Brown Howlers in
urban and peri-urban regions (e.g., establishment
of urban protected areas/biological corridors,
installation of wildlife crossings, and insulating of
power lines) are crucial for the long-term survival
of these animals (Jerusalinsky et al. 2010, Printes
et al. 2010, Alfaya et al. 2020, Chaves et al. 2022).
Although some local population census data
are available, the total remaining population is
unknown but certainly declining. In Argentina, the

122
situation is even worse, and only few populations and dog attack in the urban and peri-urban
persist with no more than 20–50 adult individuals areas they inhabit (Chaves et al. 2022). Finally,
(Agostini et al. 2014, Moreno et al. 2015). the potential impact of COVID-19 and other
associated infectious diseases on the survival of
The Southern Brown Howler is listed as Vulnerable both Brown Howler subspecies is unknown, but it
on the Brazilian list of threatened fauna (Brazil, cannot be discounted. Further investigations on
MMA 2014, Bicca-Marques et al. 2018) and the topic are crucial.
as Critically Endangered on the Argentinian
Red list of mammals (Agostini et al. 2019). On There are protected areas in the Northern Brown
the IUCN Red List of Threatened Species, it is Howler’s range in the state of Bahia and north-
currently categorized as Vulnerable (Jerusalinsky eastern Minas Gerais, all created since 1980.
et al. 2021), but it may be a candidate for the Nevertheless, the only strictly protected area
Endangered category after the 2016–2021 yellow where the species has been confirmed is the
fever outbreak (Bicca-Marques et al. 2017). Mata Escura Biological Reserve (51,046 ha), just
north of the middle Rio Jequitinhonha (Melo
Recently a National Conservation Plan for the 2004, 2005).
Primates of Argentina was officially recognized
by the government. One of its specific objectives The two subspecies of Alouatta guariba are
includes, “Evaluating and reducing the impacts included in the Brazilian National Action Plan
of yellow fever on primates in Argentina.” for Conservation of the Atlantic Rainforest
Evaluation of the recovery of A. g. clamitans Primates and Maned Sloth (Brazil MMA 2018),
populations through an ex situ conservation and the southern subspecies also in the Primate
program is given high priority – analyzing the Conservation Action Plan of Argentina (Argentina,
possibility of starting a program of reproduction MADS 2021). These plans provide measures to
and reintroduction and/or translocation of identify important areas for conservation in order
specimens with the coordination and guidance to (a) restore, maintain and increase habitat and
of IUCN experts. its connectivity, (b) mitigate the impact of roads
and power lines, and (c) assess and mitigate the
The Northern Brown Howler Monkey (Alouatta impact of epizootics on the species.
guariba guariba) inhabits lowland, submontane,
and montane Brazilian Atlantic Forest. It has The National Center for Research and
a considerably more restricted range than Conservation of Brazilian Primates (ICMBio/
A. g. clamitans and is classified as Critically CPB) is planning to establish the “Population
Endangered both on the Brazilian list of Management Program for Alouatta guariba”
threatened fauna (Brazil MMA 2014, Neves et during 2022, as part of the Brazilian action plan
al. 2018) and the IUCN Red List of Threatened and following the new national official guidelines
Species (Neves et al. 2021). It has been listed as (Brazil ICMBio 2021). The main aim of this
one of the world’s 25 most threatened primates program is to promote population restoration,
since 2012 (Neves et al. 2017, Buss et al. 2019). to counteract recent population declines and
Adding the locations in the lower reaches of suspected local extinctions following the Yellow
the Jequitinhonha basin reported by Rylands Fever outbreaks. This program may result in
et al. (1988) and the small populations of A. g. outcomes contributing to the restoration of the
guariba discovered in the last decade, the total population in Misiones Province, Argentina, now
population is unlikely to sum more than 250 reduced to less than 100 individuals.
mature individuals, and no subpopulation is
believed to have more than 50 mature individuals
(Neves et al. 2017, 2018). Overall, the main
threats to the wild populations of this subspecies Agostini, I., Holzmann, I., Di Bitetti, M.S., Oklander, L.I.,
Kowalewski, M.M., Beldomenico, P.M., Goenaga, S.,
are habitat fragmentation, hunting, and the very
Martínez, M., Moreno, E.S., Lestani, E., Desbiez, A.L.J.
small sizes of the scattered populations (Neves and Miller, P. (2014). Building a species conservation
et al. 2021) and, like the southern subspecies, strategy for the brown howler monkey (Alouatta guariba
probably risk of electrocution, vehicle collision,

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clamitans) in Argentina in the context of yellow fever de 2021 – Instituto Chico Mendes de Conservação da
outbreaks. Trop. Conserv. Sci. 7: 26–34. Biodiversidade (ICMBio). Diário Oficial da União, Seção
Agostini, I., Holzmann, I., Oklander, L.I., Peker, S., Pavé, R. 1, número 122: 51.
and Kowalewski M. (2019). Alouatta guariba. In: SAyDS– Brazil, MMA. (2014). Lista Nacional Oficial de Espécies da
SAREM (eds.), Categorización 2019 de los mamíferos de Fauna Ameaçadas de Extinção. Portaria n° 444, de 17
Argentina según su riesgo de extinción. Lista Roja de los de dezembro de 2014 – Ministério do Meio Ambiente
mamíferos de Argentina. Accessed: 14 March 2022. (MMA). Diário Oficial da União, Seção 1, número 245:
Aguiar, L.M., Mellek, D.M., Abreu, K.C., Boscarato, T.G., 121–126.
Bernadi, I.P., Miranda, J.M.D. and Passos, F.C. (2007). Brazil, MMA. (2018). Aprova o Plano de Ação Nacional para
Sympatry between Alouatta caraya and Alouatta a Conservação dos Primatas da Mata Atlântica e da
clamitans and the rediscovery of free-ranging potential Preguiça-de-Coleira. Portaria n° 702, de 07 de agosto de
hybrids in southern Brazil. Primates 48: 245–248. 2018 – Ministério do Meio Ambiente. Diário Oficial da
Alfaya, L.B., Ilha, J.G., Pairet Jr., M.C., Trigo, T.C. and Jardim, União, Seção 1, número 153: 57.
M.M.A. (2020). Distribuição espacial e composição social Buss, G., Oklander, L.I., Bicca-Marques, J.C., Hirano,
de grupos de bugios-ruivos (Alouatta guariba clamitans) Z.B., Chaves, Ó.M., Mendes, S.L., Neves, G., Melo,
em fragmentos florestais no sul do Brasil. Neotrop. F.R., Rylands, A.B. and Jerusalinsky, L. (2019). Brown
Primates 26 (2): 33–39. howler monkey Alouatta guariba Humboldt, 1812. In: C.
Almeida, M.A.B., Santos, E., Cardoso, J.C., Fonseca, Schwitzer et al. (eds.), Primates in Peril: The World’s 25
D.F., Noll, C.A., Silveira, V.R., Maeda, A.Y., Souza, R.P., Most Endangered Primates 2018–2020, pp. 24–27. Bristol,
Kanamura, C. and Brasil, R.A. (2012). Yellow fever UK and Washington, DC: IUCN SSC Primate Specialist
outbreak affecting Alouatta populations in southern Group, International Primatological Society, Global
Brazil (Rio Grande do Sul State) 2008–2009. Am. J. Wildlife Conservation, and Bristol Zoological Society.
Primatol. 74: 68–76. Chaves, Ó.M. and Bicca-Marques, J.C. (2013). Dietary
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© Felipe Alfonso/Proyecto Washu

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 BROWN-HEADED SPIDER
MONKEY
Ateles fusciceps fusciceps Gray, 1866

Colombia, Ecuador, Panama

(2006, 2012, 2014, 2016, 2022)

Diego G. Tirira, Nathalia Fuentes, Felipe Alfonso-Cortes, Citlalli Morelos-Juárez,

Pedro G. Méndez-Carvajal, Karol M. Gutiérrez-Pineda, Sebastián Montilla,
Alma Hernández-Jaramillo and Alba-Lucía Morales-Jiménez

Brown-headed Spider Monkeys are found in insects. They eat flowers and shoots to a lesser
Central and South America, from southeast degree (Konstant and Rylands 2013, Morelos-
Panama to Ecuador, west of the Andes along Juárez et al. 2015, Alfonso-Cortes et al. 2022).
the Chocó Ecoregion (Konstant and Rylands As is true for the genus, Brown-headed Spider
2013). They occur mostly in evergreen humid Monkeys are effective seed dispersers (Morelos-
tropical and subtropical forests (Tirira 2017), Juárez et al. 2018a) and key for the maintenance
living in groups of up to 35 individuals with of high levels of tree diversity in Chocoan forests
fission-fusion dynamics (Eisenberg 1976, Link et (Calle-Rendón et al. 2016).
al. 2009), forming subgroups ranging from two to
ten individuals when they are searching for food Brown-headed Spider Monkeys can travel up to
(Gavilanez 2006, Estévez-Noboa 2009, Moscoso three km in a day (Morelos-Juárez et al. 2018a).
2010, Cueva and Pozo-Rivera 2010). They travel through the canopy by quadrumanous
and prehensile-tailed scrambling and brachiation
The species inhabits mainly large continuous but may also run on all fours along thick branches.
forest patches in primary or secondary forest Females becomes sexually mature at four to five
(Defler 2004, Tirira 2017), although there is a years old, but usually do not give birth before
population inhabiting fragmented landscapes seven or eight years of age. The gestation
in Ecuador (Cervera and Griffith 2016). Its period is 226 to 232 days with females normally
presence seems to be determined by its habitat giving birth to a single offspring every 2–4 years,
requirements, such as continuous canopy cover after which the youngster clings to its mother’s
and high abundance of large and tall trees (Tirira underside for the first few months (Eisenberg
et al. 2011). It prefers the highest levels of the 1973).
canopy but can also be observed at mid-levels,
and occasionally in the understory (Tirira 2017). Defler (2004) suggested that Ateles fusciceps
should be considered a subspecies of Geoffroy’s
The diets of Brown-headed Spider Monkeys Spider Monkey (Ateles geoffroyi) following
consist mainly of ripe fruits (70–90% of its diet) Collins and Dubach (2000). However, the
(van Roosmalen and Klein 1988, Konstant and taxonomic status of both species was further
Rylands 2013). Their preferred fruit is generally reviewed, and the evidence suggests that
hard with large pits, including those of various they are valid, distinct species (Rylands et al.
palms, although it also eats soft, small, and multi- 2006, Morales-Jiménez et al. 2015). Preliminary
seeded fruits (Morelos-Juárez et al. 2015, Fuentes genetic analyses from samples from the south
et al. 2018). They also eat fresh leaves, seeds, of Colombia and the north of Ecuador show two
aerial roots, and some invertebrates, such as

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NEOTROPICS

different monophyletic clades (Morales-Jiménez

et al. 2015), and two subspecies are recognized.

The nominate subspecies inhabits the Pacific

Coast of Ecuador and possibly southern
Colombia, in an altitudinal range of 20 to 2,300
m above sea level, but usually below 1,200 m
(Tirira 2017, 2021a). In Ecuador, its range includes
the northwestern Andes (Esmeraldas Province),
extending to the northwest of Pichincha and
Manabí provinces, and to the western borders
of Imbabura and Carchi provinces (Tirira 2004,
Cervera and Griffith 2016, Morelos-Juárez et al.
2018b). Its presence in Colombia is uncertain,
but it may be present south of the Río Mira, in
Nariño Department, southwestern Colombia
(Defler 2004).

Population density estimates in the buffer areas

of the Cotacachi-Cayapas National Park and the
Awa Ethnic Reserve were 0.2–13.2 individuals/
km² (Madden and Albuja V. 1989, Gavilanez 2006,
Moscoso 2010, Cueva and Pozo-Rivera 2010,
Albuquerque 2014, Urgilés-Verdugo and Gallo-
Viracocha 2016, Morelos-Juárez et al. 2018a). In
2010, a population size of 104 individuals was
calculated in an area of 18 km² located in areas
close to the Cotacachi-Cayapas National Park
(Moscoso 2010).

Due to its restricted range and the small size of the

natural populations, Ateles fusciceps fusciceps
is classified as Critically Endangered on the
IUCN Red List of Threatened Species (Moscoso
et al. 2021) and in The Red List of Mammals of
Ecuador (Tirira 2021b). Extensive deforestation,
the illegal pet trade, and hunting are the main
threats for the species in Ecuador (Tirira et al.
2011). For example, the loss of humid tropical
and subtropical rainforest in western Ecuador has
surpassed 80% of its original area (Ecuador, MAE
2012). Recent assessments indicate that the native
remaining forest for the species is from 7,231 to
8,812 km² (Tirira 2021b, Gallo-Viracocha et al. in
prep.). This is a reduction of approximately 64%
of its suitable habitat (only 34% of the remaining
forest is under protection). By 2050, the loss of
the suitable habitat may reach 73% (Tirira 2021a).

The Ecuadorian Brown-headed Spider Monkey

has been reported as extirpated in several
locations across its historical and current range,

128
including the type locality (Hacienda Chinipamba, Colombian border (Tirira et al. 2011, Morelos-
west of Ibarra, Intag Valley, Imbabura Province), Juárez et al. 2018a). However, occasional
the whole central coast of Ecuador, and the interviews with Awa elders and rapid biodiversity
surroundings of the ríos Cayapas, San Miguel, surveys conducted by the Ecuadorian NGO
Ónzole and Santiago, in the Esmeraldas Province Fundación Ecominga in neighboring areas to
(Tirira 2004). A survey conducted between 2008 the Awa Ethnic Forest Reserve, show that large
and 2009, found a solitary male of the Ecuadorian mammals, including primates, have been critically
Brown-headed Spider Monkey living with a group diminished or probably extirpated.
of Howler Monkeys (Alouatta palliata), in the area
of Los Bancos, Pichincha Province (Moscoso Conservation actions for the conservation of this
2010, Moscoso et al. 2019). subspecies began with the PRIMENET project
in 2005. This initiative performed research and
In the province of Manabí, the subspecies education activities focused on the primate
remains in several remnant forests of the Flavio species of northwestern Ecuador and led to
Alfaro, El Carmen, and Chone cantons (Cervera training courses for local guides to create groups
and Griffith 2016). Fifty-eight individuals of parabiologists in the communities (Peck et al.
were recorded in this area in at least 22 forest 2011).
fragments (each ranging in size from 1 to 1,000
ha), with population densities ranging from 7–22 Since 2012, Proyecto Washu/Fundación
individuals/km² (Alfonso-Cortes et al. 2022). Naturaleza y Arte has been working in priority
sites for the species, involving community-led
Despite the high deforestation rates that the development, scientific research, environmental
Ecuadorian Brown-headed Spider Monkey is education, and ex situ conservation activities. To
facing, and its extirpation in some areas, remaining address the drivers of habitat loss, the project
suitable habitat shows low fragmentation, with has implemented a Sustainable Matrix Model
82% of the available habitat in a single large (SMM) in the buffer zones of private and state
fragment. The average size of fragments larger reserves in Esmeraldas Province, which integrates
than 1 km² (n = 143) barely reaches 49 km² (Tirira the concepts of sustainable development,
2021a). The survival of the species in the future land sharing (Butsic and Kuemmerle 2015),
will also depend on the socio-economic situation and agroecological matrices (Perfecto and
of local human communities (Mosandl et al. 2008, Vandermeer 2010). These efforts have resulted
Pardo 2010). in 539 ha of the territory being protected by 18
farmer families through conservation agreements
The most important area for the conservation that improve their livelihoods (Abondano et al. in
of the Ecuadorian Brown-headed Spider press).
Monkey is concentrated in the Andean slopes
of the Esmeraldas Province (Cotacachi-Cayapas At the end of 2018, the Action Plan for Ecuadorian
National Park and neighboring areas), and Primate Species Conservation was published
adjacent regions of Imbabura Province (Tirira et and approved as a national guideline by the
al. 2015, Tirira 2021a). The buffer zone of this Environmental Ministry of Ecuador (Tirira et al.
national park has three small protected areas – 2018). It established a series of conservation
El Pambilar Wildlife Reserve and Río Canandé activities to implement in the next decade,
and Tesoro Escondido private protected forests mostly related to environmental policies, in situ
– mainly along the western border. Including and ex situ management, scientific research,
some surrounding unprotected forests, this is the environmental education, and control of illegal
area that is harboring the largest subpopulation wildlife trade and hunting.
of this subspecies in Ecuador (Peck et al. 2011,
Tirira 2021a). The Colombian Black Spider Money, Ateles
fusciceps rufiventris, was assessed as Critically
Another important forest where the species is Endangered in a previous iteration of the
thought to be present is the Awa Ethnic Forest IUCN Red List of Threatened Species (Cuarón
Reserve, north of the Río Mira and close to the et al. 2008, Tirira et al. 2017), but was recently

129
 NEOTROPICS

reassessed as Vulnerable (Link et al. 2021). It of the Darien National Park (Méndez-Carvajal et
remains as Critically Endangered in Panama al. 2021).
(the Panamanian List of Endangered Species;
Méndez-Carvajal 2019) and was assessed as
Endangered in the Libro Rojo de los Mamíferos
de Colombia (Defler et al. 2006). The reasons for Abondano, L., Webber, A., Valencia, L., Gómez-Posada,
C., Hending, D., Alfonso-Cortes, F. and Fuentes, N. (in
its threatened status are habitat loss and hunting
press). Community-based strategies to promote primate
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from case studies in South America. In: T. McKinney, S.
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Occidental of the Andes (Defler 2004). In Hábitat de la Comunidad de Primates en el Refugio de
Panama, its range extends from Lago Bayano Vida Silvestre “El Pambilar”, Provincia de Esmeraldas,
through the Panamanian Darien, and includes Ecuador. Bachelor’s dissertation, Universidad Nacional de
southeast Panama Province, northern San Blas, Piura, Piura, Ecuador.
Alfonso-Cortes, F., Fuentes, N., Rivera Román, E.S., Mogro, E.
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lower Río Cauca, along the west bank to south- abundances in northwestern Ecuador. Revista Biol. Trop.
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social integration in the black spider monkey (Ateles and GIS to Find Priority Areas for Conservation. Master’s
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Contrib. Zool. 213: 1–108. Morales-Jiménez, A.L., Cortés-Ortiz, L. and Di Fiore, A.
Estévez-Noboa, M.I. (2009). Estudio Poblacional y Uso de (2015). Phylogenetic relationships of Mesoamerican
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K., Cervera, L., Macías, Q., Loor, A., Kincez, K. and Urbina, spider monkey (Ateles fusciceps fusciceps) in the
S. (2018). Patrón de actividad, uso de estrato y dieta Ecuadorian Chocó: conflict between primates and
del mono araña de cabeza café (Ateles fusciceps) en loggers over fruiting tree species. PeerJ 3: e1574v1.
fragmentos de bosque del cantón Flavio Alfaro, provincia Morelos-Juárez, C., Moscoso, P., Alfonso-Cortes, F., Cervera,
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Congreso Ecuatoriano de Mastozoología, pp. 91–92. M.R., Tapia, A. and Tirira, D.G. (2018a). Mono araña de
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Técnica Particular de Loja. Conservación de los Primates del Ecuador, pp. 172–183.
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del Ecuador, Quito. D. and Peck, M.R. (2018b). Distribución actual, ecología
Konstant, W.R. and Rylands, A.B. (2013). Brown-headed y estrategias para la conservación de un primate
spider monkey Ateles fusciceps. In: R.A. Mittermeier, críticamente amenazado (Ateles fusciceps fusciceps) en
A.B. Rylands and D.E. Wilson (eds.), Handbook of the el Ecuador. In: B. Urbani, M. Kowalewski, R.G.T. Cunha,
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Link, A., Di Fiore, A. and Spehar, S.N. (2009). Female-directed 2. Tomo II. Costa Rica–Venezuela, pp. 441–452. Caracas:
aggression and social control in spider monkeys. In: M.N. Ediciones IVIC, Instituto Venezolano de Investigaciones
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on Male Aggression against Females, pp. 157–183. Ecuador suffers the highest deforestation rate in South
Cambridge, MA: Harvard University Press. America. In: E. Beck, J. Bendix, I. Kottke, F. Makeschin
Link, A., Defler, T.R., Méndez-Carvajal, P.G., Rodríguez, V., and R. Mosandl (eds.), Gradients in a Tropical Mountain
Cortés-Ortiz, L., Shanee, S. and Guzmán-Caro, D.C. Ecosystem of Ecuador, pp. 37–40. Berlin: Springer.
(2021). Ateles fusciceps ssp. rufiventris (amended version Moscoso, P. (2010). Estado Poblacional del Mono Araña
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Species 2021. e.T39921A191688429. del Ecuador, con Notas Ecológicas de Una Relación
Madden, R.H. and Albuja V.L. (1989). Estado actual de Ateles Interespecífica con Alouatta palliata. Bachelor’s
fusciceps fusciceps en el noroccidente ecuatoriano. Rev. dissertation, Pontificia Universidad Católica del Ecuador,
Politécnica 14: 113–157. Quito.
Méndez-Carvajal, P.G. (2012). Preliminary primate survey at Moscoso, P., Shanee, S., Burneo, S.F., Fuentes, N., Alfonso-
the Chucanti Nature Reserve, Darien Province, Republic Cortes, F., Obando, M. and Tirira, D.G. (2019). Prolonged
of Panama. Mesoamericana 16: 22–29. inter-specific association between Ateles fusciceps
Méndez-Carvajal, P.G. (2019). A Long-Term Monitoring fusciceps and Alouatta palliata aequatorialis (Atelidae)
Study to Evaluate the Primate Conservation Status in a forest fragment in north western Ecuador. Neotrop.
in Panama Using Species Distribution Modelling and Primates 25: 11–20.
Complementary Information. Doctoral dissertation, Moscoso, P., Link, A., de la Torre, S., Shanee, S. and Cortes-
Durham University, Durham, UK. Ortíz, L. (2021). Ateles fusciceps ssp.fusciceps (amended
Méndez-Carvajal, P.G., Batista, A., Jaramillo, O., Rodríguez- version of 2020 assessment). The IUCN Red List of
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Pardo, M. (2010). Cambio climático y pobreza: una mala Tirira, D.G., de la Torre, S. and Zapata Ríos, G. (eds.). (2018).
combinación. In: M. Pardo and M. Rodríguez (eds.), Plan de Acción para la Conservación de los Primates
Cambio Climático y Lucha Contra la Pobreza, pp. 1–24. del Ecuador. Quito: Ministerio del Ambiente del
Madrid: Fundación Carolina and Siglo XXI de España Ecuador, Grupo de Estudio de Primates del Ecuador and
Editores, S.A. Asociación Ecuatoriana de Mastozoología.
Peck, M.R., Thorn, J., Mariscal, A., Baird, A., Tirira, D.G. and Urgilés-Verdugo, C.A. and Gallo-Viracocha, F. (2016).
Kniveton, D. (2011). Focusing conservation efforts for Ocupación del mono bracilargo (Ateles fusciceps) en
the critically endangered brown-headed Spider Monkey la Reserva Ecológica Cotacachi-Cayapas. In: D.G. Tirira
(Ateles fusciceps) using remote sensing, modeling, and (ed.), Libro de Resúmenes, III Congreso Ecuatoriano
playback survey methods. Int. J. Primatol. 32: 134–148. de Mastozoología, pp. 141–142. Santa Elena, Ecuador:
Perfecto, I. and Vandermeer, J. (2010). The agroecological Asociación Ecuatoriana de Mastozoología, Ministerio del
matrix as alternative to the land-sparing/agriculture Ambiente del Ecuador and Universidad Estatal Península
intensification model. PNAS 107: 5786–5791. de Santa Elena.
Rylands, A.B., Groves, C.P., Mittermeier, R.A., Cortés-Ortiz, Van Roosmalen, M.G.M. and Klein, L.L. (1988). The spider
L. and Hines, J.J. (2006). Taxonomy and distributions of monkeys, genus Ateles. In: R.A. Mittermeier, A.B. Rylands,
Mesoamerican primates. In: A. Estrada, P. Garber, M. A.F. Coimbra-Filho and G.A.B. da Fonseca (eds.), The
Pavelka and L. Luecke (eds.), New Perspectives in the Ecology and Behavior of Neotropical Primates, Volume
Study of Mesoamerican Primates, pp. 29–79. New York: 2, pp. 455–537. Washington, DC: World Wildlife Fund –
Springer. U.S.
Tirira, D.G. (2004). Present status of the brown-headed
spider monkey (Ateles fusciceps Gray, 1866) (Primates:
Atelidae) in Ecuador. Lyonia 13: 1–8.
Tirira, D.G. (2017). A Field Guide to the Mammals of
Ecuador. Publicación Especial sobre los Mamíferos del
Ecuador 10. Quito, Ecuador: Asociación Ecuatoriana de
Mastozoología and Editorial Murciélago Blanco.
Tirira, D.G. (2021a). Primates del Ecuador: Aportes
al Conocimiento de su Diversidad, Distribución y
Conservación. Doctoral dissertation, Universidad de
Salamanca, Salamanca, Spain.
Tirira, D.G. (ed.) (2021b). Lista Roja de los mamíferos del
Ecuador. In: Libro Rojo de los Mamíferos del Ecuador.
3rd ed. Publicación especial sobre los mamíferos del
Ecuador 13. Quito, Ecuador: Asociación Ecuatoriana de
Mastozoología, Fundación Mamíferos y Conservación,
Pontificia Universidad Católica del Ecuador and
Ministerio del Ambiente, Agua y Transición Ecológica
del Ecuador.
Tirira, D.G., Gavilanez, M.M., Moscoso, P., Arcos, R., de la
Torre, S., Pozo-Rivera, W.E. and Peck, M.R. (2011). Mono
araña de cabeza marrón (Ateles fusciceps). In: D.G. Tirira
(ed.), Libro Rojo de los Mamíferos del Ecuador, 2nd ed.,
pp. 73–75. Publicación Especial sobre los Mamíferos
del Ecuador 8. Quito, Ecuador: Fundación Mamíferos
y Conservación, Pontificia Universidad Católica del
Ecuador and Ministerio del Ambiente del Ecuador.
Tirira, D.G., Morales-Jiménez, A.L. and Moscoso, P. (2015).
Ecuadorian Brown-headed Spider Monkey Ateles
fusciceps fusciceps Gray, 1866. In: C. Schwitzer et al.
(eds.), Primates in Peril: The World’s 25 Most Endangered
Primates, 2014–2016, pp. 76–77. Arlington, VA, Bristol, UK:
IUCN SSC Primate Specialist Group (PSG), International
Primatological Society (IPS), Conservation International
(CI), and Bristol Zoological Society.
Tirira, D.G., Méndez-Carvajal, P.G. and Morales-Jiménez,
A.L. (2017). Brown-headed spider monkey (Ateles
fusciceps Gray, 1866). In: C. Schwitzer et al. (eds.),
Primates in Peril: The World’s 25 Most Endangered
Primates, 2016–2018, pp. 83–87. Arlington, VA, Bristol,
UK: IUCN SSC Primate Specialist Group, International
Primatological Society, Conservation International and
Bristol Zoological Society.

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© Diego López Flores

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 GEOFFROY’S SPIDER MONKEY
Ateles geoffroyi Kühl, 1820

Belize, Costa Rica, El Salvador, Guatemala, Honduras, Mexico, Nicaragua,

Panama, and possibly Colombia
(2016, 2018, 2022)

Pedro G. Méndez-Carvajal, Melissa E. Rodríguez, Gilberto Pozo-Montuy, Óscar M. Chaves,

Ronald Sánchez-Porras, Karol M. Gutiérrez-Pineda, Denise Spaan, Eduardo J. Pinel Ramos
and Karla Zaldaña-Orantes

Geoffroy’s Spider Monkey, Ateles geoffroyi, occurs vulnerable to habitat loss, fragmentation, and
in Belize, Costa Rica, El Salvador, Guatemala, human presence than other Mesoamerican
Honduras, Mexico, Nicaragua, and Panama primates such as Alouatta, Saimiri, and Cebus
(Rylands et al. 2006). It is listed as Endangered (Méndez-Carvajal 2013, Solano-Rojas 2018,
(EN) on the IUCN Red List of Threatened Species Mansell and McKinney 2021). This species has one
because of a suspected population decline of of the largest distributions of the Mesoamerican
more than 50% over the next 45 years (three primates, and its populations are threatened by
generations), that is by 2063. Kellogg and high rates of deforestation, habitat loss, livestock
Goldman (1944) identified nine subspecies of A. farming, and hunting (Méndez-Carvajal et al.
geoffroyi, but three have been synonymized: A. 2019, Cortés-Ortiz et al. 2021).
g. pan (syn. of A. g. vellerosus; Silva-López et al.
1996); A. g. yucatanensis (syn. of A. g. vellerosus; Due to their high degree of fission-fusion
Silva-López et al. 1996, Morales-Jiménez et al. dynamics, surveying populations of Geoffroy´s
2015); and A. g. panamensis (syn. of A. g. ornatus; Spider Monkey is inherently difficult because
Napier, 1976; Morales-Jiménez et al. 2015). More subgroups change in size and membership
molecular information and sample locations are throughout the day. The situation is further
needed (Ruiz-García et al. 2016) to confirm the complicated by the diversity of landcover types
current taxonomy: A. g. azuerensis (Critically that these spider monkeys occur in, limiting the
Endangered; Méndez-Carvajal and Cortez-Ortíz, use of a single method for all sites. As such, a
2020); A. g. frontatus (Vulnerable; Solano et al. wide range of methods have been employed
2020a); A. g. geoffroyi (Critically Endangered; to locate populations and determine their size,
Williams-Guillén et al. 2020); A. g. grisescens including line-transect surveys (Serio-Silva et
(Data Deficient; Méndez-Carvajal and Cortez- al. 2006), camera trapping (Méndez-Carvajal
Ortíz, 2020), A. g. ornatus (Vulnerable; Solano 2014), and the use of drones (Spaan et al. 2019).
et al. 2020b), and A. g. vellerosus (Endangered; Although they provide valuable information on
Rosales-Meda et al. 2020). the species´ presence in an area, population
density estimates obtained from different survey
The genus Ateles has long been considered methods cannot be compared. Large gaps
to be the most threatened primate taxon in remain in our understanding of the distribution
the Neotropics (Weghorst et al. 2016). Ateles and numbers of Geoffroy´s Spider Monkeys
geoffroyi has a long gestation period (226–232 across their geographic range, hampering
days) compared to other Mesoamerican primates the identification of priority areas for their
(Melin et al. 2020) and 69–91% of its diet consists conservation. Funds dedicated to improving
of fruit (González-Zamora et al. 2009). Spider survey methods, training in survey techniques,
monkeys have large home ranges and are more

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and the interchange of expertise are urgently

needed to fill these gaps.

The Azuero Spider Monkey, A. g. azuerensis,

was studied for the first time in the regions of La
Vaca and Coto in Panama (Carpenter 1935). Its
range and population size have been assessed
by the NGO Fundación Pro-Conservación de
los Primates Panameños (FCPP), which has been
monitoring its populations since 2001. It has
been extirpated in Chiriqui Province and western
and northern Veraguas province, and appears to
be present only in the Azuero Peninsula – south-
west of Veraguas Province and the provinces
of Herrera and Los Santos. More precisely, it
is found in the southern areas near the Cerro
Hoya National Park, and in the fragmented
landscape between Punta Duarte, La Barra,
Guanico, Quema, La Tronosa Forest Reserve, La
Miel, and Pedasi. Only ten subgroups and five
complete groups have been detected, with a
mean of 3.8 individuals/subgroup (SE ±0.6, range
2–7) and a mean of 12.5 individuals/group (SE
±3.7, range 10–22). Their density in fragmented
habitats is estimated at 1.4 individuals/km²
(Méndez-Carvajal 2019), with an approximate
total population of <150 individuals (Méndez-
Carvajal and Ruiz-Bernard 2009, Méndez-Carvajal
2013). Conservation measures led by FCPP
involve community volunteers from Azuero, and
include environmental education, the creation of
an educational guide for the primates of Azuero,
primate surveys, and biodiversity monitoring
in the Azuero Peninsula (Méndez-Carvajal et al.
2013).

The Black-handed Spider Monkey, A. g. frontatus,

ranges from northern Nicaragua to northwestern
Costa Rica. In Costa Rica, they are found in the
Santa Rosa, Rincón de la Vieja, and Palo Verde
national parks and the Lomas Barbudal Biological
Reserve (Matamoros and Seal 1997). There are
some reports in the Murciélago and the Bahía
Salinas communities in Costa Rica (Sánchez-
Porras et al. 2018), Caño Negro National Wildlife
Refuge (Ó. M. Chaves, pers. obs.), and the basins
of the ríos Princapolca, Tuma and Ulice (Sánchez-
Porras et al. 2021). It has also been recorded in
Metagalpa and the Nicaraguan highlands (Allen
1914, Solano et al. 2020a).

136
The Nicaraguan Spider Monkey, A. g. geoffroyi, Maje, alongside other forests in the region where
was described from San Juan del Norte, the subspecies is believed to occur (Méndez-
Martina Bay, southeastern Nicaragua (Kellogg Carvajal 2021).
and Goldman 1944). Based on phenotypical
characteristics, it could be present in Los Chiles, The Ornate Spider Monkey, A. g. ornatus, also
El Concho, Upala, and Las Delicias in Costa Rica, known as the Panamanian Spider Monkey,
but further molecular analysis is needed (Morales- occurs in Costa Rica and Panama. In Costa Rica,
Jiménez et al. 2015, Williams-Guillén et al. 2020, it is known to be in the Osa Peninsula, Carara
Sánchez-Porras et al. 2021). Biological Reserve, and Corcovado National
Park (Matamoros and Seal 1997). There is an
Philip Sclater described the Hooded Spider introduced population of about 15 individuals
Monkey, A. g. grisescens, in a manuscript on Nancital Island, in the Isla Negritos Biological
catalogue of the mammals in the London Reserve, Golfo de Nicoya and Puntarenas, Costa
Zoological Gardens that he wrote in 1865. Gray Rica (Chaves et al. 2022). In Panama, this species
(1866) quoted Sclater’s diagnosis of the ‘Grizzled occurs on the northern side of the Caribbean
Spider Monkey’: “Fur moderately long, black, coast, Bocas del Toro, the northern coast of
with many silvery-white hairs interspersed; tail Veraguas Province. Although the species is also
black, underside greyish; hair of the forehead found in the Coclé and north Colón provinces,
moderately long; thumb none.” (p.732). He is it rarer due to Ngobe indigenous hunting and
attributed the authorship to Sclater. The type deforestation from mining (Méndez-Carvajal et
specimen is preserved in the British Museum al. 2019). It is present in the following reserves in
of Natural History, London (Gray 1866, Napier Panama: Palo Seco Protected Forest, La Amistad
1976). The type locality is unknown. Kellogg and International Park, the national parks of Santa
Goldman (1944) suggested it might hail from the Fe, Omar Torrijos, Chagres, Portobelo, and San
Río Tuyra basin, Panama, probably extending Lorenzo, and the San Blas Mountain Range and
southeastward through the Serranía del Sapo in some areas near the northern of Darien province.
extreme southeastern Panama and the Cordillera There is an introduced population of around
de Baudó of northwestern Colombia. Hernández- 20 individuals on Barro Colorado Island in the
Camacho and Cooper (1976) indicated that Panama Canal watershed (Milton and Hopkins
Ateles g. grisescens occurs in Colombia: “[In 2006).
Colombia, it] is known only from the vicinity of
Juradó very near the Panamanian border on the The Mexican Spider Monkey, A. g. vellerosus, is
Pacific coast. It is undoubtedly restricted by the present in Mexico, Belize, Guatemala, Honduras,
Baudó mountains to a narrow coastal strip that and El Salvador (Cortés-Ortiz et al. 2021). In
may extend as far south as Cabo Corrientes.” Mexico, the subspecies occurs in the states of
(p.66). Méndez-Carvajal and Cortés-Ortiz (2020) Veracruz, Oaxaca, Chiapas, Tabasco, Campeche,
confirmed that it had never been seen in the Quintana Roo, and Yucatán. The population
wild. In 2021, the FCPP team located a group of density of A. g. vellerosus is between 2.9 and
16 spider monkeys at Homenaje, between Ipetí- 9.3 individuals/km² in Montes Azules Biosphere
Maje and the Chucantí Private Natural Reserve Reserve in Marqués Comillas ejido, Chiapas,
on the Pacific side of eastern Panama, which Mexico (Chaves et al. 2011). Population densities
Méndez-Carvajal (2021) believes could be A. g. have been recorded as being between 2.14 to
grisescens. It is largely black, with a fringe of 40.8 ind/km2 in the Cañón del Sumidero National
whitish hairs on the chin and cheeks. FCPP and Park and Selva El Ocote Biosphere Reserve in
the Grupo de Investigación de Primatología from Chiapas (Pozo-Montuy et al. 2016) and between
Universidad de Panamá (GIP-UP) are carrying out 41.5 and 122.7 ind/km2 in the Areas Voluntarily
a genetic study to confirm the identity of these Dedicated to Conservation (ADVC) Cerro
spider monkeys. Conservation proposals and Chango and Soledad Vista Hermosa in Chinantla,
measures are underway together with Panama’s Oaxaca (Pozo-Montuy et al. 2016, Pozo-Montuy
Environment Ministry, including the possibility and Pinacho-Guendulain 2021). There are other,
of creating a protected area in the Serranía del difficult-to-access regions, where encounter
rates range between 0.5 and 1 individuals/

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 NEOTROPICS

km (in the El Triunfo, La Sepultura, and La industrial agricultural (e.g., oil palm plantations)
Encrucijada Biosphere Reserves in Chiapas and and infrastructure development (e.g., road and
in the Yum Balam Flora and Fauna Protection rail construction), and the primate pet trade. It is
Area in Quintana Roo; Pozo-Montuy et al. hunted for food by indigenous people at Marquéz
2018, Pinacho-Guendulain and Pozo-Montuy in de Comillas, Chiapas (Ó.M. Chaves pers. comm.).
press). The subspecies is believed to have been Conservation activities on its behalf include
extirpated in the Pantanos de Centla Biosphere environmental education, community science
Reserve (Pozo-Montuy et al. 2021), but is found and awareness campaigns, human-wildlife
in the Calakmul Biosphere Reserve, Bala´an conflict mitigation and the improvement of survey
K´aax and Otoch Ma´ax Yetel Kooh Fauna and methods (Spaan et al. 2019). Some educational
Flora Protected Areas and in human-modified materials are available (<www.cobius.org>).
landscapes throughout the Yucatán Peninsula These activities have been implemented by
(Spaan et al. 2020). the NGOs ConMonoMaya, Conservación de la
Biodiversidad del Usumacinta A.C., and Miku
In Guatemala it occurs in Petén, Alta Verapaz, Conservación A.C. in Mexico and Asociación
Baja Verapaz, Izabal, Sololá, Huehuetenango Territorios Vivos El Salvador (ATVES) in El Salvador.
and Quiché in Guatemala (Ponce-Santizo et al. Additionally, community monitoring with the
2009) and in Belize in the Runaway Creek Nature involvement of more than 200 local people has
Reserve (Hartwell et al. 2021). In El Salvador, its been carried out throughout southeast Mexico by
presence has been registered in Chaguantique, Conservación de la Biodiversidad del Usumacinta
Normandía, El Nacascolo and El Tercio (Usulután (COBIUS AC).
Department) and Montecristo, Cerro el Mono
and Conchagua in (Rodríguez-Menjívar 2007).
New records have been obtained of spider
monkeys at the Laguna Olomega and in Cerro El Allen, J.A. (1914). New South American monkeys. Bull. Am.
Caballito, Jucuarán, also in El Salvador (Pineda- Mus. Nat. Hist. 33: 647–655.
Carpenter, C.R. (1935). Behaviour of red spider monkeys in
Peraza et al. 2017; Pineda-Peraza et al. 2020).
Panama. J. Mammal. 16: 171–180.
The phylogenetic study by Morales-Jiménez Chaves, Ó.M., Stoner, K.E. and Arroyo-Rodríguez, V. (2011).
et al. (2015) using mtDNA failed to identify the Seasonal differences in activity patterns of Geoffroy’s
subspecies for El Salvador. It seems to form a spider monkeys (Ateles geoffroyi) living in continuous and
separate clade from Ateles geoffroyi vellerosus. fragmented forests in southern Mexico. Int. J. Primatol.
32: 960–973.
Chaves, Ó.M., Retana, M., Chacón, E., Obregón, A.,
The Maya Biosphere Reserve (MBR) in the north Mena, L. and Gómez, E. (2022). Evaluación Preliminar
of Guatemala, covering 2.2 million ha, is the de la Población de Monos Araña (Ateles geoffroyi) en
largest and most important protected habitat la Reserva Biológica Islas Negritos, Golfo de Nicoya,
for the subspecies (68.6% of its original forests). Puntarenas. Informe técnico. Escuela de Biología,
Universidad de Costa Rica, San José. 20pp. doi 10.13140/
Conservation actions are maintained by several
RG.2.2.19774.77121
organizations to preserve this important forest Cortés-Ortiz, L., Solano-Rojas, D., Rosales-Meda, M.,
block in Guatemala (Ponce-Santizo et al. 2009, Williams-Guillén, K., Méndez-Carvajal, P.G., Marsh,
Rosales-Meda et al. 2020). In Honduras, A. g. L.K., Canales-Espinosa, D. and Mittermeier, R.A. (2021).
vellerosus is in several protected areas, mainly Ateles geoffroyi (amended version of 2020 assessment).
The IUCN Red List of Threatened Species 2021:
those in the northern part of the country in the e.T2279A191688782.
departments of Atlantida, Colon, Olancho and Espinal, J.M., Mora, J.M., Ruedas, L.A., López, L.-I. and
Gracias a Dios (Hines 2005, Zambrano 2008, Marineros, L. (2016). A case of albinism in the Central
Guillén and Guillén 2013, Espinal et al. 2016). American spider monkey, Ateles geoffroyi, in Honduras.
Mastozool. Neotrop. 23: 63–69.
There are no data on population densities
Gray, J.E. (1866). Notice of some new species of spider
for Honduras. Hines (2005) reported a mean monkeys (Ateles) in the collection of the British Museum.
subgroup size of 6.93 ± 4.53 individuals in Pico Proc. Zool. Soc. Lond. 1865: 732–733.
Bonito National Park. González-Zamora, A., Arroyo-Rodríguez, V., Chaves, Ó.M.,
Sánchez-López, S., Stoner, K.E. and P. Riba-Hernández.
Ateles g. vellerosus is threatened by habitat (2009). Diet of spider monkeys (Ateles geoffroyi) in
Mesoamerica: current knowledge and future directions.
loss due to forest fires, subsistence farming and Am. J. Primatol. 71: 8–20.

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 OTHER SPECIES CONSIDERED
MADAGASCAR – AYE-AYE
Daubentonia madagascariensis (Gmelin, 1788)
Madagascar
(2016, 2018)
Timothy M. Sefczek, Jeannin Nicolas Rakotondrazandry, Doménico R. Randimbiharinirina, Faranky
Ravelomandrato, Herinandrasana Pollensio, Noa E. Rasoanaivo, Romeo Rakotondrina, Jonah
Ratsimbazafy, Brigitte M. Raharivololona, Solofonirina Rasoloharijaona, Herimalala Raveloson, John
Zaonarivelo, Aristide Andrianarimisa, George Perry, Felix Rakotondraparany and Edward E. Louis Jr.
The Aye-aye (Daubentonia madagascariensis) is the only extant species in the family Daubentoniidae
(Simpson 1945). Aye-ayes have the widest distribution of any Malagasy primate – they are absent
only from southwest of the island (Sterling and McCreless 2006). Their ecological flexibility means
they can persist in different forest types, such as moist evergreen and dry deciduous forests, and of
varying quality, including primary, disturbed, and heavily degraded habitats (Ancrenaz et al. 1994,
Randimbiharinirina et al. 2018, Sefczek et al. 2020a). Nevertheless, their huge individual home ranges
(Sefczek et al. 2020b) and presumed low population densities make them susceptible to continued
anthropogenic disturbance (Perry et al. 2012), i.e., forest degradation and fragmentation, slash-and-
burn agriculture, and hunting associated with local superstitions (Randimbiharinirina et al. 2021).
Ancrenaz, M., Lackman-Ancrenaz, I. and Mundy, N. (1994). Field observations of aye-ayes (Daubentonia madagascariensis)
in Madagascar. Folia Primatol. 62: 22–36.
Perry, G.H., Reeves, D., Melsted, P., Ratan, A., Miller, W., Michelini, K., Louis, Jr., E.E., Pritchard, J.K., Mason, C.E., and
Gilad, Y. (2012). A genome sequence resource for the aye-aye (Daubentonia madagascariensis), a nocturnal lemur from
Madagascar. Genome Biol. Evol. 4: 126–135.
Randimbiharinirina, D.R., Raharivololona, B.M., Hawkins, M.T.R., Frasier, C.L., Culligan, R.R., Sefczek, T.M., Randriamampionona,
R. and Louis, Jr., E.E. (2018). Behaviour and ecology of male aye-ayes (Daubentonia madagascariensis) in the Kianjavato
Classified Forest, south-eastern Madagascar. Folia Primatol. 89: 123–137.
Randimbiharinirina, D.R., Richter, T., Raharivololona, B.M., Ratsimbazafy, J.H. and Schüßler, D. (2021). To tell a different
story: unexpected diversity in local attitudes towards endangered aye-ayes Daubentonia madagascariensis offers new
opportunities for conservation. People and Nature 3: 484–498.
Sefczek, T.M., Randimbiharinirina, D.R., Raharivololona, B.M., Razafimahaleo, H., Randrianarison, O. and Louis, Jr., E.E.
(2020a). Re-assessing the applicability of the Jarman/Bell model and Kay’s threshold to the insectivorous aye-aye
(Daubentonia madagascariensis). Am. J. Phys. Anthropol. 171: 336–341.
Sefczek, T.M., Hagenson, R.A., Randimbiharinirina, D.R., Rakotondrazandry, J.N. and Louis Jr., E.E. (2020b). Home range size
and seasonal variation in habitat use of aye-ayes (Daubentonia madagascariensis) in Torotorofotsy, Madagascar. Folia
Primatol. 91: 558–574.
Simpson, G.G. (1945). The principles of classification and classification of mammals. Bull. Am. Mus. Nat. Hist. 85: 62.
Sterling, E.J. and McCreless, E. (2006). Adaptations in the aye-aye: a review. In: L. Gould and M.L. Sauther (eds.), Lemurs:
Ecology and Adaptation, pp. 161–186. New York, NY: Springer.

MADAGASCAR – LAKE ALAOTRA GENTLE LEMUR

Hapalemur alaotrensis Rumpler, 1975
Madagascar
(2000, 2014, 2016, 2018)
Lena M. Reibelt, Herizo T. Andrianandrasana, Richard Lewis, Fidy Ralainasolo, Jonah Ratsimbazafy,
Lucile Mialisoa Raveloarimalala and Patrick O. Waeber
The Critically Endangered Lake Alaotra Gentle Lemur (Hapalemur alaotrensis) is restricted to the
wetlands fringing Lake Alaotra, a Ramsar Site and protected area in north-eastern Madagascar.
This limited geographical range and ever-increasing pressures have brought the “bandro” close
to extinction. The first published census estimated some 7,500–11,000 individuals (Mutschler and

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Feistner 1995) but the population has since declined with only an estimated 2,500 individuals in 2005
(Ralainasolo et al. 2006). Subpopulations are isolated due to a high degree of habitat fragmentation
and the permanent conversion of marshes for agricultural production. A case in point is the priority
conservation zone “Park Bandro” (Ratsimbazafy et al. 2013, Raveloarimalala and Reibelt 2016), where
more than half of the area was lost due to a fire in 2021. Very few lemurs have been spotted since.
The principal threats to Hapalemur alaotrensis remain habitat loss, habitat degradation, and hunting,
which are exacerbated with extended droughts that result in an increased need for people to take
water from the lake and marsh system (Ralainasolo et al. 2006, Copsey et al. 2009a, 2009b, Guillera-
Arroita et al. 2010). Conservation and management of the Alaotra Protected Area (IUCN category V)
is challenging for many reasons but primarily due to overstretched resources which are insufficient to
fund the running of this vast protected area. In close partnership with the authorities, Durrell Wildlife
Conservation Trust and Madagascar Wildlife Conservation implement community-based projects
around the lake, focusing on supporting the local resource-management associations, participatory
ecological monitoring, and environmental education (Andrianandrasana et al. 2005, Rendigs et al.
2015, Waeber et al. 2018). For more information on the biology, ecology, and conservation efforts on
behalf of Hapalemur alaotrensis see Reibelt et al. (2017, 2019). Since the onset of COVID-19 in early
2020, eco-tourism has become obsolete, putting further strain on the monitoring and patrolling of
the vast conservation area.
Andrianandrasana, H.T., Randriamahefasoam J., Durbin, J., Lewis, R.E. and Ratsimbazafy, J.H. (2005). Participatory ecological
monitoring of the Alaotra wetlands in Madagascar. Biodiv. Conserv. 14: 2757–2774.
Copsey, J.A., Jones, J.P.G., Andrianandrasana, H., Rajaonarison, L.H. and Fa, J.E. (2009a). Burning to fish: local explanations
for wetland burning in Lac Alaotra, Madagascar. Oryx 43: 403–406.
Copsey, J.A., Rajaonarison, L.H., Ranriamihamina, R. and Rakotoniaina, L.J. (2009b). Voices from the marsh: livelihood
concerns of fishers and rice cultivators in the Alaotra wetland. Madagascar Conserv. Develop. 4: 25–30.
Guillera-Arroita, G., Lahoz-Monfort, J.J., Milner-Gulland, E.J., Young, R.P. and Nicholson, E. (2010). Monitoring and
conservation of the Critically Endangered Alaotran gentle lemur Hapalemur alaotrensis. Madagascar Conserv. Develop.
5: 103–109.
Mutschler, T. and A.T.C. Feistner. (1995). Conservation status and distribution of the Alaotran gentle lemur Hapalemur griseus
alaotrensis. Oryx 29: 267–274.
Ralainasolo, F.B., Waeber, P.O., Ratsimbazafy, J., Durbin, J. and Lewis, R. (2006). The Alaotra gentle lemur: population
estimation and subsequent implications. Madagascar Conserv. Develop. 1: 9–10.
Ratsimbazafy, J.R., Ralainasolo, F.B., Rendigs, A., Mantilla Contreras, J., Andrianandrasana, H., Mandimbihasina, A.R.,
Nievergelt, C.M., Lewis, R. and Waeber, P.O. (2013). Gone in a puff of smoke? Hapalemur alaotrensis at great risk of
extinction. Lemur News 17: 14–18.
Raveloarimalala, L.M. and Reibelt, L.M. (2016). Update on management of Park Bandro and population numbers of
Hapalemur alaotrensis. Lemur News 20: 2.
Reibelt L.M., Ratsimbazafy J. and Waeber P.O. (2017). Lac Alaotra bamboo lemur Hapalemur alaotrensis (Rumpler, 1975). In:
C. Schwitzer et al. (eds.), Primates in Peril: The World’s 25 Most Endangered Primates 2016–2018, pp. 32–34. Arlington,
VA, and Bristol, UK: IUCN SSC Primate Specialist Group, International Primatological Society, Conservation International
and Bristol Zoological Society.
Reibelt L.M., Andrianandrasana, H.T., Ralainasolo, F., Raveloarimalala, L.M., Lewis, R., Ratsimbazafy, J. and Waeber, P.O.
(2019). Lake Alaotra Gentle Lemur Hapalemur alaotrensis (Rumpler, 1975). In: C. Schwitzer et al. (eds.), Primates in Peril:
The World’s 25 Most Endangered Primates 2018–2020, pp. 9–11. Washington, D.C. and Bristol, UK: IUCN SSC Primate
Specialist Group, International Primatological Society, Global Wildlife Conservation and Bristol Zoological Society.
Rendigs, A., Reibelt, L.M., Ralainasolo, F.B., Ratsimbazafy, J.H. and Waeber, P.O. (2015). Ten years into the marshes –
Hapalemur alaotrensis conservation, one step forward and two steps back? Madagascar Conserv. Develop. 10: 13–20.
Waeber, P.O., Ratsimbazafy, J.H., Andrianandrasana, H., Ralainasolo, F.B. and Nievergelt, C.M. (2018). Hapalemur alaotrensis,
a conservation case study from the swamps of Alaotra, Madagascar. In: A.A. Barnett, I. Matsuda and K. Nowak (eds.),
Primates in Flooded Habitats: Ecology and Conservation, pp. 293–296. Cambridge, UK: Cambridge University Press.

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MADAGASCAR – GOLDEN-CROWNED SIFAKA
Propithecus tattersalli Simons, 1988
Madagascar
(2000)
Brandon Semel, Rio Heriniaina, Jordi Salmona and Meredith Semel
The Golden-Crowned Sifaka (Propithecus tattersalli) is Critically Endangered and endemic to forest
patches encompassed by the Loky and Manambato rivers in northeastern Madagascar (Vargas et al.
2000). Surveys conducted in 2006–2008 suggested a population of about 18,000 (estimated range of
11,000-26,000) (Quéméré et al. 2010). A more recent study estimated that 10,222-12,631 sifakas remain
and found fewer sifakas in all four forest patches surveyed in 2006–2008 (B. Semel et al. in review a).
Genetic studies suggest a small effective population size of 430–795 and a significant population
decline since the mid-Holocene transition about 4,200 years ago (Quéméré et al. 2012, Salmona et
al. 2017). Surprisingly, staff at Fanamby – the Malagasy NGO, that manages the Loky-Manambato
Protected Area – recently confirmed the presence of several groups of sifakas in the Andrafiamena
Andavakoera Protected Area, about 50 km to the west of the closest known populations, therefore
extending their known distribution. Threats include habitat loss due to slash-and-burn agriculture
and grassland fires, mining, and potentially hunting, all of which are likely to be exacerbated by
improvements to the national road that bisects their range (Anania et al. 2018, M. Semel et al. in
review b).
Anania, A., Salmona, J., Rasolondraibe, E., Jan, F., Chikhi, L., Fichtel, F., Kappeler, P.M., Rasoloarison, R. (2018). Taboo
adherence and presence of Perrier’s sifaka (Propithecus perrieri) in Andrafiamena forest. Madag. Conserv. Dev. 13(1):
6–14.
Quéméré, E., Champeau, J., Besolo, A., Rasolondraibe, E., Rabarivola, C., Crouau-Roy, B. and Chikhi, L. (2010). Spatial
variation in density and total size estimates in fragmented primate populations: the golden-crowned sifaka (Propithecus
tattersalli). Am. J. Primatol. 72: 72–80.
Quéméré, E., Amelot, X., Pierson, J., Crouau-Roy, B. and Chikhi, L. (2012). Genetic data suggest a natural prehuman origin
of open habitats in northern Madagascar and question the deforestation narrative in this region. PNAS 109(32): 13028–
13033.
Salmona, J., Heller, R., Quéméré, E. and Chikhi, L. (2017). Climate change and human colonization triggered habitat loss and
fragmentation in Madagascar. Mol. Ecol. 26: 5203–5222.
Semel, B.P., Karpanty, S.M, Semel, M.A., Stauffer, D.F., Quéméré, E., Walters, J.R., Andrianiaina, A.F., Rakotonanahary, A.N.,
Ranaivoson, T., Rasolonirina, D.V. and Vololonirina, F.F. (in review a). Highly variable densities and a decline in critically
endangered golden-crowned sifaka (Propithecus tattersalli) population abundance from 2008–2018.
Semel, M.A., Abernathy, H.N., Semel, B.P., Cherry, M.J., Ratovoson, T.J.C. and Moore, I.T. (in review b). Environmental and
anthropogenic influences on movement and foraging in a critically endangered lemur species, Propithecus tattersalli:
implications for habitat conservation planning.
Vargas, A., Jiménez, I., Palomares, F. and Palacios, M.J. (2002). Distribution, status, and conservation needs of the golden-
crowned sifaka (Propithecus tattersalli). Biol. Conserv. 108: 325–334.

MADAGASCAR – PERRIER’S SIFAKA

Propithecus perrieri Lavauden, 1931
Madagascar
(2000, 2002, 2004, 2014, 2016)
Rio Heriniaina, Hosnah Bezara Hortensia and Jhon Rigobert
Perrier’s Sifaka (Propithecus perrieri) is Critically Endangered (Ranaivoarisoa et al. 2006). The species
has a very restricted home range in northern Madagascar that includes the Analamerana Special
Reserve and Andrafiamena-Andavakoera Protected Area (Petter et al. 1977). It has been estimated
that only 100 to 2,000 individuals remain in the wild, with an effective population unlikely to exceed 230
(Bank et al. 2007). This species can be found in semi-evergreen forest. Perrier’s Sifaka is threatened by
hunting, selective logging, habitat destruction to make way for agriculture, heavy fires set to increase
pasture for livestock, and charcoal production (Schwitzer et al. 2006). More recently, mining has come
to be an important threat to this species.

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Banks, M.A., Ellis, E.R., Antonio and Wright, P.C. (2007). Global population size of a critically endangered lemur, Perrier’s
sifaka. Anim. Conserv. 10: 254-262.
Petter, J.-J., Albignac, R. and Rumpler, Y. (1977). Mammifères lémuriens (Primates prosimiens). Faune de Madagascar. 44:
1–513.
Ranaivoarisoa, J.F., Ramanamahefa, R., Louis, E.E. Jr. and Brenneman, R.A. (2006). Range extension of Perrier’s Sifaka,
Propithecus perrieri, in the Andrafiamena Classified Forest. Lemur News 11: 17–21.
Schwitzer, C., Arnoult, O. and Rakotosamimanana, B. (2006). An international conservation and research programme for
Perrier’s sifaka (Propithecus perrieri Lavauden, 1931) in northern Madagascar. Lemur News 11: 12–14.

AFRICA – KIPUNJI
Rungwecebus kipunji (Ehardt, Butynski, Jones & Davenport, 2005)
Tanzania
(2006, 2008, 2018)
Tim R.B. Davenport
The kipunji (Rungwecebus kipunji) was included in Primates in Peril: The World’s 25 Most Endangered
Primates 2018–2020 (Davenport 2019), based on its small global population, narrow range and threats.
After a decade and a half of conservation in their forest habitat, Davenport et al. (2022) reassessed
the population size, demography, and distribution of R. kipunji across Tanzania’s Southern Highlands,
employing the identical sweep census methods as of 2007 (Davenport et al. 2008). A habituated
group was also monitored daily over the same period. A total of 1,866 individuals in 59 groups (μ =
31.63±SE1.2) were recorded in Livingstone Forest (in Kitulo National Park), Mt Rungwe Nature Reserve,
and Madehani Village Forest. This corresponded to a 65% increase in individuals, a 59% increase in
group numbers, and a 19% increase in Area of Occupancy since the previous census. The ratio of
subadults/juveniles to adult females, a proxy for survival, was good (1.77), but higher in Livingstone
(2.61) than Mt Rungwe (1.11). A 121% increase in group size was recorded in the habituated group.
Signs of human activity fell by 81%, with a 100% and 98% reduction in the number of charcoal pits and
timber felling in Mt Rungwe. Whilst the total remains low by global standards, both temporal and
spatial data demonstrated that long-term holistic conservation had led to a significant increase in the
numbers of R. kipunji (Davenport et al. 2022).
Davenport, T.R.B. (2019). Kipunji (Rungwecebus kipunji). In: C. Schwitzer et al. (eds.), Primates in Peril: The World’s 25 Most
Endangered Primates 2018–2020, pp. 32–36. Washington, D.C., and Bristol, UK: IUCN SSC Primate Specialist Group,
International Primatological Society, Global Wildlife Conservation and Bristol Zoological Society.
Davenport, T.R.B., Stanley, W.T., Sargis, E.J., De Luca, D.W., Mpunga, N.E., Machaga, S.J. and Olson, L.E. (2006). A new
genus of African monkey, Rungwecebus. Morphology, ecology and molecular phylogenetics. Science 312: 1378–1381.
Davenport, T.R.B., De Luca, D.W., Jones, T., Mpunga, N.E., Machaga, S.J. and Picton Phillipps, G. (2008). The kipunji
Rungwecebus kipunji of southern Tanzania: first census and assessments of distribution and conservation status. Oryx
42(3): 352–359.
Davenport, T.R.B., Machaga, S.J., Mpunga, N.E., Kimiti, S.P., Mwalwengele, W., Mwaipungu, O. and Makumbule, P.M. (2022).
A reassessment of the population size, demography and distribution of Tanzania’s endemic kipunji Rungwecebus kipunji
13 years on – demonstrating conservation success. Int. J. Primatol. doi.org/10.1007/s10764-022-00281-3

AFRICA – WHITE-THIGHED COLOBUS

Colobus vellerosus (I. Geoffroy Saint-Hilaire, 1834)
Benin, Côte d’Ivoire, Ghana, Togo
(2016, 2018)
Reiko Matsuda Goodwin
The White-thighed Colobus (Colobus vellerosus) is a Critically Endangered species. Unregulated
hunting and destruction, degradation, and fragmentation of its habitats throughout its range have
reduced the species’ population by more than 80% in the past three decades (Matsuda Goodwin
et al. 2020). In the absence of hunting, C. vellerosus can thrive in a variety of habitats, from lowland
rainforests to savannah-forest mosaics, and from high forests to moderately disturbed forests (Wong

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and Sicotte 2007, Oates 2011). This species used to be widely distributed from the area between the
Sassandra-Bandama rivers in Côte d’Ivoire to southwestern Nigeria, traversing Ghana and Togo, but
it has probably been extirpated from Nigeria. It is now distributed in extremely fragmented ranges,
occurring only in a few protected areas and community forests in the remaining range countries.
Although a lack of systematic surveys precludes us from making a precise estimate, less than 1,500
individuals probably remain in the habitat countries. The two strongholds of this species are the
Boabeng-Fiema Monkey Sanctuary (BFMS) in Ghana and Comoé National Park in Côte d’Ivoire.
Deforestation, hunting, and mining continue to negatively influence the long-term survival of the
species. Additionally, climate change would exacerbate the rate of contraction and fragmentation
of the suitable habitat of this species and many other forest-living primates (Korstjens et al. 2010,
Stewart et al. 2020). Robust conservation measures and range-wide systematic surveys are urgently
needed to protect this species.
Korstjens, A.H., Lehmann, J. and Dunbar, R.I.M. (2010). Resting time as an ecological constraint on primate biogeography.
Anim. Behav. 79(2), 361–374.
Matsuda Goodwin, R., Gonedelé Bi, S., Nobimè, G., Koné, I., Osei, D., Segniagbeto, G. and Oates, J.F. (2020). Colobus
vellerosus. The IUCN Red List of Threatened Species 2020: e.T5146A169472127.
Oates, J.F. (2011). Primates of West Africa: A Field Guide and Natural History. Arlington, VA: Conservation International.
Stewart, B.M., Turner, S.E. and Matthews, H.D. (2020). Climate change impacts on potential future ranges of non-human
primate species. Clim. Change 162(4): 2301–2318.
Wong, S.N. and Sicotte, P. (2007). Activity budget and ranging patterns of Colobus vellerosus in forest fragments in central
Ghana. Folia Primatol. 78(4): 245–254.

AFRICA – CROSS RIVER GORILLA

Gorilla gorilla diehli Matschie, 1904
Cameroon, Nigeria
(2000, 2002, 2004, 2006, 2008)
Inaoyom Imong, Andrew Dunn and Richard Bergl
The Cross River Gorilla (Gorilla gorilla diehli) is endemic to the hilly rainforest region along the
Nigeria-Cameroon border and Critically Endangered (Bergl et al. 2016). The population is small and
fragmented, estimated to number fewer than 300 individuals (Dunn et al. 2014) concentrated in about
15 small hilly areas interspersed among large areas of unoccupied but suitable habitat (Imong et al.
2014) across a landscape of approximately 12,000 km2 spanning the Afi Mountain Wildlife Sanctuary
in the west to the Kagwene Gorilla Sanctuary in the east, with an isolated outlying locality in the
Tofala Hills (Dunn et al. 2014), of which approximately 8,000 km2 is tropical forest (Bergl et al. 2012).
The population is threatened by hunting and habitat loss from agriculture, illegal logging, and road
construction. Although hunting of Cross River Gorillas has declined very considerably over the last
two decades, they are still occasionally killed by hunters as snare by-catch. While much of the species’
population lives in protected areas, a number of subpopulations exist on community land with no
formal protection (Dunn et al. 2014). Insecurity resulting from the ongoing political and armed conflict
in the Southwest and Northwest regions of Cameroon prevents the implementation of conservation
activities in the Cameroon side of the landscape. The presence of refugees in and around key habitat
areas in Nigeria are an additional threat to this population.
Bergl, R.A., Warren, Y., Nicholas, A., Dunn, A., Imong, I., Sunderland-Groves, J.L. and Oates, J.F. (2012). Remote sensing
analysis reveals habitat, dispersal corridors and expanded distribution for the Critically Endangered Cross River gorilla,
Gorilla gorilla diehli. Oryx 46: 278–289.
Bergl, R.A., Dunn, A., Fowler, A., Imong, I., Ndeloh, D., Nicholas, A. and Oates, J.F. (2016). Gorilla gorilla ssp. diehli (errata
version published in 2016). The IUCN Red List of Threatened Species 2016: e.T39998A102326240.
Dunn, A., Bergl, R., Byler, D., Eben-Ebai, S., Etiendem, D.N., Fotso, R., Ikfuingei, R., Imong, I., Jameson, C., Macfie, L.,
Morgan, B., Nchanji, A., Nicholas, A., Nkembi, L., Omeni, F., Oates, J., Pokempner, A., Sawyer, S. and Williamson, E.A.
(2014). Revised Regional Action Plan for the Conservation of the Cross River Gorilla (Gorilla gorilla diehli): 2014–2019. New
York, NY: IUCN/SSC Primate Specialist Group and Wildlife Conservation Society.
Imong, I., Robbins, M.M., Mundry, R., Bergl, R. and Kühl, H.S. (2014). Distinguishing ecological constraints from human activity
in species range fragmentation: the case of Cross River gorillas. Anim. Conserv. 17: 323–331. doi.10.1111/acv.12100

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AFRICA – WESTERN CHIMPANZEE
Pan troglodytes verus Schwarz, 1934
Côte d’Ivoire, Ghana, Guine, Guinea-Bissau, Liberia, Mali, Senegal, Sierra Leone
(2018)
Information summarized by the editors from the IUCN Red List (Humle et al. 2016)
The Western Chimpanzee (Pan troglodytes verus) is Critically Endangered due to high levels of
poaching and habitat loss and degradation. It is suspected that the population has declined at an
average annual rate of 6.53% between 1990 and 2014, with a total population of 18,000 to 56,000
remaining in the wild (Sop et al. in prep., cited in Humle et al. 2016). The population is considered
almost certainly extinct in Benin, Burkina Faso, and Togo (Ginn et al. 2013, Campbell and Houngbedji
2015) and has seen a decline of 90% in its population in Côte d’Ivoire (Campbell et al. 2008). Although
it can be found in several protected areas, most of the population is found in areas that remain
unprotected. Conservation actions and research gaps are detailed in the IUCN Red List profile (Humle
et al. 2016).
Campbell, G. and Houngbedji, M. (2015). Conservation Status of the West African Chimpanzee (Pan troglodytes verus) in
Togo and Benin. Arlington, VA: Primate Action Fund.
Campbell, G., Kühl, H., N’Goran Kouamé, P. and Boesch, C. (2008). Alarming decline of West African chimpanzees in Côte
d’Ivoire. Curr. Biol. 18: R903–R904.
Ginn, L.P., Robison, J., Redmond, I. and Nekaris, K.A.I. (2013). Strong evidence that the West African chimpanzee is extirpated
from Burkina Faso. Oryx 47: 323–327.
Humle, T., Boesch, C., Campbell, G., Junker, J., Koops, K., Kuehl, H. and Sop, T. (2016). Pan troglodytes ssp. verus (errata
version published in 2016). The IUCN Red List of Threatened Species 2016: e.T15935A102327574.

ASIA – JAVAN SURILI

Presbytis comata Desmarest, 1822
Indonesia (Java)
Never included on the Top 25 List
Arif Setiawan and Vincent Nijman
The Javan Surili (Presbytis comata) is listed as Endangered on the IUCN Red List of Threatened Species
and is endemic to the western part of the island of Java (Nijman 1997, Nijman and Setiawan 2020), an
area it shares with some 70 million humans, who are living at a population density of over 1,500 people
per km². With the Javan Surili confined to forest, there is considerable pressure on the remaining areas
where the species is still found, and the cumulative effect of small amounts of forest loss is a clear
impediment to its survival (Supartono et al. 2016). With the recognition by IUCN of the Rekrekan (P.
fredericae) from central Java as a distinct species (formerly considered P. comata), there is considerable
confusion about the exact distribution of two species, as individuals from the intervening area show
morphological affinities of them both. Further research (morphological, behavioral and molecular) is
urgently needed to clarify their taxonomic arrangement and also their range distribution. The surili is
largely known from well-trodden volcanoes where it is found mostly in the lower montane or montane
zone (up to 2,500 m asl) but where lowland forest remains it can occur down to sea-level. It is found in
a number of protected areas, including the Ujung Kulon, Gunung Halimun-Salak, and Gunung Gede-
Pangrango national parks. The total population of Presbytis comata in Ujung Kulon, Rawa Danau, Mts
Halimun-Salak, Mt Gede-Pangrango, Kawah Kamojang, Mt Patuha, Ciremay-Kuningan-Pemabarisan
is estimated at 7,500–10,200 individuals. Potentially sizeable populations, such as at Sanggabuana,
Masigit-Kareumbi, Mt Tangkuban Perahu-Burangrang, and Cibanteng-Cikepuh (Supriatna et al. 1994)
were last surveyed more than two decades ago and therefore need renewed attention. Densities
differ somewhat by altitude, from 0.4-2.4 groups/km2 in the lowlands below 1,000 m asl to 0.1-5.8
groups/km2 in montane forests. Group sizes are negatively correlated with altitude – in the lowlands
averaging around eight individuals and in the mountains around six individuals (Nijman 2017).

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Nijman, V. (1997). Occurrence and distribution of grizzled leaf monkey Presbytis comata (Desmarest, 1822) (Mammalia:
Primates: Cercopithecidae) in Java, Indonesia. Contrib. Zool. 66: 247–256.
Nijman, V. (2017). Group composition and monandry in grizzled langurs, Presbytis comata, on Java. Folia Primatol. 88(2):
237–254.
Nijman, V. and Setiawan, A. (2020). Presbytis comata. The IUCN Red List of Threatened Species 2020: e.T18125A17955175.
Supriatna, J., Tilson, R.L. Gurmaya, K.J., Manansang, J., Wardojo, W., Sriyanto, A., Teare, A., Castle, K. and Seal, U.S. (eds.).
(1994). Javan Gibbon and Javan Langur: Population and Habitat Viability Analysis Report. Apple Valley, Minnesota: IUCN/
SSC Conservation Breeding Specialist Group (CBSG).
Supartono, T.T., Prasetyo, L.B., Hikmat, A. and Kartono, A.G. (2016). Spatial distribution and habitat use of Javan Langur
(Presbytis comata): case study in District of Kuningan. Procedia Environ. Sci. 33: 340–353.

ASIA – BORNEAN BANDED LANGUR

Presbytis chrysomelas Müller, 1838
Brunei Darussalam, Indonesia (Kalimantan), Malaysia (Sarawak)
Never included on the Top 25 List
Andie Ang, Chien Lee, Erik Meijaard, Vincent Nijman and Noel Rowe
The Bornean Banded Langur or Cross-marked Langur (Presbytis chrysomelas) is endemic to the
northwestern corner of Borneo and is Critically Endangered (Nijman et al. 2020). Two subspecies
are recognized: P. c. chrysomelas and P. c. cruciger. In the past it could be found in protected areas
such as the Tanjung Datu, Maludam, Similajau and Niah national parks, and the Lanjak Entimau and
Samunsam wildlife sanctuaries, all in Sarawak, and the Danau Sentarum Wildlife Sanctuary in West
Kalimantan (Groves 2013, Phillipps and Phillipps 2018). However, the most recent records of their
occurrence in all but the first two parks date back more than a decade. While Nijman et al. (2008)
conservatively estimated that 200–500 individuals remained, more recent research suggests this
might have been an underestimate. However, their habitat is declining, partially because of cash crop
plantations and forest fires. Hunting is a continuing threat in their habitat, and even in the protected
areas.
Groves, C.P. (2013). Cross-marked langur Presbytis chrysomelas. In: R.A. Mittermeier, A.B. Rylands and D.E. Wilson (eds.),
Handbook of the Mammals of the World. Volume 3. Primates, p. 716. Barcelona: Lynx Edicions.
Nijman, V., Cheyne, S., Traeholt, C. and Setiawan, A. (2020). Presbytis chrysomelas. The IUCN Red List of Threatened Species
2020: e.T39803A17955321.
Nijman, V., Hon, J. and Richardson, M. (2008). Presbytis chrysomelas. The IUCN Red List of Threatened Species 2008:
e.T39803A10268236.
Phillipps, Q. and Phillipps, K. (2018). Phillipps’ Field Guide to the Mammals of Borneo and Their Ecology. 2nd edition.
Oxford, UK: John Beaufoy Publishing.

ASIA – NATUNA ISLAND SURILI

Presbytis natunae (Thomas & Hartet, 1894)
Indonesia (Sumatra)
(2002)
Arif Setiawan and Vincent Nijman
The Natuna Island Surili (Presbytis natunae) is confined to Bunguran, the largest island in the Natuna
archipelago, halfway between northwestern Borneo and the Thai-Malay Peninsula. The species has
been the subject of very few studies; in 86 years only four field surveys were conducted (Indrawan
and Rangkuti 2001, Lammertink et al. 2003, Firman Aldi 2007 [unpubl. report], Handayani et al. 2021).
In 2001, Lammertink et al. (2003) conducted the first systematic population survey which found that
the species had a preference for primary lowland forest and avoided logged forest. There were 2.3
groups per km2, and groups comprised only three to four individuals on average, probably because of
reduced predator pressure on the island. The population was estimated at 4,500-12,500 individuals.
Setiandari (2018) obtained data on the status of the Natuna Island Surili from 30 informants, who noted
a steady decline in numbers and identified poaching, deforestation, an increase in human population
size, and a lack of effective conservation management as the main reasons for this apparent decline.

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In 2020, research suggested that the total remaining habitat for the Natuna Island Surili is about 48,274
ha or 29.83% of the island (K. Latifiana, K. Putri Handayani and E. Yuni Agustin), with forest cover having
declined by 17.93% from 2001 to 2020. The species is found in a wide range of habitats, from lowland
and montane forest to heath forest, mixed forest gardens and rubber plantations. While a substantial
part of the island is still forested, more and more is affected by logging, fragmentation and conversion
to other land uses, and the largest area of relatively untouched forest is found on Mt Bedung. Conflict
with humans is greatest near the coast where most of the villages are situated. There are no forest
areas that are formally protected on Bunguran. Populations are likely to be experiencing continued
decline due to habitat loss and capture for pets. Data are lacking for strengthening conservation
measures.
Handayani, K.P., Latifiana, K., and Agustin, I.Y. (2021). Conservation Status of Natuna Leaf Monkey (Presbytis natunae). Final
report (no. 1667). Charlestown, RI, USA: Primate Conservation Inc.
Indrawan, M. and Rangkuti, F. (2001). Development, biodiversity and the conservation status of banded langur in Natuna
Islands, Indonesia. Trop. Biodiv. 7: 151–163.
Lammertink, M., Nijman, V. and Setiorini, U. (2003). Population size, Red List status and conservation of the Natuna leaf
monkey Presbytis natunae endemic to the island of Bunguran, Indonesia. Oryx 37(4): 472–479.
Setiandari, Y.C. (2018). Studi tingkat penurunan populasi hewan endemik Kekah Natuna (Presbytis natunae) di desa Ceruk,
kabupaten Natuna ditinjau dari penyebabnya. BSc thesis, Universitas Pasundan Bandung, Indonesia.

ASIA – EAST SUMATRAN BANDED LANGUR

Presbytis percura Lyon, 1908
Indonesia (Sumatra)
Never included on the Top 25 List
Rizaldi and Andie Ang
Presbytis percura is found in just a few isolated and unprotected lowland areas in Riau Province,
Sumatra, and faces extinction in the wild due to forest loss on a large scale (Rizaldi et al. 2019, Ang
et al. 2020). Riau experienced the highest rate of deforestation in Sumatra, and 63% of the natural
forest was lost between 1985 and 2008 (Uryu et al. 2010). Additionally, forest fires linked to the ENSO
events, and open burning of forest land for agricultural purposes destroy millions of hectares of land
in Indonesia every year, and Riau is often one of the worst impacted areas, owing in part to its high
concentration of peatland (World Bank 2016). A preliminary survey in 2018 found P. percura only in
rubber plantations, subsisting on rubber leaves and seeds (Rizaldi et al. 2019). The scattered forest
remnants and agroforests where P. percura were found might no longer remain without conservation
action on the ground. Individuals of this species are also killed by local communities when they
feed on fruit crops of such as rambutans and durians. Manpower and financial support are urgently
needed for the research and protection of not only this species, but also other poorly studied langurs
(P. bicolor and P. siamensis cana) and their remaining habitats in Riau Province in Sumatra.
Ang, A., Roesma, D.I., Nijman, V., Meier, R., Srivathsan, A. and Rizaldi. (2020). Faecal DNA to the rescue: shotgun sequencing
of non-invasive samples reveals two subspecies of Southeast Asian primates to be Critically Endangered species. Sci.
Rep. 10: 9396.
Rizaldi, Ilham, K., Prasetio, I., Lee, Z.H., Jabbar, S. and Ang, A. (2019). Preliminary study on the distribution and conservation
status of the East Sumatran banded langur (Presbytis femoralis percura) in Riau Province, Sumatra, Indonesia. Asian
Primates J. 8(1): 25–36.
Uryu, Y., Purastuti, E., Laumonier, Y., Sunarto, S.B.Y.S.A., Hadian, O., Kosasih, D.A. and Stüwe, M. (2010). Sumatra’s Forests,
Their Wildlife and the Climate, Windows in Time: 1985, 1990, 2000, and 2009. Technical Report. Jakarta, Indonesia: WWF–
Indonesia.
World Bank (2016). The Cost of Fire: An Economic Analysis of Indonesia’s 2015 Fire Crisis. Indonesia Sustainable Landscapes
Knowledge Note 1. Washington, D.C.: The World Bank.

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ASIA – TONKIN SNUB-NOSED MONKEY
Rhinopothecus avunculus Dollman, 1912
Vietnam
(2000–2018)
Tilo Nadler
The Tonkin Snub-nosed Monkey, Rhinopithecus avunculus, is a Critically Endangered species endemic
to Vietnam (Quyet et al. 2020). It is now confined to only two areas of the far northwest (Nadler et
al. 2003, Nadler and Brockman 2014). Its distribution has been drastically reduced in recent decades
due to massive deforestation and intensive hunting. As a result, the population has become severely
fragmented. The species was thought to be extinct until its rediscovery near the town of Na Hang,
Tuyen Quang Province in 1989. Conservation activities there were unsuccessful, and it is most likely
now extirpated. A population of 20 to 40 individuals was estimated for the Cham Chu Nature Reserve,
Tuyen Quang Province in 1992, but subsequent surveys provided no sightings, and the population
is also most likely extirpated. In 2001, a population was discovered in Khau Ca, close to Du Gia
Nature Reserve, Ha Giang Province. A census in 2015 confirmed 125–130 individuals. The area was
subsequently declared as the Tonkin Snub-nosed Monkey Species/Habitat Conservation Area. Based
on recent surveys the population is estimated to comprise about 160 individuals. During a survey in
March 2022, two groups were observed, one group with 84 and another with 11 individuals. It is the
only population that is not immediately threatened. In 2007, a population of about 20 Tonkin Snub-
nosed Monkeys was discovered in Tung Vai, Ha Giang Province, close to the border with China (Le
Khac Quyet and Covert 2010). This population is threatened by hunting and habitat loss and through
intensive cardamom cultivation, besides other human activities. The total population of the Tonkin
Snub-nosed Monkey is currently believed to be fewer than 200 individuals.
Le Khac Quyet and Covert, H.H. (2010). Another population of the Tonkin snub-nosed monkey (Rhinopithecus avunculus)
discovered in Ha Giang Province, Vietnam. Vietnam. J. Primatol. 1(4): 19–25.
Quyet, L.K., Rawson, B.M., Duc, H., Nadler, T., Covert, H. and Ang, A. 2020. Rhinopithecus avunculus. The IUCN Red List of
Threatened Species. 2020: e.T19594A17944213.
Nadler, T., Momberg, F., Nguyen Xuan Dang, and Lormee, N. (2003). Vietnam Primate Conservation Status Review 2002.
Part 2: Leaf Monkeys. Hanoi, Vietnam, Frankfurt, Germany: Fauna & Flora International–Vietnam Program and Frankfurt
Zoological Society.
Nadler, T. and Brockman, D. (2014). Primates of Vietnam. Vietnam: Endangered Primate Rescue Center.

ASIA – GOLDEN LANGUR

Trachypithecus geei Khajuria, 1956
(2016, 2018)
Bhutan, India (Assam)
Dilip Chetry
The Golden Langur (Trachypithecus geei) is endemic to India and Bhutan. Its range in India covers
more than 2.500 km² and in Bhutan 4,782 km². Wet evergreen and tropical semi-evergreen forests are
its primary habitats in India. In Bhutan, it is found in both warm broadleaf and tropical forests. The
Assamese Macaque, Rhesus Macaque and Bengal Slow Loris are its cohabitants in these forests. It
spends 99% of its active time in trees and primarily exploits the top and middle strata of the forest.
It lives in diverse social settings, including uni-male:multi-female, bi-male:multi-female and multi-
male:multi-female groups, besides all male bands and lone males. Annually, Golden Langurs spend
12.8–33% of time in feeding, 40–63.1% in resting, 6.3–19% in locomotion, 5–11.5% in monitoring,
2–3.7% in playing and 0.3–6% in grooming. They eat green leaves, their staple food, and other food
items from more than 200 plant species. They spend their nights in tall trees of a few selected species.
Leopard, wild dogs and python are the prominent predators. Domestic and feral dogs attack Golden
Langurs in fringe villages. It is categorized as Endangered on the IUCN Red List of Threatened Species
(Das et al. 2020), and is listed as a Schedule-I species in the Wildlife (Protection) Act of India, 1972,

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and the Forest and Nature Conservation Act of Bhutan, 1995. It is an Appendix-I species in CITES.
Along with habitat loss, habitat fragmentation and shrinkage, recent hybridization between Golden
Langurs and Capped Langurs (Trachypithecus pileatus) across the suspension bridges constructed
over the Chamkhar River has emerged as the severest threat (Wangchuk et al. 2003). Electrocution,
roadkill, and dog predation are also serious threats. Conservation challenges are likely to increase in
the coming years despite current conservation initiatives. More effective landscape conservation and
management, such as reforestation and corridors (both forested and artificial), are vital for the survival
of this species (Ghosh 2009, Chetry et al. 2010).
Chetry, D., Chetry, R., Ghosh, K., and Bhattacharjee, P.C. (2010). Status and conservation of Golden Langur in Chakrashila
Wildlife Sanctuary, Assam, India. Primate Conserv. 25: 81–86.
Das, J., Chetry, D., Medhi, R. and Choudhury, A. (2020). Trachypithecus geei. The IUCN Red List of Threatened Species 2020:
e.T22037A17960997.
Ghosh, S. (2009). Report on the Distribution and Population Status of Golden Langur (Trachypithecus geei) in Bodoland.
Territorial Council, Guwahati: Assam, India.
Wangchuk, T., Inouye, D.W. and Hare, M.P. (2003). A new subspecies of Golden Langur (Trachypithecus geei) from Bhutan.
Folia Primatol. 74(2): 104–108.

NEOTROPICS – BUFFY-TUFTED-EAR MARMOSET

Callithrix aurita (É. Geoffroy in Humboldt, 1812)
Brazil (Rio de Janeiro, São Paulo, Minas Gerais)
(2018)
Rodrigo Salles de Carvalho, Fabiano R. de Melo, Mônica Mafra Valença-Montenegro, Claudia
Igayara-Souza, Marcio Port-Carvalho and Leandro Jerusalinsky
The Buffy-tufted-ear Marmoset (Callithrix aurita) is endemic to the montane Atlantic Forest of
southeastern Brazil, in the southern part of the state of Minas Gerais, the state of Rio de Janeiro,
and the east and northeast of the state of São Paulo. The historically widespread destruction of
forests in its range has resulted in extremely reduced and severely fragmented remnant habitats,
with diminished and isolated populations (Melo et al. 2018, 2021). Even as these factors persist as a
major threat to the species, the anthropogenic introduction of congeneric species has led to invasive
populations and ecological competition, and genetic erosion is now an alarming threat to the survival
of this species (Carvalho et al. 2018, 2019, Malukiewicz et al. 2021). Although yet to be confirmed,
it is suspected that a recent yellow fever outbreak has also impacted the species’ populations. The
cumulative and synergistic effects of these threats led to the species being categorized as Critically
Endangered on the IUCN Red List of Threatened Species (Melo et al. 2021). Since 2014, the Mountain
Marmosets Conservation Program (MMCP) has been developing measures for the implementation
of conservation strategies, as established by the National Action Plan for the Conservation of the
Atlantic Forest Primates and the Maned Sloth (Brazil, ICMBio 2018). They include research and
management in small fragments as well as important protected areas where the species occurs (the
Serra dos Órgãos, Itaiaia, and Serra da Bocaina national parks) and an ex situ management program
coordinated by the Brazilian Association of Zoos and Aquaria (AZAB) and the National Center for
Research and Conservation of Brazilian Primates (ICMBio/CPB), involving the Guarulhos Zoo and the
Mountain Marmosets Conservation Center at the Federal University of Viçosa (CCSS-UFV) in Minas
Gerais, Brazil.
Brazil, ICMBio. (2018). Aprova o Plano de Ação Nacional para a Conservação dos Primatas da Mata Atlântica e da Preguiça-
de-Coleira. Portaria n° 702, de 07 de agosto de 2018 – Instituto Chico Mendes de Conservação da Biodiversidade
(ICMBio). Diário Oficial da União, Seção 1, número 153: 57.
Carvalho, R.S. et al. (2018). Callithrix aurita: a marmoset species on its way to extinction in the Brazilian Atlantic Forest.
Neotrop. Primates 24(1): 1–8.
Carvalho, R.S. et al. (2019). Buffy-tufted-ear marmoset – Callithrix aurita É. Geoffroy Saint-Hilaire, 1812. In: C. Schwitzer et al.
(eds.), Primates in Peril: The World’s 25 Most Endangered Primates 2018–2020, pp. 78–81. Washington, DC, Bristol, UK:
IUCN SSC Primate Specialist Group, International Primatological Society, Global Wildlife Conservation, Bristol Zoological
Society.

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Malukiewicz, J. et al. (2021). An introduction to the Callithrix genus and overview of recent advances in marmoset research.
ILAR Journal 61(2–3): 110–138. doi.org/10.1093/ilar/ilab027
Melo, F.R, Hilário, R.R., Ferraz, D.S., Pereira, D.G., Bicca-Marques, J.C., Jerusalinsky, L., Mittermeier, R.A., Ruiz-Miranda, C.R.,
Oliveira, L. and Valença-Montenegro, M.M. (2021). Callithrix flaviceps (amended version of 2020 assessment). The IUCN
Red List of Threatened Species 2021: e.T3571A191700879.

NEOTROPICS – PIED TAMARIN

Saguinus bicolor Spix, 1823
Brazil (Amazonas)
(2018)
Marcelo Gordo, Diogo Lagroteria, Renata Azevedo and Leandro Jerusalinsky
The Pied Tamarin is a small callitrichid (450–550g; 66–74cm) endemic to the Central Amazon, in
the vicinity of Manaus, Brazil (Gordo et al. 2017). Groups are extremely territorial, and range from
2 to 13 individuals. Population densities are low throughout their range (1–2 groups/km2) (Gordo
2012, Gordo et al. 2017). Recent research estimates its geographic distribution at about 8,000 km2,
from Itacoatiara, Rio Urubu to the Rio Cuieiras, north of Manaus (Lagroteria and Gordo, in prep.).
Expansion of the city of Manaus and the increasing occupation of rural areas has caused a severe loss
and fragmentation of its habitat. Adding to habitat loss, are roadkill, electrical shocks, and predation
by dogs (Gordo et al. 2013, 2021), and its already tiny range is contracting, giving way to the invasive
Saguinus midas with evidence of hybridization (Gordo et al. 2017). Saguinus bicolor is classified as
Critically Endangered on the IUCN Red List of Threatened Species (Gordo et al. 2021) and the Brazilian
Official National List of Threatened Species (Brazil, MMA 2014, Vidal et al. 2018) due to a projected
population reduction of 80% or more in the next 18 years (three generations) resulting from the threats
described above. More studies are needed on the interactions between the two tamarins. Strategies
for the species’ conservation were established in the National Action Plan for the Conservation of the
Pied Tamarin, including as goals: a) reduction of habitat loss; b) creation of protected areas, and the
maintenance and adequate management of those already existing; c) maintenance and expansion
of habitat connectivity both in urban and rural areas; d) reduction of the loss of individuals due to
electrocution, roadkill, and attacks by domestic animals; e) restoration of areas to improve habitat
quality; f) an understanding of the distribution of Saguinus bicolor and its potential relationship with
Saguinus midas; g) strengthening environmental education for the conservation of the species; and
h) promotion of adequate population management for the species’ conservation (Jerusalinsky et al.
2017, Brazil, ICMBio 2018). The Brazilian Association of Zoos and Aquaria (AZAB) and the National
Center for Research and Conservation of Brazilian Primates (ICMBio/CPB) are coordinating an ex situ
management program.
Brazil, ICMBio. (2018). Aprova o 2º ciclo do Plano de Ação Nacional para a Conservação do Sauim-de-coleira. Portaria nº
281, de 4 de abril de 2018. Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio), Ministério de Meio
Ambiente (MMA). Diário Oficial da União, Seção 1, número 66: 70.
Brazil, MMA. (2014). Lista Nacional Oficial de Espécies da Fauna Ameaçadas de Extinção. Portaria n° 444, de 17 de dezembro
de 2014. Ministério do Meio Ambiente (MMA). Diário Oficial da União, Seção 1, número 245: 121–126.
Gordo, M. (2012). Ecologia e Conservação do Sauim-de-Coleira, Saguinus bicolor (Primates, Callitrichidae). Doctoral thesis,
Museu Paraense Emílio Goeldi, Universidade Federal do Pará (UFPA), Belém, Brazil.
Gordo, M., Calleia, F.O., Vasconcelos, S.A., Leite, J.J.F. and Ferrari, S.F. (2013). The challenges of survival in a concrete jungle:
conservation of the pied tamarin (Saguinus bicolor) in the urban landscape of Manaus, Brazil. In: L.K. Marsh and C.A.
Chapman (eds.), Primates in Fragments: Complexity and Resilience, pp. 357–370. New York, NY: Springer.
Gordo, M., Subirá, R.J., Vidal, M.D., Röhe, F., Spironello, W.R., Valente, L.M., Oliveira, J.B., Pissinatti, A., Wormell, D. and
Jerusalinsky, L. (2017). Contextualização do sauim-de-coleira. In: L. Jerusalinsky, R.B. Azevedo and M. Gordo (eds.), Plano
de Ação Nacional para a Conservação do Sauim-de-Coleira, Série Espécies Ameaçadas 29: 25–44. Brasília, Brazil: Instituto
Chico Mendes de Conservação da Biodiversidade (ICMBio), Ministério de Meio Ambiente (MMA).
Gordo, M., Röhe, F., Vidal, M.D., Subirá, R., Boubli, J.P., Mittermeier, R.A. and Jerusalinsky, L. (2021). Saguinus bicolor
(amended version of 2019 assessment). The IUCN Red List of Threatened Species 2021: e.T40644A192551696.
Jerusalinsky, L., Azevedo, R.B. and Gordo, M. (eds.). (2017). Plano de Ação Nacional para a Conservação do Sauim-de-
Coleira. Série Espécies Ameaçadas (29). Brasília, Brazil: Instituto Chico Mendes de Conservação da Biodiversidade
(ICMBio), Ministério de Meio Ambiente (MMA).

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Vidal, M.D., Gordo, M. and Röhe, F. (2018). Saguinus bicolor (Spix, 1823). In: Instituto Chico Mendes de Conservação da
Biodiversidade (ed.), Livro Vermelho da Fauna Brasileira Ameaçada de Extinção, 1st edition. Volume 2, pp. 44–49. Brasília,
Brazil: Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio).

NEOTROPICS – OLALLA BROTHERS’ TITI MONKEY

Plecturocebus olallae Lönnberg, 1939
Bolivia
(2018)
Jesus Martinez and Robert Wallace
The Olalla Brothers’ Titi Monkey (Plecturocebus olallae) is endemic to Bolivia and Critically Endangered
(Martinez and Wallace 2021a). Its range is restricted to an area of naturally fragmented forests, and
the latest assessment reported a population of 2,855 individuals in an area of 383.4 km² (Martinez
and Wallace 2021b). Although this represents some increase in both abundance and distribution
(previously 2,000 individuals across 267 km2), the high forest fragmentation levels in the region –
less than half of the range is suitable forest (Wallace et al. 2013) – is an important spatial limitation
for these primates. The improvement of a major road will promote new human settlements and
potentially increase intensive agriculture and livestock management in the area, which, together,
represent the main risks for P. olallae populations (Martinez and Wallace 2021a, 2021b). Two municipal
protected areas, Rhukanrhuka and Pampas del Yacuma, include most of the range of P. olallae (70%)
and research and outreach there are important to raise awareness with local people about these
endemic primates (Martinez et al. 2015). Current work is focused on consolidating the management
of these conservation areas to minimize the impacts of the threats to P. olallae populations, and to
develop monitoring programs for these range-restricted primates.
Martinez, J., Wallace, R.B., Arnez, A., Barreta, J., Carvajal-Bacarreza, P., Domic, E., Turdera, C.F., Jurado, C., Lopez, L., Lopez-
Strauss, H., Morrison, L., Porcel, Z.R., Reinaga, A. and Siles Lazzo, T.M. (2015). Línea base para la conservación de los
monos lucachi de Bolivia: Callicebus olallae y C. modestus. Agrocienc. Amazonia 5: 1–11.
Martinez, J. and Wallace, R. (2021a). Plecturocebus olallae. The IUCN Red List of Threatened Species 2021:
e.T3554A17975516.
Martinez, J. and Wallace, R.B. (2021b). An update on the distribution and abundance of the endemic and threatened Olalla’s
titi monkey (Plecturocebus olallae). Primate Conserv. (35): 13–20.
Wallace, R.B., Martinez, J., López-Strauss, H., Barreta, J., Reinaga, A. and López, L. (2013). Conservation challenges facing
two threatened endemic titi monkeys in a naturally fragmented Bolivian forest. In: L.K. Marsh and C.A. Chapman (eds.),
Primates in Fragments: Complexity and Resilience, pp. 493–501. New York, NY: Springer.

NEOTROPICS – CAATINGA TITI MONKEY

Callicebus barbarabrownae Hershkovitz 1990
Brazil (Bahia, Sergipe)
(2010)
André C. Alonso, Hamilton F. Barreto, Eduardo Marques, Mônica M. Valença-Montenegro, Raone
Beltrão-Mendes, Sidney Gouveia, Stephen F. Ferrari and Leandro Jerusalinsky
The Caatinga (or Blond) Titi Monkey (Callicebus barbarabrownae) is the only endemic primate of
the Caatinga (dry forest and xerophilic scrub) in northeastern Brazil, occurring in the states of Bahia
and Sergipe (Printes et al. 2018, 2021). As for all titi monkeys, it lives in small family groups, with a
monogamous reproductive couple and its offspring, occupying a territory of 20–30 ha defended daily
by duet vocalizations. The species’ range has suffered widespread deforestation resulting in habitat
reduction and fragmentation, with scarce, diminished, and isolated remnant populations exposed to
synergistic genetic and demographic risks. Cattle ranching, agriculture, and continuing urbanization
are the main threats, besides rapid infrastructure development facilitated by an extensive network
of highways and power lines (Printes et al. 2018, 2021). The extent of hunting needs investigation,
although it is probably moderate due to their small size, and few individuals are kept as pets (Printes
et al. 2013, 2018, 2021). The combination of these threats, however, has resulted in the species being

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assessed as Critically Endangered (Brazil, MMA 2014, Printes et al. 2018, 2021). A new survey method
using standardized vocalization playbacks suggests that the species abundance may be higher than
previously thought (Coelho et al. 2020). Also, a recent reassessment of the species distribution, ten
years after its first evaluation (Estrela et al. 2011, Printes et al. 2011), revealed more areas and individuals
than were previously known (Alonso et al. 2022). On the other hand, species distribution modelling
combined with population viability analysis under different climate change scenarios and simulated
forest loss, predicted a reduction of one-third of the currently suitable area for C. barbarabrownae
(Barreto et al. 2022). These models also indicate that viable populations in large forest remnants
represent only a quarter of all known populations, with a decline in climatic suitability, and that smaller
populations may be extirpated regardless of the level of forest recovery (Barreto et al. 2022). This
situation highlights the need to effectively implement the National Action Plan for the Conservation
of the Northeastern Primates (Brazil, ICMBio 2018).
Alonso, A.C. et al. (2022). On the occurrence of the Critically Endangered blond titi (Callicebus barbarabrownae): reassessment
of occupied areas and minimum population size. Int. J. Primatol. doi.org/10.1007/s10764-021-00269-5
Barreto, H.F. et al. (2022). Viability meets suitability: distribution of the extinction risk of an imperiled titi monkey (Callicebus
barbarabrownae) under multiple threats. Int. J. Primatol. 43, 114–132. doi.org/10.1007/s10764-021-00259-7
Brazil, ICMBio. (2018). Aprova o Plano de Ação Nacional para a Conservação dos Primatas do Nordeste. Portaria n° 242,
de 27 de março de 2018 – Instituto Chico Mendes de Conservação da Biodiversidade. Diário Oficial da União, Seção 1,
número 61: 252.
Brazil, MMA. (2014). Lista Nacional Oficial de Espécies da Fauna Ameaçadas de Extinção. Portaria n° 444, de 17 de dezembro
de 2014. Ministério do Meio Ambiente. Diário Oficial da União, Seção 1, número 245: 121–126.
Coelho, I.P., Collins, S.J., Santos Junior, E.M., Valença-Montenegro, M.M., Jerusalinsky, L. and Alonso, A.C. (2020).
Playbackpoint counts and N-mixture models suggest higher than expected abundance of the critically endangered blond
titi monkey in Northeastern Brazil. Am. J. Primatol. 82(5): e23126.
Estrela, A.R., Nogueira, E.M.S. and Porfírio, S. (2011). Callicebus barbarabrownae (Hershkovitz, 1990) (Primates: Pitheciidae)
de Lamarão/BA: resultados preliminares. In: F.R. Melo and I. Mourthé (eds.), A Primatologia no Brasil – 11, pp. 93–102.
Jataí, GO, Brazil: Sociedade Brasileira de Primatologia, Universidade de Goiás.
Printes, R.C., Rylands, A.B. and Bicca-Marques, J.C. (2011). Distribution and status of the Critically Endangered blond titi
monkey Callicebus barbarabrownae of north-east Brazil. Oryx 45: 439–443.
Printes, R.C., Jerusalinsky, L., Cardoso de Sousa, M. and Rodrigues, L.R.R. and Hirsch, A. (2013). Zoogeography, genetic
variation and conservation of the Callicebus personatus Group. In: A.A. Barnett, L.M. Veiga, S.F. Ferrari and M.A. Norconk
(eds.), Evolutionary Biology and Conservation of Titis, Sakis and Uacaris, pp. 43–49. Cambridge, UK: Cambridge University
Press.
Printes, R.C., Alonso, A.C. and Jerusalinsky, L. (2018). Callicebus barbarabrownae Hershkovitz, 1991. In: ICMBio, MMA (ed.)
Livro Vermelho da Fauna Brasileira Ameaçada de Extinção: Volume II – Mamíferos, pp. 283–287. Brasília, Brasil: Instituto
Chico Mendes de Conservação da Biodiversidade.
Printes, R.C., Jerusalinsky, L., Alonso, A.C. and Mittermeier, R.A. (2021). Callicebus barbarabrownae (amended version of
2020 assessment). The IUCN Red List of Threatened Species 2021: e.T39929A191703041.

NEOTROPICS – WHITE-CHEEKED SPIDER MONKEY

Ateles marginatus (É. Geoffroy Saint-Hilaire, 1809)
Brazil (Mato Grosso, Pará)
Never included on the Top 25 List
Gustavo R. Canale, André L. Ravetta, Gerson Buss and Leandro Jerusalinsky
The White-cheeked Spider Monkey (Ateles marginatus) is endemic to the eastern Brazilian Amazon,
south of the Rio Amazonas, with its southernmost populations occupying the transition zone with
the Brazilian Cerrado, between the right bank of the Rio Tapajós and its tributary, the Rio Teles Pires,
and the left bank of the Rio Xingu (Ravetta et al. 2018, Lazari et al. 2020, Lima-Silva et al. 2022). Most
of its Extent of Occurrence lies in the Amazonian ‘arc of deforestation’, where forest loss is at the
highest rates of the entire tropics (Garcia et al. 2019, Lazari et al. 2020). The species is categorized
as Endangered on the IUCN Red List of Threatened Species (Ravetta et al. 2021) due to suspected
population reduction upwards of 50% over the last 45 years (three generations), which is due to habitat
declines and hunting in protected and unprotected areas (Brazil, MMA 2014, Ravetta et al. 2018, 2021).
Its range is cut by major highways – the Transamazônica (BR 230) and Cuiabá-Santarém (BR 163) – and
a new railway (Ferrogrão) is projected to cross the species range accelerating the deforestation and

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promoting the expansion of soybean plantations, cattle ranching, and urbanization. The construction
plans for the Tapajós Hydroelectric Complex, which includes a number of dams in the range of Ateles
marginatus (Ravetta 2008, Buss et al. 2017), might lead to the isolation of populations inhabiting
riparian forests. Moreover, highways and human settlement expansion facilitate hunting, which is
likely to have already extirpated the species in parts of its range (Ravetta and Ferrari 2009). Late
maturation (4–5 years of age) and long inter-birth intervals (up to 30 months) make it difficult for
this species to recover from hunting and other threats. The savannization of the Amazonian forests
due to climate change is predicted to be sooner and more intense in the region occupied by the
White-cheeked Spider Monkeys, one of the least-studied species of the genus. Species conservation
strategies, as outlined by the National Action Plan for the Conservation of the Amazonian Primates,
include the following goals: a) improve territorial planning; b) mitigate and compensate the impacts
of development projects; c) reduce hunting pressure; d) assess and mitigate the impacts of epizootics;
and e) minimize the effects of climate change (Brazil, ICMBio 2017). As part of the implementation of
this plan, the Brazilian Association of Zoos and Aquaria (AZAB) and the National Center for Research
and Conservation of Brazilian Primates (ICMBio/CPB) are coordinating an ex situ management
program.
Brazil, ICMBio. (2017). Aprova o Plano de Ação Nacional para a Conservação dos Primatas Amazônicos. Portaria n° 792, de
1º de dezembro de 2017 – Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio). Diário Oficial da União
Seção 1, número 232: 24.
Brazil, MMA. (2014). Lista Nacional Oficial de Espécies da Fauna Ameaçadas de Extinção. Portaria n º 444, de 17 de dezembro
de 2014. Ministério do Meio Ambiente (MMA), Diário Oficial da União Seção 1, número 245: 121–126.
Buss, G., Ravetta, A.L., Fialho, M.S., Rossato, R.S., Sampaio, R., Printes, R.C., Pinto, L.P. and Jerusalinsky, L. and. (2017).
Primatas do Parque Nacional do Jamanxim/PA: riqueza, distribuição e ameaças. Biodiv. Brasil. 7(2): 34–46.
Garcia, A.S., Vilela, V.M., Rizzo, R., West, P., Gerber, J.S., Engstrom, P.M. and Ballester, M.V. (2019). Assessing land use/
cover dynamics and exploring drivers in the Amazon’s arc of deforestation through a hierarchical, multi-scale and multi-
temporal classification approach. Remote Sens. Appl.: Soc. Environ. 15: 100233.
Lazari, P.R., Martins Oliveira, A.T., Sandmann, P.H.D., Miguel, B.V., Fontes, M.C., Filho, N.R.A., Guirau, G.V.F., Henicka, T.S.,
da Silva, L.F., Canale, G.R. and Bernardo, C.S.S. (2020). Parâmetros demográficos de populações de duas espécies
ameaçadas de macacos-aranha, Ateles chamek e Ateles marginatus, no ecótono Cerrado-Amazônia. Neotrop. Primates
26: 97–103.
Lima-Silva, L.G., de Mendonça, R.F.B., da Silva Dutra, L. and Rossi, R.V. (2022). New records and geographic distribution
extension of two primate species in the Amazonia-Cerrado transition area, Brazil. Mammalia doi.org/10.1515/
mammalia-2021-0034
Ravetta, A.L. (2008). Ateles marginatus (É Geoffroy, 1809). In: Instituto Chico Mendes de Conservação da Biodiversidade
(ed.), Livro Vermelho da Fauna Brasileira Ameaçada de Extinção. 1st edition, Volume 2: 728–730. Brasília: Instituto Chico
Mendes de Conservação da Biodiversidade.M
Ravetta, A.L. and Ferrari, S.F. (2009). Geographic distribution and population characteristics of the endangered white-fronted
spider monkey (Ateles marginatus) on the lower Tapajós River in central Brazilian Amazonia. Primates 50(3): 261–268.
Ravetta, A.L., Buss, G. and Rylands, A.B. (2018). Ateles marginatus (É. Geoffroy Saint-Hilaire, 1809). In: Instituto Chico Mendes
de Conservação da Biodiversidade (ed.), Livro Vermelho da Fauna Brasileira Ameaçada de Extinção. 2nd edition. Volume
2: 182–185. Brasília: Instituto Chico Mendes de Conservação da Biodiversidade.
Ravetta, A.L., Buss, G. and Mittermeier, R.A. (2021). Ateles marginatus (amended version of 2019 assessment). The IUCN Red
List of Threatened Species 2021: e.T2282A191689524.

NEOTROPICS – PERUVIAN YELLOW-TAILED WOOLLY MONKEY

Lagothrix flavicauda (Humboldt, 1812)
Peru
(2000, 2006, 2008, 2010)
Sam Shanee
The Yellow-tailed Woolly Monkey, Lagothrix flavicauda, is endemic to Peru and Critically Endangered
(Shanee et al. 2021). It is found largely in the northern Peruvian departments of Amazonas and San
Martín, but also in small areas of neighboring departments, as well as in a newly discovered isolated
population in the department of Junin (Shanee 2011, Aquino et al. 2016a, 2016b, McHugh et al. 2019).
It has suffered massive habitat loss and fragmentation throughout almost all of its range in the last
decades, leaving many populations isolated (Shanee 2016, Shanee et al. 2021). The species is also

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hunted for the illegal pet trade and for food throughout its range, and it is generally difficult to find
near human settlements (Shanee, N. 2012). It was thought to be extinct until its rediscovery in the
1970’s (Mittermeier et al. 1975). There are no precise estimates regarding the number of individuals
remaining, but the species is declining rapidly across its range, with an estimated population reduction
of 93% since the early 1980s (Shanee, N. and Shanee 2014). A number of conservation initiatives
now exist for the species, including the creation of new state protected areas, and rapid growth in
private and communal reserves (Shanee et al. 2017, 2018). In one community area, in Yambrasbamba,
Amazonas, community-based conservation efforts have resulted in a significant increase in the
population of Yellow-tailed Woolly Monkeys there (Shanee and Shanee 2015).
Aquino, R., Charpentier, E., García, G. and López, L. (2016a). First record of Lagothrix flavicauda on the eastern side of the
Río Huallaga: an expansion of its known geographic distribution. Primate Conserv. (30): 15–20.
Aquino, R, Garcia, G. and Charpentier, E. 2016b. Distribution and current status of the Peruvian yellow-tailed woolly monkey
(Lagothrix flavicauda) in montane forests of the Región Huánuco, Peru. Primate Conserv. (30): 31–37.
McHugh, S.M., Cornejo, F.M., McKibben, J., Zarate, M., Tello, C., Jiménez, C.F. and Schmitt, C.A. (2019). First record of the
Peruvian yellow-tailed woolly monkey Lagothrix flavicauda in the Región Junín, Peru. Oryx 54(6): 814–818.
Mittermeier, R.A., de Macedo-Ruiz, H. and Luscombe, A. 1975. A woolly monkey rediscovered in Peru. Oryx 13(1): 41–46.
Shanee, N. (2012). Trends in local wildlife hunting, trade and control in the Tropical Andes Hotspot, northeastern Peru.
Endang. Species Res. 19(2): 177–186.
Shanee, N. and Shanee, S. (2014). Yellow-tailed woolly monkey (Lagothrix flavicauda): conservation status, anthropogenic
threats, and conservation initiatives. In: T.R. Defler and P.R. Stevenson (eds.), The Woolly Monkey, pp. 283–299. New York,
NY: Springer.
Shanee, S. (2011). Distribution survey and threat assessment of the yellow-tailed woolly monkey (Oreonax flavicauda
Humboldt 1812), northeastern Peru. Int. J. Primatol. 32(3): 691–707.
Shanee, S. (2016). Predicting future effects of multiple drivers of extinction risk in Peru’s endemic primate fauna. In: T.M.
Waller (ed.), Ethnoprimatology: Primate Conservation in the 21st Century, pp. 315–349. Cham, Switzerland: Springer
International Publishing.
Shanee, S. and Shanee, N. (2015). Measuring success in a community conservation project: local population increase in a
critically endangered primate, the yellow-tailed woolly monkey (Lagothrix flavicauda) at la Esperanza, northeastern Peru.
Trop. Conserv. Sci. 8(1): 169–186.
Shanee, S., Shanee, N., Monteferri, B., Allgas, N., Alarcon Pardo, A. and Horwich, R.H. (2017). Protected area coverage of
threatened vertebrates and ecoregions in Peru: comparison of communal, private and state reserves. J. Environ. Manag.
202(1): 12–20.
Shanee, S., Allgas, N. and Shanee, N. (2018). Community conservation as a tool for primate conservation in Peru. In: Romens,
D. and Pulido Mata, J. (eds.), Primatology, Biocultural Diversity and Sustainable Development in Tropical Forests, pp.
320–329. Mexico: UNESCO.
Shanee, S., Cornejo, F.M., Aquino, R., Mittermeier, R.A. and Vermeer, J. (2021). Lagothrix flavicauda. amended version of
2019 assessment). The IUCN Red List of Threatened Species 2021: e.T39924A192307818.

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157
 AFFILIATIONS
EDITORS Alzamora, Mónica
Fundación Yunkawasi
Mittermeier, Russell A. Lima, Peru
Re:wild
Austin, TX, USA Andriajaona, Aubin
Montange des Français
Reuter, Kim E. Madagascar Biodiversity Partnership
Lemur Love Madagascar
San Diego, CA, USA
Andrianandrasana, Herizo T.
Rylands, Anthony B. Durrell Wildlife Conservation Trust
Re:wild Antananarivo, Madagascar
Austin, TX, USA
Andrianarimisa, Aristide
Jerusalinsky, Leandro Zoology and Animal Biodiversity
Centro Nacional de Pesquisa e Conservação de Primatas University of Antananarivo
Brasileiros Antananarivo, Madagascar
Instituto Chico Mendes de Conservação da Biodiversidade
(ICMBio/CPB) Ang, Andie
Cabedelo, PB, Brazil Mandai Nature
Singapore
Schwitzer, Christoph
Dublin Zoo Assis Jardim, Márcia M.
Dublin, Ireland Museu de Ciências Naturais do Rio Grande do Sul
Secretaria do Meio Ambiente e Infraestrutura
Strier, Karen B. Porto Alegre, RS, Brazil
Department of Anthropology
University of Wisconsin-Madison Azevedo, Renata B.
Madison, WI, USA Centro Nacional de Pesquisa e Conservação de Primatas
Brasileiros
Ratsimbazafy, Jonah Instituto Chico Mendes de Conservação da Biodiversidade
Groupe d’étude et de recherche sur les primates (Gerp) (ICMBio/CPB)
Antananarivo, Madagascar Cabedelo, PB, Brazil

Humle, Tatyana Bailey, Carolyn A.

Durrell Institute of Conservation and Ecology Omaha’s Henry Doorly Zoo and Aquarium
School of Anthropology and Conservation Omaha, NE, USA
University of Kent
Canterbury, Kent, UK Barreto, Hamilton F.
Programa de Pós-graduação em Ecologia e Conservação
CONTRIBUTORS Universidade Federal de Sergipe
São Cristóvão, SE, Brazil
Abwe, Ekwoge
Ebo Forest Research Project Beltrão-Mendes, Raone
San Diego Zoo Wildlife Alliance Programa de Pós-graduação em Ecologia e Conservação
Cameroon Universidade Federal de Sergipe
São Cristóvão, SE, Brasil
Alfonso-Cortes, Felipe
Grupo de Estudio de Primates del Ecuador (GEPE) Bergl, Richard
Proyecto Washu North Carolina Zoological Park
Fundación Naturaleza y Arte Asheboro, North Carolina, USA
Ecuador
Bezara Hortensia, Hosnah
Alonso, André C. Parc Andrafiamena Andavakoera
Independent Researcher Madagascar
Cabedelo, PB, Brazil

158
Bicca-Marques, Júlio César Cotton, Sam
Laboratório de Primatologia Bristol Zoological Society
Escola de Ciências da Saúde e da Vida Bristol Zoo Gardens
Pontifícia Universidade Católica do Rio Grande do Sul, Bristol, UK
Porto Alegre, RS, Brazil
Davenport, Tim R.B.
Boubli, Jean P. Re:wild
School of Science, Engineering and the Environment USA/Tanzania
University of Salford
Salford, UK De Jong, Yvonne A.
Eastern Africa Primate Diversity and Conservation
Buss, Gerson Program
Centro Nacional de Pesquisa e Conservação de Primatas Nanyuki, Kenya
Brasileiros
Instituto Chico Mendes de Conservação da Biodiversidade de la Torre, Stella
(ICMBio/CPB) Grupo de Estudio de Primates del Ecuador (GEPE)
Cabedelo, PB, Brazil Universidad San Francisco de Quito
Colegio de Ciencias Biológicas y Ambientales Ecuador
Butynski, Thomas M.
Eastern Africa Primate Diversity and Conservation Program Dempsey, Andrea
Nanyuki, Kenya West African Primate Conservation Action
Zoological Society of Hertfordshire
Canale, Gustavo Rodrigues Broxbourne, UK
Centre of Biodiversity Studies of the Southern Amazon
(NEBAM) Dinsmore, Mary P.
Federal University of Mato Grosso (UFMT) School of Environmental Sustainability
Sinop, MT, Brazil Loyola University Chicago
Chicago, IL, USA
Cardoso, Tatiane dos Santos
Museu Paraense Emilio Goeldi do Carmo, Sarisha Trindade
Belém, PA, Brazil. Centro de Conservação dos Saguis-da-Serra, Vila
Universidade Federal de Viçosa
Carvalho, Rodrigo Salles Viçosa, MG, Brasil
Programa de Educação Ambiental - PREA
Juiz de Fora, MG, Brazil Doumbé, Osiris
Sekakoh Association
Cervera, Laura Ntui, Cameroon
Grupo de Estudio de Primates del Ecuador (GEPE)
Asociación Ecuatoriana de Mastozoología AEM Dunn, Andrew
Quito, Ecuador Wildlife Conservation Society
Calabar, Nigeria
Chaves, Óscar M.
Universidad de Costa Rica (UCR) Dutton, Paul
San José, Costa Rica Independent researcher
Waikato, New Zealand
Cheyne, Susan M.
Oxford Brookes University Eppley, Timothy M.
Oxford, UK, and Conservation Science and Wildlife Health
Borneo Nature Foundation San Diego Zoo Wildlife Alliance
Central Kalimantan, Indonesia Escondido, CA, USA, and
Department of Anthropology
Chikhi, Lounès Portland State University
Laboratoire Évolution and Diversité Biologique - UMR5174 Portland, OR, USA
French National Centre for Scientific Research - IRD – UPS
Toulouse, France, and Instituto Gulbenkian de Ciência, Fernanda Solórzano, María
Oeiras, Portugal Grupo de Estudio de Primates del Ecuador (GEPE)
Asociación Ecuatoriana de Mastozoología AEM
Cornejo, Fanny Quito, Ecuador
Fundación Yunkawasi
Lima, Peru Ferrari, Stephen F.
Department of Ecology
Universidade Federal de Sergipe
Aracaju, SE, Brazil

159
Fotang, Chefor Hending, Daniel
Brandenburg University of Technology School of Biological Sciences
Cottbus-Senftenberg, Germany University of Bristol
Bristol, UK
Frasier, Cynthia L.
Omaha’s Henry Doorly Zoo and Aquarium Hernández-Jaramillo, Alma
Omaha, NE, USA Neotropical Primate Conservation
Bogotá, Colombia
Fuentes, Nathalia
Grupo de Estudio de Primates del Ecuador (GEPE) Hirano, Zelinda B.
Proyecto Washu / Fundación Naturaleza y Arte Biological Research Center (CEPESBI)
Ecuador Fundação Universidade Regional de Blumenau (FURB)
Indaial, SC, Brazil
Garber, Paul A.
Department of Anthropology, Program in Ecology, Igayara-Souza, Claudia
Evolution, and Conservation Biology Guarulhos Zoo
University of Illinois Guarulhos, SP, Brazil
Urbana, IL, USA
Ikemeh, Rachel Ashegbofe
Gonder, Mary Katherine SW/Niger Delta Forest Project
Bioko Biodiversity Protection Programme New Garki, Abuja, Nigeria
College of Arts and Sciences
Drexel University Imong, Inaoyom
Philadelphia, PA, USA Wildlife Conservation Society
Calabar, Nigeria
Gonedelé Bi, Sery
UFR Biosciences Jabbar, Sabrina
Université Félix Houphouët-Boigny Mandai Nature
Abidjan, Côte d’Ivoire Singapore

Gordo, Marcelo Jerusalinsky, Leandro

Laboratório Biologia da Conservação Departamento de Centro Nacional de Pesquisa e Conservação de Primatas
Biologia/ICB, Universidade Federal do Amazonas Brasileiros
Manaus, AM, Brazil Instituto Chico Mendes de Conservação da Biodiversidade
(ICMBio/CPB)
Gouveia, Sidney Cabedelo, PB, Brazil
Department of Ecology
Federal University of Sergipe Kamgang, Serge A.
São Cristóvão, SE, Brazil Waza National Park
Far North Region, Cameroon
Gutiérrez-Pineda, Karol M.
Fundación Pro-Conservación de los Primates Panameños Kappeler, Peter M.
(FCPP) German Primate Center
Panama City, Panama Göttingen, Germany

Hadisiswoyo, Panut Koné, Inza

Orangutan Information Centre Centre Suisse de Recherche Scientifiques en Côte d’Ivoire
Sumatra Utara, Indonesia (CSRS)
Abidjan, Côte d’Ivoire
Hanlan, Fei
College of Life Science Kowalewski, Martin
China West Normal University Estación Biológica Corrientes (EBCo)
Nanchong, China Museo Argentino de Ciencias Naturales BR CONICET
Consejo Nacional de Investigaciones Científicas y Técnicas
Hart, John A. Corrientes, Argentina
FZS-ICCN Lomami National Park Project
Kinshasa, Democratic Republic of the Congo Kuswanda, Wanda
Life Sciences and Environment Research Organization
Heriniaina, Rio National Research and Innovation Agency (BRIN) Central
Primate Biology, Behaviour and Conservation Jakarta, Indonesia
University of Roehampton
London, UK

160
Lagroteria, Diogo Melo, Fabiano R. de
Centro Nacional de Pesquisa e Conservação da Forestry Engineering Department
Biodiversidade Amazônica Viçosa Federal University
Instituto Chico Mendes de Conservação da Biodiversidade Viçosa, MG, Brazil
(ICMBio/CEPAM)
Manaus, AM, Brazil Mendes, Sérgio Lucena
Instituto Nacional da Mata Atlântica (INMA) and
Lee, Chien Universidade Federal do Espírito Santo (UFES), Vitória, ES,
Sarawak, Borneo Brazil

Lee, Zan Hui Mendes-Oliveira, Ana Cristina

Universiti Sains Malaysia Laboratório de Ecologia e Zoologia de Vertebrados
Pulau Pinang, Malaysia Instituto de Ciências Biológicas
Universidade Federal do Pará
Leonard, Neahga Belém, PA, Brazil
Cat Ba Langur Conservation Project
Ca Ba National Park Meijaard, Erik
Cat Ba Island, Cat Ba District Borneo Futures
Hai Phong Province, Vietnam Brunei

Lewis, Richard Méndez-Carvajal, Pedro G.

Durrell Wildlife Conservation Trust Fundación Pro-Conservación de los Primates Panameños
Antananarivo, Madagascar (FCPP)
Panamy City, Panama
Louis, Jr., Edward E.
Omaha’s Henry Doorly Zoo and Aquarium Montilla, Sebastián
Omaha, NE, USA Departamento de Ciencias Biológicas
Universidad de Los Andes y Fundación Proyecto Primates
Lu, Zhang Bogotá, Colombia
School of Life Sciences
Sun Yat-Sen University Morales-Jiménez, Alba Lucía
Guangzhou, Guangdong, China Fundación Biodiversa Colombia
Bogotá, Colombia
Markolf, Matthias
University of Göttingen Morelos-Juárez, Citlalli
Göttingen, Germany Fundación Reserva Tesoro Escondido
Ecuador
Marques, Eduardo
Instituto Chico Mendes de Conservação da Biodiversidade Morgan, Bethan Jane
Cabedelo, PB, Brazil Ebo Forest Research Project
Messa, Cameroon, and
Martinez, Jesus Institute for Conservation Research
Wildlife Conservation Society San Diego Zoo Global
La Paz, Bolivia Escondido, CA, USA

Matsuda Goodwin, Reiko Myers Thompson, Jo A.

Department of Sociology and Anthropology Lukuru Wildlife Research Foundation
Fordham University Marion, OH, USA
New York, USA
Nadler, Tilo
McGraw, W. Scott Three Monkeys Wildlife Conservancy
The Ohio State University Cuc Phuong Commune, Nho Quan District
Department of Anthropology Ninh Binh Province, Vietnam
Columbus, OH, USA
Nekaris, K. Anna I.
McLester, Edward Nocturnal Primate Research Group
Department for the Ecology of Animal Societies Oxford Brookes University
Max Planck Institute of Animal Behavior Oxford, UK
Konstanz, Germany

161
Neves, Leonardo G. Port-Carvalho, Márcio
Instituto de Estudos Socioambientais do Sul da Bahia Environmental Research Institute, Conservation and
(IESB) Biodiversity Nucleus
Universidade Estadual de Santa Cruz (UESC) Infrastructure and Environment Secretariat of São Paulo,
Ilhéus, BA, Brazil SP, Brazil, and
National School of Tropical Botany
Nijman, Vincent Rio de Janeiro Botanical Garden
Oxford Brookes University Rio de Janeiro, RJ, Brazil
Oxford, UK
Pozo-Montuy, Gilberto
Ngouh, Amadou Facultad Maya de Estados Agropecuarios
Université de Dschang Universidad Autónoma de Chiapas
Cameroon México

Oates, John F. Prasetyo, Didik

Dept of Anthropology Yayasan Orangutan Tapanuli Indonesia (YOTI)
Hunter College CUNY Jakarta, Indonesia
New York, NY, USA
Queiroz, Helder
Oklander, Luciana I. Mamirauá Institute for Sustainable Development
Grupo de Investigación en Genética Aplicada Tefé, AM, Brazil
IBS Consejo Nacional de Investigaciones Científicas y
Técnicas Raffel, Martina
Universidad Nacional de Misiones Allwetter Zoo, Germany
Argentina
Raharivololona, Brigitte M.
Passaro, Richard L. Mention Zoologie et Biodiversité Animale
Previously associated with the Cat Ba Langur Conservation Ecole Doctorale Sciences de la Vie et de l’Environnement
Project Université d’Antananarivo
Vietnam Antananarivo, Madagascar

Pengfei, Fan Rakotoarisoa, Gilbert

School of Life Sciences Parc Botanique et Zoologique de Tsimbazaza
Sun Yat-Sen University Antananarivo, Madagascar
Guangzhou, Guangdong, China
Rakotondraparany, Felix
Perkin, Andrew Département de Zoologie et Biodiversité Animale
Independent Researcher Université d’Antananarivo
Tanzania Antananarivo, Madagascar

Perry, George (PJ) Rakotomalala, Valérie F.

Departments of Anthropology and Biology Pennsylvania Faculté des Sciences, de Technologie et de
State University l’Environnement
University Park, PA, USA Université de Mahajanga
Mahajanga, Madagascar
Pinel Ramos, Eduardo J.
Instituto de Neuroetología Rakotondrazandry, Jeannin Nicolas
Universidad Veracruzana Ecole Doctorale des Ecosystèmes Naturels
Honduras Université de Mahajanga
Mahajanga, Madagascar
Pollensio, Herinandrasana
Département des Sciences de la Nature et de Rakotondrina, Romeo
l’Environnement Mention Zoologie et Biodiversité Animale
Université d’Antsiranana Université d’Antananarivo
Antsiranana, Madagascar Antananarivo, Madagascar

Ralainasolo, Fidy
Durrell Wildlife Conservation Trust
Antananarivo, Madagascar

162
Randimbiharinirina, Doménico R. Rigobert, Jhon
Groupe d’Etude et de Recherche sur les Primates de University of Antsiranana
Madagascar (GERP) Antsiranana, Madagascar
Antananarivo, Madagascar
Rivera, Esteban
Randriatahina, Guy H. Grupo de Estudio de Primates del Ecuador (GEPE)
Association Européenne pour l’Étude et la Conservation Proyecto Washu / Fundación Naturaleza y Arte
des Lémuriens (AEECL) Ecuador
Madagascar
Rizaldi
Rasoanaivo, Noa E. Andalas University
Département Zoologie et Biodiversité Animale Faculté des Padang City, West Sumatra, Indonesia
Sciences
Université d’Antananarivo Rodríguez-Menjívar, Melissa E.
Antananarivo, Madagascar Asociación Territorios Vivos El Salvador (ATVES)
El Salvador
Rasoamazava, Lucile
Faculté des Sciences, de Technologies et de Rowe, Noel
l’Environnement Primate Conservation Inc.
Université de Mahajanga Charlestown, RI, USA
Mahajanga, Madagascar
Rudran, Rasanayagam
Rasoloharijaona, Solofonirina Smithsonian’s Conservation Biology Institute
Faculté des Sciences, de Technologies et de Front Royal, VA, USA
l’Environnement
Université de Mahajanga Ruppert, Nadine
Mahajanga, Madagascar School of Biological Sciences
Universiti Sains Malaysia
Ratsimbazafy, Jonah Pulau Pinang, Malaysia
Groupe d’Etude et de Recherche sur les Primates de
Madagascar (GERP) Rylands, Anthony B.
Antsakaviro, Antananarivo, Madagascar Re:wild
Austin, TX, USA
Raveloarimalala, Lucile Mialisoa
Madagascar Wildlife Conservation Sánchez-Porras, Ronald
Atsimondrova Universidad de Costa Rica
Ambatondrazaka San José, Costa Rica
Madagascar
Salmona, Jordi
Ravelomandrato, Faranky Laboratoire Évolution and Diversité Biologique - UMR5174
Mention Zoologie et Biodiversité Animale Université Toulouse III Paul Sabatier - IRD - CNRS
Université d’Antananarivo Toulouse, France
Antananarivo, Madagascar
Schäffler, Livia
Raveloson, Herimalala Leibniz Institute for the Analysis of Biodiversity Change
Ecole Doctorale des Ecosystèmes Naturels (LIB)
Université de Mahajanga Museum Koenig, and
Mahajanga, Madagascar Centre for Biodiversity Monitoring and Conservation
Science
Ravetta, André Luis Section Conservation Ecology
Programa de Capacitação Institucional Bonn, Germany
Museu Paraense Emílio Goeldi (PCI-MPEG)
Belém, PA, Brazil Schrudde, Daniela
Previously associated with the Cat Ba Langur Conservation
Ren, Baoping Project
Ministry of Education Key Laboratory for Ecology of Vietnam
Tropical Islands
College of Life Sciences Schwitzer, Christoph
Hainan Normal University Dublin Zoo
Haikou, China Dublin, Ireland

163
Sefczek, Timothy M. Tinsman, Jen
Omaha’s Henry Doorly Zoo and Aquarium Conservation Genetics Department
Omaha, NE, USA Omaha’s Henry Doorly Zoo and Aquarium
Omaha, NE, USA
Seibokuro, Tukuru
Apoi Community Conservation Association Tirira, Diego G.
Nigeria Grupo de Estudio de Primates del Ecuador (GEPE)
Fundación Mamíferos y Conservación
Semedo, Thiago B.F. Capelo, Rumiñahui, Ecuador
Instituto Nacional de Pesquisa do Pantanal (INPP) Museu
Paraense Emílio Goeldi (MPEG), and Valença-Montenegro, Mônica M.
Programa de Capacitação Institucional Centro Nacional de Pesquisa e Conservação de Primatas
Cuiabá, MT, Brazil Brasileiros
Instituto Chico Mendes de Conservação da Biodiversidade
Semel, Brandon (ICMBio/CPB)
Fish and Wildlife Sciences Cabedelo, PB, Brazil
Virginia Tech University
Blacksburg, VA, USA Vital, Orlando
Programa de Pós-Graduação em Ecologia
Semel, Meredith Centro de Conservação dos Saguis-da-Serra Universidade
Biological Sciences Federal de Viçosa
Virginia Tech University Viçosa, MG, Brazil
Blacksburg, VA, USA
Volampeno, Sylviane
Setiawan, Arif Mikajy Natiora
SwaraOwa Madagascar
Yogyakarta, Indonesia
Waeber, Patrick O.
Shanee, Sam ETH Zurich, Ecosystems Management, Forest Management
Neotropical Primate Conservation and Development Group
Moyobamba, San Martín, Peru Zurich, Switzerland
Shekelle, Myron
Biology Department Wallace, Robert
Western Washington University Wildlife Conservation Society
Bellingham, WA, USA La Paz, Bolivia

Spaan, Denise Wich, Serge

Universidad Veracruzana de México Liverpool John Moores University
Xalapa, México Liverpool, UK

Steffens, Travis S. Wilmet, Leslie

Department of Sociology and Anthropology Durrell Wildlife Conservation Trust
University of Guelph Antananarivo, Madagascar
Guelph, ON, Canada, and
Planet Madagascar Zaldaña-Orantes, Karla P.
Guelph, ON, Canada Asociación Territorios Vivos El Salvador (ATVES)
San Salvador, El Salvador
Stenke, Roswitha
Previously associated with the Cat Ba Langur Conservation Zaonarivelo, John
Project Département des Sciences de la Nature et de
Vietnam l’Environnement
Université d’Antsiranana
Supriatna, Jatna Antsiranana, Madagascar
Department of Biology, FMIPA
University of Indonesia Zhou, Jiang
Indonesia, and School of Karst Science
Tapanuli Orangutan Foundation Guizhou Normal University
Indonesia Guiyang, China

Thuc, Phan Duy

Previously worked on the Golden-headed Langur

164
Back cover photo: The Ka’apor Capuchin Monkey, Cebus kaapori, in the Gurupí Biological
Reserve, Maranhão, Brazil. Photograph by Fabiano R. de Melo.

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PRIMATES IN PERIL
The World’s 25 Most Endangered Primates
2022–2023
ISBN 978-1-7372851-5-1

90000>

9 781737 285151