Lemurs of Madagascar and the Comoros The IUCN Red Data Book Prepared by The World Conservation Monitoring Centre IUCN — The World Conservation Union ¢ = a 2 4 é 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 . o 00 ‘i 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. a | ae 2 ae ee - : — ; i 2%) j ? : vf K = a if j ; a \r ae een , 8 a e ollie hy Fincet, Chen, Swineerientd and Canibicidge: a Ee . with | cin super Leer Aaijol Zao, Conservation in , ud 8D 5) Prater Pragrnad eee Cap ' Wie . es . $e BRIS WH i: og ie MAVALION = - r DENS > i nae i abate a = S mr 2 . ; 2 4 +> 3 ae < rhs ae Capetight. iad ‘ - te noni pissce rol Mature end Na ion A j ) “4 - a sat cee a Das enithics tion for edix ational ume ry ~ my SE Dee ig ssion trong We cop | ae | =) (0891-2001). — uae si: + if ptt fare eeialie or wth r dosamen: ial P urpio maka: =o - t. oe = witkivsn — ww vi deerme priate ty bs ; ~ eh nat ae ie $ eo te : : wate Besabees, <7 aited, Te a f c - "T- wR Lt ae . (ee Sarees eiries Stes peaks ies ti many tty tity Nai, CON Peotacatame § or ae ~ f E > 1 af Sa, os }:; eats Wore ee pictues Nencaitn, D Can ye he Reh roms & — onset. ds 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 ‘s cag bagi Somat y s Couserwalion needs we Gy imerraed re es. ecg (ayy “ ia ay nes i ’ es ies petrn : , ~ lee =~ (ert, BFA. uel Sugeno prem Deloresuptinn distor oft a ¢eoqgemna Of bailey ip FRA ci To Sen. Selene. (heerbees A. 0708 La soporte S dc Saath wae feee titres A Madags : 7 i e ' ore (ind a - > ee os : ee regress of fin rising Ge Tene : ae! Ss Socsttiae sa Rind vwig.J . Gitean, fo vie, Bd i bh hee meer 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 = ® =; » < cy) < < Q 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 | hs — sa own 7 lemieeadl of hapa pelaninm & ‘ Atos Sania a + Revised rele " ie fru s : Gary , TenetHnA ¢ Ss a Ag eee 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. +4 i® - él ¥ ; bata * . , he + ie erally , daz: a that di a a aa | i eSB LH “aioe ht a tae nt mca ; wie Mis mune nn hoo meat | % : ‘ dint. Wee E are Oe all rs] ee ce 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 " wee 4 xX “ka i ' “ry t af th Uy , : Tv . ae at A Fe eC ' ‘ >. . e ~ ¢ 1 pF ‘ ‘ aie. Oty ’ ivi ale a | : ae, ‘ Pimel , GPs pits Aun sat i v ? } \ ] ‘ 1 ra Ue «fel yo eae s “att sihis xia: ot Sia iB pases Litt fis Mbit ty thane rie’ vid ee Bid ms ; & Py oe Peay arn | wey: i J : oy e ‘sy! FAM i Specs 2 Pp axe 3. onan Hy Fea a pesanenaslagtth ch = oe Tot Ue actin Cnaggealiges Spotty a eiohis ot maimence and the rsdn the IEEE VCS. ‘ 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 +) few tarrpta, a a tae mt Dede toon Se vital o me's 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 | . re Ms Ce Seowmuidy adTaw: s( Tea ) cx aac rsa te otis eh avkrw Wp alates’ mirpeisderobaA'b sv1923H al ve lavivs ue exisog2 MDUI-8d) Yo 210989 IsnoienooO rota B sot Al ii urohica fue listed inl 135 A of the Affican t ‘unven 3 j anoiiomdbexd ited COPRIED Mccligtgastas®t nba eemgiellite ine [: COM pele met EAMOMOTEAGs nad jOliogusingnbe A dome STR AN wEiaddold oft to nove vilizzalD tere At eb) LE A-r9ik me 4 ‘Simobhe7A sOivils&. aoimdzy4 i) v2.34 im 3 = Dy . All lemurs are protested’ fram Sie ort nauth boriees eunere Be nl D. g Nene AEA) Apel Saree TEE-OEE . v obno! caiwn Gail toi BA ee AST Gata yea OTT We SAOT es! merger i Beenie Ritdaroy vides Lita (oormaniy jak Linitert eRAN, ie onl inuan) io ruse enotratA ay: © eed losigh logosain EMARKES The Small-toothed Spartive Vemar was recognised by N bees eg olen vines aloe: mutorD. Auswapy wBaté 4 anoint wt Des mustélinus by some authors (ep. Tallessall, 1982; 1984), & a distinct speci ce (Pctet- ov ‘oseegnbo, sb -aruin4 Aensimi HTT VD DRBLL pane bap rmibtidenistan] \ueye-oes oft 3 Sn ak AD Yh ie ot meno DootteN) Ate ate aagahiedin irs Sauindiniedr ausasei@otiay Resio (EHeL sie metal hav hae yess AidighoRe ti AME Mpa yas pem dhaljuacnw/inan bd pe eee kro 199 Jee werpemedT, | OBA) iA wunneais st raphe Ravin wh a Sod NON TADYMDD Sd wod Hit "Ne AtOG-2QI ; pi Rep ae 6 mean ie: ? -tiaeookd DRED ch sarsbveet habe mala hk cai toa) H.alemaincorngolanen giro yin) 979 06 abst 16 coordinate a Sreeding programme [Oe animais of ihis species being HER ebat bokin@G id inge Miser ngiritf emo: mogsA yoniumilsy4 88h) Et alos ; W oF noqe hevziiduga 222) wdmaiqe? ben maguA ni ralias ~ tpreeh AR Kato ri2teD was Bee one majemraay ih “AE vq sidetinn Dy. gabaM ~~ “BER ORE td *bnalal rome ods Yo nurmol es “die ryote A ~ ——s a! > x ee —_> >> meas . S a : Xe i Pry oy 7 se ag é : Some, es te Da el 3 } >, Ss ey 7 ~! Pa ‘ “7 © 3 au , a - = p + 5 a me : ee rt 4 ‘a, sd A \ ben j - i : i i. ae > i — m ~ b ts AS a - nt "vr har? (ag¥ van a rae = * ~~ Ye | pan “* ‘ ae " \ wd H PN Tr. Ss ¢ rel, er hr %~ * 3 a partg i ~T. (9 vy = a y Lo ch Se eh f -_s ee ~~ ee a Peake 24 ee a “ ye A 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 Vi “f barley a = ee ee ~ il : | enae Moeiiee Laman V heard ab Merete Lancs 14? | Frere po gy oF wa ADAPIAL 7) 1c - Grey tacked Wl «*reke aapwry ~ — “Mi n = ome uly q aa =. crniow Sgrveive Liczatp sat Ee Faowegeli Svfabe 205 “< a ss ing Ay cones \ eee t) idee TS ; eT vu 2. “errr! “ ¥- Mengiows Lio im Moat wero 5 \ ~~ =a @ : - r . Ciey ° SC ae Tattersall's 7 aoe | Wale Varecie Seegieers Wreeer i rte: 11 Vorech A ee, ec ' White fo {Sportive Lemsenii me Whi-trontod Lem 14) ie Wal ly Ler, Eastern 189 (7 Western 391) oe = = — : pe p ; ( | ' : | « . ’ i. Se) apes rie #3 0 AG: is i eye Srey ging tg jt oe AL ayy gh - oY ond ae tyre ere ie ap a as 4a = a. a6 ar Pgh et hlle- , gee" ies » t ‘ age tao eet eae oh % . me <~ 0) ae Kaha AD bd) Bie emote) prs rape Place ia Ee oe oS ee ~— alos ike =~ ee sa EE ete hor op 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