Lemurs of Madagascar
and the Comoros
The IUCN Red Data Book
Prepared by The World Conservation Monitoring Centre
IUCN — The World Conservation Union ¢
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LEMURS OF MADAGASCAR
and the Comoros
The IUCN Red Data Book
IUCN - THE WORLD CONSERVATION UNION
Founded in 1948, IUCN - The World Conservation Union - is a membership organisation
comprising governments, non-governmental organisations (NGOs), research institutions,
and conservation agencies in 120 countries. The Union's objective is to promote and
encourage the protection and sustainable utilisation of living resources.
Several thousand scientists and experts from all continents form part of a network
supporting the work of its six Commissions: threatened species, protected areas, ecology,
sustainable development, environmental law and environmental education and training. Its
thematic programmes include tropical forests, wetlands, marine ecosystems, plants, the
Sahel, Antarctica, population and natural resources, and women in conservation. These
activities enable IUCN and its members to develop sound policies and programmes for the
conservation of biological diversity and sustainable development of natural resources.
WCMC - THE WORLD CONSERVATION MONITORING CENTRE
The World Conservation Monitoring Centre (WCMC) is a joint venture between the three
partners in the World Conservation Strategy, the World Conservation Union (IUCN), the
World Wide Fund for Nature (WWF), and the United Nations Environment Programme
(UNEP). Its mission is to support conservation and sustainable development by collecting
and analysing global conservation data so that decisions affecting biological resources are
based on the best available information.
WCMC has developed a global overview database of the world's biological diversity that
includes threatened plant and animal species, habitats of conservation concer, critical sites,
protected areas of the world, and the utilisation and trade in wildlife species and products.
Drawing on this database, WCMC provides an information service to the conservation and
development communities, governments and United Nations agencies, scientific institutions,
the business and commercial sector, and the media. WCMC produces a wide variety of
specialist outputs and reports based on analyses of its data.
oY .
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A HY
LEMURS OF MADAGASCAR
and the Comoros
The IUCN Red Data Book
Compiled by the
World Conservation Monitoring Centre
Cambridge, U.K.
by
Caroline Harcourt
with assistance from
Jane Thornback
(Project coordinator and editor)
Financial support from
Bristol Zoo
Conservation International
World Wildlife Fund (U.S.) Primate Program
Madagascar Fauna Captive Propagation Group
Jersey Wildlife Preservation Trust
and the
Parc Zoologique et Botanique de la Ville de Mulhouse
IUCN - THE WORLD CONSERVATION UNION
Gland, Switzerland and Cambridge, U.K.
1990
Published by IUCN, Gland, Switzerland and Cambridge, U.K.
with financial support from Bristol Zoo, Conservation International, the World Wildlife
Fund (U.S.) Primate Program, Madagascar Fauna Captive Propagation Group, Jersey
Wildlife Preservation Trust and the Parc Zoologique et Botanique de la Ville de Mulhouse.
A contribution to GEMS - The Global Environment Monitoring System.
BRISTOL
CEN Ki ee ZOO CAS
WWE CONSERVATION GARDENS —
INTERNATIONAL
Copyright: 1990 International Union for Conservation of Nature and Natural Resources.
Reproduction of this publication for educational or other non-commercial
purposes is authorised without prior permission from the copyright holder{s].
Reproduction for resale or other commercial purposes is prohibited without
the prior written permission of the copyright holders[s].
Citation: Harcourt, C., and Thornback, J. (1990) Lemurs of Madagascar and the
Comoros. The IUCN Red Data Book. IUCN, Gland, Switzerland and
Cambridge, U.K.
ISBN: 2-88032-957-4
Printed by: Unwin Brothers Limited, The Gresham Press, Old Woking,
Surrey, U.K.
Cover illustration: Indri by: Brian Groombridge
Typesetting by: Richard Maling, IUCN Publications Services Unit
Available from: IUCN Publications Services,
219c Huntingdon Road, Cambridge CB3 ODL, U.K.
The designations of geographical entities in this book, and the presentation of the material,
do not imply the expression of any opinion whatsoever on the part of IUCN, or other
participating organisations concerning the legal status of any country, territory, or area, or of
its authorities, or concerning the delimitation of its frontiers or boundaries.
The views of the authors expressed in this publication do not necessarily reflect those of
IUCN or other participating organisations.
Dedicated to
Sir Peter Scott
(1909-1989)
who initiated Red Data Books and who gave boundless
enthusiasm and committment to the cause of conservation.
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CONTENTS
PRERACES << sa Sey Heel Dota Pails. . eae aoe Sate: 3
ACKNOWLEDGEMENTS). Wo. 22, Di Se. eS eee. 5
CATEGORIES OF THREAT (200. cee a 6
ENT RODUCTION Fro oo cc cn suis: sheep on sh cn terey ce se os SAREE EE 7
THE LEMURS OF MADAGASCAR AND THER... (ees 15
DEGREE OF THREAT
PROTECTED AREAS OF MADAGASCAR ............. 17
AND THEIR LEMURS
THE IUCN/SSC PRIMATE SPECIALIST GROUP ......... 27
IDDATVAISHEET SS rac cceccke aves ay sp os swede. ce geet ae oy 2 eee ee 29
sbhie'organisationof the datasheets >>) .02i.. shark sion tees ct) Se 29
Requesftor further, informations a) 5) -)-0- en oiey eet ee het emer 29
fiaxonomic'classification followed 7s) -).)-0-) -) ner <1) tee ee 29
DATA SHEETS OF THREATENED LEMURS ......... 31-231
OF MADAGASCAR
Family CHETROGALEIDAE ...............-..-.. 32-70
IVAECTOCEDUSIINUTINUS, os o/s, 2) cack ss ce ob os Sb lce <0j0) Shon elohben Oh auc NMR 32
IMIGCTOCEDUS TAGUS) x 5i 05 3h o. ee se oe Sesh 3) ne 00 cn'ce on ERD ete ae 40
IMAE7 ZC COGQUETELE
6. Soe anos x) Saw fog, eevee) eee ps ia see 121
PEAT ATT 01 ee Me eee oes bee on eo 126
LCN UDTAVENIET, sco lo en ene Oe ee Ro oa ciash Mans: isi clase 132
Temunyulvus.” s @ Bis Gs Sea eta Se Be ee ee ee 138
SPRGIDETONS 3 oes 55.5) SRR OO Seats Le ed an a ae 141
PS QIDOCOL GNIS 4. cio ofS ois, 8: 80 Soa A Oe ee ee 143
NERC OL ANAS 55-985 D EAST (si Cnt ST See ee 145
Jo FUIVUS occ oo Se lsnes bv ok eth cae te ow ee oni NE Oe 147
TOURISTS, CPs tas ae mee es ot Bee Reo 149
JETUSUS css Sosa b1os oe ev ay leone ooo 2 2 DO A, Cn 150
WSGMJONGL: 3s Gleam lig! eS ae A ae eT ae nN ee 153
VETO CIA VATE BAIA. ooo io oe anne oy sy Sut open ee do oe sees MOLT QUGOR 158
WRVGMIC SOLA mre che ts, 5 as ate ees rete e- Sasa Gs 3: eT 161
vi rubra, .......: IES IAA BAIRAD LAM ID. CALM So 163
Hapalemur griseus... ee oe es ss SRR 168
PAOTISCUS Tee 5. ocr NS SOME ek Sa hue) RE 171
g.alaotrensis. .. .. x. RNIGRQRGAM AD 2ZAARA BSTIATO 173
g.occidentalis ...30. cs tess. « « « CUM RES 174
HON GICMUNQUICUS, Omer eye ioge teie, ; is. wd, Se. 6 Oe 178
Hapalemur simus _.. WIORA TERIAL Ae ATAMIAG OEE SS 182
amily TINDRIEDAE o.oo. a5 Si ou oi 5. os cu oe oh ce on of bs cv ns OE 186-225
AVQRUNGNISER iccc oho. ice ee Chaves ss MRO ota a to pong 186
EIGNIB OR oo shor Ano ch a nk back ce bane as 0 « DORR Seed 30) Fea 189
Voccidentalis, ..... .. i ommenranliois {OF cons UHAg eT ars Lacoast Roy a '
Wcngucniatvah RAK eM arbaunra ht abt (ABQ): ati
wy work —oal -doo8 eed > ladold sa troy qué sod a8
A ceeater of proposals. for the conser vallnw of biglogital divarsity in Mad aguacar:
at é
ine
poe %
pet (areant by 1. . Mile miiet (ISkSjin a prelicimacy Acuen Piag for the: =A
ii to fhe survey ¢ aeBAr: aid oabneti i protected areas, ihe iodlowing ane i,
cee ettetibehoe oi ieee haan ewvation daia ventre and's byolugica) laveatory @i sets
(ieee Dat pos y im be not uD im Py Paitnbazaxe nm Mac Lig asus e ite :
iananarivs, “ay «J rts wiih he ‘University of M scat,¢ programmes
it awe eens end OCT a Hon edpcation + iv int cy ; ii 2 training of
wehveiem professsonals, the di velop of inter nal wildlife tous 30 ‘ioe es
Mipdape niet: tae .puengihening of the sociogical park and botanic k ae 2.
Were ce durvars he MOst eedangerid ie aur specte: and the develananienl Bh ig
‘wenden Prag iota Let boy strbvic< pacite.. -
As Oot need Reied ny times, Ue sexiva) af Madagaectr’s unl pe Biota: ie
, Penis, Bad whinialy the well-being of tm peopic, depends on the comune ge
Ms Gpibe Gounpp, “Server tho politeal wil) m Madaysrar, the o« verse eal wit
CS Oak Rar wre Kinane! aii (Fo. chy ror Cmts ics BO Wasi 20g, shige i
whee + is wt Me geared. te ei-ho ot, rogulre great offork pee
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a ¢eoqgemna Of bailey ip FRA ci To Sen. Selene.
(heerbees A. 0708 La soporte S dc Saath wae feee titres A Madags
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ore (ind a - > ee os
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ae! Ss Socsttiae sa Rind vwig.J .
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THE LEMURS OF MADAGASCAR AND THEIR DEGREE OF THREAT
Family CHEIROGALEIDAE
Microcebus murinus
Microcebus rufus
Mirza coquereli
Cheirogaleus medius
Cheirogaleus major
Allocebus trichotis
Phaner furcifer
Family MEGALADAPIDAE
Lepilemur dorsalis
Lepilemur edwardsi
Lepilemur leucopus
Lepilemur microdon
Lepilemur mustelinus
Lepilemur ruficaudatus
Lepilemur septentrionalis
Family LEMURIDAE
Lemur catta
Lemur coronatus
Lemur macaco
m. macaco
m. flavifrons
Lemur mongoz
Lemur rubriventer
Lemur fulvus
f. albifrons
f. albocollaris
f. collaris
f. fulvus
f. mayottensis
f. rufus
f. sanfordi
Varecia varie gata
v. variegata
v. rubra
Hapalemur griseus
g. griseus
g. alaotrensis
g. occidentalis
Hapalemur aureus
Hapalemur simus
Family INDRIIDAE
Avahi laniger
l. laniger
l. occidentalis
Indri indri
Propithecus diadema
d. diadema
d. candidus
d. edwardsi
d. perrieri
Grey Mouse Lemur
Brown Mouse Lemur
Coquerel's Dwarf Lemur
Fat-tailed Dwarf Lemur
Greater Dwarf Lemur
Hairy-eared Dwarf Lemur
Fork-marked Dwarf Lemur
Grey-backed Sportive Lemur
Milne-Edwards' Sportive Lemur
White-footed Sportive Lemur
Microdon Sportive Lemur
Weasel Sportive Lemur
Red-tailed Sportive Lemur
Northern Sportive Lemur
Ring-tailed Lemur
Crowned Lemur
Black Lemur
Black Lemur
Sclater's Lemur
Mongoose Lemur
Red-bellied Lemur
Brown lemur
White-fronted Lemur
White-collared Lemur
Collared Lemur
Brown Lemur
Mayotte Lemur
Red-fronted Lemur
Sanford's Lemur
Ruffed Lemur
Black and White Ruffed Lemur
Red-ruffed Lemur
Grey Gentle Lemur
Grey Gentle Lemur
Alaotran Gentle Lemur
Western Gentle Lemur
Golden Bamboo Lemur
Greater Bamboo Gentle Lemur
Woolly Lemur
Eastern Woolly Lemur
Western Woolly Lemur
Indri
Diademed Sifaka
Diademed Sifaka
Silky Sifaka
Milne-Edwards' Sifaka
Perrier's Sifaka
15
Abundant
Abundant
V
Abundant
Abundant
E
R
Zahamena (73,160ha) Tropical evergreen forest
Microcebus rufus Varecia variegata
Cheirogaleus major Propithecus diadema diadema
Lepilemur mustelinus Indri indri
Hapalemur griseus Avahi laniger
Lemur fulvus albifrons Daubentonia madagascariensis
Lemur rubriventer
R.N.1.4 Tsaratanana (48,622 ha) Tropical evergreen forest
Cheirogaleus major Lemur fulvus
Phaner furcifer Lemur macaco
Hapalemur griseus Lemur rubriventer
Lepilemur mustelinus
20
The IUCN Red Data Book
R.N.I.5 Andringitra (31,160 ha) Rain forest and some dry forest
Microcebus rufus Lemur fulvus fulvus
Lepilemur microdon Varecia variegata variegata
Lemur catta Avahi laniger laniger
R.N.1I.6 Lokobe (740 ha) Humid forest
Microcebus rufus Lemur macaco macaco
Lepilemur dorsalis
R.N.I.7 Ankarafantsika (60,520 ha) Dry western forest
Microcebus murinus Lemur mongoz
Cheirogaleus medius Propithecus verreauxi coquereli
Lepilemur edwardsi Avahi laniger occidentalis
Lemur fulvus fulvus
R.N.1I.8 Namoroka (21,742 ha) Dense dry forest
Microcebus murinus Lemur fulvus rufus
Lepilemur edwardsi Propithecus verreauxi deckeni
R.N.I.9 Bemaraha (152,000 ha) Dense dry forest
Microcebus murinus Hapalemur griseus occidentalis
Phaner furcifer Lemur fulvus rufus
Mirza coquereli Propithecus verreauxi deckeni
Lepilemur edwardsi
R.N.I.10 Tsimanampetsotsa (43,200 ha) Dry Didiereaceae brush
Microcebus murinus Lepilemur leucopus (probably)
Lemur catta Propithecus verreauxi verreauxi
R.N.I.11 Andohahela (76,020 ha) Eastern rain forest and spiny forest
Microcebus murinus Hapalemur griseus
Microcebus rufus Lemur fulvus collaris
Cheirogaleus medius Lemur catta
Cheirogaleus major Propithecus verreauxi verreauxi
Lepilemur leucopus Propithecus diadema
Lepilemur mustelinus (microdon) Avahi laniger laniger
Phaner furcifer Daubentonia madagascariensis
R.N.I.12 Marojejy (60,150 ha) Rain forest
Microcebus rufus Lemur rubriventer
Cheirogaleus major Varecia variegata (reported)
Lepilemur mustelinus Propithecus diadema candidus
Hapalemur griseus Avahi laniger laniger
Lemur fulvus Daubentonia madagascariensis
Special Reserves:
Ambohijanahary (24,750 ha) Dry western forest
Propithecus verreauxi deckeni (possibly others, the fauna is unknown)
Ambohitantely (5,600 ha) Rain forest in the central plateau
Microcebus rufus Lemur fulvus fulvus
21
Lemurs of Madagascar and the Comoros
Analamazaotra (Perinet) (810 ha) Eastern rain forest
Microcebus rufus Varecia variegata variegata (sometimes)
Cheirogaleus major Propithecus diadema diadema
Lepilemur microdon Indri indri
Hapalemur griseus Avahi laniger laniger
Lemur fulvus fulvus Daubentonia madagascariensis
Lemur rubriventer
Analamera (34,700 ha) Mostly dry, but some rain forest
Microcebus sp. (probably rufus) Lemur coronatus
Lepilemur septentrionalis Propithecus diadema perrieri
Lemur fulvus sanfordi Daubentonia madagascariensis
Andranomena (6,420 ha) Dry western forest
Microcebus murinus Lepilemur ruficaudatus
Cheirogaleus medius Lemur fulvus rufus
Mirza coquereli Propithecus verreauxi verreauxi
Phaner furcifer
Anjanaharibe-Sud (32,100 ha) Eastern rain forest
Microcebus rufus Propithecus diadema candidus
Hapalemur griseus Avahi laniger laniger
Lemur fulvus Indri indri
Ankarana (18,220 ha) Dry western forest
Microcebus sp. (maybe rufus) Lemur fulvus sanfordi
Cheirogaleus medius Lemur coronatus
Phaner furcifer Propithecus diadema perrieri
Lepilemur septentrionalis Daubentonia madagascariensis
Hapalemur griseus
Beza Mahafaly (600 ha) Spiny forest and gallery forest
Microcebus murinus Lemur catta
Cheirogaleus medius Propithecus verreauxi verreauxi
Lepilemur leucopus
Bora (4,780 ha) Dry western forest
Lemur fulvus Propithecus verreauxi
Forét d'Ambre (4,810 ha) Rain forest
Microcebus rufus Lemur fulvus sanfordi
Cheirogaleus major Lemur coronatus
Phaner furcifer Daubentonia madagascariensis
Lepilemur septentrionalis.
Kalambatritra (28,250 ha) Rain forest
Lemur fulvus rufus (and others)
Manongarivo (34,250 ha) Lowland to Montane rain forest
Microcebus rufus Lemur fulvus
Cheirogaleus major Lemur macaco
Phaner furcifer Avahi laniger occidentalis
Hapalemur griseus occidentalis Daubentonia madagascariensis
Lepilemur dorsalis
22
The IUCN Red Data Book
Manombo (5,020 ha) Lowland eastern rain forest
Hapalemur griseus Daubentonia madagascariensis
Lemur fulvus albocollaris
Nosy Mangabe (520 ha) Lowland eastern rain forest
Microcebus rufus Varecia varie gata variegata
Lemur fulvus albifrons Daubentonia madagascariensis
Private Reserves:
Analabe Dry western forest
Microcebus murinus Lepilemur ruficaudatus
Cheirogaleus medius Lemur fulvus rufus,
Mirza coquereli Propithecus verreauxi verreauxi
Phaner furcifer.
Berenty (200 ha) Spiny and gallery forest
Microcebus murinus Lemur fulvus collaris (introduced)
Cheirogaleus medius Lemur catta
Lepilemur leucopus Propithecus verreauxi verreauxi
Lemur fulvus rufus (introduced)
There are other protected areas shown on the map but not mentioned above. This is because
no information has been found on the lemur species within those areas.
REFERENCES
Andriamampianina, J. (1972). Les réserves naturelles intégrales de Madagascar. In:
Comptes rendus de la Conférence internationale sur la Conservation de la Nature et de ses
Ressources a Madagascar. Tananarive, Madagascar 7-11 October. Publication IUCN
Nouvelle Series Document supplémentaires No. 36. Pp.103-123
Hawkins, A. F. A., Ganzhorn, Bloxam, Q.M.C., Barlow, S.C., Tonge,
S.J. and Chapman, P. (in press). A survey and assessment of the conservation
Status and needs of lemurs, birds, lizards and snakes in the Ankarana Special Reserve,
Antseranana, Madagascar: with notes on the lemurs and birds of the nearby Analamera
Special Reserve. Biological Conservation.
IUCN/UNEP/WWE (1987). Madagascar, an environmental profile. Edited by M.D.
Jenkins. IUCN, Gland, Switzerland and Cambridge, U.K.
Nicoll, M. and Langrand, O. (1989). Revue Generale d’Aires Protegees et de la
Conservation a Madagascar. Unpublished report to WWF. Project number 3746.
O'Connor, S., Pidgeon, M. and Randria, Z. (1986). Conservation Program for
the Andohahela Reserve, Madagascar. Primate Conservation 7: 48-52.
Pollock, J. I. (1984). Preliminary Report on a Mission to Madagascar by Dr J. I.
Pollock in August and September 1984. Unpublished report to WWF.
Raxworthy, C.J. and Rakotondraparany, F. (1988). Mammals report. In:
Quansah, N. (Ed.), Manongarivo Special Reserve (Madagascar), 1987/88 Expedition
Report. Unpublished. Pp. 122-130.
Safford, R.J., Durbin, J.C. and Duckworth, J.W. (1989). Cambridge
Madagascar Rainforest Expedition to R.N.J. 12 - Marojejy. Unpublished preliminary
report.
Now available: Nicoll, M.E. et Langrand, O. (1989) Madagascar: Revue de la
Conservation et des Aires Protégées. WWF, Gland, Suisse.
23
Lemurs of Madagascar and the Comoros
Ampasindava
Peninsula
Narinda Bay
Mahajamba Bay,
Peninsula
Bombetoka Bay,
Antogil Bay 16°S
f
ta”)
L. Kinkony
Yo Alaotra
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c
i)
Cc
a
Ivgndro
Capricorn.
Figure 4: Map of some of Madagascar's major rivers, including most of those mentioned in
the text.
24
The IUCN Red Data Book
COMORO ISLANDS
Ngazidja
Ndzouani
Cap d'Ambre
aS
Anivorano Nord
Mayotte t ,
Mahajanga
So 16°S
Cap St Andre Mananara
Nosy
Boraha
Maintirano
Toamasina
Antsalova
e
Se
Belo-sur-Tsiribihina ks
Morondava
Ranomafand O ananjary
e lfanadiana
Fianarantsoa
Manakara
lvohibe
e
Wandioze
Sakaraha
Toliara\
Capricorn
ra
aolanaro
Figure 5: Map of some towns in Madagascar, including most of those mentioned in the
text.
25
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THE IUCN/SSC PRIMATE SPECIALIST GROUP
The IUCN/SSC Primate Specialist Group (PSG) has been in existence since the late 1960s
and has been under the leadership of its present Chairman, Dr Russell A. Mittermeier, since
1977. The group is one of numerous specialist groups of the Species Survival Commission
of IUCN and its membership has grown to nearly 200 primate scientists and
conservationists from 45 different countries. The PSG is organised into six main
subdivisions, corresponding to the four regions where primates occur, together with a
captive breeding division and a special division for miscellaneous activities (i.e.
conservation education, satellite imagery analysis, veterinary medicine, wildlife trade).
The goal of the PSG is to maintain the current diversity of the order Primates, with dual
emphasis placed on:
* ensuring the survival of threatened species wherever they occur;
* providing effective protection for large numbers of primates in areas of high primat2
diversity and/or abundance.
Activities underway in many parts of the world make it inevitable that a certain portion of the
world's forests and the primate species which reside in them will disappear. The role of the
PSG is to minimize this loss whenever possible by:
* setting aside special protected areas for threatened species;
* creating large national parks and reserves in areas of high primate diversity and/or
abundance;
* maintaining parks and reserves that already exist and enforcing protective legislation in
them;
* creating public awareness of the need for primate conservation and the importance of
primates as a natural heritage in the countries where they occur.
The PSG places particular emphasis on conservation of habitat and furtherance of
conservation education as both these measures are considered essential and in large part
inseparable. Regardless of how broadly one wishes to define conservation, long-term
survival of natural populations will not be possible if habitats are not preserved and if local
people in the areas where primates occur do not fully support conservation efforts.
Additional measures taken by the PSG include:
* determining ways in which man and his fellow primates can coexist in multiple use areas;
* establishing conservation orientated captive breeding programmes for "Endangered
species";
* ending all illegal and otherwise destructive traffic in primates;
* ensuring that research institutions using primates are aware of the conservation problems
and that they are using primates as prudently as possible, without threatening the survival
of any wild populations.
Among the many functions of the PSG are production of the newsletter/journal Primate
Conservation (formerly the IUCN/SSC Primate Specialist Group Newsletter), edited by
Isabel Constable, which is a major means of communication among the world's primate
conservationists and is distributed free of charge to PSG members. The PSG is also
responsible for the production of Action Plans for Primate Conservation, which update the
original Global Strategy for Primate Conservation, produced in 1977. These Action Plans
are intended to determine priorities in global primate conservation, to estimate the costs of
conserving the world's primate fauna and to serve as tools in the fundraising efforts to make
27
Lemurs of Madagascar and the Comoros
these projects possible. The first two regional Action Plans, the Action Plan for African
Primate Conservation 1986-1990 by John Oates and the Action Plan for Asian Primate
Conservation 1987-1991 by Ardith Eudey, have already been published. Action Plans for
Madagascar and the Neotropical region are in preparation.
For further information on the IUCN/SSC Primate Specialist Group, please contact:
Dr. Russell A. Mittermeier
Conservation International
1015 Eighteenth Street, N.W.
Washington D.C. 20036
U.S.A.
Tel (202) 429-5660, Fax (202) 887-5188.
28
DATA SHEETS
The organisation of the data sheets
Each data sheet refers to one species and, if relevant, the subspecies within it. There are
eight sections (summary, distribution, population, habitat and ecology, threats, conservation
measures, captive breeding and remarks) within each sheet, which may be repeated for the
subspecies, followed by a reference list for that species. The distribution maps have been
adapted from those in Petter et al (1977), Petter and Petter Rousseaux (1979) and Tattersall
(1982) with some additional information of new sightings from other reports. It is not, of
course, suggested that the species concerned occurs throughout the relevant shaded part of
the map. If that were the case, there would probably be no cause for concern about the
status of the lemurs. Each is found in the, probably, small patches of suitable habitat within
the range shown. Population numbers and/or densities and information as to whether the
species is declining is given in the section on population. The habitat and ecology section
has been expanded to make it more detailed than in previous Red Data Books. It is hoped
that this will make it a useful reference for anybody attempting a study of the lemurs. The
threats to each species are essentially similar throughout Madagascar, more so than when a
species is found in many different countries. Similarly for the conservation measures.
Information on captive breeding has come mainly from ISIS sheets, which mostly list only
USA institutions, and from Wilde et al (1988) who have reported on only European
institutions. Obviously this information is, as a result, probably far from complete, but it
should at least give an idea of what numbers of each species are held and which breed well
in captivity. The remarks section contains brief information on taxonomy, a short
description of the species (size and colour) and its Malagasy name.
Request for further information
Information such as that presented here needs to be continually revised and updated. While
the data sheets are as accurate as possible they do rely on information from those in the field.
It is hoped that anybody in the position to provide the required data will do so, preferably to
both IUCN, at the address and in the format of the sample inventory sheet given in
Appendix A, and to Ian Tattersall as requested in Appendix B. This help will be greatly
appreciated.
The taxonomic classification followed
The taxonomy of the lemurs is a constantly changing and much debated subject. There is no
level of classification of this group that is accepted by all authors. The system followed here
for all levels above the species is that of Schwartz and Tattersall (1985). The most
controversial aspect of this classification is, undoubtedly, the positioning of Lepilemur in the
family Megaladapidae. For species and subspecies, Jenkins (1987) is mostly followed
except in the case of subspecies of Propithecus verreauxi and P. diadema where Tattersall
(1986, in litt) is used. There have been two new species discovered since the publication of
Jenkin's catalogue and the descriptions of these follow Meier et al, (1987) for Hapalemur
aureus and Simons (1988) for Propithecus tattersalli.
a
Lemurs of Madagascar and the Comoros
Taxonomy References
Jenkins, P.D. (1987). Catalogue of Primates in the British Museum (Natural History)
and elsewhere in the British Isles. Part IV: Suborder Strepsirrhini, including the
subfossil Madagascan lemurs and Family Tarsiidae. British Museum (Natural History),
London.
Meier, B., Albignac, R., Peyriéras, Rumpler, Y. and Wright, P. (1987). A
new species of Hapalemur (Primates) from south east Madagascar. Folia Primatologica
48: 211-215.
Schwartz, J.H. and Tattersall, I. (1985). Evolutionary relationships of the living
lemurs and lorises (Mammalia, Primates) and their potential affinities with European
Eocene Adapidae. Anthropological Papers of the American Museum of Natural History
60 (1): 1-100.
Simons, E.L. (1988). A new species of Propithecus (Primates) from northeast
Madagascar. Folia Primatologica 50: 143-151.
30
LEMURS OF MADAGASCAR and the Comoros:
The IUCN Red Data Book
DATA SHEETS OF THREATENED LEMURS OF
MADAGASCAR
31
Lemurs of Madagascar and the Comoros
a
The Grey Mouse Lemur, Microcebus murinus, is a tiny nocturnal species found in
Madagascar's dry western forests. It is common in secondary vegetation.
Photo by Mark Pidgeon.
The IUCN Red Data Book
GREY or LESSER MOUSE LEMUR ABUNDANT
Microcebus murinus (J. F. Miller. 1777)
Order PRIMATES Family CHEIROGALEIDAE
SUMMARY The small, nocturnal Grey Mouse Lemur is found throughout the dry
deciduous forest of the west and south of Madagascar. It is common in secondary forest,
possibly more so than in undisturbed areas. It can reach high population densities and
appears unlikely to be severely threatened at present. However, destruction of its habitat is
occurring and it is probable that its numbers are declining. It has been the subject of several
short term studies. It is usually seen alone while foraging at night but sleeps in groups
during the day, the composition of which depends on the season. M. murinus is an
omnivore, its diet includes fruit, insects, flowers, seeds, gums and leaves. There are over
370 animals in captivity and most were bred there. It occurs in most of the protected areas in
the west and south, from Andohahela to Analamera. Listed in Appendix 1 of CITES, Class
A of the African Convention and is protected by Malagasy law.
DISTRIBUTION Occurs throughout the dry deciduous forested areas of western and
southern Madagascar from Taolanaro (Fort Dauphin) to, at least, the Sambirano River,
though its precise northern limit is not known (Tattersall, 1982). It has recently been
reported in Ankarana Special Reserve (Fowler et al, 1989), but other authors are not sure
which species of Mouse Lemur is present in either Ankarana or Analamera Special Reserves
(Hawkins et al, in press; Nicoll and Langrand, 1989; Ganzhorn, in litt.). It appears to be
replaced by M. rufus in the Sambirano Region as it is this species that is reported in
Manongarivo Special Reserve (Raxworthy and Rakotondraparany, 1988; Nicoll and
Langrand, 1989). M. murinus is found in drier areas of Andohahela Special Reserve (M.
Pidgeon, in litt.) and in the littoral forest in the area around Taolanaro (Martin, 1972).
POPULATION Population numbers are unknown, but the Grey Mouse Lemur must be
one of the most numerous of the lemurs. However, Richard et al (1985) suspect that its
numbers are "probably" declining. Population density has been estimated by Petter (1978)
and Hladik et al (1980) in Marosalaza Forest, north of Morondava; the former estimated 3-4
individuals per ha (i.e. 300-400 per sq. km) and the latter, 400 animals per sq. km.
Ganzhorn (1988) found much lower densities at Ampijoroa in Ankarafantsika Forest, only
42 + 19 (mean and 95% confidence limits) individuals per sq. km. However, Martin's
(1972) observations that this species occurs in "population nuclei" implies that it would be
difficult to estimate accurate densities when extrapolating from a small to a large area. In
addition, Grey Mouse Lemurs can be very difficult to find at some times of the year,
particularly during long dry periods, and this causes another problem in estimating densities
(M. Pidgeon, in litt.).
HABITAT AND ECOLOGY Grey Mouse Lemurs are reported to be far more common
in secondary forest than in primary forest; they were even found in gardens and in patches of
waste land round the port at Taolanaro (Martin, 1973) and have been seen in very degraded
roadside bush and scrub habitat (M. Pidgeon, in /itt.). They occupy the "fine branch" niche
and, as a result, the height at which they are seen depends on the height at which fine
branches, lianes and dense foliage are found (Martin, 1972, 1973). In secondary forest and
along paths they are generally observed at 0-10m above the ground, whereas they are found
at 15-30m in the canopy of primary forest (Martin, 1973). M. murinus is found in the spiny
forest parcels of Andohahela Special Reserve and is more numerous in this habitat than in
the gallery forest (M. Pidgeon, in /itt.). In the area around Taolanaro, Martin (1972) found
33
Lemurs of Madagascar and the Comoros
M. coquereli
Y M. murinus
FE M. rufus
Capricorn
Figure 6: Distribution of Mirza and both species of Microcebus. Shaded areas represent
approximate limits of ranges.
34
The IUCN Red Data Book
this species in the drier, littoral forest, while the very similar Brown Mouse Lemur was in
the inland, rain forest area.
The Grey Mouse Lemur is omnivorous; invertebrates and fruit appear to be the most
important components of its diet, but it has also been seen eating flowers, nectar, leaves
(Uapaca sp), sap and gum (from Euphorbia and Terminalia trees), secretions from
Homopteran larvae, and small vertebrates such as tree frogs, geckos and chameleons
(Martin, 1972, 1973; Petter, 1978; Hladik, 1979; Barre er al, 1988). Its insect prey was
frequently caught on the ground (Martin, 1972, 1973). Though there is no period of
dormancy, as there is in Cheirogaleus spp, the Mouse Lemur does lay down some fat in its
tail (its volume varies from 5 to 20 cu cm through the year) and under its skin and this is
probably used to make up for the reduced food available in the dry season from June to
September (Martin, 1972; Petter, 1978; Hladik, 1979; Petter-Rousseaux, 1980). Grey
Mouse Lemurs are most often seen alone at night, but, during the day, they are frequently
seen asleep in groups. Their nests are either spherical constructions made from leaves or are
in tree hollows, it appears that the latter are preferred (Martin, 1972, 1973). The minimum
extemal diameter of trees in which Mouse Lemur nests were found was 5 cm and the median
was 13 cm; it may be that trees of this size are a necessary part of a healthy habitat for
M. murinus. (Martin, 1973). In the non-mating season in Mandena, males were always
found either singly or in pairs at nest sites while females were in groups of 1-15 (with a
median of four), the sexes were usually separate (Martin, 1972). However, group
composition was very different during the mating season at the same study site; mixed sex
groups were common, frequently a single male was found sharing a nest site with between
three and seven females, though single females were also found with between one and three
males (Martin, 1973). During a brief radio-tracking study in Ankarafantsika Forest, Pagés-
Feuillade (1989) confirmed that male Grey Mouse Lemurs usually sleep alone, whereas
females are often in groups. Martin (1972, 1973) found that, in apparently homogenous
belts of forest, Grey Mouse Lemurs tended to occur in localised concentrations ("population
nuclei"). In his study, the sex ratio was biased towards females (three or four females to
one large male) in the population nucleus core, while smaller, adult males were found on the
periphery. Marked individuals were found no more than 50m from their original sighting
point, which suggested that home ranges were quite small (Martin, 1972, 1973). Two brief
radio-tracking studies in Ankarafantsika Forest found that males tended to have bigger home
ranges than females and that the males travelled further during the night (Barre et al, 1988;
Pagés-Feuillade, 1989). During a six week study, Pagés-Feuillade (1989) found that her
four radio collared males had home ranges of 3.2 + 0.22 ha, while those of four females
were 1.8 + 0.24 ha (Pagés-Feuillade, 1989). The ranges of nine individuals overlapped,
occupying a total of 7 ha, with the central portion being shared by all (Pagés-Feuillade,
1989). There was 66% overlap in the males' ranges and 44% overlap between females’
ranges (Pagés-Feuillade, 1989). Neither of the studies in Ankarafantsika found any sign of
a biased sex ratio, as Martin had done (1972, 1973), nor were central and peripheral males
found (Barre et al, 1988; Pagés-Feuillade, 1989).
In Mandena, mating of the Grey Mouse Lemurs begins around mid-September and infants
are born in November (Martin, 1972). Gestation period is 59-62 days (Petter-Rousseaux,
1964). The infants are born in a leaf nest or in a tree hole, litter size is usually two although
singletons, triplets or, very occasionally, quadruplets can be produced (Petter-Rousseaux,
1964, 1988). Offspring up to three weeks of age (in captivity) are carried in their mother's
mouth, rather than clinging to her fur (Petter-Rousseaux, 1964; Martin, 1972). Within two
months, the young are behaving much like adult Mouse Lemurs, females are sexually mature
within a year but, in captivity, they were at least 18 months old before they gave birth
(Petter-Rousseaux, 1964).
THREATS The Grey Mouse Lemur is very small and nocturnal, it exists in areas of
secondary forest and brush and, therefore, it seems unlikely that it could be severely
35
Lemurs of Madagascar and the Comoros
threatened. However, Richard et al (1985) suspect that its numbers are "probably" declining
as a result of habitat destruction. Sussman and Richard (1986) point out that since this
species is dependent upon areas of dense undergrowth, it is possible that the extensive
grazing by cattle and goats in southern Madagascar is destroying some of its optimal habitat.
Martin (1972, 1973) also notes that trees of a certain age and size containing hollows of
appropriate dimensions may be necessary for the long-term maintenance of a thriving Mouse
Lemur population. Heavy tree-felling occurred in Martin's study area between 1968 and
1970 and he found that this was having an effect on M. murinus. For instance, individuals
tended to be lighter in 1970, they were using smaller trees as nesting sites and the maximum
female nesting group size was only seven as opposed to 15 in the earlier study (Martin,
1973):
CONSERVATION MEASURES Found in Andohahela, Ankarafantsika, Namoroka,
Bemaraha and Tsimanampetsotsa Nature Reserves, in Andranomena and Beza Mahafaly
Special Reserves, in Berenty and Analabe Private Reserves and may be in Analamera and
Ankarana Special Reserves (Nicoll and Langrand, 1989)
At present, no specific measures are needed to conserve this Mouse Lemur. However, a
range wide survey to determine which Mouse Lemur is where would be useful. These could
be done in conjunction with surveys of more threatened species.
All species of Cheirogaleidae are listed in Appendix 1 of the 1973 Convention on
International Trade in Endangered Species of Wild Fauna and Flora. Trade in them, or their
products, is subject to strict regulation and may not be carried out for primarily commercial
purposes.
All Lemuroidea are listed in Class A of the African Convention, 1969. They may not,
therefore, be hunted, killed, captured or collected without the authorization of the highest
competent authority, and then only if required in the national interest or for scientific
purposes.
Malagasy law protects all lemurs from unauthorised capture and from hunting.
CAPTIVE BREEDING The Grey Mouse Lemur breeds well in captivity, though it is
not as common in zoos as the larger lemurs, presumably because it is small, grey, nocturnal
and generally not so interesting as an exhibit. ISIS (June 1989) lists 171 individuals (their
M. murinus [no subspecies] and M. m. murinus) in 14 institutes. Over 97% of these Mouse
Lemurs are captive born. The largest single colony is at Duke Primate Center. Over 97% of
these Mouse Lemurs are captive born. Wilde et al (1988) list 172 individuals in 15
European institutes that are not included in ISIS lists and there are a further 30 animals at
Paris Zoo (J.-J. Petter, in litt.)
REMARKS The Grey Mouse Lemur is one of the smallest primates, its mean body
weight is 60g , though there are considerable seasonal fluctuations in this (Martin, 1972).
Fur on its back is grey to grey-brown and greyish-white below. It has large membranous
ears. For a more detailed description see Tattersall (1982), Jenkins (1987) or Petter et al
(1977). The Malagasy names of this species are tsidy, pondiky, vakiandri, titilivaha and
koitsiky (Paulian, 1981; Tattersall, 1982).
REFERENCES
Barre, V., Lebac, A., Petter, J.-J. and Albignac, R.(1988). Etude du Microcébe
par radiotracking dans la forét de l'Ankarafantsika. In: Rakotovao, L., Barre, V. and
Sayer, J. (Eds), L’Equilibre des Ecosystémes forestiers ad Madagascar: Actes d'un
séminaire international. IUCN, Gland and Cambridge. Pp. 61-71.
36
The IUCN Red Data Book
Fowler, S.V., Chapman, P., Hurd, S., McHale, M., Ramangason, G.-S.,
Randriamsy, J.-E., Stewart, P., Walters, R. and Wilson, J.M. (1989).
Survey and management proposals for a tropical deciduous forest reserve at Ankarana in
northern Madagascar. Biological Conservation 47: 297-313.
Ganzhorn, J. U. (1988). Food partitioning among Malagasy primates. Oecologia
(Berlin) 75: 436-450.
Hawkins, A.F.A., Ganzhorn, J.U., Bloxam, Q.M.C., Barlow, S.C.,
Tonge, S.J., and Chapman, P. (in press). A survey and assessment of the
conservation status and needs of lemurs, birds, lizards and snakes in the Ankarana
Special Reserve, Antseranana, Madagascar: with notes on the lemurs and birds of the
nearby Analamera Special Reserve. Biological Conservation.
Hladik, C.M. (1979). Diet and ecology of prosimians. In: Doyle, G.A. and Martin,
R.D. (Eds), The Study of Prosimian Behavior. Academic Press, New York. Pp. 307-
a7.
Hladik, C.M., Charles-Dominique, P. and Petter, J.-J. (1980). Feeding
strategies of five nocturnal prosimians in the dry forest of the west coast of Madagascar.
In: Charles-Dominique, P., Cooper, H.M., Hladik, A., Hladik, C.M., Pagés, E.,
Pariente, G.F., Petter-Rousseaux, A., Petter, J.-J. and Schilling, A. (Eds), Nocturnal
Malagasy Primates: Ecology, Physiology and Behavior. Academic Press, New York.
Pp. 41-73.
ISIS (1989). ISIS Species Distribution Report Abstract for Mammals, 30 June 1989.
International Species Information System, 12101 Johnny Cake Ridge Road, Apple
Valley, MN, U.S.A.
Jenkins, P.D. (1987). Catalogue of Primates in the British Museum (Natural History)
and elsewhere in the British Isles. Part IV: Suborder Strepsirrhini, including the
Subfossil Madagascan Lemurs and Family Tarsiidae. British Museum (Natural History),
London.
Martin, R.D. (1972). A preliminary field study of the lesser mouse lemur (Microcebus
murinus J. F. Miller 1777). Zeitschrift fiir Tierpsychologie, Supplement 9: 43-89.
Martin, R.D. (1973). A review of the behaviour and ecology of the lesser mouse lemur
(Microcebus murinus J.F. Miller 1777). In: Michael, R.P. and Crook, J.H.
(Eds),Comparative Ecology and Behaviour of Primates. Academic Press, London. Pp.
Nicoll, M.E. and Langrand, O. (1989). Revue Générale du Systéme d’Aires
Protégées et de la Conservation a Madagascar. Unpublished report to WWF.
Pagés-Feuillade, E. (1989). Modalités de l'occupation de l'espace et relations
interindividuelles chez un prosimien nocturne malagache (Microcebus murinus). Folia
Primatologica 50 (3/4): 204-220.
Paulian, R. (1981). Les mammiféres: vestiges d'un monde disparu. In: Oberlé, P. (Ed.),
Madagascar, Une Sanctuaire de la Nature. Le Chevalier, Paris. Pp. 75-94.
Petter, J-J., Albignac, R. and Rumpler, Y. (1977). Mammiféres lémuriens
(Primates prosimiens). Faune de Madagascar No. 44. ORSTOM-CNRS, Paris.
Petter, J.-J. (1978). Ecological and physiological adaptations of five sympatric
nocturnal lemurs to seasonal variation in food production. In: Chivers, D.J. and Herbert,
J. (Eds), Recent Advances in Primatology 1: Behaviour. Academic Press, London.
Pp.211-223.
Petter-Rousseaux, A. (1964). Reproductive physiology and behavior of the
Lemuroidae. In: Buettner-Janusch, J. (Ed.), Evolution and Genetic Biology of the
Primates.. Academic Press, New York. Pp. 91-132.
Petter-Rousseaux, A. (1980). Seasonal activity rhythms, reproduction, and body
weight variations in five sympatric nocturnal prosimians, in simulated light and climatic
conditions. In: Charles-Dominique, P., Cooper, H.M., Hladik, A., Hladik, C.M.,
Pagés, E., Pariente, G.F., Petter-Rousseaux, A., Petter, J.-J. and Schilling, A. (Eds),
Nocturnal Malagasy Primates: Ecology, Physiology and Behavior. Academic Press, New
York. Pp.137-152.
37
Lemurs of Madagascar and the Comoros
Petter-Rousseaux, A. (1988). Photopériode et réproduction de Microcebus murinus.
In: Rakotovao, L., Barre, V. and Sayer, J. (Eds), L'Equilibre des Ecosystémes forestiers
a Madagascar: Actes d'un séminaire international. IUCN, Gland, Switzerland and
Cambridge, UK. Pp. 72-77.
Raxworthy, C.J. and Rakotondraparany, F. (1988). Mammals report. In:
Quansah, N. (Ed.), Manongarivo Special Reserve (Madagascar), 1987/88 Expedition
Report. Unpublished report, Madagascar Environmental Research Group, U.K.
Richard, A.F., Sussman, R.W. and Ravelojaona, G. (1985). Madagascar:
current projects and problems in conservation. Primate Conservation 5: 53-59.
Susman, R.W. and Richard, A.F. (1986). Lemur conservation in Madagascar: the
status of lemurs in the south. Primate Conservation 7: 86-92.
Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New
York.
Wilde, J., Schwibbe, M.H. and Arsene, A. (1988). A census for captive primates
in Europe. Primate Report 21: 1-120.
38
Lemur, ana this is.pura trably related tothe pnerbiararhn. Gil forene 7
ih ol] 13 flame set SN ee sentabasaas i
town seu! BB on Se Leuuaa aa ncjae.
; baa : i,
hight. During. a brief study ac Ran
Lemurs of Madagascar and the Comoros
‘
The Brown Mouse Lemur, Microcebus rufus, is the smallest of the lemurs. It is found in the
eastern rain forests where it can survive in scrub secondary vegetation.
Photo by Mark Pidgeon.
The IUCN Red Data Book
BROWN or RUFOUS MOUSE LEMUR ABUNDANT
Microcebus rufus (Lesson, 1840)
Order PRIMATES Family CHEIROGALEIDAE
SUMMARY The smallest of the Malagasy primates, the Brown Mouse Lemur is found
throughout the eastern rain forest and across to the Sambirano Region in the north-west of
Madagascar. It is found in primary forest but is more common in secondary forest. Though
it is vulnerable to habitat destruction and its numbers are probably declining, it is unlikely to
be severely threatened. M. rufus has not been studied in any detail. It is a nocturnal, mostly
solitary species which feeds principally on fruit and insects. There are very few individuals
in captivity. It is found in several reserves. Listed in Appendix 1 of CITES, Class A of the
African Convention and is protected by Malagasy law.
DISTRIBUTION Brown Mouse Lemurs are found throughout the rain forest of eastem
Madagascar from Taolanaro (Fort Dauphin) to Montagne d'Ambre and across to the
Sambirano Region and Nosy Bé (Tattersall, 1982, Petter and Petter-Rousseaux, 1979;
Petter et al, 1977). A Brown Mouse Lemur (its specific status is unclear) also exists south
of the Sambirano River, though it occurs only sparsely there; specimens have been collected
near Morondava and have been reported from Ankarafantsika where they are sympatric with
the Grey Mouse Lemur (Petter et al, 1971; Petter et al, 1977; Tattersall, 1982; Petter and
Andriatsarafara, 1987).
POPULATION Population numbers are not known but Sussman et al (1985) report that
its numbers are "probably" declining. Density has been estimated in Analamazaotra Forest
(Perinet) as 110 + 34 individuals per sq. km (mean and 95% confidence limits) (Ganzhorm,
1988). However, the apparent population density in Analamazaotra was five times lower in
1985/86 than in 1984 (mean 0.11 per 100m compared to 0.52 per 100m respectively) and
Ganzhorn suggests this was due to the availability of fruiting shrubs and trees (Ganzhorn,
1987, 1988). This implies that, for M. rufus at least, figures for population density which
are based on transect walks can be very misleading. Petter and Petter-Rousseaux (1964)
found 250-262 individuals per sq. km at Mahambo on the east coast.
HABITAT AND ECOLOGY There are few, other than incidental, observations on this
species. It, like the Grey Mouse Lemur, appears to be more common in secondary
vegetation than in primary forest. During a short trapping study near Ranomafana in south-
eastern Madagascar, the Brown Mouse Lemur was seen most frequently in an old plantation
of the introduced Chinese guava, Psidium cattleyanum. Here an 80m trap line with ten traps
caught 24 individuals, while seven other traps spaced over 110m in a much less disturbed
area of primary forest caught only four individuals (Harcourt, 1987 and unpubl. data). At
Analamazaotra (Perinet) M. rufus can be found in old eucalyptus plantations, though at
greatly reduced densities compared to those in "natural" forest (Ganzhorn, 1987).
The diet of this species seems to be very similar to that of the Grey Mouse Lemur, they have
been seen eating fruit, insects and flowers (Martin, 1972; Harcourt, 1987) and, very
occasionally, young leaves (Ganzhorn, 1988). They are normally seen eating in shrubs and
little trees but they have also been seen in the tops of the tallest trees in Analamazaotra Forest
(Ganzhorn, 1988). They appear to be much less prone to storing fat in their tail than the
Grey Mouse Lemur, and this is probably related to the less marked seasonal differences in
food availability (Martin, 1972).
Little is known about the Brown Mouse Lemurs' social organisation, they are mostly seen
alone during the night. During a brief study at Ranomafana 23 males and only five females
41
Lemurs of Madagascar and the Comoros
were caught, but this biased sex ratio could not be explained (Harcourt, 1987). M. rufus
sleeps in tree holes and leaf nests in the daytime (Martin, 1972) and has been seen using old
birds' nests as well (Pollock, 1979). There are no reports as to the sex or numbers of
animals sleeping together during the day.
THREATS There are no recognised threats specific to the Brown Mouse Lemur, but all
Madagascar's primates are declining to a greater or lesser degree due to habitat destruction.
However, this small, nocturnal species must be one of the least threatened, especially as it
appears to thrive in secondary vegetation providing that fruit and insects are available there.
CONSERVATION MEASURES M. rufus is probably present in most of the protected
areas throughout its range. It is reported in Montagne d'Ambre National Park, in Marojejy,
Zahamena, Betampona, Andringitra and Andohahela Nature Reserves and in Manongarivo,
Analamazaotra, Anjanaharibe-Sud and Nosy Mangabe Special Reserves (Pollock, 1984;
Safford et al, 1989; Nicoll and Langrand, 1989; O'Connor et al, 1986; Raxworthy and
Rakotondraparany, 1988; Constable et al, 1985). Mouse Lemurs have been reported in
Ankarana and Analamera Special Reserves but it is not clear whether these are the Grey or
Brown species (Nicoll and Langrand, 1989; Ganzhorn, in litt.).
There are no conservation measures suggested specifically for the Brown Mouse Lemur, but
a range wide survey, to determine which Mouse Lemur is where, would be useful.
All species of Cheirogaleidae are listed in Appendix 1 of the 1973 Convention on
International Trade in Endangered Species of Wild Fauna and Flora. Trade in them, or their
products, is subject to strict regulation and may not be carried out for primarily commercial
purposes.
All Lemuroidea are listed in Class A of the African Convention, 1969. They may not,
therefore, be hunted, killed, captured or collected without the authorization of the highest
competent authority, and then only if required in the national interest or for scientific
purposes.
All lemurs are protected from unauthorised capture and from hunting by Malagasy law.
CAPTIVE BREEDING ISIS (June 1989) does not list any of this species in captivity.
Wilde et al (1988) report 18 individuals in Rotterdam Zoo and there is one pair at
Tsimbazaza in Madagascar (G. Rakotoarisoa and M. Pidgeon, in litt.). In captivity, they do
not breed as well as M. murinus (E. Simons, in litt.).
REMARKS This species is even smaller than the Grey Mouse Lemur, average weight is
around 50 g (Harcourt, 1987). It is distinguished from M. murinus by its slightly smaller
ears and the red tinge to its coat. For a more detailed description see Petter et al (1977),
Tattersall (1982) and Jenkins (1987). The taxonomic status of the Brown Mouse Lemurs
south of the Sambirano River is unclear (Tattersall, 1982). The Malagasy names of this
species are tsidy and tsitsihy or tsitsidy (Paulian, 1981; Tattersall, 1982).
REFERENCES
Constable, I.D., Mittermeier, R.A., Pollock, J.I., Ratsirarson, J. and
Simons, H. (1985). Sightings of aye-ayes and red ruffed lemurs on Nosy Mangabe
and the Masoala Peninsula. Primate Conservation 5: 59-62.
Ganzhorn, J.U. (1987). A possible role of plantations for primate conservation in
Madagascar. American Journal of Primatology 12: 205-215.
Ganzhorn, J.U. (1988). Food partitioning among Malagasy primates. Oecologia
(Berlin) 75: 436-450.
42
The IUCN Red Data Book
Harcourt, C.S. (1987). Brief trap/retrap study of the brown mouse lemur (Microcebus
rufus). Folia Primatologica 49: 209-211.
ISIS (1989). ISIS Species Distribution Report Abstract for Mammals, 30 June 1989.
International Species Information System, 12101 Johnny Cake Ridge Road, Apple
Valley, MN, U.S.A. Pp 17-22.
Jenkins, P.D. (1987). Catalogue of Primates in the British Museum (Natural History)
and elsewhere in the British Isles. Part IV: Suborder Strepsirrhini, including the
Subfossil Madagascan Lemurs and Family Tarsiidae. British Museum (Natural History),
London.
Martin, R.D. (1972). A preliminary field study of the lesser mouse lemur (Microcebus
murinus J. F. Miller 1777). Zeitschrift fiir Tierpsychologie, Supplement 9: 43-89.
Nicoll, M.E. and Langrand, O. (1989). Revue Générale du Systéme d’Aires
Protégées et de la Conservation a Madagascar. Unpublished report to WWF.
O'Connor, S., Pidgeon, M. and Randria, Z. (1986). Consexvation program for
the Andohahela Reserve, Madagascar. Primate Conservation 7: 48-52.
Paulian, R. (1981). Les mammiféres: vestiges d'un monde disparu. In: Oberlé, P.
(Ed.), Madagascar, Une Sanctuaire de la Nature. Le Chevalier, Paris. Pp. 75-94.
Petter, J-J., Albignac, R. and Rumpler, Y. (1977). Mammiféres lémuriens
(Primates prosimiens). Faune de Madagascar No. 44. ORSTOM-CNRS, Paris.
Petter, J.-J. and Andriatsarafara, F. (1987). Conservation status and distribution
of lemurs in the west and northwest of Madagascar. Primate Conservation 8: 169-171
Petter, J.-J. and Petter-Rousseaux, A. (1964). Premiére tentative d'estimation des
densités de peuplement des lémuriens malagaches. La Terre et la Vie 18: 427-435.
Petter, J.-J., Schilling, A. and Pariente, G. (1971). Observations éco-
éthologiques sur deux lemuriens malagaches noctumes: Phaner furcifer et Microcebus
coquereli. LaTerre etla Vie 25: 287-327.
Pollock, J.I. (1979). Spatial distribution and ranging behavior in lemurs. In: Doyle,
G.A. and Martin, R.D. (Eds), The Study of Prosimian Behavior. Academic Press, New
York. Pp. 359-409.
Pollock, J.I. (1984). Preliminary report on a mission to Madagascar by Dr J. I.
Pollock in August and September 1984. Unpublished report to WWF.
Raxworthy, C.J. and Rakotondraparany, F. (1988). Mammals report. In:
Quansah, N. (Ed.), Manongarivo Special Reserve (Madagascar), 1987/88 Expedition
Report. Unpublished report, Madagascar Environmental Research Group, U.K.
Safford, R.J., Durbin, J.C. and Duckworth, J.W. (1989). Cambridge
Madagascar rainforest expedition to R.N.I. No. 12 - Marojejy. Unpublished preliminary
report.
qecnan, R.W., Richard, A.F. and Ravelojaona, G. (1985). Madagascar:
current projects and problems in conservation. Primate Conservation 5: 53-59.
Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New
York.
Wilde, J., Schwibbe, M.H. and Arsene, A. (1988). A census for captive
primates in Europe. Primate Report 21: 1-120.
43
Lemurs of Madagascar and the Comoros
Coquerel's Dwarf Lemur, Mirza coquereli, is a nocturnal species which survives the dry
season in the western forests by eating carbohydrate-rich secretions from insect larvae.
Photo by Jean-Jacque Petter/WWF
The IUCN Red Data Book
COQUEREL'S DWARF LEMUR VULNERABLE
Mirza coquereli (A. Grandidier, 1867)
Order PRIMATES Family CHEIROGALEIDAE
SUMMARY Cogquerel's Dwarf Lemur is a nocturnal species found in some areas of dry
deciduous forest and in the more humid Sambirano Region in the west of Madagascar. The
extent of its distribution is not clear. Population numbers are not known and estimates of
density vary considerably. It appears to thrive in secondary forest, it has been reported at
densities as high as 385 individuals per sq. km in an area dominated by cashew nut trees. It
is, however, thought to be declining in number due to destruction of its habitat. The only
studies of M. coquereli have been brief and more are needed, though surveys to determine
its distribution would be more valuable in determining its conservation status. Its diet
includes insects, flowers, fruit, small vertebrates and, particularly, the secretions from
Homopteran larvae. It, like all the lemurs, is threatened by habitat destruction. This Dwarf
Lemur is found in only three protected areas. A colony of about 50 individuals at Duke
Primate Center breeds well, but the species is kept in few other institutes. Listed in
Appendix 1 of CITES, Class A of the African Convention and is protected by Malagasy
Law.
DISTRIBUTION Discontinuously distributed along the west coast of Madagascar. The
maps of its range given by Petter et al (1977), Petter and Petter-Rousseaux (1979) and
Tattersall (1982) differ quite considerably, those of the first two authors being more
extensive than the latter's. Tattersall (1982) shows its range from near Ankazoabo (i.e. just
south of the Mangoky River) northwards to around Antsalova and also in the Sambirano
region (the Ampasindava Peninsula and Ambanja area). Petter et al (1977) and Petter and
Petter-Rousseaux (1979) show it also extending along the west coast from around Cap St
André to Narinda’ Bay. However, in their text, Petter et al (1977) say that it only probably
exists in this stretch of west coast. M. Pidgeon (in litt.) reports seeing an individual of this
species as far south as the north bank of the Onilahy River, about 40 km due east from the
coast. His sighting was confirmed by M. Nicoll (in litt. from M. Pidgeon).
POPULATION Numbers are unknown and cannot be estimated until the range of
Coquerel's Dwarf Lemur is much better known. Petter et al (1971) estimated densities of 50
individuals per sq. km in Marosalaza Forest (50 km north of Morondava), but they recorded
as many as 210 per sq. km when counting in a strip of forest 5 m in width along either side
of a river in the region. In the same forest, Hladik et al (1980) estimated that there were 30
individuals per sq. km. The density (and biomass) of this species is the lowest of the five
nocturnal lemurs found in Marosalaza Forest (Hladik et al, 1980). Much higher densities of
M. coquereli have been reported from an area of secondary forest near Ambanja which was
dominated by Anacardium occidentalae (cashew nut) trees. Here there were as many as 385
individuals per sq. km (Andrianarivo, 1981). In 1975, Richard and Sussman considered
M. coquereli to be extremely rare and probably on the brink of extinction. Later, in 1985,
Sussman et al recorded the species as probably declining in number due to habitat
destruction.
HABITAT AND ECOLOGY Within the dry deciduous forest of western Madagascar,
Coquerel's Dwarf Lemur is generally found along rivers and near semipermanent ponds,
where the forest is thicker and slightly taller than in the drier areas (Petter et al, 1971). In
Marosalaza forest, M. coquereli has been briefly studied using radio tracking equipment
(Pages, 1978, 1980). This was done during part of the dry season (June-July) in 1974.
M. coquereli feeds on a variety of food resources including insects, spiders, frogs,
chameleons, small birds, fruits, flowers, buds, gums, and insect secretions (Pages, 1980,
45
Lemurs of Madagascar and the Comoros
Andrianarivo, 1981). During the dry season, the secretions of cochineals and homopteran
larvae are particularly important (Petter et al, 1971; Hladik et al, 1980) and, during June in
Marosalaza Forest these accounted for 50% of the feeding observations (Pages 1980).
However, these secretions are low in protein (Hladik et al, 1980) and it appears that the
distribution of Coquerel's Dwarf Lemur depends more on the availability of other insects
than on these colonies (Pages, 1980). Andrianarivo (1981) found that in secondary forest
dominated by cashew nut trees, the cashew fruits were a very important food source during
the dry season. In Marosalaza, feeding usually occurred at heights between 1.5 and 3m,
though M. coquereli may also forage on the ground (Pages, 1980).
The species is nocturnal, spending the day in a spherical nest made of interlaced lianas,
branches and leaves which is usually located in the fork of a large branch or among dense
lianas at a height of 2-10m (Petter et al, 1971; Pages 1980, Andrianarivo, 1981). In
Marosalaza Forest a female and her offspring could be found sharing a nest, but males were
never seen with them (Pages, 1980). This was not the case at a study site in secondary
forest near Ambanja. Here, out of four occupied nests two contained an adult male and
female, one contained two young individuals and in the fourth there was an adult female and
a young animal (Andrianarivo, 1981). In the area near Ambanja, nests were found clustered
together in "villages" of about 1 ha in size (Andrianarivo, 1981). Six to ten individuals lived
in each of these "villages" and though they changed nests within the area they did not move
between nesting areas (Andrianarivo, 1981). Individuals left their nests around dusk and
generally spent the first half of the night feeding, self-grooming or resting, while the second
half of the night was devoted more to social activities such as vocalisations, mutual
grooming and play (Pages, 1978, 1980). There was also an increase in distance travelled
during this second part of the night (Pages, 1978, 1980).
Pages (1978, 1980) found that the home range of adults of both sexes appeared to contain a
heavily used and defended central area (1.5 ha for males and 2.5-3.0 ha for females),
surrounded by a large peripheral area (a maximum of 4 ha for males and 4.5 ha for females),
which was less frequently visited. There was a much greater degree of overlap in the
peripheral area, however even the central core could be overlapped by an adult individual of
the opposite sex or, in the case of females, by their offspring (Pages, 1980). The males
generally made longer incursions into distant areas than did females (Pages, 1980).
Meetings between individuals were rare, males encountered other animals only every other
night on average and prolonged contact was even less frequent (Pages, 1980). Although
adult males met a number of adult females, their periods of prolonged contact appeared to be
restricted to just one of these females; Pages (1978, 1980) suggested a loose pair bonding
social system in this species.
Mating takes place in October, gestation lasts three months (Petter-Rousseaux, 1980) and
normally two infants are born (Pages, 1978). Infants initially stay in the nest, they leave
this for the first ttme when they are about three weeks old (Pages, 1980). They, like most
of the other members of the family Cheirogaleidae, do not ride on their mother, but are
carried in her mouth (Pages, 1980).
THREATS M. coquereli may be threatened by habitat destruction though its high density
in secondary forest suggests that it may survive the disappearance of its natural habitat. The
forests in the Sambirano Region are being cleared for cultivation, while the dry deciduous
forests are mostly being destroyed by fire. These are frequently set to encourage new grass
growth for the large numbers of livestock that are kept in western Madagascar. In 1981,
pert NEP estimated net degradation of the western forests to be perhaps 200,000 ha since
1 :
CONSERVATION MEASURES Found in Bemaraha Nature Reserve, Andranomena
Special Reserve and the Private Reserve of Analabe (Nicoll and Langrand, 1989). All three
reserves would benefit from more guards with better equipment to protect them, particularly
46
The IUCN Red Data Book
from fires. Signposting of the Reserve boundaries would be an asset, as would an
education programme for the local villagers to emphasise the uniqueness of the reserves and
their biota. Analabe has great tourist potential and this could be developed for the benefit of
both the wildlife and the local people (Nicoll and Langrand, 1989)
Surveys are desirable to determine the actual distribution and numbers of M. coquereli so
that an effective conservation strategy can be developed.
All species of Cheirogaleidae are listed in Appendix 1 of the 1973 Convention on
International Trade in Endangered Species of Wild Fauna and Flora. Trade in them, or their
products, is subject to strict regulation and may not be carried out for primarily commercial
purposes.
All Lemuroidea are listed in Class A of the African Convention, 1969. They may not,
therefore, be hunted, killed, captured or collected without the authorization of the highest
competent authority, and then only if required in the national interest or for scientific
purposes.
All lemurs are protected from unauthorised capture and from hunting by the laws of
Madagascar. However, its is very difficult to enforce these regulations.
CAPTIVE BREEDING This species breed wells in captivity (Petter et al, 1977; E.
Simons, in litt.), but it is not kept by many institutions. According to the ISIS records (June
1989) there are 62 individuals in captivity, most of which (45) are at Duke Primate Center;
Cincinnati holds eight animals, San Francisco has six and Paris has three. It is reported that
95% of the 62 animals are captive born; all those in American insitutions are descendents of
six individuals imported by Duke in 1982 (E. Simons, in litt.). Duke Primate Center is now
coordinating a captive breeding programme for this species (E. Simons, pers. comm.).
REMARKS Tattersall (1982) puts this species in its own genus; however, it is still
frequently referred to as belonging to the genus Microcebus. M. coquereli, at 300g (Pages,
1978), is considerably larger than the Mouse Lemurs. Dorsally, the fur of this species is
brown or grey-brown, sometimes with rosy or yellowish tinges; ventrally the grey base
colour of the downy hair shows through beneath the yellowish or slightly russet tips
(Tattersall, 1982). The ears of Coquerel's Dwarf Lemur are long and hairless. For a more
detailed description see Petter et al (1977) or Tattersall (1982). In the southern area of its
range, this species is called tsiba or tilitilivaha and setohy or fitily in its northern range.
REFERENCES
Andrianarivo, A.J. (1981). Etude comparee de l'organisation sociale chez Microcebus
coquereli. Unpublished dissertation, University of Madagascar, Antananarivo.
FAO/UNEP (1981). Tropical Forest Resources Assessment Project. Forest Resources
of Tropical Africa. Part II Country Briefs. FAO, ROME.
Hladik, C.M., Charles-Dominique, P. and Petter, J.-J. (1980). Feeding
strategies of five nocturnal prosimians in the dry forest of the west coast of Madagascar.
In: Charles-Dominique, P., Cooper, H.M., Hladik, A., Hladik, C.M., Pages, E.,
Pariente, G.F., Petter-Rousseaux, A., Petter, J.-J. and Schilling, A. (Eds), Nocturnal
Malagasy Primates: Ecology, Physiology and Behavior. Academic Press, New York.
. 41-73.
Iss (1989). ISIS Species Distribution Report Abstract for Mammals, 30 June 1989.
International Species Information System, 12101 Johnny Cake Ridge Road, Apple
Valley, MN, U.S.A.
Nicoll, M.E. and Langrand, O. (1989). Revue Générale du Systéme d’Aires
Protégées de la Conservation a Madagascar. Unpublished report to WWF.
47
Lemurs of Madagascar and the Comoros
Pages, E. (1978). Home range, behaviour and tactile communication in a nocturnal
malagasy lemur Microcebus coquereli. In: Chivers, D.A. and Joysey, K.A. (Eds),
Recent Advances in Primatology, Vol 3. Academic Press, London. Pp. 171-177.
Pages, E. (1980). Ethoecology of Microcebus coquereli during the dry season. In:
Charles-Dominique, P., Cooper, H.M., Hladik, A., Hladik, C.M., Pages, E., Pariente,
G.F., Petter-Rousseaux, A., Petter, J.-J. and Schilling, A. (Eds), Nocturnal Malagasy
Primates: Ecology, Physiology and Behavior. Academic Press, New York. Pp. 97-116.
Petter, J-J., Albignac, R. and Rumpler, Y. (1977). Mammiféres lémuriens
(Primates prosimiens). Faune de Madagascar No. 44. ORSTOM-CNRS, Paris.
Petter, J.-J. and Petter-Rousseaux, A. (1979). Classification of the Prosimians
In: Doyle, G.A. and Martin, R.D. (Eds), The Study of Prosimian Behavior. Academic
Press, London. Pp. 1-44.
Petter, J.-J., Schilling, A. and Pariente, G. (1971). Observations éco-
éthologiques sur deux lemuriens malagaches nocturnes: Phaner furcifer et Microcebus
coquereli. La Terre et la Vie 25: 287-327.
Petter-Rousseaux, A. (1980). Seasonal activity rhythms, reproduction, and body
weight variations in five sympatric nocturnal prosimians, in simulated light and climatic
conditions. In: Charles-Dominique, P., Cooper, H.M., Hladik, A., Hladik, C.M.,
Pages, E., Pariente, G.F., Petter-Rousseaux, A., Petter, J.-J. and Schilling, A. (Eds),
Nocturnal Malagasy Primates: Ecology, Physiology and Behavior. Academic Press,
New York. Pp. 97-116.
Pages, E., Pariente, G.F., Petter-Rousseaux, A., Petter, J.-J. and
Schilling, A. (Eds), Nocturnal Malagasy Primates: Ecology, Physiology and
Behavior. Academic Press, New York. Pp. 137-152.
Richard, A.F. and Sussman, R.W. (1975). Future of the Malagasy lemurs:
conservation or extinction? In: Tattersall, I. and Sussman, R.W. (Eds), Lemur Biology,
Plenum Press, New York. Pp. 335-350.
Sussman, R.W., Richard, A.F. and Ravelojaona (1985). Madagascar: current
projects and problems in conservation. Primate Conservation 5: 53-58.
Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New
York.
; uyuin shy beter rs tuk WY auger!
Lemurs of Madagascar and the Comoros
The Fat-tailed Dwarf Lemur, Cheirogaleus medius, stores fat in its tail which enables it to
survive long periods of food shortage in the dry western forests.
Photo by Russell Mittermeier.
50
The IUCN Red Data Book
FAT-TAILED DWARF LEMUR ABUNDANT
Cheirogaleus medius KE. Geoffroy, 1812
Order PRIMATES Family CHEIROGALEIDAE
SUMMARY The Fat-tailed Dwarf Lemur is one of the smaller, nocturnal lemurs. It has a
wide distribution in the dry forests, both primary and secondary, in the west and south of
Madagascar. Population numbers are unknown, but the species is considered to be
declining as the dry forests are being reduced in area. However, it can be found at densities
as high as 300-400 per sq. km which, along with its wide distribution, suggests that it may
not be severely threatened. The species has been studied briefly but little is known of its
social organisation. Its diet consists of fruit, flowers and insects. The most characteristic
feature of Cheirogaleus medius is its ability to become torpid for six to eight months in the
dry season. It is found in at least seven protected areas. There is a minimum of 120
individuals in captivity and most are captive bred. Listed in Appendix 1 of CITES and Class
A of the African Convention and protected by Malagasy law.
DISTRIBUTION Found in the dry forests of southern and western Madagascar from
Narinda Bay to Taolanaro (Fort Dauphin) (Tattersall, 1982; Petter and Petter-Rouseaux,
1979). Locality information on museum specimens collected late last century and in 1929-
1931 indicate that the Fat-tailed Dwarf Lemur was then found in eastern and northern
Madagascar and in the Sambirano region, in sympatry with C. major (Tattersall, 1982), but
there are no recent reports of its occurrence in these areas except in Ankarana, where
Cheirogaleus sp. was seen and members of the expedition identified it provisionally as
C. medius (Hawkins et al, in press).
POPULATION Numbers are unknown but the species is considered to be declining due
to habitat destruction (Richard and Sussman, 1975, 1987). It is, however, found at
densities as high as 300-400 animals per sq. km in deciduous forest near Morondava
(Petter, 1978; Hladik, 1979), which, along with its wide distribution, suggests that it may
not be severely threatened as yet. Estimates of population densities at other areas are lower,
37 individuals per sq. km at Berenty (Russell, in Jolly, 1987, 1988), and either 12
(Albignac, 1981) or 81 + 36 individuals per sq. km (mean and 95% confidence limits based
on 10 census walks along 1.7 km of trail [Ganzhorn, 1988]) at Ankarafantsika.
HABITAT AND ECOLOGY Fat-tailed Dwarf Lemurs are found in both primary and
well-established secondary forest (Martin, 1984). In 1935, Rand reported that he found
them in gallery forest through savanna and dry brush. The animals are active for half the
year or less, they avoid the seasonal shortage of food by becoming torpid during the dry
season (May-October). At this time individuals can be found alone or in groups of up to
five individuals in hollow tree trunks (Petter, 1978, Hladik, 1979; Hladik et al, 1980). The
age/sex composition of the dormant groups varies. Petter (1988) found solitary adults of
both sexes; two young females; an adult male and female with two young females; an adult
male and female with one young female; and a group of two adult males, one adult female
and two young females during observations at his study site near Morondava. Adults can
become torpid as early as March, while offspring born that year tend to become dormant
slightly later, thereby suffering less competition for food (Hladik et al, 1980). They re-
emerge in November at the beginning of the rainy season (Petter, 1978).
In Marosalaza Forest, the diet of C. medius includes fruit (Operculicarya gummifera,
Grewia glanulosa, Strychnos decussata and Diospyros aculeata) and flowers (Baudouina
fluggeiformis), the nectar of some flowers (e.g. Delonix floribunda), insects (especially
beetles), a few leaf buds, gums and some small vertebrates (the skin of a chameleon was
51
Lemurs of Madagascar and the Comoros
C. medius
a C. major
Capricorn
Figure 7: Distribution of both species of Cheirogaleus. Shaded areas represent approximate
limits of ranges.
52
The IUCN Red Data Book
found in one faecal sample) (Hladik, 1979, Hladik et al, 1980). Flowers and nectar are
used at the beginning of the rainy season (November), fruits and an increasing proportion of
insects are taken from December to February, while from then onwards fruits are the staple
food (Hladik, 1979; Hladik et al, 1980). During the rains, the animals lay down fat under
their skin and in their tail, their body weight increases by approximately 75 g, to about
220 g, and the volume of their tail triples, from a mean of 15 cc in November to a mean of
42 cc in May (Hladik et al, 1980).
In a study of 31 marked individuals, C. medius had a home range with a maximum diameter
of 200 m (about 4 ha) and the ranges of adult animals overlapped (Hladik et al, 1980). In
Captivity, adult animals of the same sex are intolerant of each other (Hladik et al, 1980), but
there is no information available on their social organisation in the wild.
In Marosalaza Forest, mating was observed at the beginning of November and infants were
born in January (Petter, 1978; Hladik et al, 1980). Gestation in this species is 61-64 days;
litter size varies from one to four, but twins are most frequently produced (Foerg, 1982). In
captivity, offspring reached adult weight between the 14th and 16th week of life and they
attained sexual maturity in their first year (Foerg, 1982).
THREATS There appear to be no threats specific to C. medius. However, the dry forests
of the west are being reduced in area every year. Even in the early 1970s few large areas of
western forest remained, most persist only in small isolated residual patches
(IUCN/UNEP/WWF, 1987). FAO/UNEP (1981) estimated net degradation of these forests
to have been perhaps 200,000 ha since 1955. Fires are set each year to encourage new
grass growth for grazing and this is the principal cause of forest destruction in the west.
Similarly, the southern forests are being degraded, collection of wood for charcoal is one of
the major threats. Though the Fat-tailed Dwarf Lemur has been seen in brush vegetation
(Rand, 1935), it is likely that it needs hollow trees of a certain size to be able to survive the
dry season. Nevertheless, the wide distribution of the species and its ability to survive in
secondary forest makes it unlikely that it is severely threatened.
CONSERVATION MEASURES C. medius is reported in two Nature Reserves,
Ankarafantsika and Andohahela, it is also found in the Special Reserves of Andranomena,
Beza Mahafaly and, probably, Ankarana and the Private Reserves of Berenty and Analabe
(Richard, 1975; Nicoll and Langrand, 1989; O'Connor et al, 1986, 1987;
Andriamampianina, 1981; Hawkins et al, in press).
The Department of Water and Forests (Direction des Eaux et Foréts) and the World Bank are
developing a management programme for Ankarafantsika Reserve and the Classified Forest
of Ampijoroa (Nicoll and Langrand, 1989). More money is required for effective guarding
of the Reserve and a fire break is needed around the area (Nicoll and Langrand, 1989). In
addition, a reafforestation programme is necessary, to provide the local people with fuel and
building material, as is education as to the importance of the forest and the protected area
(Nicoll and Langrand, 1989).
The other reserves in which C. medius is found suffer from more or less the same problems:
cattle grazing within them, fires destroying more each year, cutting and collecting of wood
and some illegal hunting. Similar conservation measures are needed for each, more
protection, education of the local people and development of alternatives to using the forest
for fuel and building materials. If tourists can be encouraged, as has happened at Berenty,
this provides money and employment for the local people.
All species of Cheirogaleidae are listed in Appendix 1 of the 1973 Convention on
International Trade in Endangered Species of Wild Fauna and Flora. Trade in them, or their
products, is subject to strict regulation and may not be carried out for primarily commercial
purposes.
53
Lemurs of Madagascar and the Comoros
All Lemuroidea are listed in Class A of the African Convention, 1969. They may not,
therefore, be hunted, killed, captured or collected without the authorization of the highest
competent authority, and then only if required in the national interest or for scientific
purposes.
Malagasy law protects all lemurs from unauthorised capture or killing, but this is very
difficult to enforce.
CAPTIVE BREEDING Breeds well in captivity. In June 1989, there were 59 males,
45 females and 12 unsexed individuals held in nine institutes, of which 94 % were captive
born (ISIS). The majority (49) are at Duke Primate Center. Wilde et al (1988) report
another nine individuals in two institutes not included in the ISIS list. Many more could be
bred, but few zoos are interested as a small, dull, nocturnal species does not make a good
exhibit (E. Simons, in litt.).
REMARKS This species is sometimes divided into two subspecies, C. m. medius and
C.m. samati, but Petter and Petter (1971), Petter et al (1977) and Tattersall (1982) consider
this distinction unwarranted. Fur is short and dense, grey with rosy or brownish tints on
the upperparts and white to light brown on the underparts (Tattersall, 1982). Body weight
changes seasonally. In Marosalaza Forest, adult mean body weight varied from 142 g in
November to 217g in March (Hladik et al, 1980), an individual can weigh as much as 400 g
(Petter et al, 1977). See Tattersall (1982) for a more detailed description of the species. In
the north-west, the Malagasy name for this species is matavirambo or matavrambo; kely be-
ohy or kelibehohy in the Morondava region and tsidy or tsitsihy in the far south (Tattersall,
1982; Petter et al, 1977).
REFERENCES
Albignac, R. (1981). Lemurine social and territorial organisation in a north-western
Malagasy forest (restricted area of Ampijoroa). In: Chiarelli, A.B. and Corruccini, R.S.
(Eds), Primate Behavior and Sociobiology. Springer Verlag, Berlin. Pp. 25-29.
Andriamampianina, J. (1981). Les réserves naturelles et la protection de la nature a
Madagascar. In Oberlé (Ed.), Madagascar, un Sanctuaire de la Nature. Pp. 105-111.
FAO/UNEP (1981). Tropical Forest Resources Assessment Project. Forest Resources
of Tropical Africa. Part II Country Briefs. FAO, Rome.
Foerg, R. (1982). Reproduction in Cheirogaleus medius. Folia Primatologica 39: 49-
62
Ganzhorn, J.U. (1988). Food partitioning among Malagasy primates. Oecologia 75:
436-450.
Hawkins, A.F.A., Ganzhorn, J.U., Q.M.C. Bloxham, Barlow, S.C., Tonge
S.J. and Chapman, P. (in press). A survey and assessment of the conservation
status and needs of lemurs, birds, lizards and snakes in Ankarana Special Reserve,
Antseranana, Madagascar: with notes on the lemurs and birds of the nearby Analamera
Special Reserve. Biological Conservation
Hladik, C.M. (1979). Diet and ecology of prosimians. In: Doyle, G.A. and Martin,
R.D. The Study of Prosimian Behavior. Academic Press, New York. Pp. 307-357.
Hladik, C.M. and Charles-Dominique, P. and Petter, J.-J. (1980). Feeding
strategies of five nocturnal prosimians in the dry forest of the West coast of Madagascar.
In: Charles-Dominique, P., Cooper, H. M., Hladik, A., Hladik, C. M., Pages, E.,
Pariente, G. F., Petter-Rousseaux, A., Petter, J.-J. and Schilling, A. (Eds), Nocturnal
Malagasy Primates: Ecology, Physiology and Behavior. Academic Press, New York.
Pp. 41-73.
ISIS (1989). [SIS Species Distribution Report Abstract for Mammals, 30 June 1989.
International Species Information System, 12101 Johnny Cake Ridge Road, Apple
Valley, MN, U.S.A.
54
The IUCN Red Data Book
IUCN/UNEP/WWE (1987). Madagascar, an Environmental Profile. Edited by M.D.
Jenkins. IUCN, Gland, Switzerland and Cambridge, U.K.
Jolly, A. (1987). Priorités dans l'étude des populations de Lémuriens. In: Priorités en
Matiére de Conservation des Espéces a Madagascar. Occasional Papers of the IUCN
Species Survival Commission, Number 2.
Jolly, A. (1988). Lemur survival. In: Benirschke, K. (Ed.), Primates: The Road to
Self-Sustaining Populations. Springer-Verlag, New York. Pp. 71-98.
Martin, R.D. (1984). Dwarf and mouse lemurs. In: Macdonald, D. (Ed.), The
Encyclopaedia of Mammals:1. George Allen and Unwin, London. P. 331.
Nicoll, M.E. and Langrand, O. (1989). Revue Générale du Systéme d’Aires
Protégées et de la Conservation a Madagascar. Unpublished report to WWF.
O'Connor, S., Pidgeon, M. and Randria, Z. (1986). Conservation program for
the Andohahela Reserve, Madagascar. Primate Conservation 7: 48-52.
O'Connor, S, Pidgeon, M. and Randria, Z. (1987). Un programme de
conservation pour la Réserve d'Andohahela. In: Priorités en Matiére de Conservation des
Espéces a Madagascar. Occasional Papers of the IUCN Species Survival Commission,
Number 2.
Petter, A. and Petter, J.-J. (1971). Part 3.1 Infraorder Lemuriformes. In: Meester,
J. and Setzer,H.W. (Eds), The Mammals of Africa: An Identification Manual.
Smithsonian Institution Press, City of Washington. Pp. 1-10.
Petter, J-J., Albignac, R. and Rumpler, Y. (1977). Mammiféres lémuriens
(Primates prosimiens). Faune de Madagascar No. 44. ORSTOM-CNRS, Paris.
Petter, J.-J. and Petter-Rousseaux, A. (1979). Classification of the Prosimians
In: Doyle, G.A. and Martin, R.D. (Eds), The Study of Prosimian Behavior. Academic
Press, London. Pp. 1-44.
Petter, J.-J. (1988). Contribution a l'étude du Cheirogaleus medius dans la féret de
Morondava. In: Rakotovao, L., Barre, V. and Sayer, J. (Eds), L’Equilibre des
Ecosystémes forestiers @ Madagascar: Actes d'un séminaire international. IUCN Gland,
Switzerland and Cambridge, U.K. Pp. 57-60.
Rand, A.L. (1935). On the habits of some Madagascar mammals. Journal of
Mammalogy, 16: 89-104.
Raxworthy, C.J. and Rakotondraparany, F. (1988). Mammals report. In:
Quansah, N. (Ed.), Manongarivo Special Reserve (Madagascar), 1987/88 Expedition
Report. Madagascar Environmental Research Group, U.K.
Richard, A. (1975). Patterns of mating in Propithecus verreauxi verreauxi. In: Martin,
R.D., Doyle, G.A. and Walker, A.C. (Eds), Prosimian Biology. Duckworth, London.
. 49-74,
einai. A.F. and Sussman, R.W. (1975). Future of the Malagasy lemurs;
conservation or extinction? In: Tattersall, I. and Sussman, R.W. (Eds), Lemur Biology.
Plenum Press, New York. Pp. 335-350.
Richard, A.F. and Sussman, R.W. (1987). Framework for Primate Conservation
in Madagascar. In: Marsh, C.W. and Mittermeier, R. (Eds), Primate Conservation in the
Tropical Forest. Alan R. Liss, Inc., New York. Pp. 329-341.
Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New
York.
Wilde, J., Schwibbe, M.H. and Arsene, A. (1988). A census for captive
primates in Europe. Primate Report 21: 1-120.
55
Lemurs of Madagascar and the Comoros
The nocturnal Greater Dwarf Lemur, Cheirogaleus major, is found in the eastern rain
forests.
Photo by Phillip Coffrey/Jersey Wildlife Preservation Trust.
56
The IUCN Red Data Book
GREATER DWARF LEMUR ABUNDANT
Cheirogaleus major E. Geoffroy, 1812
Order PRIMATES Family CHEIROGALEIDAE
SUMMARY The Greater Dwarf Lemur is a small, nocturnal species that is widely
distributed in the eastern rain forest. There are no estimates of total population number, but
it is reported to occur at high densities in some areas. It is possible that Cheirogaleus major
is one of the least threatened of the lemur species but, nevertheless, it is less widespread
than it was even a few decades ago. There have been no detailed studies of this species and
little is known about its habitat requirements or its ecology. It is a nocturnal species, usually
sighted alone at night. Its diet includes ripe fruit and invertebrates. It has a three month
torpid period from July to October. It occurs in at least nine protected areas. Only three
individuals, all female, are held in captivity. Listed in Appendix 1 of CITES, in Class A of
the African Convention and protected by Malagasy law.
DISTRIBUTION Found throughout the eastern rain forest, from Taolanaro (Fort
Dauphin) in the south to Montagne d'Ambre in the far north and extending westwards to the
Tsaratanana Massif and the Sambirano region (Tattersall, 1982). As recently as a few
decades ago the range of this species extended onto the central plateau (Tattersall, 1982).
Petter et al, (1977) and Petter and Petter-Rouseaux (1979) show a population of C. major on
the Bongolava Massif, near the Manambolo River in the west of Madagascar. In a 1987
publication Petter and Andriatsarafara report that C. major have recently been caught on the
Bongolava Massif but they gives no details. They also think it likely that the species is
present in Ankarafantsika.
POPULATION Numbers are unknown but Richard and Sussman (1975, 1987) report
that the species is declining. It is found at high densities in some areas (Petter et al, 1977),
as many as 75-110 per sq. km were reported at Mahambo (Petter and Petter-
Rousseaux,1964), while Ganzhorn estimated 68 + 38 individuals per sq. km (mean and
95% confidence limits based on 25 census walks along 3.2 km of trails) in Analamazaotra
Forest. The wide distribution of the species suggests that total population number may be
quite high.
HABITAT AND ECOLOGY A small, nocturnal species, which lives in primary rain
forest and well-established secondary forest (Martin 1984). Little is known about its social
organisation but it is invariably sighted alone (Petter et al, 1977). During the day it may
sleep in a tree hollow ora nest. Petter et al (1977) report finding two adults together in one
nest. There are few data on ranging patterns of this species though Martin (1972) found that
Dwarf Lemurs rarely descended below 3 m in the trees and that they preferred large
branches to fine ones. He describes them as slow moving, essentially quadrupedal forms.
The diet of the Greater Dwarf Lemur consists of ripe fruit, nectar and pollen with insects
and, probably, small vertebrates also being taken, they have never been seen to eat leaves
(Petter et al, 1977). A peculiarity of both this species and the Fat-tailed Dwarf Lemur is
their ability to store fat in their tail, which enables them to survive periods of dormancy in
winter. C. major is reported to have a three month torpid period between July and October
(Petter et al, 1977). During this time they hide in the leaf litter at the foot of a big tree
(Paulin, 1981; Petter et al, 1977).
Length of gestation in C. major is 70 days and the infants are born in January, in
Madagascar's summer (Petter-Rousseaux, 1964). A litter of iwo or three offspring is
generally produced (Petter et al, 1977). They cannot cling to their mother, instead she
carries them in her mouth when necessary (Petter-Rousseaux, 1964). Within a month of
37
Lemurs of Madagascar and the Comoros
birth the infants can follow their mother when she goes to feed (Petter-Rousseaux, 1964).
Lactation lasts about 1.5 months, but the young start eating fruit at about 25 days of age
(Petter-Rousseaux, 1964).
THREATS Destruction of the rain forests for timber, fuel and agricultural land is a threat
to this species. However its small size and nocturnal habits will ensure that it survives
longer than its larger, diurnal relatives. Its wide distribution within the rain forest suggests
that it may be one of the less threatened of the lemurs, though more information on its
habitat requirements is needed before its true status can be ascertained. Both E. Simons and
I. Tattersall (in litt.) consider this species more threatened than C. medius, the Fat-tailed
Dwarf Lemur found in the west of Madagascar. Tattersall's (1982) report that Greater
Dwarf Lemurs used to be found in areas of the central plateau as recently as a few decades
ago does indicate that it is susceptible to changes in its habitat. C. major is hunted for food
by local people using long sticks to poke around in holes where it might be resting (Petter er
al, 1977).
CONSERVATION MEASURES The Greater Dwarf Lemur occurs in a number of
protected areas including the Montagne d'Ambre National Park, the Nature Reserves of
Betampona, Zahamena, Tsaratanana and Andohahela and the Special Reserves of
Analamazaotra, Nosy Mangabe and Manongarivo (Andriamampianina and Peyrieras, 1972;
Nicoll and Langrand, 1989; Constable et al, 1985; O'Connor et al, 1986, Raxworthy and
Rakotondraparany, 1988). Several new areas which contain C. major have been proposed
for protection (Nicoll and Langrand, 1989). These are Ranomafana and Masoala (both
proposed as National Parks) and Mananara (proposed as a Biosphere Reserve). All the
protected areas need adequate funding and guards to ensure the survival of the lemurs within
them. In addition, education projects for the local people to demonstrate the importance of
the reserves would be useful, as would development programmes that provide alternatives to
harmful exploitation of the protected areas.
Studies of the habitat requirements of the Greater Dwarf Lemur would help to ascertain its
conservation status. E. Simons (in litt.) considers it important to import males to join the
females at Duke Primate Center so that a captive breeding colony of this species can be
established.
All species of Cheirogaleidae are listed in Appendix 1 of the 1973 Convention on
International Trade in Endangered Species of Wild Fauna and Flora. Trade in them, or their
products, is subject to strict regulation and may not be carried out for primarily commercial
purposes.
All Lemuroidea are listed in Class A of the African Convention, 1969. They may not,
therefore, be hunted, killed, captured or collected without the authorization of the highest
competent authority, and then only if required in the national interest or for scientific
purposes.
All lemurs are protected under Malagasy law. The national legislation is based primarily on
the 1933 London Convention and on Ordonnance No. 60-126 of 3rd October 1960. It is,
however, difficult to enforce the laws preventing capture or killing of the lemurs.
CAPTIVE BREEDING Three wild born individuals, all females, are held at Duke
Primate Center (ISIS, June 1989). It appears that this species survives in captivity but it is
difficult to breed (Petter et al, 1977). However, Petter et al (1977) reports the birth of
triplets to one female and twins to two others in Paris. There are no longer any captive
individuals in Paris.
REMARKS Some authors recognise two subspecies of the Greater Dwarf Lemur, C. m.
crossleyi, with reddish fur, occurring north of the Masoala Peninsula and the browner C.
58
The IUCN Red Data Book
m. major to the south (Petter and Petter, 1971; Petter et al, 1977; Petter and Petter-
Rousseaux, 1979; Jenkins, 1987). Tattersall (1982) considers the species variable but
monotypic. Fur short and dense, grey brown to reddish above with paler underparts; dark
rings around the eyes (Tattersall, 1982; Martin, 1984). Body weight varies seasonally,
mean is about 450 g (Petter et al, 1977, Tattersall, 1982; Martin, 1984). Malagasy names
for this species are tsitsihy, tsidy and hataka (Tattersall, 1982).
REFERENCES
Constable, I.D., Mittermeier, R.A., Pollock, J.I., Ratsirarson, J. and
Simons, H. (1985). Sightings of aye-ayes and red-ruffed lemurs on Nosy Mangabe
and the Masoala Peninsula. Primate Conservation 5: 59-62.
Jenkins, P.D. (1987). Catalogue of Primates in the British Museum (Natural History)
and elsewhere in the British Isles. Part IV: Suborder Strepsirrhini, including the
pees Madagascan lemurs and family Tarsiidae. British Museum (Natural History),
London.
Martin, R.D. (1984). Dwarf and mouse lemurs. In: Macdonald, D. (Ed.), The
Encyclopaedia of Mammals:1. George Allen and Unwin, London. P. 331.
O'Connor, S., Pidgeon, M. and Randria, Z. (1986). Conservation program for
the Andohahela Reserve, Madagascar. Primate Conservation 7: 48-52.
Paulin, R.R. (1981). Les mammiféres: vestiges d' un monde disparu. In: Oberlé, P.
(Ed.), Madagagascar un sanctuaire de la nature. Libraire de Madagascar, Antananarivo.
Pp. 75-94.
Petter, J.-J. and Andriatsarafara, F. (1987). Conservation status and distribution
of lemurs in the west and northwest of Madagascar. Primate Conservation 8: 169-171.
Petter, A. and Petter, J.-J. (1971). Part 3.1 Infraorder Lemuriformes. In: Meester,
J. and Setzer, H.W. (Eds), The Mammals of Africa: An Identification Manual.
Smithsonian Institution Press, City of Washington. Pp. 1-10.
Petter, J-J., Albignac, R. and Rumpler, Y. (1977). Mammiféres lémuriens
(Primates prosimiens). Faune de Madagascar No. 44. ORSTOM-CNRS, Paris.
Petter, J.-J. and Petter-Rousseaux, A. (1979). Classification of the Prosimians
In: Doyle, G.A. and Martin, R.D. (Eds), The Study of Prosimain Behavior. Academic
Press, London. Pp. 1-44.
Petter, J.J. and Petter-Rousseaux, A. (1964). Premiére tentative d'estimation des
densités de peuplement des Iémuriens malagaches. La Terre et La Vie 18: 427-435.
Petter-Rousseaux, A. (1964). Reproductive physiology and behavior of the
Lemuroidae. In: Buettner-Janusch, J. (Ed.), Evolution and Genetic Biology of the
Primates. Academic Press, New York. Pp. 91-132.
Raxworthy, C.J. and Rakotondraparany, F. (1988). Mammals report. In:
Quansah, N. (Ed.), Manongarivo Special Reserve (Madagascar), 1987/88 Expedition
Report. Madagascar Environmental Research Group, U.K.
Richard, A.F. and Sussman, R.W. (1975). Future of the Malagasy Lemurs;
Conservation or Extinction? In: Tattersall, I. and Sussman, R.W. (Eds), Lemur
Biology. Plenum Press, New York. Pp. 335-350.
Richard, A.F. and Sussman, R.W. (1987). Framework for Primate Conservation
in Madagascar. In: Marsh, C. W. and Mittermeier, R. (Eds), Primate Conservation in
the Tropical Forest. Alan R. Liss, Inc., New York. Pp. 329-341.
Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New
York.
59
Lemurs of Madagascar and the Comoros
The Hairy-eared Dwarf Lemur, Allocebus trichotis, is possibly the least known of all the
lemur species. It has been found in only one area of north-eastern Madagascar, where it was
rediscovered in 1989.
Photo by Bernhard Meier.
The IUCN Red Data Book
HAIRY-EARED DWARF LEMUR ENDANGERED
Allocebus trichotis (Giinther, 1875)
Order PRIMATES Family CHEIROGALEIDAE
SUMMARY Until 1989 the Hairy-eared Dwarf Lemur was known from only five
museum specimens, all but one of which were collected in the last century. However, it
was rediscovered in 1989 in lowland forest in north-east Madagascar. Little or no
information is available on its distribution or numbers, nor on any aspects of its ecology. It
may be very rare or simply very cryptic, it is nocturnal. It must be threatened, however, by
destruction of its habitat, the eastern rain forests. Three individuals are in captivity in
Madagascar. Listed in Appendix 1 of CITES, Class A of the African Convention and is
protected by Malagasy law.
DISTRIBUTION Until its rediscovery in 1989, the Hairy-eared Dwarf Lemur was
known from only five museum specimens. The holotype was collected by Crossley in
1874, but the information on its label, stating that it was collected in S. Madagascar, differs
from Gunther's (1875) statement that it came from between "Tamantave" (i.e. Toamasina,
on the east coast) and "Murundava" (i.e. Morondava, on the west coast) (Tattersall, 1982).
The provenance of the two collected by Humblot around 1880 is unknown (Tattersall,
1982). A fourth specimen has recently been discovered in the collections of the
Naturhistoriska Rijksmuseet in Stockholm by G. H. Albrecht (pers. comm. to P. Jenkins,
1987). No date is given for this specimen. Its locality is either "Nanaka" or "Namaka"
(Jenkins, 1987). Jenkins suggests that this may be equivalent to Nanakara (24°17'S,
45°53'E), but as this village is not in an area of rain forest, it seems an unlikely site for the
Hairy-eared Dwarf Lemur ever to have been located. Peyrieras, in 1965, captured the fifth
specimen in Andranomahitsy forest, near the village of Ambavala, 16 km from the town of
Mananara, which is on the east coast of Madagascar (Peyrieras pers. comm. to Meier and
Albignac, in press). An expedition to the same forest in 1975 failed to find any Allocebus
trichotis (Petter et al, 1977). However, it was relocated there by Bernhard Meier and
Ronald Albignac early in 1989.
Meier and Albignac (in press) consider that the distribution of this species may be restricted
and patchy. They report seeing a number of Allocebus in the area around Mananara. Meier
saw one individual close to the village of Ambavala (16°12'S, 49°37'E) and one at
16°26'S, 49°38 E, 1.5 km from the Bedinta mountain, which is 34 km from Mananara. In
addition, three individuals were caught near the village of Andranombazaha (16°28'S,
49°38'E).
Tattersall (1982) suggests that the species once occurred quite widely in the eastern humid
forests, but the paucity of either specimens or sightings makes it difficult to confirm this.
POPULATION Numbers are not known, but Tattersall, in 1982, considered this the
rarest of the surviving lemurs and one which probably never existed at high densities. Meier
and Albignac (in press) say that the population density may be very low. Its numbers are
almost certainly declining as the eastern rain forests are reduced in size (Richard and
Sussman, 1975, 1987). Meier and Albignac (in press) also consider that its numbers are
probably declining.
HABITAT AND ECOLOGY Comparatively little is known about this species but it
appears to occur only in lowland rain forest. One of the individuals seen by Meier was in
degraded primary lowland forest, while the other was in virgin primary forest (Meier and
61
Lemurs of Madagascar and the Comoros
P. furcifer
™ A. trichotis
Tropic of Capricorn
Figure 8: Distribution of Phaner and Allocebus. Shaded areas represent approximate limits
of ranges.
62
The IUCN Red Data Book
Albignac, in press). The three captured individuals were all in primary lowland forest
(Meier and Albignac, in press). It is a nocturnal species, becoming active at dusk and
remaining so until the very first light of dawn (Meier and Albignac, in press). It jumps a lot,
in a manner similar to Microcebus rather than Cheirogaleus (Meier and Albignac, in press).
There is no information on the diet of the Hairy-eared Dwarf Lemur in the wild. Meier and
Albignac (in press) suggest that this species may feed on nectar, it has a very long tongue
and, in captivity, eats honey. Caged animals ate locusts, which were jumped on and caught
with both hands; fruit was also eaten (Meier and Albignac, in press). In May, Allocebus
has a considerable fat deposit which is not stored in the tail, as in Cheirogaleus, but is
distributed all over the body (Meier and Albignac, in press). Local people reported that they
did not see active Hairy-eared Dwarf Lemurs between May and September and it appears
that they are in some type of hibernation during that time (Meier and Albignac, in press). In
Captivity, activity is drastically reduced from June to September (Meier and Albignac, in
press). The animals are usually found sleeping in tree holes. One individual was caught in
a hole in a small dead tree that was broken off 4 m above the ground (Meier and Albignac, in
press). This was a juvenile male and there were two other individuals in the same hole.
Local people reported that usually two or three individuals are found in a tree hole but thet
up to six animals could be together (Meier and Albignac, in press). It is possible that infants
are born in January or February as some Malagasy tree cutters saw half grown individuals in
March (Meier and Albignac, in press).
THREATS The main threat to this species must be destruction of the rain forest for
agriculture and fuel and by timber companies. It has recently been estimated that 111,000 ha
of eastern rain forest have been cleared each year between 1950 and 1985, most of this has
been the lowland forest (Green and Sussman, in press). If the cutting continues, forests on
only the steepest slopes will survive the next thirty-five years (Green and Sussman, in
press) and this will probably mean the extinction of Allocebus if it does, indeed, live only in
the lowland rain forest.
CONSERVATION MEASURES There are no measures suggested specifically for
conserving the Hairy-eared Dwarf Lemur and, until more is known about its ecology and
range, none can be made. Its chances of survival will be increased by preservation of the
eastern rain forest. An area around Mananara, the only known location of Allocebus, has
been proposed as a Biosphere Reserve (Nicoll and Langrand, 1989). It is suggested that the
protected area should have the status of a National Park and that a buffer zone be set up
surrounding it (Nicoll and Langrand, 1989). Extensive surveys are needed to try to locate
any remaining populations of the Hairy-eared Dwarf Lemur.
All species of Cheirogaleidae are listed in Appendix 1 of the 1973 Convention on
International Trade in Endangered Species of Wild Fauna and Flora. Trade in them, or their
products, is subject to strict regulation and may not be carried out for primarily commercial
purposes.
All Lemuroidea are listed in Class A of the African Convention, 1969. They may not,
therefore, be hunted, killed, captured or collected without the authorization of the highest
competent authority, and then only if required in the national interest or for scientific
purposes.
Malagasy law protects all lemurs from unauthorised capture and from hunting, but this is
very difficult to enforce.
CAPTIVE BREEDING Until recently, none had been kept in captivity except the one
collected by Peyrieras, which he had for only a few days (Petter et al, 1977). Now there are
three individuals, an adult pair and a juvenile, being held in captivity in Madagascar (Meier
and Albignac, in press).
63
Lemurs of Madagascar and the Comoros
REMARKS The adult female of this species caught by Meier and Albignac (in press)
weighed 80 g, while the adult male weighed 75 g. A juvenile of unknown sex weighed 58g
Body length of the female was 145 mm, tail length was 165 mm; body length of the male
was 125 mm and tail length was 195 mm. Pelage on the dorsal side was rosy brownish-
grey, while the ventral fur was grey. There are narrow dark rings round the eyes. Its ears
are very short and concealed in fur but long wavy hairs form the eartufts from which this
lemur gets its common name. Its nails are kneeled, except on the hallux, but the apex of the
nails are rounded, not pointed (Meier and Albignac, in press). For a more complete
description of Allocebus trichotis, see Giinther (1875), Petter-Rousseaux and Petter (1967),
Tattersall (1982), Jenkins (1987) and Meier and Albignac (in press). The species was
initially classified in the genus Cheirogaleus (Giinther, 1875), but is now considered to be in
its own genus (Petter-Rousseaux and Petter, 1967; Tattersall, 1982; Jenkins, 1987). The
Malagasy name of this species is tsidy ala, meaning mouse lemur of the big forest, as
opposed to tsidy savoka (Microcebus rufus), meaning mouse lemur of the brush (Meier and
Albignac, in press).
REFERENCES
Giinther, A. (1875). Notes on some mammals from Madagascar. Proceedings of the
Zoological Society, London: 78-80.
Jenkins, P. D. (1987). Catalogue of Primates in the British Museum (Natural History)
and elsewhere in the British Isles. Part IV: Suborder Strepsirrhini, including the
Subfossil Madagascan Lemurs and Family Tarsiidae. British Museum (Natural History),
London.
Meier, B. and Albignac, R. (in press). Rediscovery of Allocebus trichotis Giinther
1875 (Primates) in North East Madagascar. Folia Primatologica.
Nicoll, M.E. and Langrand, O. (1989). Revue Générale du Systéme d’Aires
Protégés et de la Conservation d Madgascar. Unpublished report to WWF.
Petter, J-J., Albignac, R. and Rumpler, Y. (1977). Mammiféres lémuriens
(Primates Prosimiens) Faune de Madagascar No. 44. ORSTOM-CNRS, Paris.
Petter, J.-J. and Petter-Rousseaux, A. (1979). Classification of the Prosimians.
In: Doyle, G.A. and Martin, R.D. (Eds), The Study of Prosimian Behavior. Academic
Press, London. Pp. 1-44.
Petter-Rousseaux, A. and Petter, J.-J. (1967). Contribution 4 la systématique des
Cheirogaleinae (lémuriens malagaches). Allocebus, gen. nov. pour Cheirogaleus
trichotis Giinther 1875. Mammalia 31: 574-582.
Richard, A.F. and Sussman, R.W. (1975). Future of the Malagasy Lemurs;
Conservation or Extinction? In: Tattersall, I. and Sussman, R.W. (Eds), Lemur
Biology. Plenum Press, New York. Pp. 335-350.
Richard, A.F. and Sussman, R.W. (1987). Framework for Primate Conservation
in Madagascar. In: Marsh, C.W. and Mittermeier, R. (Eds), Primate Conservation in the
Tropical Rain Forest. Alan R. Liss, Inc., New York. Pp. 329-341.
Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New
York.
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Lemurs of Madagascar and the Comoros
The nocturnal Fork-marked Lemur, Phaner furcifer, is found widely but patchily distributed
in Madagascar. There are probably several subspecies.
Photo by Russell Mittermeier.
The IUCN Red Data Book
FORK-MARKED LEMUR RARE
Phaner furcifer (Blainville, 1839)
Order PRIMATES Family CHEIROGALEIDAE
SUMMARY The Fork-marked Lemur is a small nocturnal species with a wide, but
discontinuous, distribution in Madagascar. It is found in both humid and dry deciduous
forests and is common in secondary forest. Several subspecies have recently been
described. Population numbers are unknown but they are considered to be declining.
Figures for densities vary from 850 individuals per sq. km to 50 per sq. km. Phaner
furcifer is a territorial species and is frequently seen in pairs during the night. The main
component of its diet is gum though some insects, fruit and nectar are also taken. It is found
in several reserves, but is threatened by habitat destruction in most areas. Only two are
recorded in captivity. Listed in Appendix 1 of CITES, in Class A of the African
Convention and is protected under Malagasy law
DISTRIBUTION Phaner has a wide but discontinuous distribution in Madagascar. It is
found mainly in the dry deciduous forest in the west of the country, where it extends from
about the latitude of Toliara (Tulear) northwards to near Antsalova. Petter et al (1977) and
Petter and Petter-Rousseaux (1979) show another portion of Phaner's range from Cap St
André southwards along the coast to north of the Bay of Bombetoka, whereas Tattersall
(1982) shows two smaller, disjunct populations, one south of Soalala and the other around
the Bay of Bombetoka. Phaner also occurs in the Sambirano region (on the Ampasindava
Peninsula), in the far north around Montagne d'Ambre and in the eastern rain forest on the
Masoala Peninsula (Tattersall, 1982, Petter et al, 1977; Petter and Petter-Rousseaux, 1979).
Again, the range shown by Tattersall (1982) in the east is much smaller than that shown by
the other authors. Petter ef a/ (1977) and Petter and Petter-Rousseaux (1979) show Phaner
extending north from the Masoala Peninsula to beyond Sambava, while Tattersall (1982)
considers that it extends only as far as Antalaha. Two other populations have also been
reported, one on the Tsaratanana Massif (Andriamampianina and Peyrieras, 1972) and the
other in the arid Didierea bush in the south in Andohahela reserve (Russell and McGeorge,
1977). The presence of Phaner in parcel 2 of Andohahela Reserve is confirmed by M.
Pidgeon (in litt.) but only in the gallery forest. The presence of the species in Bemaraha
Nature Reserve has been reported (Petter and Andriatsarafara, 1987; Nicoll and Langrand,
1989).
Groves and Tattersall (in press) consider that there are the five principal populations of
Phaner existing today and that there are four different subspecies of the Fork-marked
Lemur. These are mentioned in "Remarks". They consider that reports of the presence of
Phaner in Andohahela, Tsaratanana and around the Bay of Bombetoka need further
confirmation, as does its existence between Tsiribihina River and Namoroka.
POPULATION Total numbers are unknown but they are considered to be probably
declining (Richard and Sussman, 1975, 1987). Petter et al (1971) counted 17 Phaner in 2
ha of Marosalaza Forest (in the west, 50 kms north of Morondava), thereby estimating a
density of 850 individuals per sq. km. In another area, near Mangoky 200 km south of
Marosalaza, they estimated population densities of at least 550 individuals per sq. km.
Charles-Dominique and Petter (1980) found 14 individuals in their 25 ha study area in
Marosalaza Forest and, from this, estimate densities of 50-60 individuals per sq. km. They
consider that the densities estimated by Petter et al (1971) reflect observations in small, gum
rich areas and that these high numbers cannot be extrapolated to mean population densities.
67
Lemurs of Madagascar and the Comoros
HABITAT AND ECOLOGY This species is found in the humid forests of the east and
the Sambirano region as well as in the dry deciduous forests of the west (Petter et al, 1971).
It has also been reported in the Didiereaceae bush in the south (Russell and McGeorge,
1977) though more recent observations suggest that it is confined to the gallery forest there
(M. Pidgeon, in litt.). Petter et al (1975), working in Marosalaza Forests, found that P.
furcifer was most often seen in areas of secondary forest, although it appeared that this
species did not occupy zones in which a continuous canopy was missing (Petter et al,
1975). Between March and May, gum, particularly from Terminalia trees, was the principal
food of Phaner in Marosalaza Forest, but insects, sap and bud exudate were also taken
(Charles-Dominique and Petter, 1980). In November, Terminalia gum was still providing
the bulk of the Fork-marked Lemur's diet, but flowers were licked and the "syrup" from
insect larvae of the family Machaerotidae was also eaten (Charles-Dominique and Petter,
1980). Phaner spends the day in holes, usually in large trees such as Baobabs (Adansonia
Spp), or sometimes in the abandoned nests of Mirza coquereli (Petter et al, 1971, 1975); it
leaves these at nightfall and feeds most actively for the first hour of the night (Petter et al,
1975). Locomotion consists of rapid quadrupedal running, climbing and leaping, generally
at 3-4m (where most horizontal branches occur in Marosalaza) though they were also seen
on the ground and at above 10m (Petter et al, 1975).
Charles-Dominique and Petter (1980) found three territories, each containing one adult male
and one adult female Phaner, though they also found a solitary male in a territory and one
male with a range overlapping those of two females. Four of these territories also contained
juvenile individuals. The pairs were in close proximity for about half of the night and were
in almost continuous vocal contact throughout the night; in addition, they slept together
during the day (Charles-Dominique and Petter, 1980). The male whose range overlapped
those of two females divided his active and rest time between the two of them (Charles-
Dominique and Petter, 1980). Mean size of females’ territories was about 4 ha, while that
of the males was 3.8 ha.; there was little overlap between the territories of same sex
individuals (Charles-Dominique and Petter, 1980). The overlap zones appeared to be
"meeting areas", between three and nine neighbours coming together and emitting
simultaneous calls for 10-20 minutes; no aggression was seen on these occasions (Charles-
Dominique and Petter, 1980). Allogrooming was frequent between males, females and
juveniles (Charles-Dominique and Petter, 1980). Charles-Dominique (1978) described this
social stystem as "pre-gregarious".
The Fork-marked Lemur is very vocal, a mean of 30 loud calls an hour (emitted only by the
males) have been counted in a radius of about 200m in Marosalaza forest; olfactory signals
appear to be much less important in this species than in other nocturnal prosimians (Charles-
Dominique and Petter, 1980).
Mating occurs in June (Charles-Dominique and Petter, 1980). A single infant is born in
November or December. It is initially left in a tree hole and then carried on the front of its
mother, later moving to her back (Petter et al, 1971, 1975; Charles-Dominique and Petter,
1980).
THREATS Nothing specific recorded for this species, but its habitat is being destroyed
by fires, clearing for pasture land and for crops.
CONSERVATION MEASURES Found in Ankarana, Manongarivo and
Andranomena Special Reserves, the Nature Reserves of Bemaraha, Tsaratanana and
Andohahela, Mt d'Ambre National Park and Analabe Private Reserve (Raxworthy and
Rakotondraparany, 1988; Hawkins et al, in press; Nicoll and Langrand, 1989). In general,
all these areas would benefit from better protection and conservation awareness programmes
would help the local people understand the importance of the reserves and their wildlife
(Nicoll and Langrand, 1989).
68
The IUCN Red Data Book
It has been proposed that a National Park is established on the Masoala Peninsula and this
would protect what is probably a distinct subspecies of Phaner.
Surveys are needed to find out the range and numbers of this species. Without this
information, the conservation status of Phaner cannot be assessed, nor can the populations
requiring protection be identified.
All species of Cheirogaleidae are listed in Appendix 1 of the 1973 Convention on
International Trade in Endangered Species of Wild Fauna and Flora. Trade in them, or their
products, is subject to strict regulation and may not be carried out for primarily commercial
purposes.
All Lemuroidea are listed in Class A of the African Convention, 1969. They may not,
therefore, be hunted, killed, captured or collected without the authorization of the highest
competent authority, and then only if required in the national interest or for scientific
purposes.
Malagasy law protects all lemurs from unauthorised capture and from killing. This is,
however, very difficult to enforce.
CAPTIVE BREEDING ISIS (June, 1989) does not record any Phaner in captivity. At
least two females and one male have been kept in Brunoy, France for “several years"
(Petter-Rousseaux, 1980; Cooper, 1980), but it is not clear if they ever bred there. Two
individuals are now in Paris Zoo (J.-J. Petter, in litt.).
REMARKS A small lemur of 360-500g (Petter et al, 1977), characterised by a broad
dorsal stripe which bifurcates on the crown, the two stripes continuing to the eyes. Fur on
the back is grey-brown, underparts are white to pale brown (Tattersall, 1982). Though
traditionally viewed as monotypic it was suggested in 1975 that the populations in the north
and east should be regarded as distinct subspecies (Petter et al, 1975) and four subspecies
have now been described by Groves and Tattersall (in press). These are: P. f. furcifer,
found on the Masoala Peninsula; P. f. pallescens found in western Madagascar from just
south of Fiherenana River to the region of Soalala (though the authors consider it absent
between the Tsiribihina River and Namoroka); P. f. parienti in the Sambirano region; and
P. f. electromontis found in the area of Mt d'Ambre. Malagasy names for P. furcifer are
tanta, tantaraolana, vakiandrina and vakivoho (Petter et al, 1977; Tattersall, 1982; Paulian,
1981).
REFERENCES
Charles-Dominique, P. (1978). Solitary and gregarious prosimians: evolution of
social structure in primates. In: Chivers, D.J. and Joysey, K.A. (Eds), Recent Advances
in Primatology, Volume 3: Evolution. Academic Press, London. Pp. 139-149.
Charles-Dominique, P. and Petter, J.-J. (1980). Ecology and Social life of
Phaner furcifer. In: Charles-Dominique, P., Cooper, H.M., Hladik, A., Hladik, C.M.,
Pages, E., Pariente, G.F., Petter-Rousseaux, A., Petter, J.-J. and Schilling, A. (Eds),
Nocturnal Malagasy Primates: Ecology, Physiology and Behavior. Academic Press,
New York. Pp. 75-95.
Cooper, H.M. (1980). Ecological correlates of visual learning in nocturnal prosimians.
In: Charles-Dominique, P., Cooper, H.M., Hladik, A., Hladik, C.M., Pages, E.,
Pariente, G.F., Petter-Rousseaux, A., Petter, J.-J. and Schilling, A. (Eds), Nocturnal
Malagasy Primates: Ecology, Physiology and Behavior. Academic Press, New York.
Pp. 191-203.
Grores, C.P. and Tattersall, I. (in press). Geographical variation in the Fork-
marked lemur Phaner furcifer (Mammalia, Primates). Folia primatologica.
69
Lemurs of Madagascar and the Comoros
ISIS (1989). SIS Species Distribution Report Abstract for Mammals, 30 June 1989.
International Species Information System, 12101 Johnny Cake Ridge Road, Apple
Valley, MN, U.S.A. Pp. 17-22.
Nicoll, M.E. and Langrand, O. (1989). Revue Générale du Systéme d’Aires
Protégées et de la Conservation a Madagascar. Unpublished report to WWF.
Paulian, R.R. (1981). Les mammiféres: Vestiges d'un monde disparu. In: Oberlé, P.
(Ed.), Madagascar un Sanctuaire de la Nature. La Societe Malagache d'Edition,
Antananarivo. Pp. 75-94.
Petter, J-J., Albignac, R. and Rumpler, Y. (1977). Mammiféres lémuriens
(Primates prosimiens). Faune de Madagascar No. 44. ORSTOM-CNRS, Paris.
Petter, J.-J. and Andriatsarafara, F. (1987). Conservation status and distribution
of lemurs in the west and northwest of Madagascar. Primate Conservation 8: 169-171.
Petter, J.-J. and Petter-Rousseaux, A. (1979). Classification of the Prosimians
In: Doyle, G.A. and Martin, R.D. (Eds), The Study of Prosimian Behavior. Academic
Press, London. Pp. 1-44.
Petter, J.-J., Schilling, A. and Pariente, G. (1975). Observations on behavior
and ecology of Phaner furcifer. In: Tattersall, I. and Sussman, R.W. (Eds), Lemur
Biology. Plenum Press, New York. Pp. 209-218.
Petter, J.-J., Schilling, A. and Pariente, G. (1971). Observations éco-
éthologiques sur deux lémuriens malagaches nocturnes: Phaner furcifer et Microcebus
coquereli. La Terre et la Vie 3: 287-327.
Petter-Rousseaux, A. (1980). Seasonal activity rhythms, reproduction, and body
weight variations in five sympatric nocturnal prosimians, in simulated light and climatic
conditions. In: Charles-Dominique, P., Cooper, H.M., Hladik, A., Hladik, C.M.,
Pages, E., Pariente, G.F., Petter-Rousseaux, A., Petter, J.-J. and Schilling, A. (Eds),
Nocturnal Malagasy Primates: Ecology, Physiology and Behavior. Academic Press,
New York. Pp. 137-152.
Richard, A.F. and Sussman, R.W. (1975). Future of the Malagasy lemurs;
conservation or extinction? In: Tattersall, I. and Sussman, R.W. (Eds), Lemur Biology,
Plenum Press, New York. Pp. 335-350.
Richard, A.F. and Sussman, R.W. (1987). Framework for Primate Conservation
in Madagascar. In: Marsh, C.W. and Mittermeier, R. (Eds), Primate Conservation in the
Tropical Rain Forest, Alan R. Liss Inc., New York. Pp. 329-341.
Russell, R.J. and McGeorge, L.W. (1977). Distribution of Phaner (Primates,
Lemuriformes, Cheirogaleidae, Phanerinae) in southeast Madagascar. Journal of
Biogeography 4: 169-170.
Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New
York.
70
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Lemurs of Madagascar and the Comoros
The Grey-backed Sportive Lemur, Lepilemur dorsalis, has a very limited distribution in
north-western Madagascar. It is threatened by destruction of its habitat.
Photo by Chris Raxworthy.
12
The IUCN Red Data Book
GREY-BACKED or NOSY BE SPORTIVE LEMUR VULNERABLE
Lepilemur dorsalis Gray, 1871
Order PRIMATES Family MEGALADAPIDAE
SUMMARY The Grey-backed Sportive Lemur is one of the least widely distributed of the
Sportive Lemurs. It is found only in the Sambirano Region of north-west Madagascar and
on the island of Nosy-Bé. Population numbers are unknown. Only very brief studies have
been made of this species and little is known of its ecology and social organisation. It is a
solitary, nocturnal species which feeds principally on leaves, though fruit and bark are also
taken. Lepilemur dorsalis is certainly threatened by habitat destruction. It is found in two
Reserves, Manongarivo on the mainland and Lokobe on Nosy Bé, neither of which is safe
from encroachment. None is in captivity. Listed in Appendix 1 of CITES, Class A of the
African Convention and is protected under Malagasy law.
DISTRIBUTION Inhabits the Sambirano Region on the north-west coast of Madagascar
and the island of Nosy Bé (Tattersall, 1982; Petter et al, 1977; Petter and Petter-Rousseaux,
1979). The latter two authors show a more northerly extension to the range of the Grey-
backed Sportive Lemur than does Tattersall.
POPULATION Population numbers are unknown but the limited distribution of this
species suggest that it must be one of the least common of the Sportive Lemurs. Numbers
are declining as the forest is being cleared throughout its range.
HABITAT AND ECOLOGY The forests on Nosy Bé and in the Sambirano Region
where this species occurs are humid forests quite similar in structure, though not in floristic
composition, to those in the east. On Nosy Bé, the nocturnal L. dorsalis apparently does
not sleep in tree holes instead it spends the day curled up in foliage (Petter and Petter,
1971). However, it has been observed in Manongarivo Reserve asleep in tree holes
(Raxworthy and Rakotondraparany, 1988). It feeds on leaves, fruit and bark (Petter and
Petter, 1971). One infant is born between September and November (Petter and Petter,
1971).
THREATS The main threat to this species is habitat destruction due to agricultural
encroachment and clearance for settlement. Most of the forests on the coast have been
destroyed and the remainder are patchy in distribution. Even the protected areas are not safe
from this encroachment. An expedition to Manongarivo Special Reserve in 1988 found that
a large area of the reserve had already been cleared and that this was continuing. The
disturbance extended to about 6 km into the reserve (Quansah, 1988). In the Nature
Reserve on Nosy Bé where L.dorsalis is also found, the main threat is illegal exploitation of
the forest, trees are cut for making into canoes and for building material and the land is
cleared for coffee and rice plantations (Nicoll and Langrand, 1989).
CONSERVATION MEASURES Found in Manongarivo Special Reserve (Raxworthy
and Rakotondraparany, 1988) and in Lokobe Nature Reserve on Nosy Bé (Petter et al,
1977).
Neither of the reserves in which this species is found is adequetely protected from
encroachment. Conservation awareness programmes would be helpful so that the local
people could learn about the existence and the importance of the reserves. Around
Manongarivo, slash and burn clearing of the forest will cease only if alternative agricultural
practices are developed so that the land already cleared can produce enough food on a
sustainable basis to support the villagers (Quansah, 1988). The reserve on Nosy Bé could
73
Lemurs of Madagascar and the Comoros
20°S
septentrionalis
. ruficaudatus
. Mustelinus
microdon
. edwardsi
. leucopus
. dorsalis
Figure 9: Distribution of all species of Lepilemur. Shaded areas represent approximate
limits of ranges.
74
The IUCN Red Data Book
be developed as a tourist attraction, this would provide some employment and income to the
islanders (Nicoll and Langrand, 1989).
All species of Lemuridae (which is taken to include the genus Lepilemur) are listed in
Appendix 1 of the 1973 Convention on International Trade in Endangered Species of Wild
Fauna and Flora.
Trade in them, or their products, is subject to strict regulation and may not be carried out for
primarily commercial purposes.
All Lemuroidea are listed in Class A of the African Convention, 1969. They may not,
therefore, be hunted, killed, captured or collected without the authorization of the highest
competent authority, and then only if required in the national interest or for scientific
purposes.
Malagasy law protects all lemurs from unauthorised capture and from hunting. It is,
however, very difficult to enforce this.
CAPTIVE BREEDING None is known to occur in captivity. Breeding of all the
Sportive Lemurs appears to be difficult (Rumpler, 1975, Petter et al, 1977).
REMARKS The Grey-backed Sportive Lemur is one of the smaller species within the
genus. Its head and body measure 250-260 mm (Tattersall, 1982) and it probably weighs
around 500 g. Upperparts are medium to dark brown and underparts are also brown, only a
little paler than the dorsum (Tattersall, 1982). See Jenkins (1987) and Tattersall (1982) for
more measurements of this species.
The taxonomy of this group is very confused. It is common for all the types to be
considered as subspecies of L. mustelinus (e.g. Tattersall, 1982; Richard, 1984), while the
names and numbers of the forms vary with the author. Tattersall (in /itt.) now considers that
the genus should contain several species. The Grey-backed Sportive Lemur was described
as a distinct species by Rumpler (1975) on the basis of karyotype studies. There is also
considerable disagreement over which family the genus should be in. It is traditionally put
in Lemuridae, now it is commonly placed in Lepilemuridae but Schwartz and Tattersall
(1985) have placed it in Megaladapidae, along with some of the extinct lemurs. More
details of the taxonomy can be found in Tattersall (1982) and Jenkins (1987). The Malagasy
name for this species is apongy (Tattersall, 1982).
REFERENCES
Jenkins, P. D. (1987). Catalogue of Primates in the British Museum (Natural History)
and elsewhere in the British Isles. Part IV: Suborder Strepsirrhini, including the
Subfossil Madagascan Lemurs and the Family Tarsiidae. British Museum (Natural
History), London.
Nicoll, M.E. and Langrand, O. (1989). Revue Générale du Systéme d’Aires
Protégées de la Conservation a Madagascar. Unpublished report to WWF.
Petter, A. and Petter, J.J. (1971). Part 3.1 Infraorder Lemuriformes. In: Meester,
J. and Setzer, H.W. (Eds), The Mammals of Africa: An Identification Manual.
Smithsonian Institution Press, City of Washington.
Petter, J-J., Albignac, R. and Rumpler, Y. (1977). Mammiféres lémuriens
(Primates prosimiens). Faune de Madagascar No. 44. ORSTOM-CNRS, Paris.
Petter, J.-J. and Petter-Rousseaux, A. (1979). Classification of the Prosimians.
In: Doyle, G.A. and Martin, R.D. (Eds), The Study of Prosimian Behavior. Academic
Press, London. Pp. 1-44.
a
Lemurs of Madagascar and the Comoros
Quansah, N. (Ed.), (1988). Conclusions and Recommendations. Manongarivo
Special Reserve (Madagascar): 1987/88 Expedition Report. Madagascar Environmental
Research Group, U.K.
Raxworthy, C.J. and Rakotondraparany, F. (1988). Mammals report. In:
Quansah, N. (Ed.), Manongarivo Special Reserve (Madagascar): 1987/88 Expedition
Report. Madagascar Environmental Research Group, U.K.
Richard, A.F. (1984). Lemurs. In: Macdonald, D. (Ed.), The Encyclopaedia of
Mammals:1. George Allen and Unwin, London. Pp. 330-331.
Rumpler, Y. (1975). The significance of chromosomal studies in the systematics of the
Malagasy lemurs. In: Tattersall, I. and Sussman, R.W. (Eds), Lemur Biology. Plenum
Press, New York. Pp. 25-40.
Schwartz, J.H. and Tattersall, I. (1985). Evolutionary relationships of the living
lemurs and lorises (Mammalia, Primates) and their potential affinities with European
Eocene Adapidae. Anthropological Papers of the American Museum of Natural History
60 (1): 1-100.
Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New
York.
76
wh ky AS) Vn: Lahosh ®» z ae +)
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Lemurs of Madagascar and the Comoros
Milne-Edwards' Sportive Lemur, Lepilemur edwardsi, is found in the dry, deciduous
forests of western Madagascar. Its main diet is leaves.
Photo by Russell Mittermeier.
78
The IUCN Red Data Book
MILNE-EDWARDS' SPORTIVE LEMUR RARE
Lepilemur edwardsi Major, 1894
Order PRIMATES Family MEGALADAPIDAE
SUMMARY Milne-Edward's Sportive Lemur is found in the dry deciduous forests of
western Madagascar but its precise distibution is not known. There are no estimates of
population number but the species may be found at high densities in some areas. Lepilemur
edwardsi has been studied only briefly and there is little information available on its ecology
and social organisation. It is a solitary, nocturnal species and its main diet is leaves.
Probably threatened by habitat destruction. The species is found in at least three protected
areas. None occurs in captivity. Listed in Appendix 1 of CITES, Class A of the African
Convention and is protected by Malagasy law.
DISTRIBUTION This species is found in the dry deciduous forests of western
Madagascar. Petter and Petter-Rousseaux (1979) show a very discontinuous distribution
from approximately 14°40'S to just north of Manambolo River (this discontinuity appears
to be largely a function of mapping its occurrence in the remaining forests in the west,
whereas it is usual to show more approximate ranges, including whole areas rather than
particular habitats). Petter et al (1977) show two large populations, a southern one between
Manambolo and Manambao Rivers, and a more northerly one from approximately 17°S to
15°S, with a small disjunct population between them. Tattersall (1982) gives the
distribution from the Bay of Mahajamba south to at least Antsalova, and possibly as far as
Tsiribihina River.
POPULATION No estimations of population numbers have been made, though Albignac
(1981) suggests that they can exist at quite high densities; he caught 25 individuals in a 16
ha study site at Ampijoroa in Ankarafantsika Forest. However, Ganzhorn (1988) reports
densities of only 57+ 22 individuals per sq. km (mean and 95% confidence limits, based on
10 census walks along 1.7 km of trail) at Ampijoroa.
HABITAT AND ECOLOGY Two or three individuals (males and females) sleep
together in tree holes but they generally move separately at night (Albignac, 1981; Petter et
al, 1977). The home range of this species is reported to be about 1 ha, with the ranges of
"certain males” being larger (Albignac, 1981). Overlap of the ranges can be quite extensive
(Albignac, 1981). L.edwardsi has aggresive auditory territorial behaviours involving
howling and shaking of branches (Albignac, 1981). The main diet of this sportive lemur is
leaves, including mature ones, but it also takes fruit, flowers and fleshy seeds
(Razanahoera-Rakotomalala, 1981; Albignac, 1981; Ganzhorn, 1988).
THREATS No specific information, but habitat loss must be a threat. Fires, set each year
to encourage new grass growth for domestic stock, are the main cause of the destruction of
the dry deciduous forests. For instance, 500 ha of forest in Namoroka Nature Reserve was
burnt in 1984 (Nicoll and Langrand, 1989). In 1981, FAO/UNEP estimated net
degradation of of the western forests to have been perhaps 200,000 ha since 1955. The
map in Petter and Petter-Rousseaux (1979) suggests that the populations are becoming
increasingly small and isolated as the forest disappears.
CONSERVATION MEASURES Milne-Edwards' Sportive Lemur is found in the
Nature Reserves of Ankarafantsika, Namoroka and Bemaraha and it may also be in some of
the Special Reserves that are in its range.
More guards with better equipment wouid be an asset in all the reserves in the west as it is
essential that the fires are controlled (Nicoll and Langrand, 1989). Education projects that
79
Lemurs of Madagascar and the Comoros
demonstrate the danger of the fires and the benefits of the reserves to the local people would
be helpful for the conservation of the protected areas. In addition, notices on the boundaries
of the reserves could be used to inform people that hunting and tree cutting is illegal (Nicoll
and Langrand, 1989). The Department of Water and Forests, with the World Bank, is
developing a management programme for Ankarafantsika and this will benefit by the
inclusion of a reafforestation plan to provide wood for fuel and building material (Nicoll and
Langrand, 1989).
All species of Lemuridae (which is taken to include the genus Lepilemur) are listed in
Appendix 1 of the 1973 Convention on International Trade in Endangered Species of Wild
Fauna and Flora. Trade in them, or their products, is subject to strict regulation and may not
be carried out for primarily commercial purposes.
All Lemuroidea are listed in Class A of the African Convention, 1969. They may not,
therefore, be hunted, killed, captured or collected without the authorization of the highest
competent authority, and then only if required in the national interest or for scientific
purposes.
Malagasy law protects all lemurs from hunting or unauthorised capture, but it is difficult to
enforce this.
CAPTIVE BREEDING None is known to be held in captivity. Members of the genus
Lepilemur do not survive well in captivity (Petter et al, 1977).
REMARKS The dimensions of this species are quite similar to those of L. ruficaudatus
(Tattersall, 1982; Jenkins, 1987), it weighs between 800 and 1000 g (Razanahoera, 1988).
Its dorsal fur is grey-brown with a red-brown wash, while its underparts are grey, flecked
with cream (Tattersall, 1982; Jenkins, 1987).
The taxonomy of Lepilemur is very confused. It is common for all the forms within the
genus to be considered as subspecies of L. mustelinus (e.g. Tattersall, 1982; Richard,
1984) and the number and names of the types vary with the author. Though, Tattersall (in
litt.) now considers that the genus does contain several species, he remains unconvinced
that edwardsi warrants separation from ruficaudatus, the species found in the dry forests to
the south of the range of edwardsi. There is also considerable disagreement over which
family the genus should be in. Traditionally it is put in Lemuridae, now it is commonly
placed in Lepilemuridae but Schwartz and Tattersall (1985) have placed it in Megaladapidae,
along with some of the extinct lemurs. Further details of the taxonomy of this group can be
found in Tattersall (1982) and Jenkins (1987). In Ankarafantsika and north of there, the
Malagasy name for this species is repahaka; south of Ankarafantsika, it is called boenga
(Tattersall, 1982).
REFERENCES
Albignac, R. (1981). Lemurine social and territorial organisation in a north-western
Malagasy Forest (restricted area of Ampijoroa). In: Chiarelli, A.B. and Corruccini, R.S.
(Eds), Primate Behavior and Sociobiology. Springer Verlag, Berlin. Pp. 25-29.
FAO/UNEP (1981). Tropical Forest Resources Assessment Project. Forest Resources
of Tropical Africa. Part II Country Briefs. FAO, Rome.
Ganzhorn, J. (1988). Food partitioning among Malagasy primates. Oecologia (Berlin)
75: 436-450.
Jenkins, P.D. (1987). Catalogue of Primates in the British Museum (Natural History)
and elsewhere in the British Isles. Part IV: Suborder Strepsirrhini, including the
Subfossil Madagascan Lemurs and the Family Tarsiidae. British Museum (Natural
History), London.
80
The IUCN Red Data Book
Nicoll, M.E. and Langrand, O. (1989). Revue Générale du Systéme d’Aires
Protégées de la Conservation a Madagascar. Unpublished report to WWF.
Petter, J-J., Albignac, R. and Rumpler, Y. (1977). Mammiféres lémuriens
(Primates prosimiens). Faune de Madagascar No. 44. ORSTOM-CNRS, Paris.
Petter, J.-J. and Petter-Rousseaux, A. (1979). Classification of the Prosimians.
In: Doyle, G.A. and Martin, R.D. (Eds), The Study of Prosimian Behavior. Academic
Press, London. Pp. 1-44.
Razanahoera-Rakotomalala, M. (1981). Les adaptations alimentaires comparés de
deux lémuriens folivores sympatriques: Avahi Jourdan, 1934 - Lepilemur I. Geoffroy,
1851. Unpublished PhD thesis, University of Madagascar, Antananarivo.
Razanahoera, M.R. (1988). Comportement alimentaire de deux espéces sympatriques
dans la forét d'Ankarafantsika (nord-ouest de Madagascar): Lepilemur edwardsi et Avahi
laniger (Lémuriens Nocturnes). In: Rakotovao, L., Barre, V. and Sayer, J. (Eds),
L’Equilibre des Ecosystémes Forestiers @ Madagascar Actes d'un séminaire
international. IUCN, Gland, Switzerland and Cambridge, U.K. Pp. 96-99.
Schwartz, J.H. and Tattersall, I. (1985). Evolutionary relationships of the living
lemurs and lorises (Mammalia, Primates) and their potential affinities with European
Eocene Adapidae. Anthropological Papers of the American Museum of Natural History
60 (1): 1-100.
Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New
York.
81
Lemurs of Madagascar and the Comoros
The White-footed Sportive Lemur, Lepilemur leucopus, is found in southern Madagascar.
Its numbers are probably declining as the forests there are being reduced in area.
Photo by Chris Raxworthy.
82
The IUCN Red Data Book
WHITE-FOOTED SPORTIVE LEMUR RARE
Lepilemur leucopus Major, 1894
Order PRIMATES Family MEGALADAPIDAE
SUMMARY The White-footed Sportive Lemur is found only in the primary Didiereaceae
forest and the remaining gallery forests of southern Madagascar. Its precise distribution is
not known. Overall population figures are unknown, but it is reported to occur at very high
densities in some habitats, as many as 810 individuals per sq. km. It is a nocturnal, mainly
solitary, folivore. There have been at least two brief studies of Lepilemur leucopus but their
findings are contradictory. Its numbers are probably declining as the dry forests are being
reduced in area. It is present in four reserves. None is in captivity. Listed in Appendix 1 of
CITES, Class A of the African Convention and protected by Malagasy law.
DISTRIBUTION Found in the primary Didiereaceae and gallery forests of southern
Madagascar (Sussman and Richard,1986) from Taolanaro (Fort-Dauphin) westwards to
Onilahy River (Petter and Petter-Rousseaux, 1979; Petter et al, 1977). Tattersall (1982)
considers its westwards range to be at least as far as Ejeda, and only possibly to Onilahy
River.
POPULATION Total population figures are not known, but numbers as high as 810
individuals per sq. km have been reported in dense areas of gallery forest bordering the river
in Berenty Private Reserve (Charles-Dominique and Hladik, 1971). Density throughout the
gallery forest at Berenty was calculated to be 450 individuals per sq. km (Charles-
Dominique and Hladik, 1971; Hladik and Charles Dominique, 1974). The densities were
estimated to be somewhat lower in the Didiereaceae bush in the same area, here there were
200-350 individuals per sq. km (Charles-Dominique and Hladik, 1971; Hladik and Charles-
Dominique, 1974).
HABITAT AND ECOLOGY This medium sized nocturnal lemur has been studied
briefly by Charles-Dominique and Hladik (1971, 1974) in Didierea bush near Berenty. In
the dry months of September and October, the diet of the White-footed Sportive Lemur was
primarily the tough foliage of Alluaudia procera and A. ascendens (Charles-Dominique and
Hladik, 1971; Hladik and Charles-Dominique, 1974). At the driest time of the year, when
no leaves were available, the White-footed Sportive Lemur survived on a diet of Alluaudia
flowers (Charles-Dominique and Hladik, 1971; Hladik and Charles-Dominique, 1974).
These authors report that the Sportive Lemur injests its faeces (caecotrophy), thereby
gaining extra nutrition from its poor quality, folivorous diet.
White-footed Sportive Lemurs live in small territories. Mean range size of adult females
was 0.18 ha (between 0.15 and 0.32 ha), adult males had bigger ranges, of 0.3 ha
(between 0.20 and 0.46 ha), and juvenile females ranged over 0.19 ha (0.18-0.20 ha)
(Hladik and Charles-Dominque, 1974). There was little range sharing within the sexes, but
males’ territories overlapped those of females (Hladik and Charles-Dominique, 1974). The
territory of one male (the largest) overlapped the ranges of five females, while those of the
three other males overlapped two, one and one females (Hladik and Charles-Dominique,
1974). Females shared their ranges with their immature offspring and it was possible that
adult daughters remained in their natal range (Charles-Dominique and Hladik, 1971).
Territories were defended by mutual surveillance, especially between males, by
vocalisations and by displays rather than by olfactory cues (Hladik and Charles-Dominique,
1974).
Males and females slept separately, either in a tree hole or on a bundle of lianes or other
vegetation (Hladik and Charles-Dominique, 1974). They emerged, frequently after a bout
83
Lemurs of Madagascar and the Comoros
of calling (pers. obs.), when the sun set, but their activity throughout the night was
minimal, one male Lepilemur travelled only 270m during a full night's follow (Charles-
Dominique and Hladik, 1971). Mode of locomotion is principally vertical clinging and
leaping.
A later study by Russell (1980) at the same site produced results contradictory to those of
Hladik and Charles-Dominique. He suggests that Lepilemur leucopus is as active as other
lemurs, spending only 13% of the night at rest, and travelling an average of 400m a night.
He also found that male/female or female/female pairs often shared most or all of a range and
these were rarely defended by vocalisations or surveillance behaviour (Russell, 1977 in
Tattersall, 1982). Russell (1980) found no evidence of caecotrophy.
Mating occurs between May and July and after a gestation period of about four and a half
months, females give birth to a single infant in September to November (Petter et al, 1977).
Sexual maturity is attained at 18 months (Richard, 1984) and females can probably have one
offspring each year thereafter (Petter et al, 1977).
THREATS The dry forest in the south is being destroyed by fire, over grazing and poor
land use and this is probably a threat to L. leucopus. However, no specific information is
available. Though the two small reserves in which it is found, Beza-Mahafaly Special
Reserve and Berenty Private Reserve, are well demarcated, fenced and guarded, this is not
so for the Natural Reserves of Andohahela and Tsimanampetsotsa (Sussman et al, 1987).
The principal threat to the forests in the south is probably the collection of wood for
conversion into charcoal (M. Pidgeon, in litt.).
CONSERVATION MEASURES The White-footed Sportive Lemur is found in two
Nature Reserves, Andohahela (O'Connor et al, 1986,1987) and Tsimanampetsotsa
(Andriamampianina and Peyrieras, 1972) and in the Special Reserve of Beza-Mahafaly as
well as in the Private Reserve of Berenty (Sussman and Richard, 1986; Sussman et al,
1987).
The Department of Water and Forests, the University of Madagascar, WWF and USAID
have jointly proposed and financed a management plan for Andohahela Nature Reseve
(Nicoll and Langrand, 1989). Under this plan it is suggested that more guards with better
equipment are employed to protect the area from illegal tree felling, agricultural
encroachment, overgrazing by domestic stock and hunting. In addition, it would be useful
to clearly mark the boundaries of the Reserve and to patrol them. The needs of the local
people for food, fuel and building material have to be satisfied without encroaching on the
protected area. Education programmes could be started so that the villagers understood the
importance of the reserve and what they could do to protect it. Similar measures are
required to protect Tsimanampetsotsa Nature Reserve.
Surveys of the remaining vegetation and the condition of all the forests within the range of
this species need to be carried out. A census of the distribution and numbers of the White-
footed Sportive Lemur is needed so that its conservation status can be assessed so that any
necessary protective measures can be taken.
All species of Lemuridae (which is taken to include the genus Lepilemur) are listed in
Appendix 1 of the 1973 Convention on International Trade in Endangered Species of Wild
Fauna and Flora. Trade in them, or their products, is subject to strict regulation and may not
be carried out for primarily commercial purposes.
All Lemuroidea are listed in Class A of the African Convention, 1969. They may not,
therefore, be hunted, killed, captured or collected without the authorization of the highest
competent authority, and then only if required in the national interest or for scientific
purposes.
84
The IUCN Red Data Book
In Madagascar, all lemurs are legally protected from killing or unauthorised capture. This
is, however, impossible to enforce.
CAPTIVE BREEDING None is recorded in captivity (ISIS, June 1989). Members of
this genus are generally difficult to keep alive in captivity for very long (Petter et al, 1977).
REMARKS L. leucopus is possibly the smallest of the Lepilemur species, it weighs about
550 g. Upper parts are medium to light grey, underparts are very pale grey or white. Its
tail is very light brown. The Malagasy name of the White-footed Sportive Lemur is songiky
(Tattersall, 1982).
The taxonomy of this group is very confused. It is common for all the types, the names and
numbers of which depend on the writer, to be considered as subspecies of L. mustelinus
(e.g. Tattersall, 1982; Richard, 1984). Contrary to his earlier work, Tattersall (in litt.) now
considers that the genus should contain several species. There is also considerable
disagreement over which family the genus should be in. It is traditionally put in Lemuridae
now it is commonly placed in Lepilemuridae but Schwartz and Tattersall (1985) have placed
it in Megaladapidae, along with some of the extinct lemurs. For a more detailed description
of this species and its taxonomy see Tattersall (1982) or Jenkins (1987).
REFERENCES
Andriamampianina, J. and Peyrieras, A. (1972). Les réserves naturelles intégrales
de Madagascar. In: Comptes Rendus de la Conférence Internationale sur la
Conservation de la Nature et de ses Ressources ad Madagascar, Tananarive, Madagascar
7-11 Octobre 1970. YUCN Gland, Switzerland and Cambridge, U.K. Pp. 103-123.
Charles-Dominique, P. and Hladik, C.M. (1971). Le Lepilemur du sud de
Madagascar: écologie, alimentation et vie sociale. La Terre et la Vie 25: 3-66.
Hladik, C.M. and Charles-Dominique, P. (1974). The behavior and ecology of
the sportive lemur (Lepilemur mustelinus) in relation to its dietary peculiarities. In:
Martin, R.D., Doyle, G.A. and Walker, A.C. (Eds), Prosimian Biology. Duckworth,
London. Pp. 23-37.
Hladik, C.M., Charles-Dominique, P. and Petter, J.-J. (1980). Feeding
strategies of five nocturnal prosimians in the dry forest of the west coast of Madagascar.
In: Charles-Dominique, P., Cooper, H.M., Hladik, A., Hladik, C.M., Pages, E.,
Pariente, G.F., Petter-Rousseaux, A., Petter, J.J. and Schilling, A. (Eds), Nocturnal
Malagasy Primates: Ecology, Physiology and Behavior. Academic Press, New York.
Pp. 41-73.
ISIS (1989). /S/S Species Distribution Report Abstract for Mammals, 30 June 1989.
International Species Information System, 12101 Johnny Cake Ridge Road, Apple
Valley, MN, U.S.A. Pp. 17-22.
Jenkins, P. D. (1987). Catalogue of Primates in the British Museum (Natural History)
and elsewhere in the British Isles. Part IV: Suborder Strepsirrhini, including the
Subfossil Madagascan Lemurs and the Family Tarsiidae. British Museum (Natural
History), London.
Nicoll, M.E. and Langrand, O. (1989). Revue Générale du Systéme d’Aires
Protégées et de la Conservation a Madagascar. Unpublished report to WWF.
O'Connor, S., Pidgeon, M. and Randria, Z. (1986). Lemur conservation in
Andohahela Reserve, Madagascar. Primate Conservation 7: 48-52.
O'Connor, S., Pidgeon, M. and Randria, Z. (1987). Un programme de
conservation pour la Réserve d'Andohahela. In: Priorités en Matiére de Conservation des
Espéces a@ Madagascar. Occasional Papers of the IUCN Species Survival Commission,
Number 2.
85
Lemurs of Madagascar and the Comoros
Petter, J-J., Albignac, R. and Rumpler, Y. (1977). Mammiféres lémuriens
(Primates prosimiens). Faune de Madagascar No. 44. ORSTOM-CNRS, Paris.
Petter, J.-J. and Petter-Rousseaux, A. (1979). Classification of the Prosimians.
In: Doyle, G.A. and Martin, R.D. (Eds), The Study of Prosimian Behavior. Academic
Press, London. Pp. 1-44.
Richard, A.F. (1984). Lemurs. In: Macdonald, D, (Ed.), The Encyclopaedia of
Mammals: 1. George Allen and Unwin, London. Pp. 330-331.
Russell, R.J. (1980). The environmental physiology and ecology of Lepilemur
ruficaudatus (=L. leucopus) in arid southern Madagascar. American Journal of Physical
Anthropology 52: 273-274.
Schwartz, J.H. and Tattersall, I. (1985). Evolutionary relationships of living
lemurs and lorises (Mammalia, Primates) and their potential affinities with European
Eocene Adapidae. Anthropological Papers of the American Museum of Natural History
60(1): 1-100.
Sussman, R.W. and Richard, A.F. (1986). Lemur conservation in Madagascar: the
status of lemurs in the south. Primate Conservation 7: 85-92.
Sussman, R.W., Richard, A.F. and Rakotomanga, P. (1987). La conservation
des lémuriens 4 Madagascar: leur statut dans le sud. In: Priorités en Matiére de
Conservation des spéces @ Madagascar. Occasional Papers of the IUCN Species
Survival Commission, Number 2.
Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New
York.
86
Pie “peat.
oceaynin.t v8 vid? vo
Lemurs of Madagascar and the Comoros
The small-toothed Sportive Lemur, Lepilemur microdon, occurs in the eastern rain forests.
Photo by Olivier Langrand/BIOS
88
The IUCN Red Data Book
SMALL-TOOTHED SPORTIVE LEMUR RARE
Lepilemur microdon Major, 1894
Order PRIMATES Family MEGALADAPIDAE
SUMMARY The Small-toothed Sportive Lemur is probably the most widely distributed
of the sportive lemurs. It is found in the south and central areas of the eastern rain forest.
Some authors do not distinguish between this species and Lepilemur mustelinus, which is
found in the rain forest north of the range of L. microdon and overlaps with it in some areas.
There are no estimates of population number. L. microdon has not been studied in any
detail and there is very little information on its ecology or social organisation. It is
nocturnal, mainly solitary and it eats leaves, fruit and flowers. Found in several reserves.
None is in captivity. Protected by Malagasy law and listed in Appendix 1 of CITES and
Class A of the African Convention.
DISTRIBUTION In the eastern forests from just south of Toamasina (Tamatave) at
around 18°S, to near Taolanaro (Fort-Dauphin) (Petter et al, 1977; Petter and Petter-
Rousseaux, 1979; Jenkins, 1987). It is sympatric with L. mustelinus in the north of its
range.
POPULATION Population numbers unknown but densities at Analamazaotra (Perinet)
were estimated to be about 100 per sq. km (Charles-Dominique and Hladik, 1971). These
authors did not distinguish between Lepilemur and Avahi in their nocturnal surveys, but
found combined densities of 200 individuals per sq. km and they considered that
approximately half of these were Lepilemur (Charles-Dominique and Hladik, 1971). The
figure estimated by Ganzhorn (1988) in the same forest is considerably lower, only 13 + 9
individuals per sq. km (mean and 95% confidence limits based on 25 census walks along
3.2 km of trails). The species is probably declining in number as the rain forest decreases in
area.
HABITAT AND ECOLOGY Little information exists on the ecology of this species. It
eats leaves, fruit and flowers and is mainly solitary (Ganzhorm, 1988).
THREATS No specific threats have been found, but the eastern rain forest, where this
species occurs, is disappearing. The principal agent of destruction of this forest is slash and
burn cultivation. FAO/UNEP (1981) gave a figure of 40,000 ha of previously undisturbed
closed forest cleared per year for the years 1976-1980, and they estimated 35,000 ha per
year for the years 1981-1985; the great majority of this is expected to be in the eastern
forests IUCN/UNEP/WWF, 1987).
CONSERVATION MEASURES Found in Andohahela (listed as L. m. mustelinus,
O'Connor et al, 1987) and Analamazaotra (Perinet) Nature Reserves (Nicoll and Langrand,
1989).
Better protection is needed for the eastern forest reserves to save at least some of the area
from destruction. This will succeed best if combined with education programmes and
development of alternative resources to replace those supplied by the forest at the moment.
Some of the protected areas could be developed as tourist attractions which would provide
some employment and income for Malagasy in the area. Analamazaotra, for instance, is
already comparatively easy of access and tourists do visit there.
All species of Lemuridae (which is taken to include the genus Lepilemur) are listed in
Appendix 1 of the 1973 Convention on International Trade in Endangered Species of Wild
89
Lemurs of Madagascar and the Comoros
Fauna and Flora. Trade in them, or their products, is subject to strict regulation and may not
be carried out for primarily commercial purposes.
All Lemuroidea are listed in Class A of the African Convention, 1969. They may not,
therefore, be hunted, killed, captured or collected without the authorization of the highest
competent authority, and then only if required in the national interest or for scientific
purposes.
All lemurs are protected from killing or unauthorised capture by Malagasy law, but this is
difficult to enforce.
CAPTIVE BREEDING None is known to be in captivity (ISIS, June 1989). Members
of this genus generally do not survive for long in captivity (Petter et al, 1977).
REMARKS The Small-toothed Sportive Lemur was recognised by Major (1894) on the
basis of the small size of its molars and pelage colour. It has been synonymised with
mustelinus by some authors (e.g. Tattersall, 1982; Richard, 1984), but others identify it as
a distinct species (Petter et al, 1977; Petter and Petter-Rousseaux, 1979). Jenkins (1987) has
re-examined Major's specimens and considers that two distinct forms do occur. Though
Tattersall (in litt.) now (in contrast to his 1982 publication) considers that the genus should
contain several species (six), he still does not separate mustelinus and microdon. Jenkins
(1987) describes L. microdon as red-brown on its upperparts, generally with a dark mid-
dorsal line and a yellowish-buff wash laterally and ventrally. Its underparts are grey and its
tail is distally dark brown. Its head and body length is given as 300-350 mm (Jenkins,
1987) and it probably weighs about 1 kg. Malagasy names of the eastern rain forest
Lepilemur are trangalavaka, kotrika, fitiliky, hataka, varikosy (Tattersall, 1982), but the two
species, microdon and mustelinus are not distinguished.
The taxonomy of the genus Lepilemur is very confused, greater details can be found in
Tattersall (1982) and Jenkins (1987). Briefly, it is common for all the species to be
considered as subspecies of L. mustelinus and the numbers and names of the subspecies
vary. There is also considerable disagreement over which family the genus should be in. It
is traditionally put in Lemuridae, now it is commonly placed in Lepilemuridae but Schwartz
and Tattersall (1985) have placed it in Megaladapidae, along with some of the extinct
lemurs.
REFERENCES
Charles-Dominique, P. and Hladik, C.M. (1971). Le Lepilemur du sud de
Madagascar: écologie, alimentation et vie sociale. La Terre et La Vie 25: 3-66.
FAO/UNEP (1981). Tropical Forest Resources Assessment Project. Forest Resources
of Tropical Africa. Part II Country Briefs. FAO, Rome.
mie ares J.U. (1988). Food partitioning among Malagasy primates. Oecologia 75:
ISIS (1989). ISIS Species Distribution Report Abstract for Mammals, 30 June 1989.
International Species Information System, 12101 Johnny Cake Ridge Road, Apple
Valley, MN. U.S.A. Pp. 17-22.
IUCN/UNEP/WWFE (1987). Madagascar, an Environmental Profile. Edited by M.D.
Jenkins. IUCN, Gland, Switzerland and Cambridge, U.K.
Jenkins, P. D. (1987). Catalogue of Primates in the British Museum (Natural History)
and elsewhere in the British Isles. Part IV: Suborder Strepsirrhini, including the
Subfossil Madagascan Lemurs and the Family Tarsiidae. British Museum (Natural
History), London.
Nicoll, M.E. and Langrand, O. (1989). Revue Générale du Systéme d’Aires
Protégées de la Conservation a Madagascar. Unpublished report to WWF.
90
The IUCN Red Data Book
O'Connor, S, Pidgeon, M. and Randria, Z. (1987). Un programme de
conservation pour la Réserve d'Andohahela. In: Priorités en Matiére de Conservation
des Espéces a Madagascar. Occasional Papers of the IUCN Species Survival
Commission, Number 2.
Petter, J-J., Albignac, R. and Rumpler, Y. (1977). Mammiféres lémuriens
(Primates prosimiens). Faune de Madagascar No. 44. ORSTOM-CNRS, Paris.
Petter, J.-J. and Petter-Rousseaux, A. (1979). Classification of the Prosimians.
In: Doyle, G.A. and Martin, R.D. (Eds), The Study of Prosimian Behavior. Academic
Press, London. Pp. 1-44.
Richard, A. (1984). Lemurs. In: Macdonald, D. (Ed.), Encyclopaedia of Mammals: 1.
George Allen and Unwin, London. Pp. 330-331.
Schwartz, J.H. and Tattersall, I. (1985). Evolutionary relationships of the living
lemurs and lorises (Mammalia, Primates) and their potential affinities with European
Eocene Adapidae. Anthropological Papers of the American Museum of Natural History
60 (1): 1-100.
Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New
York.
91
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Alaotran Gentle Lemur 173
Allocebus trichotis 60
Avahilaniger 187
Avahi laniger laniger 189
Avahi laniger occidentalis 191
Aye-aye 226
Black and White Ruffed Lemur 161
Black Lemur 118
Broad-nosed Gentle Lemur 182
Brown Lemur 139
Brown Mouse Lemur 40
CHEIROGALEIDAE 32-70
Cheirogaleus medius 50
Cheirogaleus major 56
Collared Lemur 145
Coquerel's Dwarf Lemur 44
Coquerel's Sifaka 219
Crowned Lemur 112
Daubentonia madagascariensis 226
DAUBENTONIIDAE 226-231
Decken's Sifaka 222
Diademed Sifaka 201
Dwarf Lemur,
Coquerel's 44
Fat-tailed 50
Greater 56
Hairy-eared 60
Fork-marked 66
Eastern Woolly Lemur 189
Fat-tailed Dwarf Lemur 50
Fork-marked Lemur 66
Gentle Lemur,
Alaotran 173
Broad-nosed 182
Grey 169
Western 174
Golden Bamboo Lemur 178
Golden-crowned Sifaka 210
Greater Bamboo Lemur 183
Greater Dwarf Lemur 56
Grey-backed Sportive Lemur 72
Grey Gentle Lemur 169
Grey Mouse Lemur 32
Hairy-eared Dwarf Lemur 60
Hapalemur aureus 178
Hapalemur griseus 169
SPECIES INDEX
239
Hapalemur griseus alaotrensis 173
Hapalemur griseus griseus 171
Hapalemur griseus occidentalis 174
Hapalemur simus 182
Indri 194
Indri indri 194
INDRIIDAE 186-225
Lemur,
Black 118
Black and White Ruffed 161
Brown 139
Collared 145
Crowned 112
Fork-marked 66
Golden Bamboo 178
Greater Bamboo 182
Mayotte 149
Mongoose 126
Red-bellied 132
Red-fronted 150
Red-ruffed 163
Ring-tailed 104
Rufous 150
Ruffed 159
Sanford's 153
Sclater's 121
White-collared 143
White-fronted 141
Woolly 187
LEMURIDAE 104-185
Lemur catta 104
Lemur coronatus 112
Lemur fulvus 139
Lemur fulvus albifrons 141
Lemur fulvus albocollaris 143
Lemur fulvus collaris 145
Lemur fulvus fulvus 147
Lemur fulvus mayottensis 149
Lemur fulvus rufus 150
Lemur fulvus sanfordi 153
Lemur macaco 118
Lemur macaco flavifrons 121
Lemur macaco macaco 120
Lemur mongoz 126
Lemur rubriventer 132
Lepilemur dorsalis 72
Lepilemur edwardsi 78
Lepilemur leucopus 82
Lepilemur imicrodon 88
Lepilemur mustelinus 93
Lepilemur ruficaudatus 97
Lepilemur septentrionalis 100
Lesser Mouse Lemur 32
Mayotte Lemur 149
MEGALADAPIDAE 72-103
Microcebus murinus 32
Microcebus rufus 40
Microdon Sportive Lemur 88
Milne-Edwards' Sifaka 205
Milne-Edwards' Sportive Lemur 78
Mirza coquereli 44
Mongoose Lemur 126
Mouse Lemur,
Brown 40
Grey 32
Lesser 32
Rufous 40
Northern Sportive Lemur 100
Nosy Bé Sportive Lemur 72
Perrier's Sifaka 206
Phaner furcifer 66
Propithecus diadema 201
Propithecus diadema candidus 204
Propithecus diadema diadema 203
Propithecus diadema edwardsi 205
Propithecus diadema perrieri 206
Propithecus tattersalli 210
Propithecus verreauxi 215
Propithecus verreauxi coquereli 219
Propithecus verreauxi deckeni 222
Propithecus verreauxi verreauxi 216
Red-bellied Lemur 132
Red-fronted Lemur 150
Red-ruffed Lemur 163
Red-tailed Sportive Lemur 97
Ring-tailed Lemur 104
Ruffed Lemur,
Black and White 161
Red-ruffed 163
Rufous Lemur 150
Rufous Mouse Lemur 40
Sanford's Lemur 153
Sclater's Lemur 121
Sifaka,
Coquerel's 219
Diademed 201
Decken's 222
Golden Crowned 210
Milne-Edwards' 205
Perrier's 206
Silky 204
Tattersall's 210
Verreaux's 215
240
Silky Sifaka 204
Small-toothed Sportive Lemur 88
Sportive Lemur,
Grey-backed 72
Microdon 88
Milne-Edwards' 78
Northern 100
Nosy Bé 72
Red-tailed 97
Small-toothed 88
Weasel 93
White-footed 82
Tattersall's Sifaka 210
Varecia variegata 159
Varecia variegata rubra 163
Varecia variegata variegata 161
Verreaux's Sifaka 215
Weasel Sportive Lemur 93
Western Gentle Lemur 174
Western-Woolly Lemur 191
White-collared Lemur 143
White-footed Sportive Lemur 82
White-fronted Lemur 141
Woolly Lemur,
Eastern 189
Wester 191
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The lemurs are a unique assemblage of primates that are found only on the
island of Madagascar. Isolated in their island home they have evolved and
flourished. That is until the arrival of humans on the island some 2000 years
ago. Since then several lemur species have become extinct as a direct result
of human activities. Today, the future of these fascinating creatures is even
more imperilled. Many of the habitats of Madagascar are being degraded, to
the detriment of both lemurs and humans. Lands that once supported life
are becoming increasingly barren, This book describes the plight of all lemur
species and discusses options for their conservation.
This book forms part of the IUCN Red Data Book Series, the authoritative
international register of threatened species. The IUCN Red List of
Threatened Animals is published every three years. In addition, analysis of
particular taxonomic groups are produced at periodic intervals.
Prepared by the World Conservation Monitoring Centre, a joint venture
between IUCN — The World Conservation Union, the World Wide Fund for
Nature (WWF), and the United Nations Environment Programme (UNEP).
Published by IUCN — The World Conservation Union, Gland, Switzerland
and Cambridge, U.K. with financial assistance from Bristol Zoo, Conserva-
tion International, World Wildlife Fund (U.S.) Primate Program, Madagascar
Fauna Captive Propagation Group, Jersey Wildlife Preservation Trust and
the Parc Zoologique et Botanique de la Ville de Mulhouse.
a
“j LOO
WWF CONSERVATION GARDENS
INTERNATIONAL
This book is part of
THE IUCN CONSERVATION LIBRARY
For a copy of the IUCN Publications Catalogue, please write to:
IUCN Publications Unit
219c Huntingdon Road,
Cambridge, CB3 ODL, UK