Vol. 15 - Deutsches Primatenzentrum

LEMUR NEWS 

The Newsletter of the Madagascar Section of the IUCN/SSC Primate Specialist Group 

Vol. 15, 2010 ISSN 1608-1439 

Editors 

Christoph Schwitzer (Editor-in-chief) 

Bristol Conservation and Science Foundation, Bristol Zoo Gardens, UK; cschwitzer@bcsf.org.uk 

Claudia Fichtel 

German Primate Center, Göttingen, Germany; claudia.fichtel@gwdg.de 

Jörg U. Ganzhorn 

University of Hamburg, Germany; ganzhorn@biologie.uni-hamburg.de 

Rodin M. Rasoloarison 

German Primate Center, Göttingen, Germany; kirindy@simicro.mg 

Jonah Ratsimbazafy 

GERP, Antananarivo, Madagascar; gerp@wanadoo.mg 

Anne D. Yoder 

Duke University Lemur Center, Durham, USA; anne.yoder@duke.edu 

IUCN/SSC Primate Specialist Group 

Chairman Russell A. Mittermeier, Conservation International, Arlington, VA, USA 

Deputy Chair Anthony B. Rylands, Conservation International, Arlington, VA, USA 

Coordinator – Section on Great Apes Liz Williamson, Stirling University, Stirling, Scotland, UK 

Regional Coordinators – Neotropics 

Mesoamerica – Liliana Cortés-Ortiz, Museum of Zoology & Department of Ecology and Evolutionary 

Biology, University of Michigan, Ann Arbor, MI, USA 

Andean Countries – Erwin Palacios, Conservation International Colombia, Bogotá, Colombia and 

Eckhard W. Heymann, Deutsches Primatenzentrum, Göttingen, Germany 

Brazil and the Guianas – M. Cecília M. Kierulff, Instituto para a Conservação dos Carnívoros 

Neotropicais – Pró-Carnívoros, Atibaia, São Paulo, Brazil, Fabiano Rodrigues de Melo, Universidade 

Federal de Goiás, Jataí, Goiás, Brazil, and Maurício Talebi, Universidade Federal de São Paulo, Diadema, 

São Paulo, Brazil 

Regional Coordinators – Africa 

West Africa – W. Scott McGraw, The Ohio State University, Columbus, OH, USA 

Regional Coordinators – Madagascar 

Jörg U. Ganzhorn, Hamburg University, Hamburg, Germany, and Christoph Schwitzer, Bristol Conservation 

and Science Foundation, Bristol Zoo Gardens, Bristol, UK 

Regional Coordinators – Asia 

China – Long Yongcheng, The Nature Conservancy, China 

Southeast Asia – Jatna Supriatna, Conservation International Indonesia Program, Jakarta, Indonesia, and 

Christian Roos, Deutsches Primatenzentrum, Göttingen, Germany 

IndoBurma – Ben Rawson, Conservation International, Hanoi, Vietnam 

South Asia – Sally Walker, Zoo Outreach Organization, Coimbatore, Tamil Nadu, India, and Sanjay Molur, 

Wildlife Information Liaison Development, Coimbatore, Tamil Nadu, India 

Editorial assistants 

Nicola Davies, Rose Marie Randrianarison 

Layout 

Heike Klensang, Anna Francis 

Front cover: The Endangered golden-crowned sifaka (Propithecus tattersalli) at the edge of an area devastated 

by gold mining activities in the Daraina region of north-eastern Madagascar. © Pete Oxford/naturepl.com 

Addresses for contributions 

Christoph Schwitzer 

Bristol Conservation and Science Foundation 

Bristol Zoo Gardens 

Clifton, Bristol BS8 3HA 

United Kingdom 

Fax: +44 (0)117 973 6814 

Email: cschwitzer@bristolzoo.org.uk 

Lemur News online 

All 15 volumes are available online at www.primate-sg.org, www.aeecl.org and www.dpz.eu 

This volume of Lemur News was kindly supported by the Margot Marsh Biodiversity Foundation 

(through Conservation International’s Primate Action Fund) and by WWF Madagascar. 

Printed by Goltze GmbH & Co. KG, Göttingen, Germany 


GERP 

34, Cité des Professeurs 

Antananarivo 101 

Madagascar 

Email: gerp@wanadoo.mg

Lemur News Vol. 15, 2010 Page 1 

Editorial 

I am writing this Editorial only a couple of days after another 

attempted (and failed) Coup d’Etat in Madagascar,in which a 

faction of the army tried to topple the Transition Government. 

For nearly two years now, since the start of the political 

crisis in early 2009,the country has not seen a week without 

demonstrations, tensions between different political 

parties and attempts from international mediators to get 

power-sharing agreements signed by all sides. Most donors, 

governments and multinational organisations alike, have 

frozen all non-humanitarian aid for Madagascar,which has led 

to severe funding shortages in the environmental and conservation 

sector. The political crisis has thus quickly turned 

into a full-blown environmental crisis, with large-scale illegal 

logging taking place mainly in eastern Madagascar (Marojejy, 

Masoala, Makira), and unseen levels of lemur poaching all 

across the island.To keep people aware of the seriousness of 

the situation we have decided to run another feature on 

Madagascar’s environmental crisis in this issue of Lemur News, 

with an excellent update on illegal logging by Erik Patel as 

well as a case study of ongoing threats to lemurs and their 

habitat in Sahamalaza National Park by Melanie Seiler and 

colleagues. 

The conservation situation of lemurs has also been a big concern 

in several presentations given at the most recent 23rd 

Congress of the International Primatological Society in 

Kyoto, Japan. The talk that I remember best was by Lemur 

News co-editor Jonah Ratsimbazafy, who reminded the audience 

in a very emotional way that scientists and conservationists 

working in Madagascar had a moral responsibility to 

respond to the "cries of the lemurs", as otherwise these 

would remain unheard by the Malagasy and international 

community.In the biennial discussion session of "Primates in 

Peril", the list of the world’s top 25 most endangered primates, 

issued jointly by the IUCN/SSC Primate Specialist 

Group and IPS,lemurs remained a very high priority and will 

again make up 20% of the 25 listed species in the next biennium.Sadly,Madagascar 

thus retains its first place (along with 

Vietnam) as the country harbouring the highest number of 

the top 25. It can only be hoped that the political classes of 

Madagascar come to agree a way out of the current crisis 

sooner rather than later, as otherwise we run the very serious 

risk,during the UN Decade of Biodiversity 2011-2020,of 

losing a substantial proportion of the endemic biodiversity of 

this amazing megadiversity country. 

Alison Jolly with Russ Mittermeier at the IPS Lifetime Achievement 

Award 2010 ceremony in Kyoto. (Photo: R. Mittermeier) 

For a change,on a very positive note,I am thrilled to say that 

Alison Jolly was awarded the IPS Lifetime Achievement 

Award for her long-term commitment to lemur conservation 

and environmental education in Madagascar (see News 

and Announcements). My two daughters (now 4 and 2 years 

old) and I particularly enjoy reading Alison’s children’s book 

on Bitika,the mouse lemur,as,I am sure,do lots of children in 

Madagascar and elsewhere in the world. 

It is encouraging to see that this volume of Lemur News is 

again full of articles and short reports not only on lemur species 

red-listed in one of the Threatened categories (VU, EN 

or CR),but also on Data Deficient nocturnal species such as 

Mirza zaza,Lepilemur leucopus and the recently rediscovered 

Cheirogaleus sibreei (see the articles by Rode et al., Fish, and 

Blanco, respectively). As Johanna Rode and colleagues point 

out in their short report on Mirza zaza,Madagascar is in the 

unusual situation that 45 % of its primate species are redlisted 

as Data Deficient,which is a far higher percentage than 

in any other primate habitat country and mainly derived 

from the discovery of dozens of cryptic species in the genera 

Lepilemur and Microcebus over the last couple of years. Many 

of those species are only known from their type localities 

and may in fact be highly endangered. The more research is 

conducted and published on them, the easier it will become 

to assign them a conservation status and target them with 

conservation measures. It will require a concerted effort of 

the lemur research and conservation community over the 

next decade or so to to reduce the number of Data Deficient 

species to a level comparable to other regions (or, ideally, to 

zero). 

Another encouraging development is the frenzy of research 

and conservation activities now under way for Prolemur simus 

at various locations both south and north of the Mangoro 

River,reported by Dolch et al.as well as Rajaonson et al.in this 

volume. The greater bamboo lemur undoubtedly remains 

one of the most endangered of Madagascar’s lemurs. However, 

with several additional populations having been discovered 

over the last two years, workshops having been conducted 

that have led to a joint-up approach to this species’ 

conservation,and the ex situ population having been included 

as an integral part of conservation efforts, I now think that 

we stand a real chance of saving Prolemur simus from extinction. 

As Jörg Ganzhorn announced in his editorial to Lemur News 

14, I have taken over the coordination of this newsletter 

from him after the 2009 volume, hence this is now the first 

volume that I have helped produce (which is my humble excuse 

for its slightly late publication). Jörg has been involved 

with Lemur News since its inception in 1993,first as a member 

of its Editorial Board and from volume 3 (1998) as its Editor.I 

am thus pleased to say that we will not lose his experience 

and backing,as he has kindly agreed to remain part of the editorial 

team. Likewise, Jonah Ratsimbazafy and Rodin Rasoloarison,who 

have been the newsletter’s Malagasy coordinators 

since 2006, and Anne Yoder, who represents the Duke 

Lemur Center, will carry on as editorial team members, for 

which I am grateful.I am indebted to Heike Klensang,who has 

been doing the layout for Lemur News now for more than a 

decade and is still not tired of it,and to Anna Francis,who has 

designed the beautiful new logo and front cover. Very many 

thanks also to Stephen D. Nash for the wonderful lemur silhouettes 

that we printed on the inside back cover. 

This volume of Lemur News was kindly supported by the 

Margot Marsh Biodiversity Foundation through Conservation 

International’s Primate Action Fund, and by the WWF 

Madagascar and West Indian Ocean Programme Office.

Page 2 Lemur News Vol. 15, 2010 

I very much look forward to helping to take Lemur News into 

the UN Decade of Biodiversity together with the editorial 

team and with its base of loyal contributors and readers,and I 

will do my best to ensure that the newsletter will continue to 

help promote the conservation of lemurs as it has done for 

the last 17 years. 

Christoph Schwitzer 

Feature: Madagascar’s 

Environmental Crisis 

Madagascar’s illegal logging crisis: an update 

and discussion of possible solutions 

Erik R. Patel 

Cornell University, 211 Uris Hall, Ithaca, NY 14850, USA, 

patel.erik@gmail.com 

How sure are you that your favorite rosewood or ebony 

acoustic guitar was not made from rare,illegally logged trees 

in Madagascar;an exceptional biodiversity hotspot with desperately 

little original forest remaining? What is the origin of 

the wood in the expensive oriental-style rosewood furniture 

which is heavily advertised for sale on the internet? Unfinished 

rosewood boards from Madagascar are openly sold 

even in the United States (www.gilmerwood.com/boards_ 

rosewood-exotic_unique.htm) and the United Kingdom (www. 

exotichardwoods.co.uk/Woods_List/Madagascar_Rosewood.asp). 

Can such vendors prove that the rosewood was legally (and 

ethically) obtained? The answer is usually "no".These can be 

difficult questions for consumers to answer, but purchasing 

these products can prolong the ongoing logging crisis in 

northeastern Madagascar in some of the most unique and biologically 

diverse forests in the world. 

Consumers should be suspicious since none of these rapidly 

disappearing Madagascan rosewood and ebony species are 

yet protected under CITES,the Convention on International 

Trade in Endangered Species. In November of last year, Gibson 

Guitars, one of the two largest U.S. stringed-instrument 

companies,came under federal investigation for violating the 

Lacey Act by allegedly using illegal rosewood from Madagascar 

which had first been shipped to Germany and then the 

United States (Michaels, 2009). Most of the illegally logged 

rosewood in Madagascar is used for the manufacture of furniture 

in China. Some of this is known to be sold in China as 

luxurious "Ming Dynasty style" furniture (Global Witness 

and Environmental Investigation Agency, 2009). Some may 

well be exported to western countries. China is the world’s 

leading exporter of furniture.According to the Office of the 

United States Trade Representative, the United States imported 

16 billion dollars of Chinese furniture in 2009,making 

it the USA’s fifth largest import from China. 

Illegal logging of rosewood (Dalbergia spp.) and ebony (Diospyros 

spp.) has emerged as the most severe threat to Madagascar’s 

dwindling northeastern rainforests. In 2009, a year 

of political upheaval in Madagascar due to an undemocratic 

change of power,approximately 100,000 of these trees were 

illegally cut in the UNESCO World Heritage Sites of Masoala 

National Park,Marojejy National Park,the Makira Conservation 

Site, and Mananara Biosphere Reserve (also a national 

park).Needless to say,the wood is extremely valuable.Rosewood 

can sell for US$5,000 per cubic meter,more than double 

the price of mahogany.Several hundred million dollars of 

these precious hardwoods were cut in 2009 in protected 

areas. The overwhelming majority of these profits are taken 

by a rosewood mafia of a few dozen organizing individuals, 

many of whose identities are well known.Few others benefit. 

Harvesting these extremely heavy hardwoods is a labor intensive 

activity requiring coordination between local residents 

who manually cut the trees, but receive little profit 

(about US$5/day), and a criminal network of exporters, domestic 

transporters, and corrupt officials who initiate the 

process and reap most of the enormous profits. This is a 

"tragedy with villains" unlike habitat disturbance from subsistence 

slash-and-burn agriculture which has been well described 

as a "tragedy without villains" (Barrett et al., 2010; 

Débois, 2009; Global Witness and Environmental Investigation 

Agency,2009;Patel,2007,2009;Randriamalala and Liu,in 

press;Schuurman and Lowry,2009;Schuurman,2009;Wilme 

et al., 2009; Wilme et al., in press). 

Globally, illegal logging results in an estimated US$10 billion 

lost per year to the economies of timber producing countries 

(Furones, 2006). In addition to depriving the government 

of Madagascar of millions of dollars of taxable revenue, 

illegal logging of this precious wood has decimated tourism 

in northeastern Madagascar, which had become a growing 

source of local income. Although selective logging results in 

less absolute forest loss than clear-cutting, it is often 

accompanied by substantial peripheral damage such as decreases 

in genetic diversity and increases in the susceptibility 

of the impacted areas to burning and bushmeat hunting. 

Documented long-term ecological consequences of selective 

logging in Madagascar include invasion of persistent, 

dominant non-native plant species, impaired faunal habitat, 

and a diminution of endemic mammalian species richness 

(Gillies, 1999; Cochrane and Schultze, 1998; Brown and 

Gurevitch, 2004; Stephenson, 1993). In actual practice, rosewood 

logging has turned out to be far less "selective" than 

originally believed. Often rafts made of a lighter species of 

wood (Dombeya spp.) are constructed to float the much 

more dense rosewood logs down rivers. Approximately five 

Dombeya trees are cut as "raft wood" for every one rosewood 

tree (Randriamalala and Liu,in press).Tall adult trees of 

a variety of species, that simply happen to be very close to 

rosewood trees, must often be cut simply to gain access to 

cut down a rosewood tree. This has been observed in 

Marojejy (pers. obs.). 

Red ruffed lemurs (Varecia rubra) are probably the most negatively 

impacted lemur since many were hunted by these loggers 

and this species is known to feed on ebony trees 

(Diospyros spp.) as well as pallisandre (Dalbergia spp.) in 

Masoala (Vasey, pers. comm.). Varecia rubra probably also 

feeds on the fruits and leaves of the logged "raft wood" 

Dombeya spp. trees like Varecia v. editorium in Manombo Forest 

in southeastern Madagascar (Ratsimbazafy, 2007). In 

Mantadia National Park, Indri indri and Propithecus diadema 

consume young leaves of one species of actual rosewood 

(Dalbergai baronii) which is also consumed by Milne-Edwards’ 

sifakas (Propithecus edwardsi) in Ranomafana National Park 

(Powzyk and Mowry,2003;Arrigo-Nelson,2007).Propithecus 

diadema at Tsinjoarivo consume the unripe fruit of ebony 

trees (Irwin, 2006). In Marojejy, silky sifakas (Propithecus 

candidus) not uncommonly feed on the young leaves of pallisandre 

(Dalbergia chapelieri) which is also a preferred sleeping 

tree (pers. obs.). 

When discussing the impacts of precious wood logging, it is 

important not to forget how damaging all this has been to local 

communities as well.Local residents have suffered as foreign 

and domestic elites have corrupted the forest service, 

leading to losses of sustainable employment in tourism, re-


search, and conservation. In some cases, community life has 

suddenly changed as gambling, prostitution, and crime have 

increased in rural communities. Moreover, the risks of local 

food shortages and nutritional deficiencies mount when 

farmers abandon subsistence agriculture for temporary,physically 

dangerous illegal logging work (Global Witness and 

Environmental Investigation Agency,2009;Patel,2007,2009). 

Moreover,illegal loggers trample on the beliefs and taboos of 

local people.In traditional Sakalava culture,ebony is a sacred 

wood only cut by priests who conduct traditional ceremonies 

with ebony staffs. The chief of Ankalontany, a Sakalava 

Malagasy village in the northeast, explained that "Some 

strangers from outside our village came here. They started 

cutting ebony and they clearly had no right. We asked for 

their authorization but they said they didn’t have to show us 

papers.They said they had police clearance and we can’t stop 

them." Laurent Tutu, president of the forest association of 

Ankalontany, remarked "It hurts us to see our trees cut like 

this. The forest loses its personality" (Cocks, 2005). 

Although illegal logging in Madagascar has received some media 

attention recently, confusion still remains regarding a 

number of key facts. The aim of this report is to provide an 

update (at the time of writing: May 25, 2010), dispel a few 

myths,discuss some of the possible solutions to this ongoing 

crisis, and present a comprehensive bibliography of articles, 

photos, films, and videos related to this topic. 

Four myths about illegal logging in Madagascar 

Myth #1: "Plenty of Madagascar rosewood is harvested legally…" 

says Bob Taylor, founder of Taylor Guitars. Quote 

from Gill, C. (2010). Log Jam. Guitar Aficionado. Spring Issue. 

P. 68 

This is simply not true. A vast amount of published evidence 

clearly shows that very very little,if any,of the rosewood logging 

in Madagascar is legal.The overwhelming majority of exported 

Madagascar rosewood is illegally logged within Masoala 

National Park and Marojejy National Park (which are part 

of a UNESCO World Heritage Site) as well as Mananara Biosphere 

Reserve (also a national park) and the vast Makria 

Conservation Site (Barrett et al., 2010; Débois, 2009; Global 

Witness and Environmental Investigation Agency,2009;Patel, 

2007, 2009; Randriamalala and Liu, in press; Schuurman and 

Lowry, 2009; Schuurman, 2009; Wilme et al., 2009; Wilme et 

al., in press). 

Myth #2: The current ban has stopped illegal logging. 

In late March, the government of Madagascar announced a 

new two to five year ban on export and cutting of ebony and 

rosewood. The decree #2010-141 officially passed on April 

14, 2010. Clearly this was an important and large step forward. 

However, the decree does not apparently include 

pallisandre, a precious hardwood in the same genus (Dalbergia) 

as rosewood. Illegal logging of pallisandre has heavily 

impacted some reserves such as Betampona Natural Reserve 

(Kett, 2005; Bollen, 2009). At the time of writing (May 

25, 2010), there have been no new exports since the recent 

ban.However,illegal rosewood and ebony logging still continues 

inside Mananara Biosphere Reserve and the Makira Conservation 

Site according to reliable anonymous informants. 

The clearest information has come from Mananara where at 

least several hundred, recently cut, rosewood logs were observed. 

Myth #3: Illegal logging was never a big problem in Madagascar 

until the recent political crisis. 

Illegal logging in Madagascar of rosewood (Dalbergia spp.) and 

ebony (Diospyros spp.) did not begin with the culmination of 

the political crisis in March 2009.A major illegal logging crisis 

in World Heritage Sites (Masoala National Park and Marojejy 

National Park) took place during 2004-2005,a time of political 

stability. The earliest documented case of rosewood logging 

in Madagascar and foreign export dates to 1902.Foreign 

exports of Madagascar rosewood occurred at "low" levels 

(1000 to 5000 tonnes) between 1998 and 2007. In 2008, exports 

jumped to 13,000 tonnes,and jumped again in 2009 to 

more than 35,000 tonnes (Botokely,1902;Randriamalala and 

Liu,in press;Global Witness and Environmental Investigation 

Agency, 2009). 

Myth #4: There are 43 species of rosewood trees in Madagascar. 

Some recent reports had mistakenly made this statement. It 

is not entirely clear exactly how many rosewood species are 

found in Madagascar. More botanical research is needed. 

However, currently, there are believed to be 10 species of 

rosewood in Madagascar in the genus Dalbergia which contains 

48 total species. The rosewood species are presumed 

to be Dalbergia baronii [VU], D. bathiei [EN], D. davidii [EN],D. 

louvelii [EN], D. mollis [NT], D. monticola [VU], D. normandii 

[EN], D. purpurascens [VU], D. tsiandalana [EN], and D. viguieri 

[VU] (Barrett et al., 2010). 

Rosewood stockpile solutions? 

Approximately 10,280 tonnes of illegally logged rosewood 

remain stockpiled in numerous locations in northeastern 

Madagascar, such as the ports of Vohemar and Antalaha as 

well as private residences in those cities and Sambava, 

Ampanifena,Ambohitralalana,and others.Each 150 kg log has 

an approximate market value of US$1,300 usd.As unfinished 

logs, the value of the current stockpile is therefore approximately 

US$90 million.Value increases dramatically,of course, 

after being constructed, for example, into high-end Ming 

Dynasty style furniture in China.A single armoire composed 

of only a few logs can sell for US$20,000 or more.It’s a horrid 

contrast to the annual income in Madagascar (about 

US$400) or the daily wage provided to loggers (US$5) for 

the dangerous and physically debilitating work (Randriamalala 

and Liu, in press; Global Witness and Environmental 

Investigation Agency, 2009; anonymous local informants). 

If the export ban holds (numerous other bans did not),what 

should be done with these stockpiles? Several ideas have 

been suggested. 

1. The "Forest Counterpart Fund" (Wilme et al., 2009) aims 

to create a conservation and charitable works fund to assist 

local communities and forests damaged by the illegal logging. 

The logs are not sold on the open market as in the second 

proposal below.Rather,philanthropists,conservation organizations, 

and international aid agencies pay to "adopt" a log. 

Each log can be "adopted" for its market value (about 

US$1,300). The logs themselves are given to (carefully selected) 

local residents who are victims of the illegal logging. 

The logs would then be carved,engraved,and customized for 

public display as symbols. If sufficient donors can be found, 

this proposal offers a win-win solution for Madagascar’s forests 

as well as people. 

2. The Moratorium-Conservation-Amnesty-Reforestation 

(MCAR) program (Butler, 2009). This is essentially a one-off 

actual sale with conservation benefits. Logs would be auctioned 

via a transparent market system in which the price 

and the log code would be recorded, publicly available, and 

digitally traceable.Funds generated would mainly go towards


conservation programs such as reforestation and forest 

monitoring. Criminal traders would receive amnesty from 

prosecution as well as a very small percentage of the funds. 

An export moratorium would be required.There is always a 

danger that one-off sales can encourage further logging; a 

topic which has been extensively debated with respect to 

confiscated elephant ivory stockpiles. An impressive recent 

review paper in Science (Wasser et al., 2010): 

www.sciencemag.org/cgi/content/short/327/5971/1331) 

argued that no one-off ivory sales should be approved even if 

the funds go towards conservation. 

3. Destroy the stockpile. This was recently reiterated by 

Global Witness (GW) and Environmental Investigation 

Agency (EIA). Andrea Johnson, Director of Forest Campaigns 

at EIA explained that "To end the cycle of illegal harvest 

and corruption, the government should take the step of 

destroying all stocks that are not contained in the latest official 

inventories…Traders, who are currently stockpiling illegal 

timber, hoping for another ‘exceptional’ export authorization,must 

receive a clear signal that it will be impossible to 

profit from the illegal trade in the future." Numerous examples 

can be found from around the world of simple and effective 

destruction of stockpiles of contraband such as small 

arms,drugs,and ivory.Destruction also eliminates the not insignificant 

expense of storing and guarding the items.Burning 

the rosewood stockpiles would create a lot of pollution, it 

has been argued, and might be dangerous given the high volume. 

Other ways of destroying the wood are possible however. 

The wood could be hacked into tiny unusable pieces. 

This is already done sometimes by park rangers in Madagascar. 

This would take a very long time, but would be a fitting 

punishment of hard labor for members of the rich rosewood 

mafia! Of course, destruction of the wood, whatever the 

method,would contribute no money for any conservation or 

community development funds. 

Any of these possibilities are better than what has happened 

in the past:seized wood was auctioned off to the highest bidder.Foreign 

export remains a possibility too,despite the ban. 

French shipping company CMA-CGM Delmas exported 

rosewood from Madagascar several times in 2009 and 2010. 

Long-term solutions? 

Thinking long-term, what can be done to prevent another illegal 

logging crisis in Madagascar? 

Some may argue that so little rosewood and ebony remains, 

logging on this scale could never happen again.However,this 

had been claimed before 2009 too. More surveys are clearly 

needed. One hopes that some of the more impenetrable regions 

of mountainous Marojejy National Park may still have 

rosewood. But because rosewood tends to be harvested at 

lower elevations, near rivers (where the largest individuals 

are found), it is less protected by the physical challenges of 

the massif than some other tree species. It is encouraging 

that some Dalbergia and Diospyros species can form stump 

sprouts which can grow into a new tree over many many 

years. Unfortunately, some entire rosewood stumps are removed 

either to hide evidence of logging or for wood for 

small, locally made rosewood vases. Rosewood trees are 

known to be some of the oldest trees in the eastern Malagasy 

humid forests. They can live to be more than 400 years old, 

according to local guides. Traders explain that they can be 

harvested after 50 years (Patel 2007, 2009). 

1. CITES 

The surest way to reduce the likelihood of another illegal 

logging crisis in Madagascar, is to list all species in the genera 

of Dalbergia and Diospyros on CITES Appendix 1. Currently 

none of Madagascar’s ebony or rosewood species are protected 

under any appendices within the Convention on International 

Trade in Endangered Species (CITES). Globally, 

only one species of rosewood,Brazilian rosewood (Dalbergia 

nigra), is listed under CITES Appendix 1. This is the most 

stringent category,and prohibits all commercial trade of that 

wood from the date of listing. This has generally been effective.Guitars 

in the United States made of Brazilian rosewood 

are known to have risen in price and are harder to find since 

Appendix 1 listing. Similarly, Appendix 1 listing of Alerce 

(Fitzroya cupressoides),a heavily logged South American conifer, 

has significantly reduced logging and trade (Barrett et al., 

2010; Keong, 2006). 

A few other Brazilian and Central American rosewood species 

are listed under CITES Appendix 2 and 3. These lower 

appendices aim to regulate trade,not prohibit it.Just this year, 

another species of Brazilian rosewood (Aniba rosaeodora),exported 

extensively as fragrant oil, was listed under CITES 

Appendix 2. Two additional species of Central American 

rosewood (D. retusa and D. stevensonii) are listed under Appendix 

3. Appendix 2, unlike Appendix 3, does require that 

the CITES authorities in the export nation determine that 

the species were legally obtained and that their export will 

not be detrimental to species survival.There seem to be few 

cases where Appendix 3 listing was sufficient, except as a 

means to Appendix 2 or higher listing. The well examined 

case-studies of big-leaf mahogany (Swietenia macrophylla) and 

ramin (Gonystylus spp.) both began as Appendix 3 species 

(which only requires unilateral listing by a habitat country) 

and were later voted in as Appendix 2 species by the CITES 

parties (Keong, 2006). 

To what degree can CITES regulations be implemented and 

enforced? The need for more officially trained import inspectors 

has been suggested numerous times. The agency 

chosen as the CITES management authority should be free 

of corruption and have experience in forest management. 

Insufficient trained staff has also hindered the ability of export 

authorities to determine whether an Appendix 2 species 

was legally obtained and non-detrimental to species survival.Range 

countries often require assistance in this respect. 

An unusually good example comes from Indonesia where biological 

data for ramin has been used in non-detriment findings 

to examine sustainability. Missing "certificates of origin" 

have been a problem for some Appendix 3 species. While 

ramin and big-leaf mahogany were listed on Appendix 3, the 

required ‘certificates of origin’ were not consistently issued 

by exporting nations;while importing countries were not always 

diligent about confirming that shipments arrived with 

such certificates (Blundell, 2007; Keong, 2006). 

2. Independent forest monitoring (IFM) 

In addition to CITES,actual improvements in forest monitoring 

on the ground are needed. A new system called independent 

forest monitoring (IFM) may be needed in order 

stop illegal logging, monitor implementation of REDD (Reducing 

Emissions from Deforestation and Forest Destruction) 

programs, restore the confidence of international donors, 

and ultimately to save Madagascar’s precious forests as 

well as attain social justice for Madagascar’s impoverished 

population. IFM has been defined as "the use of an independent 

third party that,by agreement with state authorities,provides 

an assessment of legal compliance, and observation of 

and guidance on official forest law enforcement systems" 

p. 18 (Global Witness,2005). IFM is similar in principle to unbiased 

international election observers. Local and international 

expertise is utilized, and monitoring teams operate


independently but with the consent of the host government. 

Independent forest monitors are strictly observers, law enforcement 

remains the responsibility of local officials and 

governments. 

Of course other nations have been faced with similar forest 

monitoring problems.IFM has already been used successfully 

in several African and Central American nations seeking to 

improve the effectiveness of their forest monitoring. Since it 

was first introduced in 1999, IFM has been established in 

Cameroon, Cambodia, and Honduras. Smaller scale feasibility 

and pilot studies have been conducted in Ghana, Peru, 

Mozambique, Republic of Congo, Tanzania, and Democratic 

Republic of Congo. In Cambodia and Cameroon, donor 

countries have been the impetus behind IFM.Though in Honduras,the 

incentive for IFM was domestic,and hosted by the 

Honduran Commission for Human Rights (CONADEH). 

Furones (2006) and Young (2007) review the results of IFM in 

these nations, and consider them to be "broadly positive". 

Specific examples of the impact of IFM in these nations include:documentation 

of hundreds of forest crimes,cancellation 

of logging concessions,moratoriums on logging and timber 

transport,and creation of new "forest crimes monitoring 

units" in the forestry administrations. In some cases,IFM has 

earned money for these governments by identifying violations 

which led to large fines against logging companies and 

individuals breaching the law and forest management regulations. 

3. Update IUCN Red List assessments 

The approximately 10 Madagascar rosewood species listed 

above have not had their official conservation status evaluated 

by the IUCN since 1998. At that time, all were threatened 

except for D. mollis.Five of the ten were already classified 

as ‘endangered’ then. Given the extreme logging since 

that time, it is likely that their Red List categories should be 

reassessed (IUCN, 2010). 

4. UNESCO World Heritage Sites "in danger" 

The majority of the illegally logged rosewood in Madagascar 

comes from two UNESCO World Heritage Sites: Masoala 

National Park and Marojejy National Park.Why have Masoala 

and Marojejy not been placed on the World Heritage Sites 

"In Danger" List? After all,2010 is the United Nations "International 

Year of Biodiversity". Nine national parks and seven 

other protected natural areas are currently on this danger 

list, mainly for extensive anthropogenic disturbance such as 

poaching, logging, and war. The extent of the logging damage 

in Masoala National Park, in particular, over the past 5 years, 

must rival that of some of the other national parks "in danger". 

Placing a site on the UNESCO "danger list" is not utter 

de-listing. It is a reversible process meant to draw attention 

to and attract possible resources which can alleviate the crisis.There 

are specific funds that can become available if a site 

is placed on the danger list. One can only speculate that the 

reasons for no change in status may well be political and 

practical. Perhaps it complicates matters that eight national 

parks (which include these two) comprise the single Atsinanana 

World Heritage Site Complex. Perhaps there are 

fears of triggering an even greater loss of tourism.Whatever 

the reasons may be, it is odd that UNESCO has not been 

more vocal or active in its support of these two national 

parks which are the biodiversity jewels of the Atsinanana 

World Heritage Site Complex (IUCN, 2007). 

5. DNA fingerprinting 

DNA fingerprinting has recently been used on confiscated 

ivory to determine which populations of African elephants 

were slaughtered. Similar genetic techniques would be of 

great assistance in determining which populations of Madagascar 

rosewood are being logged the most,and in identifying 

species. DNA testing has already been used to track timber, 

but not yet in Madagascar.One of the biggest methodological 

challenges is extracting DNA from the heartwood of dead 

tree trunks (e.g., rosewood stockpiles), which consist of 

dead cells with partly degraded DNA. In living trees, it is a 

routine process to obtain DNA from the cambium just beneath 

the bark or leaves or buds. Nevertheless, several new 

techniques have successfully extracted DNA from dry wood 

of ramin (Gonystylus spp.) and other woods including 1000 

year old beech (Fagus spp.) (Nielson and Kjaer, 2008). 

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Bohannon, J. 2010. Madagascar’s forests get a reprieve – But 

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le genre Dalbergia (Papilionaceae) à Madagascar et aux 

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Cochrane,M.A.;Schulze,M.D.1998.Forest fires in the Brazilian 

Amazon. Conservation Biology 12: 948-950. 

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Reuters. July 4. 

Débois, R.2009. La fièvre de l’or rouge saigne la forêt malgache. 

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J.; Moat, J. 2002. The Leguminosae of Madagascar. Royal 

Botanic Gardens, Kew, U.K. 

Furones, L. 2006. Independent forest monitoring: Improving 

forest governance and tackling illegal logging and corruption. 

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crisis. 

news.mongabay.com/2009/1215-rowan_madagascar.html. 

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Gill, C. 2010. Log Jam. Guitar Aficionado. Spring Issue. 

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using RAPDs. Heredity 83: 722-732. 

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sedfinalen.pdf. Downloaded on 20 November 2010. 

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on diademed sifakas (Propithecus diadema) at Tsinjoarivo, 

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landscapes. Ph.D. thesis, Stony Brook University, 

New York, USA. 

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3.1. www.iucnredlist.org. Downloaded on 25 May 2010. 

Keong,C.H.2006.The role of CITES in combating illegal logging: 

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504&it=document. Downloaded on 20 November 2010. 

Kett, G. 2005. Checking the reserve. Monthly from Madagascar. 

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373. In: S.M. Goodman; J.P. Benstead (eds.) The Natural 

History of Madagascar. University of Chicago Press, Chicago, 

USA. 

Michaels, S. 2009. Gibson guitars raided for alleged use of 

smuggled wood. www.guardian.co.uk/music/2009/nov/ 

20/gibson-guitars-raided. Downloaded on 20 November 

2009. 

Nielsen, L.R.; KjFr, E.D. 2008. Tracing timber from forest to 

consumer with DNA markers. Danish Ministry of the 

Environment, Forest and Nature Agency. 

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China trade facts. www.ustr.gov/countries-regions/china. 

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Patel, E.R. 2007. Logging of rare rosewood and pallisandre 

(Dalbergia spp.) within Marojejy National Park, Madagascar. 

Madagascar Conservation and Development 2(1): 

11-16. 

www.erikpatel.com/Logging_of_Rosewood_Patel_2007.pdf. 

Electronic Publication. 

Patel, E.R. In press. A tragedy with villains: Severe resurgence 

of selective rosewood logging in Marojejy National Park 

leads to temporary park closure. Lemur News. 

Patel,E.R.;Rasarely,E.;Tegtmeter,R.;Furones,N.;Fritz-Vietta, 

N.; Malan, S.; Waeber, P. In Prep. Beyond Ecological Monitoring: 

A proposal for "Independent Forest Monitoring" 

in Madagascar. Madagascar Conservation and Development. 

Randriamalala,H.;Liu,Z.In press.Bois de rose de Madagascar: 

Entre democratie et protection. Madagascar Conservation 

and Development. 

Ratsimbazafy, J. 2006. Diet composition, foraging, and feeding 

behavior in relation to habitat disturbance: Implications 

for the adaptability of ruffed lemurs (Varecia v.editorium) 

in Manombo forest, Madagascar. Pp. 403-422. In L. Gould; 

M.L.Sauther,(eds.) Lemurs:ecology and adaptation.Springer, 

New York. 

Rubel, A.; Hatchwell, M.; Mackinnon, J.; Ketterer, P. 2003. Masoala–L’oeil 

de la Forêt. Zoo Zurich. 

Schuurman, D.; Lowry, P.L. 2009. The Madagascar rosewood 

massacre. Madagascar Conservation and Development 

4(2): 98-102. www.mwc-info.net. Electronic Publication. 

Schuurman,D.2009.Illegal logging of rosewood in the rainforests 

of northeast Madagascar. TRAFFIC Bulletin 22(2): 

49. 

Stasse, A. 2002. La Filière Bois de Rose. Région d’Antalaha – 

Nord-est de Madagascar. Thèse de mastère non publiée, 

Université de Montpellier, France. 

Stephenson, P.J. 1993. The small mammal fauna of Reserve 

Speciale d’Analamazaotra, Madagascar: The effects of 

human disturbance on endemic species diversity. Biodiversity 

and Conservation 2: 603-615. 

Wasser, S.; Poole, J.; Lee, P.; Lindsay, K.; Dobson, A.; Hart, J.; 

Douglas-Hamilton, I.; Wittemyer, G.; Granli, P.; Morgan, B.; 

Gunn,J.;Alberts,S.;Beyers,R.;Chiyo,P.;Croze,H.;Estes,R.; 

Gobush,K.;Joram,P.;Kikoti,A.;Kingdon,J.;King,L.;Macdonald, 

D.; Moss, C.; Mutayoba, B.; Njumbi, S.; Omondi, P.; 

Nowak, K. 2010. Elephants, ivory, and trade. Science 327 

(5971): 1331-1332. 

Wilmé,L.;Schuurman,D.;Lowry II,P.P.In Press.A forest counterpart 

fund:Madagascar’s wounded forests can erase the 

debt owed to them while securing their future, with support 

from the citizens of Madagascar. Lemur News. 

Wilmé, L.;Schuurman,D.; Lowry II, P.P.; Raven, P.H. 2009. Precious 

trees pay off – but who pays? Poster prepared for 

the World Forestry Congress in Buenos Aires,Argentina. 

www.mwc-info.net/en/services/Journal_PDF%27s/ 

Issue4-2/MCD_2009_vol4_iss2_rosewood_massacre_ 

Supplementary_Material.pdf. Downloaded on 23 October 

2009. 

Young, D. 2007. Independent forest monitoring: Seven years 

on. International Forestry Review 9(1): 563-574. 

Rosewood logging photos 

Photographer Toby Smith: 

www.telegraph.co.uk/culture/photography/7625511/ 

Madagascar-undercover-slideshow.html 

Photographer Chris Maluszynsk: 

www.photoshelter.com/c/moment/gallery/ Rosewood-loggingin-Madagasar-by-Chris- 

Maluszynski/ G0000JWMAJa78LJ0/ 

Rosewood logging films 

Dan Rather Reports:Treasure Island.Episode 437.A detailed 

investigation of the impact of the recent political crisis in 

Madagascar on the unique biodiversity of this island continent. 

Filmed in high-definition, active rosewood logging 

camps are shown. The impact of such habitat disturbance on 

the silky sifaka and the World Heritage Sites of Marojejy NP 

and Masoala NP are discussed. The debates surrounding the 

Ambatovy nickel mine adjacent to Andasibe-Mantadia NP 

are also discussed. The mine may be endangering one of the 

rarest animals on earth,the greater bamboo lemur (Prolemur 

simus) which is being protected there by the NGO Mitsinjo. 

Aired on HD-NET cable television November 2009. Purchasable 

and downloadable on I-Tunes in the United States. 

DVDs can be purchased online: 

hdnet-store.stores.yahoo.net/danrare437.html 

Sample Clip 1: 

www.facebook.com/video/video.php?v= 600388589544 

Sample Clip 2: www.youtube.com/watch?v= dEi-yRlJ-mk 

Carte Blanche: Madagascar (Part 1 and Part 2). Two short 

films examining illegal rosewood logging in Madagascar and 

the impact on the critically endangered silky sifaka. They 

were produced by Neil Shaw and commissioned and funded 

by Carte Blanche which is one of the most respected television 

news programs in the Southern Hemisphere. Aired on 

South African Television in April,2010,and streams freely online 

here: 

Carte Blanche: Madagascar Part 1: 

beta.mnet.co.za/carteblanche/Article.aspx?Id= 3919&ShowId=1 

Carte Blanche: Madagascar Part 2: 

beta.mnet.co.za/mnetvideo/browseVideo.aspx?vid=25570 

506: Bois de Rose. A Documentary Film by Joseph Areddy. 

2003. RSI, Comano/Signe, Genve/GAP, Antananarivo. 

Rosewood logging videos 

Madagascar Rainforest Massacre (English): 

www.youtube.com/watch?v=FzWNPHBRrAc 

Madagascar Rainforest Massacre (French): 

www.youtube.com/watch?v=KtjmFWpGNKs&feature=related 

Madagascar Rainforest Massacre (Malagasy): 

www.youtube.com/watch?v=rHYYhhLHeQw&feature=related


Global Witness – Environmental Investigation Agency - Illegal 

logging in Madagascar – Part 1 

www.youtube.com/watch?v=T1hPviSbRcU 



www.youtube.com/watch?v=LBtsNBpWW0E 



www.youtube.com/watch?v=payUUJed0dc 



www.youtube.com/watch?v=lm6a6Hrat3o 

Rosewood logging radio programs 

BBC World Service – Africa. September 17, 2009. 

www.bbc.co.uk/worldservice/africa/2009/09/090917_ 

madge_rosewood2.shtml 

Ongoing threats to lemurs and their habitat 

inside the Sahamalaza - Iles Radama 

National Park 

Melanie Seiler 1,2, Guy H. Randriatahina 3, Christoph 

Schwitzer 1* 

1Bristol Conservation and Science Foundation, Bristol Zoo 

Gardens, Clifton, Bristol BS8 3HA, UK 

2University of Bristol, School of Biological Sciences, Woodland 

Road, Bristol BS8 1UG, UK 

3Association Européenne pour l’Etude et la Conservation 

des Lémuriens (AEECL), Lot: IVH 169 N Ambohimanandray, 

Ambohimanarina, Antananarivo 101, Madagascar 

*Corresponding author: cschwitzer@bcsf.org.uk 

The Sahamalaza - Iles Radama National Park,officially inaugurated 

in July 2007 and managed by Madagascar National 

Parks (MNP), includes both marine and terrestrial ecosystems 

and is the first park that was created under the 

"Programme Environnemental III" of the Malagasy government 

and the World Bank. In addition to the few remaining 

forest fragments of the Southern Sambirano ecoregion, the 

park is home to extensive mangrove forests, which harbour 

their own highly endangered fauna, and also includes offshore 

coral reefs. In 2003, researchers from the Cologne 

Zoo, funded by AEECL, undertook an expedition to Sahamalaza 

to explore the opportunities for the establishment of 

a permanent field station in order to study and protect the 

Critically Endangered blue-eyed black lemur (Eulemur flavifrons) 

and its habitat.In 2004 and 2005,the field station in the 

Ankarafa Forest became reality (Schwitzer et al.,2006),and it 

has since been used by both European and Malagasy scientists 

as a basis for research on E. flavifrons and other lemur 

species, especially the Sahamalaza sportive lemur (Lepilemur 

sahamalazensis) and the northern giant mouse lemur (Mirza 

zaza),occurring on the Sahamalaza Peninsula (Schwitzer and 

Randriatahina, 2009). 

Sahamalaza - Iles Radama National Park lies within a transition 

zone between the Sambirano region in the north and 

the western dry deciduous forest region in the south, harbouring 

semi-humid forests with tree heights of up to 30m 

(Schwitzer et al.,2006).The forests include a mixture of plant 

species typical of both domains (Birkinshaw, 2004), and the 

remaining primary and secondary forest fragments vary in 

their degree of degradation. There are no larger connected 

areas of intact primary forest left on the Sahamalaza Penin- 

sula, and the remaining fragments all show some degree of 

anthropogenic disturbance and/or edge effects (Schwitzer et 

al., 2007). The forests and forest fragments are separated by 

grass savannah and shrubs. Sahamalaza is the only protected 

area that harbours the blue-eyed black lemur,the Sahamalaza 

sportive lemur and the northern giant mouse lemur. Other 

lemur species in the park include the aye-aye (Daubentonia 

madagascariensis), the western bamboo lemur (Hapalemur 

occidentalis),and an as yet unidentified species of dwarf lemur 

(Cheirogaleus spec.). 

The remaining forest of the Sahamalaza Peninsula and its 

unique fauna are in grave danger of disappearing.The habitat 

is already extremely degraded, nonetheless bush fires and 

tree-felling are activities that are routinely pursued and accepted 

within the local society (Ruperti et al., 2008). During 

the first field season of a study on the impact of habitat degradation 

and fragmentation on the ecology and behaviour of 

the Sahamalaza Peninsula sportive lemur (Lepilemur sahamalazensis), 

conducted by MS in 2009, local people from the 

villages surrounding the protected area were found logging 

trees in the already small forest fragments almost on a daily 

basis. Logging activities mainly occurred in forest fragments 

where no researchers had been present in previous years. 

During walks through different forest fragments, in addition 

to large numbers of logged trees, two places where trees 

were processed for further use were found. Trees were 

felled mainly in the early morning hours,on the one hand because 

of the high temperatures later in the day, on the other 

hand probably because of the assumption that the researchers 

started observing animals later in the day and therefore 

would not realise the illegal logging activities. Nonetheless, 

trees were sometimes also felled in the afternoons. Because 

locals immediately fled when becoming aware of researchers’ 

presence, we believe that the presence of researchers 

and/or field guides, park authorities or park rangers is a crucial 

factor in stopping illegal logging in the remaining fragments.For 

the next field season (2010) we therefore plan to 

expand the observations of Lepilemur to other, not yet used 

forest fragments to help prevent their destruction. Of 

course this cannot be a long-term solution to this problem. 

The presence of park rangers and further environmental education 

of the local people will thus be extremely important 

to save the Sahamalaza forests from further degradation. 

About five times between August and October 2009, fires 

occurred near the Ankarafa field station, three times in the 

savannah and twice in the forest itself. After having extin- 

Fig. 1: Lepilemur sahamalazensis 

poached 

and roasted by locals in 

Sahamalaza - Iles Radama 

National Park“.


guished these fires it became obvious that they had all 

started right beside the fire breaks that are frequently used 

as paths by people on their way between villages. The Ankarafa 

field guides, all of them locals from the surrounding villages, 

assumed that the fires were set by villagers to show 

their dissatisfaction with the recently established national 

park that prohibits the use of the forests for collecting building 

material for their dwellings. As we followed the smoke 

that was coming from another fire, we found an area inside 

one of the core zones of the national park that was inhabited 

by a young couple. They harvested a rice field and regularly 

burned undergrowth around it. Additionally, they kept cattle 

and goats and had built 2 houses at this site,one for the cattle 

and one for themselves. As we talked to them, they claimed 

that they were allowed to stay on this site and that MNP had 

sold this part of the forest to them.They affirmed that,if they 

set fire on this site, they would keep an eye on it and would 

prevent the fire from expanding into the forest. Unfortunately 

this was not the case,however,as we later observed a 

fire around this site without anyone near it. Overall, it 

seemed that there were various people living inside the national 

park on permits given to them by what they claimed to 

have been MNP agents;we were told that there was a map of 

the park showing all the "excluded" areas available for housing 

and agriculture, which could be seen in the village of 

Marovato. If that was indeed the case (we did not have the 

opportunity to verify the information),it would be a massive 

problem for protecting Sahamalaza’s unique wildlife and forests. 

If people claiming to be MNP staff illegally sold permits 

for activities inside the national park, the destruction of the 

small forest fragments will continue rapidly. 

Another big problem comes with cattle; every day zebu cattle 

were observed in all forest fragments and on the savannah 

in Ankarafa, as people from nearby villages let their cattle 

roam freely.The abundance of zebu themselves and their excrements 

indicated that they frequently used the forest fragments 

as grazing grounds, especially those with remaining 

primary forest parts. When zebu were grazing in the forest 

rather than on the savannah, their movements were accompanied 

by crashing and breaking sounds;they were undoubtedly 

hindering the growth of many saplings,if not eating them. 

This is an additional threat to the forest fragments, and furthermore, 

the abundance of the excrements of local zebu 

has been found to negatively correlate with the density of L. 

sahamalazensis (Ruperti, 2007). Additionally, the introduced 

bush pig is responsible for considerable habitat destruction 

due to digging up large areas, thus hindering the growth of 

saplings. Unfortunately, the bush pig is reproducing wildly as 

it is regarded as fady (taboo) by the local people and therefore 

not hunted. 

Not only the activities of local people seem to be a threat to 

the endangered wildlife on the Sahamalaza Peninsula.One of 

the Ankarafa field guides encountered a foreigner,probably a 

resident living in Madagascar (since he spoke Malagasy fluently), 

with a 4x4 car and two local guides about 1 km from 

the researchers’ camp. These people had set up a tent and 

told the Ankarafa field guide that they were visiting all villages 

on the Sahamalaza Peninsula to look for fish. As we checked 

their camp site the next day, the three men were gone, but 

signs of a fire,logged branches and feathers of a harrier hawk 

were found,indicating that they had caught and killed this endangered 

bird of prey. We wrote a report about this event 

and handed it over, together with feathers of the bird, to 

MNP in Maromandia. However, as long as there are no signs, 

borders or fences indicating the national park area and its 

restrictions, these problems will continue. 

The ongoing political crisis is a further big concern that hinders 

the effective protection not only of the biodiversity of 

the Sahamalaza Peninsula, but of Madagascar and its national 

parks system as a whole. Only 10 years ago, Madagascar was 

notorious for its environmental degradation and deforestation, 

but that began to change in 2003 when then President 

Marc Ravalomanana, working with international conservation 

organizations and local groups, set aside 10 % of the 

country’s surface area as national parks and started supporting 

ecotourism, which slowed deforestation and helped to 

safeguard biodiversity.After the political events in early 2009 

that saw the ousting of the President and the installation of a 

transitional government, the majority of donor funds, which 

provided half the government’s annual budget, have been 

withdrawn, leading to major funding gaps that have affected 

protected areas and their management. There currently is 

almost no money to employ park rangers or to implement 

other measures to protect the forests inside Madagascar’s 

national parks, and forest degradation is going on without 

noticeable resistance from the relevant authorities. Despite 

the political crisis that affects most of the social and environmental 

activities of numerous NGOs, AEECL is still carrying 

out its research activities and support to the villagers surrounding 

the Sahamalaza - Iles Radama National Park. Since 

the establishment of the protected area in 2007, AEECL has 

been conducting, besides its research programme, different 

projects that aim to reduce the excessive environmental exploitation 

inside and around the park.As the major activity of 

the local population surrounding the Sahamalaza Peninsula 

National Park is rice-growing,every year AEECL organizes a 

rice-growing training course and rice-growing competition, 

using modern techniques in order to increase yield per ha 

and to decrease the use of slash and burn agriculture.To stop 

the ongoing overexploitation of the environment, environmental 

education is another important part of AEECL´s 

work.As many villages in Sahamalaza are unable to pay teachers, 

AEECL subsidizes teachers’ salaries to ensure the primary 

education of the local children. Additionally, leaflets 

about the Sahamalaza biodiversity and its importance are 

distributed. They inform and educate villagers about the importance 

of lemurs and other species for their forest ecosystem. 

To minimise bush fires and to protect the forest against 

uncontrolled fires, AEECL organizes firebreak programs 

around the Ankarafa Forest, close to the research camp, 

where during three days, hundreds of local people remove 

the grasses on a 7m wide strip around the forest fragments. 

Furthermore, several reforestation campaigns have been 

conducted, where villagers, including many teachers and 

their pupils,have planted trees around their villages with the 

help of AEECL. 

Because of all the factors described here, the protection of 

Sahamalaza’s unique flora and fauna continues to be a major 

challenge that has to be faced by the local human population 

with the help of Madagascar National Parks and foreign partners. 

Two essential parts of AEECL’s efforts to help meeting 

this challenge are to stimulate further scientific study of 

endangered lemurs and other wildlife at its research station 

in the Ankarafa Forest, especially by Malagasy students, and 

to enable the local human population around the Sahamalaza 

- Iles Radama National Park to sustainably use their natural 

resources. 

Acknowledgements 

We would like to thank Madagascar National Parks (MNP), 

especially the director of Sahamalaza - Iles Radama National 

Park, M. ISAIA Raymond, for their continuing collaboration.


Thank you also to the DGEF and CAFF/CORE for granting 

us research permits for our work in Sahamalaza,and to Prof. 

RABARIVOLA Clément for his ongoing help. Tantely Ralantoharijaona 

and Bronwen Daniel,along with all Ankarafa field 

guides, contributed substantially to fighting forest fires and 

other environmental threats in Ankarafa in 2009. MS was 

funded by Bristol Conservation and Science Foundation, 

AEECL, Conservation International Primate Action Fund, 

Margot Marsh Biodiversity Foundation, Mohamed bin Zayed 

Species Conservation Fund,International Primatological Society 

and Christian-Vogel-Fonds. 

References 

Birkinshaw, C.R. 2004. Priority areas for plant conservation. 

Ravintsara 2(1): 14-15. 

Ruperti, F. 2007. Population density and habitat preferences 

of the Sahamalaza sportive lemur (Lepilemur sahamalazensis) 

at the Ankarafa research site, NW Madagascar. 

Unpublished MSc thesis, Oxford Brookes University, UK. 

82 p. 

Ruperti,F.;Smith,J.;Ratovonasy,L.;Thorn,J.2008.Sahamalaza 

Conservation Action Plan (SCAP).Unpublished report to 

the Association Européenne pour l’Etude et la Conservation 

des Lémuriens (AEECL). 17 p. 

Schwitzer, C.; Randriatahina, G.H. 2009. AEECL: Update on 

activities. Lemur News 14: 11-12. 

Schwitzer, N.; Randriatahina, G.H.; Kaumanns, W.; Hoffmeister, 

D.; Schwitzer, C. 2007. Habitat utilization of blue-eyed 

black lemurs,Eulemur macaco flavifrons (Gray,1867),in primary 

and altered forest fragments.Primate Conservation 

22: 79-87. 

Schwitzer, C.; Schwitzer, N.; Randriatahina, G.H.; Rabarivola, 

C.; Kaumanns, W. 2006. "Programme Sahamalaza": New 

perspectives for the in situ and ex situ study and conservation 

of the blue-eyed black lemur (Eulemur macaco flavifrons) 

in a fragmented habitat.Pp.135-149.In:C.Schwitzer; 

S.Brandt;O.Ramilijaona;M.Rakotomalala Razanahoera;D. 

Ackermand; T. Razakamanana; J. U. Ganzhorn (eds.). Proceedings 

of the German-Malagasy Research Cooperation 

in Life and Earth Sciences. Berlin: Concept Verlag. 

News and Announcements 

Madagascar conservationist wins international 

environmental prize 

Mr Rabary Desiré has been awarded the 2010 Seacology 

Prize (www.seacology.org/prize/index.htm) for his his tireless 

efforts to further forest conservation in northeastern Madagascar.Mr 

Desiré will receive the US$10,000 Prize on October 

7, 2010 at a ceremony in Berkeley, California. 

Rabary Desiré is recognized by many as a major conservation 

leader in northeastern Madagascar, and is a highlysought-after 

research and eco-tourism guide. With the money 

he makes from guiding,he buys forested land in order to 

protect it.Years of work have finally culminated in the establishment 

of his own small private nature reserve called 

Antanetiambo (antanetiambo.marojejy.com/Intro_e.htm), 

which means "on the high hill". It is perhaps the only reserve 

in northern Madagascar that has been entirely created from 

start to finish by a single local resident. 

According to Mr Desiré, "I am very happy to receive this 

award and I feel very lucky for myself and Madagascar. After 

many years of hard work and political instability,finally we are 

having some local conservation success. I plan to use these 

funds for such projects as reforestation, developing tourist 

Fig. 1: Rabary Desiré next to the sign for the Antanetiambo 

Nature Reserve he created. 

infrastructure and purchasing the land around Antanetiambo 

Nature Reserve to increase the size of the reserve and the 

amount of protected land in this region. This award will help 

preserve the precious biodiversity and high endemism of 

Madagascar,as well as fight the ongoing battle against massive 

deforestation and possible extinction of many beloved species... 

Thanks Seacology for giving me this prize. The whole 

region will never forget it." 

Read the full press release: 

www.seacology.org/news/display.cfm?id=4238 

Célébration du quinzième anniversaire 

du GERP (1994-2009) 

Jonah Ratsimbazafy*, Rose Marie Randrianarison, 

Muriel Nirina Maeder 

GERP, 34, Cité des Professeurs, Antananarivo 101, 

Madagascar 

*Corresponding author: gerp@wanadoo.mg 

Quinze ans se sont écoulés depuis la création, en 1994, de la 

Société de Primatologie malgache ou Groupe d’Etude et de 

Recherche sur les Primates de Madagascar (GERP). Elle fut 

fondée par dix Primatologues dont le Professeur Berthe 

Rakotosamimanana qui occupait à la fois le poste de Secrétaire 

Général du GERP et le Co-éditeur de la revue Lemur 

News jusqu’à sa disparition en 2005. De son vivant, elle 

désirait ardemment passer le flambeau au Docteur Jonah 

Ratsimbazafy pour le poste de Secrétaire Général du GERP 

qui, en 2006, a été mandaté à l’unanimité par les membres 

nationaux et internationaux du GERP au titre de Leader du 

GERP. 

L’Association compte aujourd’hui 169 membres et 20 d’entre 

eux sont de nationalité étrangère. La multidisciplinarité 

des membres du groupe (Primatologues, Anthropologues, 

Paléontologues, Ornithologues, Herpétologues, Spécialistes 

de Micromammifères et Mammifères, Parasitologistes, Botanistes, 

Géographes, Vétérinaires, Agro-forestiers, Biochimis-


tes, Dessinateur, Financiers) apporte une importante potentialité 

dans l’accomplissement de la mission du GERP: transférer 

les compétences nécessaires à la préservation de la 

biodiversité pour les générations futures. Par ailleurs, les actions 

du GERP comprennent également la formation des 

pépinières de Primatologues, la mise en œuvre du plan de 

conservation des lémuriens, la contribution à l’amélioration 

des activités génératrices de revenu des communautés de 

base liées à la conservation,sans oublier l’éducation environnementale 

de la population cible. 

En 2007,l’attribution par le GERP du nom de Microcebus macarthurii 

à une nouvelle espèce découverte dans la forêt de 

Makira représentait un témoignage de reconnaissance au 

dévouement de la Fondation MacArthur. De plus, le GERP a 

depuis 2008 officiellement été mandaté par le MEFT/DGEF/ 

DSAP comme Gestionnaire de la forêt de Maromizaha,pour 

que cette dernière devienne une Nouvelle Aire Protégée 

(NAP).Plus récemment encore,en février 2010,le prix "lifetime" 

décerné par l’IPS a été attribué à un membre scientifique 

du GERP en la personne du Docteur Alison Jolly. 

A l’occasion de son quinzième anniversaire, le GERP aura 

l’honneur d’organiser une conférence scientifique sur les 

lémuriens, à Antananarivo en novembre 2010. 

Conservation International’s Primate 

Action Fund:Projects funded March 2009 

to March 2010 

Anthony Rylands 

Conservation International, 2011 Crystal Drive, Suite 500, 

Arlington, VA 22202, USA, a.rylands@conservation.org 

Conservation International’s Primate Action Fund awards 

small grants (up to $5,000) to support projects and initiatives 

promoting the conservation of primates worldwide,focusing 

on Critically Endangered and Endangered species in their 

natural habitats (and most especially those included in the biennial 

listing of the World’s 25 Most Endangered Primates). 

Projects should contribute to at least one of the following 

themes: (1) enhancement of scientific understanding/knowledge 

of the target species/ecosystem; (2) improved protection 

of a key species, habitat, or a reserved area; (3) demonstration 

of economic benefits achieved through conservation 

of a species and its habitat,as compared to its loss;(4) increased 

public awareness or educational impact resulting 

from the project in question; (5) improved local capacity to 

carry out future conservation efforts through training or 

practical experience obtained through project participation; 

and (6) modification of inappropriate policies or legislation 

that previously led to species or habitat decline. Awards are 

given most frequently for population and distribution surveys,and 

ecological and behavioral studies pertinent to conservation 

initiatives for threatened species. Grants are also 

given that support genetic and taxonomic studies, publications, 

workshops for action plans and suchlike, and primate 

field courses. Some awards are given to help primate habitat-country 

primatologists attend the biennial congresses of 

the International Primatological Society. The fund does not 

support participation in academic courses. 

The Primate Action Fund comes from an annual award to 

Conservation International, Arlington, Virginia, USA, made 

by the Margot Marsh Biodiversity Foundation. It is managed 

jointly by Ms Ella Outlaw and Dr Anthony B.Rylands,both of 

CI’s Office of the President. Guidelines for application can be 

obtained by writing to Anthony Rylands (see Funding and 

Training section in this issue). 

Five grants were awarded to benefit lemur conservation in 

the March 2009 – March 2010 funding cycle. They were as 

follows: (1) Halting politically-induced deforestation in the 

short term to preserve the unique primate community of 

Tsinjoarivo, eastern central Madagascar–Mitchell T. Irwin, 

Fanomezantsoa, Jean-Luc Raharison and Marina Blanco; (2) 

Rapid survey and assessment of the northern sportive lemur, 

Lepilemur septentrionalis, in the Sahafary Region, Madagascar–Edward 

Louis Jr, Jean 

Ranaivoarisoa, John Zaonarivelo and Steig Johnson; (3) Support 

for the publication of the IUCN/SSC Primate Specialist 

Group newsletter and journal Lemur News, volume 14–Jörg 

U. Ganzhorn and Christoph Schwitzer; (4) Student training 

course “Field Methods in the Study of Primate Behavior and 

Ecology”,Kirindy forest,2010–Melanie Dammhahn,Peter M. 

Kappeler, Claudia Fichtel, Cornelia Kraus and Rodin Rasoloarison; 

and (5) Comparison of habitat requirements of the 

Data Deficient northern giant mouse lemur (Mirza zaza) in 

two differently degraded habitats, in Sahamalaza,northwestern 

Madagascar–Johanna Rode and Christoph Schwitzer. 

International Technical Meeting on Prolemur 

simus, 26-28 January 2010, Antananarivo, 

Madagascar 

The greater bamboo lemur Prolemur simus has long been considered 

to be one of the rarest primate species in the world. 

Up to 2007 only 60 individuals were known from the wild, 

and another 22 were in captivity (Wright et al.,2008;Primate 

Conservation 23: 5-17). Once widespread across Madagascar, 

more recent confirmed sightings were exclusively from 

south-eastern Madagascar, which led to the assumption that 

the species was extinct on the rest of the island. In 2008, 

Dolch et al. (Lemur News 13: 14-17) rediscovered P. simus in 

the Torotorofotsy wetlands, north of the Mangoro River. 

Since then, several extensive surveys have been conducted 

north and south of the Mangoro, and evidence of greater 

bamboo lemurs was found at several sites in the Ankeniheny-Zahamena 

Corridor, in the central region of the eastern 

rainforest (King and Chamberlan, 2010; Oryx 44: 167). 

In the context of developing a conservation action plan for 

the greater bamboo lemur, the Madagascar Fauna group organised, 

from 26-28 January 2010 at the motel d’Antananarivo, 

Anosy, an international technical meeting with the 

theme "Conservation of the critically endangered greater 

bamboo lemur Prolemur simus:What we know now,what we 

need to know and potential conservation strategies".Several 

members of the PSG contributed to this. 

The objectives of the meeting were 1) to share information 

about the current situation of the various groups/populations 

of Prolemur simus in the wild and in captivity; 2) to discuss 

the threats, the solutions and the conservation strategies 

for three groups - north of the Mangoro River (Torotorofotsy 

and the Ankeniheny-Zahamena corridor CAZ), 

south of the Mangoro River (south-east and the Fandriana- 

Vondrozo corridor COFFAV), and in captivity (Madagascar 

and Europe); and 3) to make a plan, short to long term, to 

move towards a conservation action plan for the species. 

With 28 participants,the meeting was well attended.Presentations 

were given by researchers studying P. simus in the wild 

and in captivity, representatives from the Ambatovy, Madagascar 

National Parks, the University of Antananarivo and


conservation NGOs.While other potential P.simus sites still 

need to be explored, results from the most recent surveys 

suggest the total estimated size of the known population is 

between 221-346 individuals. Another 20 individuals are 

housed in one Malagasy (Parc Ivoloina Zoo) and several European 

zoos and and managed under the umbrella of an EEP. 

The following recommendations for the conservation of P. 

simus came out of the meeting: 

We need to achieve official/formal protection for all currently 

known P. simus habitat (using whatever status is appropriate 

to the site); 

Animals of the northern and southern populations (wild 

or captive) should not be mixed until the taxonomic situation 

is clarified; 

Faecal samples should be collected from all sites using a 

standard protocol (meeting participants agree to collaborate 

to achieve this); 

When animals are caught/immobilised the opportunity 

should be used to maximise the collection of samples; 

Bamboo plot data should be collected from all sites using 

a standard protocol (meeting participants agree to collaborate 

to achieve this); 

A health screening protocol should be applied whenever 

the opportunity arises; 

Sites in the Ankeniheny-Zahamena Corridor (CAZ) recently 

shown to harbour P. simus should be evaluated by 

2011 at the latest to assess population size; 

Maromiza and Lakato need to be evaluated for the presence 

of P. simus, and protected to ensure connectivity; 

We agree that assuring connectivity between Torotorofotsy 

and CAZ is a high priority,and that the area needs an 

integrated conservation plan involving all stakeholders – 

CI to drive the process under supervision of the Alaotra- 

Mangoro Forestry Commission; 

It is important to make P. simus a priority (conservation 

target) for the CAZ; 

Improved communication using a mailing list will be established, 

the "Prolemur Conservation Working Group"; 

There are other sites that need to be surveyed for P.simus 

(a list of sites has already been identified); 

Maximising connectivity between P. simus sites is important; 

Local communities should be directly involved in P. simus 

conservation wherever possible; 

In case of a crisis scenario involving potential translocation, 

a technical strategy is needed consistent with IUCN 

guidelines; 

The EEP-Ivoloina exchange of P. simus is important to 

strengthen the global captive population; 

For the time being,it is not recommended that additional 

wild P. simus be added to the global captive programme, 

except in emergency; 

In the case of emergency, we recommend that animals go 

to PBZT if upgraded facilities have been installed; if not 

then they should go to Ivoloina; 

Based on the development of the global captive programme, 

integrated (metapopulation) management of P. simus 

should be considered; 

Another technical meeting should be held in January 

2011. 

The workshop was financially and technically supported by 

the Madagascar Fauna Group with additional contributions 

from Conservation International Madagascar. 

Lemur presentations at the 23rd Congress 

of the International Primatological 

Society, Kyoto, Japan 


GERP, 34, Cité des Professeurs, Antananarivo 101, Madagascar, 

gerp@wanadoo.mg 

The 23rd Congress of the International Primatological Society 

(IPS) was held in Kyoto (Yoshida Main Campus),Japan on 

12th-18th September, 2010. This congress brought together 

more than 1,000 delegates from 56 countries. Twenty-eight 

talks and three posters were presented on lemur studies 

during that congress. 

I am also pleased to share with you the good news that the 

winner of the 2010 IPS Lifetime Award is Professor Alison 

Jolly who is an active member of GERP (Groupe d’Etude et 

de Recherche sur les Primates de Madagascar). The lemur 

lady,Prof.Jolly,has devoted her life to the conservation of the 

world’s primates. Education is one of the main activities that 

she never stops to discuss,as she found that the only chance 

to save the endangered lemurs of Madagascar is to provide 

the Malagasy children with tools enabling them to learn and 

love the creatures that exist in their backyards. 

I hope that even more lemur researchers will present the 

results of their work at the 24th IPS Congress in Mexico. 

Short Communications 

Preliminary conservation status assessment 

for the Data Deficient northern 

giant mouse lemur Mirza zaza 

Eva Johanna Rode 1,2, K. Anne-Isola Nekaris 2, Christoph 

Schwitzer 1* 

1Bristol Conservation and Science Foundation, c/o Bristol 

Zoo Gardens, Clifton, Bristol BS8 3HA, UK 

2Nocturnal Primate Research Group, School of Social Sciences 

and Law, Oxford Brookes University, OX3 0BP, UK 

*Corresponding author: cschwitzer@bcsf.org.uk 

Madagascar is one of the world’s most important biodiversity 

hotspots, underpinned by its large proportion of endemic 

species and high rates of deforestation.During the last 

decade, species diversity of Madagascar’s endemic lemurs 

has increased dramatically due to new discoveries and taxonomic 

revisions. This has resulted in the unusual situation of 

45 % of all Malagasy primate species being Red-Listed as Data 

Deficient (DD) by the IUCN. This is by far the highest such 

figure for any primate habitat country (by comparison, 13 % 

of all primates and 15 % of all mammals are Red-Listed as 

DD).The lack of species-specific knowledge makes it impossible 

to design effective conservation measures targeting 

these taxa. To help assign a conservation status to the DD 

northern giant mouse lemur Mirza zaza, described in 2005 

due to distinctive features in morphology, behaviour and 

genetics (Kappeler et al., 2005; Primate Report, 71, 3-26), we 

examined space requirements and group size of this small 

nocturnal lemur species during a three-month study (May- 

July 2010) and extrapolated our results to the taxon’s area of


occupancy in order to estimate the size of its remaining 

population. 

Mirza zaza lives in dry forests of north-western Madagascar, 

one of the fastest declining habitats of the island, with a decrease 

in forest cover of 40 % from 1975 to 2000.The area of 

occurrence of the species is limited by the Maeverano River 

in the south and the Mahavavy River in the north.Combining 

forest cover data collected by the Madagascar Vegetation 

Mapping Project (www.vegmad.org) with data on group home 

range size and group size calculated from our study and additional 

literature, we calculated minimum and maximum estimates 

of total remaining population size. Since data for the 

Madagascar Vegetation Mapping Project were collected several 

years ago,we lowered the estimate of total available habitat 

according to the estimated annual rate of decline.Habitat 

decline may have accelerated since the onset of the political 

crisis in Madagascar in early 2009, which is not yet reflected 

in our estimates.Since a previous survey failed to detect Mirza 

zaza in several regions within the species’ area of occurrence, 

we applied different estimates of the percentage of 

suitable habitat actually inhabited by the species.Our calculations 

yielded the following estimates: 

Maximum estimate: The total area covered in dry forest 

within the area of occupancy of M. zaza is approximately 

1,650 km 2 . Assuming an occupancy of 80 %, group home 

ranges of 2 ha and group size of 4 individuals there would 

be max. 177,500 animals left in total. 

Minimum estimate: In order to reflect the long-term survival 

of the species in a very fragmented area, only fragments 

< 1km 2 and smaller fragments closer than 500 m to 

other, larger fragments (total area: 955 km 2 ) were considered.We 

chose 1 km 2 to allow a minimum viable population 

of 250 animals.If only 30 % of the habitat is inhabited, 

animals use group home ranges of 4 ha and live in groups 

of on average 2.3 animals, this leads to an estimate of 

16,500 individuals left. 

Mirza zaza should be assessed as Vulnerable (VU B2ab) since 

its area of occupancy in both estimates is lower than 2,000 

km 2. With several sites within the species’ distribution area 

found to be unoccupied, the remaining habitat being extremely 

fragmented with the smallest fragments unsuitable 

to support a viable population, and habitat vanishing quickly, 

M. zaza may become Endangered (EN B2ab) in the near 

future if its area of occupancy shrinks below 500 km 2. 

Our preliminary conservation status assessment used the 

best available data for Mirza zaza. More accurate estimates 

will be possible if more data become available, especially on 

percentage of occupancy. This method might be applied to 

other DD lemur species in order to gain initial assessments 

of their conservation status. 

An observation of the hairy-eared dwarf 

lemur, Allocebus trichotis, in the Lakato 

region, eastern Madagascar 

Erwan Lagadec 1, Steven M. Goodman 2* 

1Centre de Recherche et de Veille sur les maladies émergentes 

dans l’Océan Indien (CRVOI),GIP Cyclotron Réunion 

Océan Indien, 2 rue Maxime Rivière, 97492 Sainte Clotilde, 

Ile de la Réunion, France, and Centre National de la Recherche 

Scientifique, UMR5557 Ecologie Microbienne, Bât A. 

Forel, 43 bd du 11 novembre 1918, 69622 Villeurbanne 

CEDEX, France 

2Field Museum of Natural History, 1400 South Lake Shore 

Drive, Chicago, Illinois 60605, USA, and Association Vahatra, 

BP 3972, Antananarivo 101, Madagascar 

*Corresponding author: sgoodman@vahatra.mg 

Although a few decades ago the hairy-eared dwarf lemur 

(Allocebus trichotis) was considered "unquestionably the rarest 

of surviving lemurs" (Tattersall,1982,p.131),more recent 

field work has found this species to be widely distributed 

across portions of the eastern humid forests of Madagascar 

(e.g., Meier and Albignac, 1991; Rakotoarison, 1998; Schütz 

and Goodman, 1998; Goodman and Raselimanana, 2002). 

Since more than a decade, there have been numerous records 

of this species from the central portion of the eastern 

humid forests,and information is now available on aspects of 

its ecology and natural history (e.g.Rakotoarison et al.,1997; 

Garbutt,2000;Biebouw,2009;Ralison,2010).Here we add an 

additional record from the region of Lakato, an area from 

where this species had not been previously recorded. 

From 22-28 October 2010 we were part of a research group 

that conducted a biological inventory of a forest block in the 

Lakato area and in the southern portion of the Zahamena- 

Ankeniheny forest corridor. The specific study site was centered 

at the following locality,which served as the base camp 

for all inventory activities: Province de Toamasina, Alaotra- 

Mangoro Region, 14.5 km SW of Andasibe (Périnet) village, 

Ampasipotsy-Anivonimaro/Ambalafary Forest, 19°02’38"S, 

48°20’55"E, 995 m elevation. 

During a nocturnal survey on 28 October 2010, the first author 

observed and photographed an individual of A. trichotis. 

The animal appeared not to be accompanied by any conspecifics. 

The distinctive ear-tufts, characteristic of Allocebus, 

were clearly visible (Fig.1).The lemur was observed at 21h58 

and for about five minutes.The site was in partially disturbed 

lower montane forest, about 200 m away from the research 

camp and within a few meters of the dirt road connecting the 

RN 2 (connecting Antananarivo-Toamasina) and the village 

of Lakato. The animal was not particularly active and rested 

in the upper portion of a 4 m tall tree. As it was photographed, 

including the use of flash, the individual remained 

largely stationary, until it finally turned and moved off into 

another tree and into dense vegetation. 

During the course of nocturnal observations of forest animals 

within the study site,this was the only observation of A. 

trichotis. Each night numerous individuals of Microcebus cf. 

Fig. 1: Photo of Allocebus trichotis taken during the night of 28 

October 2010 in a forest block approximately halfway between 

the turn-off of RN 2 and the village of Lakato. The 

ear-tufts of this animal, diagnostic of this species, are readily 

visible in the photo.


lehilahytsara were observed in close proximity. Although A. 

trichotis is now known to have a broad distribution across a 

good portion of the eastern humid forests, from lowland to 

montane forests (up to about 1,000 m),it occurs in low densities 

(Mittermeier et al., 2006).This factor might account for 

its absence in other forested sites surveyed within the 

Zahamena-Ankeniheny forest corridor (e.g., Schmid et al., 

1999; Randrianabinina and Rasoloharijaona, 2006). However, 

continued surveying efforts at these sites will probably result 

in the finding that it occurs across the forest corridor. 


The survey of the Lakato region was financed by a grant from 

the Volkswagen Foundation.We are grateful to the Département 

de Biologie Animale, Université d’Antananarivo and 

the Direction du Système des Aires Protégées, Direction 

Générale de l’Environnement et des Forêts for permits to 

conduct this research. 

References 

Biebouw,K.2009.Home range size and use in Allocebus trichotis 

in Analamazaotra Special Reserve, central eastern Madagascar. 

Int. J. Primatol. 30: 367-386. 

Garbutt,N.2000.Brief observations of hairy-eared dwarf lemur 

(Allocebus trichotis) in Analamazaotra Special Reserve, 

eastern Madagascar. Lemur News 6: 37. 

Goodman, S.M.; Raselimanana, A.P. 2002. The occurrence of 

Allocebus trichotis in the Parc National de Marojejy. Lemur 

News 7: 21-22. 

Meier, B.; Albignac, R. 1991. Rediscovery of Allocebus trichotis 

Günther 1875 (Primates) in northeast Madagascar. Folia 

Primatol. 56: 57-63. 

Mittermeier,R.A.;Konstant,W.R.;Hawkins,A.F.A.;Louis,E.E.; 

Langrand, O.; Ratsimbazafy, J.; Rasoloarison, R.M.; Ganzhorn, 

J.U.; Rajaobelina, S.; Tattersall, I.; Meyers, D.M. 2006. 

Lemurs of Madagascar. Second edition. Conservation International, 

Washington, D.C. 

Rakotoarison,N.1998.Recent discoveries of the hairy-eared 

dwarf lemur (Allocebus trichotis). Lemur News 3: 21. 

Rakotoarison, N.; Zimmermann, H.; Zimmermann, E. 1997. 

First discovery of the hairy-eared dwarf lemur (Allocebus 

trichotis) in a highland rain forest of Eastern Madagascar. 

Folia Primatol. 68: 86-94. 

Ralison,J.M.2010.The lemurs of the Ambatovy-Analamay region. 

Malagasy Nature 3: 178-191. 

Randrianambinina, B.; Rasoloharijaona, S. 2006. Inventaires 

des lémuriens nocturnes dans la forêt pluviale de Maromizaha 

(Est de Madagascar). Lemur News 11: 9-11. 

Schmid,J.;Fietz,J.;Rakotobe,Z.L.R.1999.Lémuriens du corridor 

Mantadia-Zahamena, Madagascar. In: J. Schmid, L.E. 

Alonso (eds.). Une évaluation biologique rapide du corridor 

Mantadia-Zahamena, Madagascar. Bulletin of Biological 

Assessment 32: 61-72. 

Schütz, H; Goodman, S.M. 1998. Photographic evidence of - 

Allocebus trichotis in the Reserve Speciale d’Anjanaharibe- 

Sud. Lemur News 3: 21-22. 

Tattersall,I.1982.The primates of Madagascar.Columbia University 

Press, New York. 

When big lemurs swallow up small ones: 

Coquerel’s dwarf lemur as a predator of 

grey mouse lemurs and endemic rodents 

Susanne Schliehe-Diecks 1, Matthias Markolf 2, Elise 

Huchard 2* 

1Courant Research Center "Evolution of Social Behavior", 

Georg-August-University of Göttingen,Kellnerweg 6,37077 

Göttingen, Germany 

2Abteilung Verhaltensökologie and Soziobiologie,Deutsches 

Primatenzentrum,Kellnerweg 4,37077 Göttingen,Germany 

*Corresponding author: ehuchard@gmail.com 

Predation has probably played a major role in the evolutionary 

history of lemurs, and specifically affects small nocturnal 

lemurs, which are heavily predated upon by a wide range of 

vertebrates, including carnivores (e.g., viverrid or domestic 

carnivores), birds (e.g. raptors, owls) or reptiles (e.g. Boidae) 

(Goodman, 2003). In contrast, lemur predation by other lemur 

species appears exceptional and highly opportunistic, 

with one observed case of predation of an infant Lemur catta 

by Eulemur fulvus (Pitts, 1995). However, such events might 

occur more regularly in other lemur species. Two indirect 

lines of evidence suggest that Coquerel’s dwarf lemur (Mirza 

coquereli) predates on closely related smaller mouse lemurs 

(Microcebus sp.) (Kappeler and Rasoloarison,2003).The first 

report is based on events where the partially eaten carcass of 

a gray mouse lemur (M. murinus) was found together with a 

live M. coquereli in a trap (Goodman, 2003). The second observation 

consists of an experimental confrontation of M. 

murinus with M. coquereli, both being kept in separate cages 

that were temporarily placed next to each other. In most 

experiments, mouse lemurs started alarm-calling at the 

Coquerel’s dwarf lemur and moved around in their cage in an 

agitated fashion (Fichtel,2009).Here,we present the first direct 

evidence of predation by wild M. coquereli upon gray 

mouse lemurs and endemic rodents (western tuft-tailed rats, 

Elliurus myoxinus). 

Study animals, study site and methods 

Coquerel’s dwarf lemurs (300 g;mean home range size:4 ha) 

occur in the western lowland forests and gray mouse lemurs 

(60 g;mean home range size:1.5 ha) can be found in most remaining 

forests in southern and western Madagascar (Kappeler 

and Rasoloarison,2003;Rasoloarison et al.,2000).Both 

species share several features.Both are nocturnal and omnivorous 

solitary foragers. They mainly feed on primary resources 

(fruits,gum,flowers,young leaves),insect secretions, 

small invertebrates and occasionally vertebrates (chameleons 

and lizards). Their diet displays seasonal fluctuation, as 

well as interspecific variation (Goodman, 1993, 2003), and 

the Coquerel’s dwarf lemur is reported to be slightly more 

carnivorous than the gray mouse lemur (Petter et al., 1977). 

In captivity, both species have been observed eating young 

rodents (Petter et al., 1977) although this has never been reported 

in the wild. Both species occur sympatrically in central 

western Madagascar with western tuft-tailed rats,a nocturnal, 

frugi- and granivorous and partially arboreal rodent 

(average body mass: 66 g) (Carleton, 2003). 

All following observations were made in the Forêt de 

Kirindy,a 12,500 ha forestry concession of the C.N.F.E.R.E.F. 

(formerly C.F.P.F.) Morondava. This dry deciduous forest is 

situated 60 km northeast of Morondava (44°39´E, 20°03´S). 

The predation of the western tufted-tail rat was witnessed 

during a focal observation of a Coquerel’s dwarf lemur which 

was equipped with a radio collar (Biotrack TW3). The observed 

mouse lemurs were similarly equipped with radio 

collars (Holohil Systems Ltd., BD-2C, 1.8 g), permitting behavioural 

observations of focal animals. 

Results and discussion 

The first observation reports the predation of a western 

tuft-tailed rat by an adult male M.coquereli in November 2006 

(Fig.1).The Coquerel’s dwarf lemur was found sitting on the 

ground at 20h17, feeding on a tufted-tail rat, and changed its 

position only to climb-up the vegetation from 1-3 m height 

and to recover the carcass when it fell to the ground. It devoured 

the whole carcass, including (cracked) bones. After 

finishing eating,the M.coquereli groomed its face and hands.


Fig. 1: An adult Coquerel’s 

dwarf lemur (Mirza coquereli) 

in the Foret de Kirindy, Madagascar. 

The second observation 

reports an unsuccessful 

attack on an adult female 

gray mouse lemur (55 g, 

approx. age: 1 year and 9 

months) by a Coquerel’s 

dwarf lemur in October 

2009.At 21h24,the mouse 

lemur had been foraging 

high up in the vegetation 

(between 6 and 15 m) for 

at least 10 minutes, licking 

sugary insect secretions 

off leaves,when a M.coquereli, 

adult size, was spotted 

at the same height, about 

10m from the focal subject, 

slowly and silently 

moving towards the 

mouse lemur. Marking 

brief and frequent pauses 

in an apparently easy pro- 

gression into the canopy, its whole attitude strongly recalled 

the hunting cat, with a low head and a flexible body, apparently 

entirely focused on its prey.In less than 30 seconds,the 

Mirza was within 5 m of the mouse lemur,who kept feeding in 

the same location. While the Mirza approached within 2 m, 

the mouse lemur suddenly disappeared in an eclipse, quickly 

fleeing among the top and tiniest branches,and jumping from 

one slim branch to the next.The Coquerel’s dwarf lemur did 

not try to chase it.After 2 minutes out of sight,the mouse lemur 

was retrieved quietly feeding on tree exudates, 25 m 

away from its previous localization in its fleeing direction,and 

15 m away from a frequently used sleeping site. Less than 10 

minutes later, the female was joined by a related female and 

both entered the tree hole together (21h37). 

Finally, an adult Coquerel’s dwarf lemur was observed feeding 

on a young male gray mouse lemur (body mass: 37 g; 

approx.age:2-3 months) in June 2010.The predation was recorded 

at 22h10,about two hours after behavioural data had 

been collected from the predated mouse lemur,which at the 

time showed no signs of injuries and displayed normal behaviour.The 

body of the gray mouse lemur was almost complete 

when the observer spotted the M. coquereli feeding on 

it,suggesting that the mouse lemur was killed shortly before. 

The Coquerel’s dwarf lemur was sitting with its prey in a tree 

of about 10m height, which stood 25 m away from the position 

where the grey mouse lemur was last spotted alive. It 

took about one hour to finish the entire carcass,interrupted 

by occasional vigilance scans of the surroundings. 

The frequency of such events is probably relatively low, and 

all reported observations happened during, or at the end of, 

the dry season in Kirindy. It is thus possible that predation 

pressure by M. coquereli increases at times of food scarcity, 

when alternative resources like fruits and invertebrates are 

rare or absent. However, it is also important to note that 

most observations took place during the dry season, when 

vegetation density is low in this dry, deciduous forest. This 

means that the timing of events reported here might simply 

reflect study methods. Nevertheless, this suit of anecdotal 

observations represents the first direct and unambiguous 

evidence for predation by the Coquerel’s dwarf lemur upon 

small nocturnal lemurs, as well as other mammals. Predation 

among other primate species is relatively rare. So far, only 

chimpanzees, orangutans, baboons, blue monkeys and capuchins 

have been observed preying upon other primates 

(Fichtel, in press). Our report provides evidence for the first 

case where a lemur species might commonly predate upon 

other lemurs. 

References 

Carleton,M.D.2003.Eliurus,Tufted-tailed rats.Pp.1373-1380. 

In:S.M.Goodman;J.Benstead (eds.). The Natural History 

of Madagascar.The University of Chicago Press,Chicago. 

Fichtel, C. 2009. Costs of alarm calling: lemur alarm calls 

attract fossas. Lemur News 14: 53-54. 

Fichtel,C.in press.Predation.In:J.Mitani:J.Call;P.M,Kappeler; 

R.Palombit;J.B.Silk;(eds.) The Evolution of Primate Societies. 

The University of Chicago Press, Chicago. 

Goodman,S.M.;O’Connor,S.;Langrand,O.1993.A review of 

predation on lemurs: implications for the evolution of 

social behavior in small, nocturnal primates. Pp. 51-66. In: 

P.M.Kappeler;J.U.Ganzhorn (eds.).Lemur social systems and 

their ecological basis. Plenum Press, New York. 

Goodman,S.M.2003.Predation on Lemurs.Pp 1221-1228.In: 

S. M. Goodman: J. Benstead (eds.). The Natural History of 

Madagascar. The University of Chicago Press, Chicago. 

Kappeler,P.M.;Rasoloarison,R.M.2003.Microcebus,mouse lemurs, 

tsidy. Pp 1310-1315. In: S. M. Goodman: J. Benstead 

(eds.).The Natural History of Madagascar.The University 

of Chicago Press, Chicago. 

Petter, J-J.; Albignac, R.; Rumpler, Y. 1977. Mammifères lémuriens 

(primates prosimiens). In: Faune de Madagascar, 

Paris. 

Pitts, A. 1995. Predation by Eulemur fulvus on an infant Lemur 

Catta at Berenty, Madagascar. Folia Primatol. 65: 169-71. 

Rasoloarison, R.M.; Goodman, S.; Ganzhorn, J.U. 2000. Taxonomic 

revision of mouse lemur (Microcebus) in the western 

portions of Madagascar.Int.J.Primatol.21:963-1019. 

Collective mobbing of a boa by a group of 

red-fronted lemurs (Eulemur fulvus rufus) 

Lennart Pyritz 1,2*, Tianasoa Andrianjanahary 1 

1Behavioral Ecology & Sociobiology Unit, German Primate 

Center, Kellnerweg 4, 37077 Göttingen, Germany 

2CRC "Evolution of Social Behavior", University of Göttingen, 

Germany 

*Corresponding author: LennartPyritz@gmx.net 

Key words: red-fronted lemurs, boa, predation, mobbing, 

anti-predator behaviour 

Introduction 

Collective anti-predator behaviour is one of the principal advantages 

of group-living (for mammals, e.g., Janzen, 1970; van 

Schaik, 1983). It can be broadly divided into two strategies 

and tactics employed before and after predator encounters 

(Caro,2005;Rahlfs and Fichtel, 2010;Fichtel, in press). While 

the former include predator-sensitive foraging and increased 

vigilance, mobbing occurs in several mammal species after 

detecting a predator (e.g., Tamura, 1989). Why animals 

engage in mobbing and who benefits from it in which way 

remains an unresolved question in animal behaviour (for 

reviews see Curio et al., 1978; Shields, 1984). Until today, 

published field observations of group-living lemurs mobbing 

a predator are rare (summarised in Scheumann et al., 2007). 

Regarding snakes, only three interactions have been described 

so far (Colquhoun,1993;Rakotondravony,1998;Burney, 

2002). Here, we report a prolonged mobbing display 

against a Madagascar ground boa (Acrantophis madagascariensis) 

by a group of red-fronted lemurs (Eulemur fulvus rufus) 

in Kirindy Forest. Observations like this may help to elucidate 

fundamental mechanisms of collective anti-predator behaviour 

by contributing to a pool of data on mobbing by particular 

pairs of prey and predators.

Lemur News Vol. 15, 2010 Page 15 

Observations 

The event was observed during regular behavioural observations 

of red-fronted lemurs in Kirindy Forest,60 km north of 

Morondava. It was the only snake-lemur interaction observed 

during the entire study period from November 2007 

to April 2010, in which four lemur groups were followed 

daily by one or two observers,respectively (> 4,000 h of observation 

data).Red-fronted lemurs live in multi-male,multifemale 

groups of 5-12 individuals (Pereira and Kappeler, 

1997;Wimmer and Kappeler,2002).The study group (B) that 

encountered the boa included 9 individually marked animals 

at the time (2 adult females, 5 adult males, 1 juvenile male, 1 

male infant). 

On March 1,2010,at 7.19 h,sudden alarm calls of several redfronted 

lemurs were heard in the study area known as CS7. 

Six individuals (2 adult females,2 adult males,1 juvenile male, 

1 male infant) could be identified after approaching the group 

to within 10 m.Three of them (2 adult females,1 adult male) 

emitted "Woofs" and "Huvvs",vocalisations typically uttered 

during predator encounters (Fichtel and Kappeler, 2002). 

The 5 individuals surrounded an approximately 2 m long 

Madagascar ground boa (Fig. 1) that was lying motionless on 

the ground. The lemurs sat at a height of 1-2 m, each about 

3 m away from the snake, wagging their tails vigorously, ex- 

Fig. 1: Madagascar ground boa in Kirindy. (Photo: Lennart 

Pyritz) 

cept the infant that kept a distance of 5 m during the entire 

event and did not display any vocalisations or tail-wagging. 

During the next 4 min,one of the adult males approached the 

front end of the boa twice, getting as close as 1-2 m. After 5 

min, he left the scene. During this time, one of the adult females 

also approached the snake up to within 2 m.When the 

male left,the second female started to quickly circuit the boa 

for 4 min,maintaining a distance of 2-3 m.After 14 min of several 

approaches and continuous alarm calls by 3-5 individuals, 

the boa moved for the first time,heading slowly away.The remaining 

adult male approached the moving snake also within 

2 m; also at its front end. About 1 min later, the boa had 

moved 15 m away, and the lemurs left in the opposite direction,still 

uttering grunts continuously.Once the boa was out 

of sight, the mobbing stopped and the lemurs` behaviour returned 

to baseline levels. 

Discussion 

The mobbing reaction of the group was strong and prolonged 

and included most of the group members.This is similar 

to the behaviour of a black lemur (Eulemur macaco 

macaco) group encountering a Madagascar boa at Ambato 

Massif, where the group mobbed the snake for 15-20 min, 

and some individuals approached it as close as 1 m before 

finally leaving the location (Colquhoun, 1993). It is also noteworthy 

that females and males mobbed and approached the 

snake in equal measure as observed in a number of other 

species (e.g., Tamura, 1989; Ferrari and Ferrari, 1990; Tello et 

al., 2002). The infant maintained a larger distance to the boa 

and did not engage in the mobbing displays, however. Similar 

infant behaviour has also been reported for other primates 

(e.g., Ferrari and Ferrari, 1990) and might be due to a higher 

susceptibility to an attack due to smaller body size or a lack 

of innate experience regarding predator encounters and 

mobbing strategies (Curio et al., 1978; Fichtel, in press). 

The strong mobbing reaction of the lemurs might be explained 

by the hunting strategy of the snake.Boas are ambush 

hunters that usually abandon an attack as soon as they have 

been detected (Montgomery and Rand, 1978; Slip and Shine, 

1988). Therefore, it seems beneficial for prey animals to signal 

the ambush hunter quickly and distinctly that it has been 

detected. As boas do not pursue their prey after an unsuccessful 

attack, it is also unsurprising that the lemurs’ behaviour 

returned to baseline levels of anxiety shortly after departing 

the site of the predator encounter. In contrast, 

groups of red-fronted lemurs showed increased vigilance behaviour 

for at least 30-60 min after encountering a fossa 

(Cryptoprocta ferox;pers.comm.Jean-Pierre Tolojanahary and 

pers.observation by LP),which is probably due to the higher 

agility and climbing abilities of the largest mammalian carnivore.Furthermore,fossas 

have been observed to hunt cooperatively 

and pursue prey up to 45 min (Lührs and Dammhahn, 

2009). 

There are no quantitative data on predation rates of lemurs 

by snakes in Kirindy, only opportunistic observations (e.g., 

Schülke, 2001; Eberle and Kappeler, 2008) that are biased by 

several factors, however. The low observation rate of boalemur 

interactions could be due to the reptiles` nocturnal 

lifestyle (Raxworthy,2003),so that most of the attacks would 

occur at night when no or only few observers are working in 

the forest. Furthermore, boas at Kirindy are only active during 

the rainy season from January to April, when observations 

are often limited by dense foliage and frequent rainfalls. 

Five of the six individuals taking part in the mobbing were related 

(1 adult female and her 4 offspring from the last 4 

years), while it is currently unknown whether the second 

adult male sired one of the two youngest group members. 

However, due to the small number of detailed observations 

of predator encounters it remains impossible to identify the 

ultimate causes of collective mobbing (kin defence/parental 

care, self-/group defence or cultural transmission of enemy 

recognition; Curio et al., 1978) in this species for the time 

being. 


LP was supported financially by the Deutsche Forschungsgemeinschaft 

(DFG; KA 1082/16-1, FuE). We thank field assistant 

Jean-Pierre Tolojanahary for sharing his long-term observation 

experiences with us. We are also grateful to Peter 

Kappeler, Claudia Fichtel and Moritz Rahlfs for constructive 

and helpful comments on earlier drafts of the manuscript. 

References 

Burney, D.A. 2002. Sifaka predation by a large boa. Folia Primatologica 

73: 144-145. 

Caro, T. 2005. Antipredator defenses in birds and mammals. 

The University of Chicago Press, Chicago, London. 

Colquhoun, I.C. 1993. The socioecology of Eulemur macaco: 

A preliminary report. Pp. 11-23. In: P.M. Kappeler; J.U. 

Ganzhorn (eds.). Lemur social systems and their ecological 

basis. Plenum Press, New York and London.


Curio, E.; Ernst, U.; Vieth, W. 1978. Cultural transmission of 

enemy recognition:one function of mobbing.Science 202: 

899-901. 

Eberle, M.; Kappeler, P.M. 2008. Mutualism, reciprocity, or kin 

selection? Cooperative rescue of a conspecific from a boa 

in a nocturnal solitary forager the gray mouse lemur. 

American Journal of Primatology 70: 410-414. 

Ferrari, S.F.; Ferrari, M.A.L. 1990. Predator avoidance behaviour 

in the buffy-headed marmoset, Callithrix flaviceps. 

Primates 31: 323-338. 

Fichtel, C. In press. Predation on primates. In: J.C. Mitani; J. 

Call;P.Kappeler;R.Palombit;J.Silk (eds.).The evolution of 

primate societies. University of Chicago Press, Chicago. 

Fichtel, C.; Kappeler, P.M. 2002. Anti-predator behaviour of 

group-living Malagasy primates: mixed evidence for a 

referential alarm call system. Behavioral Ecology and 

Sociobiology 51: 262-275. 

Janzen,D.1970.Altruism by coatis in the face of predation by 

boa constrictor. Journal of Mammalogy 51: 387-389. 

Lührs,M.L.;Dammhahn,M.2009.An unusual case of cooperative 

hunting in a solitary carnivore. Journal of Ethology 

DOI: 10.1007/s10164-009-0190-8. 

Montgomery, G.G.; Rand, A.S. 1978. Movements, body temperature 

and hunting strategy of a Boa constrictor. Copeia 

3: 532-533. 

Pereira, M.E.; Kappeler, P.M. 1997. Divergent system of agonistic 

behaviour in lemurid primates.Behavior 34:225-74. 

Rahlfs,M.;Fichtel,C.2010.Anti-predator behaviour in a nocturnal 

primate, the grey mouse lemur (Microcebus murinus). 

Ethology 116: 429-439. 

Rakotondravony, D.; Goodman, S.M.; Soarimalala, V. 1998. 

Predation on Hapalemur griseus griseus by Boa manditra 

(Boidae) in the Littoral Forest of Eastern Madagascar. 

Folia Primatologica 69: 405-408. 

Raxworthy, C.J. 2003. Boidae, Boas. Pp. 993–997. In: S.M. 

Goodman; J.P. Benstead (eds.). The Natural History of 

Madagascar. University of Chicago Press, Chicago. 

Scheumann, M.; Rabesandratana, A.; Zimmermann, E. 2007. 

Predation, communication, and cognition in lemurs. Pp. 

100-126. In: S. Gursky; K.A.I. Nekaris (eds.). Primate antipredator 

strategies. Springer, New York. 

Schülke, O. 2001. Social anti-predator behaviour in a nocturnal 

lemur. Folia Primatologica 72: 332-334. 

Shields,W.1984.Barn swallow mobbing:self-defence,collateral 

kin defence, group defence, or parental care? Animal 

Behaviour 32: 132-148. 

Slip, D.; Shine, R.1988. Feeding habits of the diamond python, 

Morelia s. spilota: ambush predation by a Boid snake. Journal 

of Herpetology 22: 323-330. 

Tamura, N. 1989. Snake-directed mobbing by the Formosan 

squirrel Callosciurus erythraeus thaiwanensis. Behavioral 

Ecology and Sociobiology 24: 175-180. 

Tello, N.S.; Huck, M.; Heymann, E.W. 2002. Boa constrictor attack 

and successful group defence in moustached tamarins, 

Saguinus mystax. Folia Primatologica 73: 146-148. 

Van Schaik, C. 1983. Why are diurnal primates living in 

groups? Behaviour 88: 120-143. 

Wimmer,B.;Kappeler,P.M.2002.The effects of sexual selection 

and life history on the genetic structure of redfronted 

lemur, Eulemur fulvus rufus, groups. Animal Behaviour 64: 

557-68. 

Response of two nocturnal lemurs (Microcebus 

murinus and Lepilemur leucopus) 

to a potential boiidae (Sanzinia madagascariensis) 

predator 

Krista Fish 

Department of Anthropology, The Colorado College, 14 E 

Cache La Poudre, Colorado Springs, CO 80903, USA, 

krista.fish@coloradocollege.edu 

Primates display an array of responses to predators, and 

differences in these responses were once thought to exist 

based on activity pattern. Solitary foraging, cryptic color- 

ation, and cryptic movements to remain hidden from predators 

were considered anti-predator adaptations among 

nocturnal primates while diurnal primates used large group 

size to enhance their ability to detect and defend against 

predators, give alarm calls to warn conspecifics of the presence 

of a predator,and flee from predators (Hill and Dunbar, 

1998;Isbell,1994;Stanford,2002).As research into nocturnal 

primate behavior expanded,results revealed that,like diurnal 

primates,nocturnal primates display a range of anti-predator 

behaviors upon encountering a predator (Fichtel, 2007). 

Nocturnal primates vary in the type of response (mobbing, 

alarm calling), height in the canopy, proximity to other 

individuals, and vigilance levels in the presence of different 

predators (Fichtel, 2007; Gursky, 2002, 2003, 2005, 2006; 

Schuemann et al.,2007;Schulke,2001).In particular,mobbing 

behaviors are well- documented in tarsiers (Tarsius sp.), 

mouse lemurs (Microcebus spp.), and fork-marked lemurs 

(Phaner furcifer) (Gursky, 2007; Eberle and Kappeler, 2008; 

Schulke, 2001). Mobbing includes close approach, touching, 

sniffing, and pouncing on the predator (Gursky, 2007). Interspecies 

mobbing of a predator occurs as well. Fork-marked 

lemurs and coquerel’s dwarf lemurs (Mirza coquereli) together 

mobbed a snake (Boa manditra) (Schulke, 2001). 

Several hypotheses have been put forth to explain the evolution 

of mobbing behavior (Eberle and Kappeler,2008):1) byproduct 

mutualism in which individuals defend others in the 

process of defending themselves, 2) reciprocity where animals 

obtain higher fitness by cooperating with others,and 3) 

kin selection whereby animals cooperate when they share 

common genes. A fourth hypothesis known as perception 

advertisement was developed as an explanation for the 

evolution of alarm calling and other mobbing behaviors in 

birds (Curio et al., 1978; Zuberbuhler et al., 1999) but has 

been extended to account for the presence of mobbing behaviors 

in primates (Gursky,2005).According to this hypothesis, 

alarm calling and mobbing are signals to the predator 

that the element of surprise has been lost. Snakes, leopards, 

and other animals that hunt by crypticity and rely on the element 

of surprise to capture prey are common recipients of 

alarm calling and mobbing.Research on both diurnal and nocturnal 

primates suggests that alarm calling and mobbing by 

these primates results in predators leaving an area (Zuberbuhler 

et al.,1999;Gursky,2006).Here I report the divergent 

responses of two species of nocturnal primates (Microcebus 

murinus and Lepilemur leucopus) to the same potential predator- 

a nocturnal boiidae snake (Sanzinia madagascariensis) 

and discuss implications for the above hypotheses based on 

these observations. 

Methods 

The observations reported here were made in the Ankoba 

gallery forest of Berenty Private Reserve in southern Madagascar. 

The encounters between the snake and primates 

were observed in the course of a six month study investigating 

the ecology of Microcebus murinus. During this study, 

trails within the reserve were walked to locate unhabituated 

mouse lemurs.When encountered,the time at which the encounter 

began, height, location, and activity of the mouse lemur 

were recorded continuously until the mouse lemur was 

out of sight for more than five minutes. The time, height, and 

activity of potential predators were also noted when they 

were encountered, but predators were not followed unless 

they were within proximity to a primate. 

Results 

While conducting walks to locate mouse lemurs in May 2009, 

a sportive lemur alarm call at 20:52 alerted me to the pres-


ence of a boiidae snake later identified as Sanzinia madagascariensis. 

I saw two white-footed sportive lemurs (Lepilemur 

leucopus),one located at 4m height and the other at 5m 

height in a tree over the trail.The sportive lemurs were barking 

and looking at the snake. The snake was moving at 

approximately 4 m high in a tree and was attempting to cross 

a gap and move into a tree nearer to the tree in which the 

sportive lemurs were located.The attempt was unsuccessful 

as the snake slipped and almost fell out of the thin, terminal 

branches. The snake then moved down to 3.5 m height and 

began crossing the canopy gap along thicker branches. The 

sportive lemurs continued to alarm bark at the snake until 

21:02 when the snake turned away from the gap in the 

canopy and began moving lower down in the tree and away 

from the sportive lemurs. The sportive lemur alarm calling 

ended by 21:04 when the snake had traveled down the tree 

trunk to 3m in height. The sportive lemur lowest in the tree 

continued to watch the snake while the other sportive lemur 

moved to 8m height in its tree and began feeding. 

At 21:15, I was preparing to leave the area when I noticed a 

mouse lemur at 0.5 m in the same tree as the snake. The 

snake was at 3 m height in the tree and moving down the 

main trunk of the tree. The mouse lemur looked at me as it 

walked up the main trunk of the tree in the direction of the 

snake. The snake faced the mouse lemur, but the mouse 

lemur did not appear to notice the snake as it alternated 

looking in my direction with looking around its immediate 

area while foraging for insects. The mouse lemur continued 

to move up the tree until it was within 0.25 m of the snake 

where it paused and looked intently at the spot where the 

snake was located and then jumped backwards away from 

the snake. The mouse lemur then began moving around the 

tree at the same height (3 m) as the snake,jumping to terminal 

branches,and running along main branches while pausing 

to look at the snake. The mouse lemur’s movements took it 

from 1m to only a few cms in distance from the snake. The 

mouse lemur continued this pattern of running and pausing 

to examine the snake for 7 mins. The mouse lemur then began 

foraging for insects in a neighboring tree at 3-4 m height 

and within 1-2 m of the snake. While foraging, the mouse 

lemur would pause to look in the direction of the snake 

which remained motionless.The mouse lemur foraged in this 

manner for 4 mins until a sportive lemur alarm called. Prior 

to the alarm call, the mouse lemur was foraging approximately 

1m from the snake and the snake began moving down 

the tree trunk.At the sound of the sportive lemur alarm call, 

the mouse lemur jumped to 2 m distance from the snake and 

paused in its foraging to watch the snake. The snake remained 

motionless.A minute later the sportive lemurs alarm 

called again. The snake began moving down the trunk of the 

tree again and the mouse lemur moved to forage insects in 

trees that were approximately 3-5 m from the snake and at 

5m height.After 5 mins of foraging at this increased distance 

from the snake, the mouse lemur moved back into the tree 

where it was initially observed and foraged within 1m of the 

snake for 2 mins. The mouse lemur then moved close to the 

snake coming within less than 1m of the boa and running and 

jumping around the snake while pausing to watch it while 

standing bipedally. After 3 mins of remaining motionless 

while the mouse lemur ran, jumped and watched the snake, 

the snake began moving back up into the dense crown of the 

tree. The mouse lemur continued running and jumping 

around the snake and watching it at a distance of 0,5-1 m 

away as the snake moved into the crown of the tree. 8 mins 

later,the mouse lemur began foraging insects at 3-4 m height 

in the canopy and within 0,5-1 m from the snake with occasional 

glances in the direction of the snake.The mouse lemur 

then moved further away from the snake and foraged insects 

at 5m in height and approximately 5 m distance from the 

snake.At 21:46 the mouse lemur was out of sight and did not 

return by 21:51 when I left and continued the mouse lemur 

walk.A return visit to the location at 10:05 for species identification 

of the species revealed the presence of no mouse lemurs 

or Lepilemur in the vicinity. 

Discussion 

Eberle and Kappeler (2008) describe the successful mobbing 

of a Sanzinia madagascariensis by two female and one male 

mouse lemur (Microcebus murinus). The mobbing caused the 

snake to release a captured male mouse lemur. Subsequent 

genetic analysis revealed that the mobbing females were related 

to the attempted victim (Eberle and Kappeler, 2008). 

Additional encounters between mouse lemurs and snakes 

without captured prey did not elicit mobbing behaviors from 

the mouse lemurs (Eberle and Kappeler, 2008). Instead, the 

mouse lemurs sat approximately 2 m from the snake and 

watched it. (Eberle and Kappeler, 2008). The mobbing of the 

snake by relatives of a captured mouse lemur and the lack of 

mobbing behaviors by solitary mouse lemurs in Eberle and 

Kappeler’s study support the kin selection hypothesis for the 

evolution of mobbing behaviors. However, the solitary 

mouse lemur observed in this study displayed mobbing-like 

behaviors in the absence of kin or other individuals. Additionally, 

under experimental conditions, solitary mouse lemurs 

monitored model snakes and even locomoted towards 

the model predators (Rahlfs and Fichtel,2010).The presence 

of mobbing-like behaviors in solitary animals lends support 

to the by-product mutualism hypothesis. Mouse lemurs may 

display mobbing-like behaviors as an individual strategy and 

then when larger numbers of mouse lemurs contact a predator, 

the appearance of a group-defense strategy may result 

from multiple individuals pursuing the same strategy. 

The observations here also lend support to the perception 

advertisement hypothesis.The mouse lemur may have benefited 

by displaying mobbing behaviors toward the snake to let 

the snake know that it was being monitored and would not 

surprise the lone mouse lemur as it foraged.However,a prediction 

of the perception-advertisement hypothesis is that 

cryptic predators should flee when faced with alarm calls and 

mobbing as they cannot surprise prey in their vicinity. The 

alarm calling by the sportive lemurs and the mobbing-like 

behaviors of the mouse lemur observed in this study did not 

cause the snake to flee. Instead, the snake remained in the 

area while both the sportive lemurs and mouse lemur left 

the area.Size of the mobbing and alarm-calling group may be 

important. Tarsiers were more likely to retreat first if their 

mobbing group consisted of four or fewer individuals (Gursky,2007).The 

small group sizes here may have not caused the 

predator to flee,but may have served as an adequate warning 

that it had lost the element of surprise and should not 

expend energy in an attack. 

Of particular interest in these observations was the opportunity 

to view the responses of two different nocturnal primate 

species to the same predator. Such observations are 

infrequent in the literature. Schulke (2001) observed Phaner 

and dwarf lemur (Mirza coquereli) together mobbed Boa 

manditra. In the case of the sportive and mouse lemurs, 

neither showed the same behavioral response to the Sanzinia 

madagascariensis. The pair of Lepilemurs maintained a 

larger distance between themselves and the snake than did 

the mouse lemur. The mouse lemur did not vocalize in the 

vicinity of the snake, however, the sportive lemurs vocalized 

during their encounter with the snake.Mouse lemurs seldom 

respond to predator models with alarm calls in experimental


conditions (Rahlfs and Fichtel,2010).Nocturnal primates and 

other mammals may not rely on alarm calls as an anti-predator 

strategy (Rahlfs and Fichtel, 2010) because early detection 

of predators at night is more difficult and solitary foraging 

may limit the usefulness of this strategy. The sportive 

lemurs however did alarm call in the presence of the snake. 

Alarm calls in nocturnal primates may function as a deterrent 

to the predator or to recruit conspecifics in defense against 

the predator (Rahlfs and Fichtel,2010).The exact function of 

the sportive lemur alarm calls in this study cannot be determined, 

but they do reveal a need for future studies to examine 

potential variation in alarm call behaviors between nocturnal 

species.Given variation in body size,diet,and sociality 

in nocturnal primates, we might expect variation in antipredator 

behaviors such as alarm calling among nocturnal 

species. 

The possibility exists that the mouse lemur may have recognized 

the alarm calls of the sportive lemur because it increased 

its distance from the snake, ceased foraging, and 

monitored the snake following an alarm call. Recognition of 

the alarm calls of other species has been documented for 

birds, small mammals, and primates including diurnal lemurs 

(Fichtel,2004;Rainey et al.,2004;Shriner,1998).However,the 

suggestion that mouse lemurs recognize sportive lemur 

alarm calls needs to be further investigated with field experiments. 

The ability of nocturnal primate species to recognize 

the alarm calls of other sympatric species would be beneficial 

to animals that frequently forage away from conspecifics. 

The few encounters documented between nocturnal primates 

and their predators describe a range of anti-predator 

responses that vary depending on the type of predator,proximity 

of conspecifics,and available vegetative cover.Such flexibility 

is interesting because nocturnal primates- especially 

mouse lemurs- are often used as living models for the ancestral 

primate. The anti-predator behaviors of nocturnal primates 

such as mouse lemurs may reflect the primitive 

primate or even primitive mammal condition (Stanford, 

2002). The range of anti-predator behavior described for 

nocturnal primates in this and other studies implies that the 

flexibility in anti-predator behaviors observed in primates 

have a more ancient origin than originally suspected or that 

mouse lemurs and sportive lemurs have developed divergent 

anti-predator behaviors that may not have been present in 

early mammals and primates. Wider use of experiments to 

explore the differing conditions which elicit variable responses 

to predators within and between nocturnal species 

will be necessary to develop a more complete understanding 

of ancestral versus derived anti-predator behaviors. Additionally, 

comparative research exploring variation in antipredator 

behaviors in other nocturnal mammal species will 

be needed. 


I would like to thank Dr.Alison Jolly for her facilitation of this 

research project and the de Heaulme family for their permission 

to work at Berenty and their support of the project.I am 

grateful also to the staff of MICET (Madagascar Institut pour 

la Conservations des Ecosystèmes Tropicaux) for their assistance 

in travel and obtaining research permits and visas. 

Funding for this project was provided by Sigma Xi, The University 

of Colorado Museum, the University of Colorado 

Graduate Student Grants, and the American Society of 

Primatologists. 

References 

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avian mobbing.II.Cultural transmission of enemy recogni- 

tion in blackbirds: effectiveness and some constraints. Z. 

Tierpsychol. 48: 184-202. 

Eberle, M.; Kappeler, P.M. 2008. Mutualism, reciprocity, or kin 

selection? Cooperative rescue of a conspecific from a boa 

in a nocturnal solitary forager the gray mouse lemur.Am.J. 

Primatol. 70: 410-414. 

Fichtel, C. 2004. Reciprocal recognition of sifaka (Propithecus 

verreauxi verreauxi) and redfronted lemur (Eulemur fulvus 

rufus) alarm calls. Anim. Cogn. 7: 45-52. 

Fichtel, C. 2007. Avoiding predators at night: antipredator 

strategies in red-tailed sportive lemurs (Lepilemur ruficaudatus). 

Am. J. Primatol. 69: 611-624. 

Gursky, S. 2002. The behavioral ecology of the spectral tarsier, 

Tarsius specturn. Evol. Anth. 11: 226-234. 

Gursky, S. 2003. Predation experiments on infant spectral 

tarsiers (Tarsius spectrum) Folia. Primatol. 74: 272-284. 

Gursky, S. 2005. Predator mobbing in Tarsius spectrum. Internat. 

J. Primatol. 26(1): 207-221. 

Gursky, S. 2006. Function of snake mobbing in spectral tarsiers. 

Am. J. Phys. Anth. 129:601-608. 

Gursky, S. 2007. The Spectral Tarsier. Upper Saddle River, NJ: 

Pearson/Prentice Hall. 

Hill, R.A.; Dunbar, R.I.M. 1998. An evaluation of the roles of 

predation rate and predation risk as selective pressures 

on primate grouping behavior.Behaviour 135(4):411-430. 

Isbell, L.A. 1994. Predation in primates: ecological patterns 

and evolutionary consequences. Evol. Anth. 3(2): 61-71. 

Rahlfs, M.; Fichtel, C. 2010. Anti-predator behavior in a nocturnal 

primate, the grey mouse lemur (Microcebus murinus). 

Ethology 116: 429-439. 

Rainey HJ,Zuberbuhler K,Slater PJB.2004.Hornbills can distinguish 

between primate alarm calls. Proc R Soc Lond B 

271: 755-759. 

Scheumann, M.; Rabesandratana, A.; Zimmermann, E. 2007. 

Predation, communication, and cognition in lemurs. Pp. 

100-126. In: S. Gursky; K.A.I Nekaris (eds.). Primate Anti- 

Predator Strategies. Springer. 

Schulke, O. 2001. Social anti-predator behaviour in a nocturnal 

lemur. Folia Primatol. 72: 332-334. 

Shriner,W.M.1998.Yellow-bellied marmot and golden-mantled 

ground squirrel responses to heterospecific alarm 

calls. Anim. Behav. 55: 529-536. 

Stanford, C.B. 2002. Avoiding predators: expectations and 

evidence in primate antipredator behavior. Int J Primatol 

23(4): 741-757. 

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function of primate alarm calls. Ethology105: 

477-490. 

Effective predation defence in Cheirogaleus 

medius 

Kathrin H. Dausmann 

Animal Ecology & Conservation, Biocentre Grindel, University 

Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, 

Germany, kathrin.dausmann@uni-hamburg.de 

Besides one’s own death, the predation of offspring is the 

most severe loss of fitness possible.In species that invest significantly 

in their offspring it is therefore natural that this "expensive" 

offspring should be guarded to avoid predation. 

Nonetheless, and particularly in small animals, it is uncertain 

whether an adult can effectively defend its offspring when it 

is attacked by a larger predator. 

The fat-tailed dwarf lemur (Cheirogaleus medius) is a small 

(130 g), strictly nocturnal primate that occurs in the dry, 

deciduous woodlands of western Madagascar, and lives in 

social monogamous small family groups consisting of a reproductive 

male-female pair and their offspring from one or 

more breeding seasons. Males and females maintain lifelong 

pair bonds and usually separate only when one partner dies 

(Fietz, 1999; Müller, 1999; Fietz and Dausmann, 2003). The


diet of C.medius consists mainly of fruits and varying proportions 

of arthropods depending on the season (Fietz and 

Ganzhorn, 1999). C. medius is unique among primates because 

it spends seven months hibernating during the cooler 

dry-season of the southern winter (April to October),when 

food and water availability are low (Petter,1978;Hladik et al., 

1980; Dausmann et al., 2004). When resting during the day 

and when hibernating,the dwarf lemurs occupy tree hollows 

either alone or with members of the family group (Dausmann 

et al., 2005). In the wild, female C. medius usually give 

birth to twins and in most cases reproduction only takes 

place every second year (Fietz and Dausmann, 2006). C. 

medius only reproduce after delayed emigration from their 

family and successful occupation of their own territory (thus 

in their third year at the earliest;Fietz et al.,2000).In addition, 

their life span is restricted by their size and is usually 

between 4 and 11 years for territory holders, and so opportunities 

to reproduce are limited (for most animals between 

one and three). Therefore, every young is a costly and valuable 

investment. Avoiding predation of their young should 

therefore be a critical part of parental care. 

Predators of C. medius include raptors (Madagascar harrier 

hawk Polyboroides radiatus, Madagascar buzzard Buteo brachypterus, 

Madagascar long-eared owl Asio madagascariensis), 

mammals (Fossa Cryptoprocta ferox, Narrow-striped mongooses 

Mungodictis decemlineata), and snakes (Madagascar 

Ground Boa Acrantophis madagascariensis, Madagascar Tree 

Boa Sanzinia madagascariensis, Malagasy Cat-eyed Snake Madagascarophis 

colubrinus) (Dausmann,submitted). The choice 

of an appropriately sized tree hollow in which to give birth 

can reduce attacks from many of these predators with the 

exception of snakes which are able to enter any hollow that 

can be used by C. medius. 

C. medius leave their tree hollows at sunset to forage alone 

but both sexes defend their shared territory. After the birth 

of their offspring, both parents take turns in guarding the 

young in the tree hollow throughout the night, while the 

other one forages.As the young get older, the proportion of 

time that parents spend guarding them gradually declines 

(Fietz and Dausmann, 2003). At the age of about two weeks, 

both parents leave the hollow and return only occasionally. 

During this time, the young are particularly vulnerable to 

predation,since they are unguarded.Later,the young accompany 

the parents during their nightly excursions. 

In this note, I want to describe evidence that adult C. medius 

can repel larger predators and therefore guarding or at least 

remaining within hearing range of the tree hollow is an effective 

measure against predation of their offspring. I report an 

observation in which a snake (Madagascarophis colubrinus) 

tried to attack two C. medius young within a tree hollow but 

was successfully repelled by the mother. 

Our observation occurred in the Kirindy C.F.P.F.forest,a dry 

deciduous forest near the west coast of Madagascar (60 km 

north east of Morondava) during a focal animal survey of a 

female C. medius on January 31st. For a more detailed description 

of the area see Ganzhorn and Sorg (1996). 

The female was a mother of two young aged two weeks.The 

male of the pair was also being observed.The female left the 

tree hollow after sunset at 18:57 hours and the male 

followed at 19:01 hours.The two young were left alone in the 

hollow within a dead tree (Malagasy name: Mapingo). The 

entrance of the hollow was 3 m above ground. The female 

started her regular patrol of the territory border, but suddenly 

abandoned the patrol at 19:29 hours when about 80 m 

from the hollow. She returned quickly to the hollow in 

almost a straight line.On approach to the hollow,it was clear 

that the two young who had been left alone within the 

hollow were making loud and constant distress calls. On a 

branch of the same tree at a height of about 1.5 m above 

ground was a large M. colubrinus (> 1 m length) eying the 

hollow. Even though this crepuscular or nocturnal snake is 

mainly terrestrial, scansorial behaviour is possible. The female 

approached the snake to within a few centimetres and 

actively attacked it, and the snake responded by striking 

towards the female.Both the adult female and the juvenile C. 

medius were loudly vocalizing constantly.After three min,the 

female seized the tail of the snake with both hands and bit it 

about 10 cm from the end. The snake tried to drop to the 

ground, but remained dangling in the female’s teeth. After 

10 s of wriggling and repeated attempts to strike the female 

the snake fell to the ground and moved away quickly. The 

female descended to about 1 m, observed the ground for a 

few minutes and then spent 15 min agitatedly observing the 

surroundings at a height of about 3 m and inspecting the tree 

hollow containing the young,who were still loudly vocalizing. 

For the next 3.5 hours the female was moving rapidly around 

within the territory, which is very unusual for a C. medius, 

frequently returning and checking the tree hollow with the 

young.She finally carried leaves into a new tree hollow about 

50 m away and separately carried both young to the new tree 

hollow. She did not return to sleep in the original tree hole 

for the next two months, even though it had been used 

frequently prior to this encounter.It seems puzzling that the 

male of the pair did not come to help during the attack.Since 

the male was followed simultaneously we know that at the 

time of the attack he was less than 20 m from the tree hollow, 

and clearly within hearing range of the distress calls. He 

returned about 30 min after the attack where he met the 

female and groomed her while she and the young continued 

making distress calls. Since reproduction in this species is a 

fairly rare event even including extra pair copulations, the 

possibility of siring offspring is restricted, and the male 

should have been highly motivated to defend his young in 

order to increase his fitness. Considering the high (obligate) 

paternal investment in guarding the young and the life-long 

pair bond, C.medius have a surprisingly high rate (ca 40 %) of 

extra pair young (Fietz et al.,2000;Schwensow et al.,2007).It 

is thought that the male cannot discriminate between intra 

pair and extra pair young and therefore cares for any offspring 

of his pair-partner,so as not to jeopardize the survival 

of his own young. Alternatively, paternal care of extra pair 

young could be an indicator for male quality or simply a tactic 

to maintain his bond with the female and so securing future 

mating possibilities in such a long-lasting relationship (Fietz 

and Dausmann, 2003). Genetic analyses showed that the 

male ("social father") of our observation was indeed only the 

genetic father of one offspring, but not the other (Schwensow 

et al., 2007). However, even if he was able to distinguish 

kin from non-kin, he should still have defended the tree hollow 

in order to protect his one own offspring.Interestingly,in 

the weeks before these observations, the male and the 

female had always spent the daily resting period together in 

the same tree hollow. However, the day after the predation 

attempt they slept apart, the male in the old, and the female 

together with the offspring in the new tree hollow. We 

cannot judge whether the male was unable to find the female 

in the new tree hollow, whether he chose to rest in the old 

hollow,or was prevented from entering the new tree hollow. 

Conclusion 

Clearly,the surveillance of offspring either directly within the 

tree hollow (additionally offering thermoregulatory advan-


tages simultaneously; Fietz and Dausmann, 2003), or by staying 

within hearing range, does offer protection from attack 

by at least some predators even in this small species of primate. 


This study was carried out under the "Accord de Collaboration" 

between Madagascar National Parks (MNP, formerly 

ANGAP), the University of Antananarivo and the University 

of Hamburg. We thank MNP, Chantal Andrianarivo, Jocelyn 

Rakotomala, Domoina Rakotomalala, the late Olga Ramilijaona 

and Daniel Rakotondravony for their collaboration and 

support. We acknowledge the authorization and support of 

this study by the Ministère de l’Environnement, des Eaux et 

Forêts et du Tourisme, MNP and the University of Antananarivo.C.Thurner 

was an invaluable observation companion. 

The study was financed by DFG (Ga 342/14) and DAAD. 

References 

Dausmann, K.H. submitted. Spoilt for choice – Choice of 

hibernacula and its influence on predation and energy expenditure 

during hibernation in Cheirogaleus medius. In: J. 

Masters; F. Génin (eds.). Leaping Ahead. 

Dausmann, K.H.; Glos, J.; Ganzhorn, J.U.; Heldmaier, G. 2004. 

Hibernation in a tropical primate. Nature 429: 825-826. 

Dausmann, K.H.; Glos, J.; Ganzhorn, J.U.; Heldmaier, G. 2005. 

Hibernation in the tropics:lessons from a primate.J Comp 

Physiol B 175: 147-155. 

Fietz, J. 1999. Monogamy as a rule rather than exception in 

nocturnal lemurs: The case of the fat-tailed dwarf lemur, 

Cheirogaleus medius. Ethology 105: 259-272. 

Fietz,J.;Dausmann,K.H.2003.Costs and potential benefits of 

parental care in the nocturnal fat-tailed dwarf lemur (Cheirogaleus 

medius). Folia Primatol 74: 246-258. 

Fietz, J.;Dausmann,K.H.2006.Big is beautiful: fat storage and 

hibernation as a strategy to cope with marked seasonality 

in the fat-tailed dwarf lemus (Cheirogaleus medius). Pp. 

97-111. In: L. Gould; Sauther, M. L. (eds.). Lemurs: Ecology 

and Adaptation. Springer, Berlin Heidelberg New York. 

Fietz, J; Ganzhorn, J. U. 1999. Feeding ecology of the hibernating 

primate Cheirogaleus medius: how does it get so fat? 

Oecologia 121: 157-164. 

Fietz,J.;Zischler,H.;Schwiegk,C.;Tomiuk,J.;Dausmann,K.H.; 

Ganzhorn,J.U.2000.High rates of extra-pair young in the 

pair-living fat-tailed dwarf lemur, Cheirogaleus medius.Behav 

Ecol Sociobiol 49: 8-17. 

Ganzhorn,J.U.;Sorg,J.P.1996.Ecology and economy of a tropical 

dry forest in Madagascar. Primate Report 46-1, Göttingen. 

Hladik, C.M.; Charles-Dominique, P.; Petter, J. J. 1980. Feeding 

strategies of five nocturnal prosimians in the dry forest of 

the west coast of Madagascar.Pp.41-73.In:P.Charles-Dominique;H.M.Cooper;A.Hladik;C.M.Hladik;E.Pages;G.F. 

Pariente;A.Petter-Rousseaux;J.J.Petter;A.Schilling (eds.). 

Nocturnal Malagasy Primates:ecology,physiology and behaviour. 

Academic Press, New York. 

Müller, A.E. 1999. Social organization of the fat-tailed dwarf 

lemur (Cheirogaleus medius) in North-western Madagascar. 

Pp. 139-157. In: B. Rakotosamimanana;H.Rasaminanana; 

J.U. Ganzhorn; S.M. Goodman (eds.). New Directions 

in Lemur Studies. Kluwer Academic/Plenum Publishers, 

New York. 

Petter, J.J. 1978. Ecological and physiological adaptations of 

five sympatric nocturnal lemurs to seasonal variations in 

food production. Pp. 211-223. In: D.J. Chivers; J. Herbert 

(eds.). Recent Advances in Primatology. Academic Press, 

New York. 

Schwensow, N.; Fietz, J.; Dausmann, K.H.; Sommer, S. 2007. 

Neutral versus adaptive variation in parasite resistance: 

importance of MHC-supertypes in a free-ranging primate. 

Heredity 99: 265-277. 

Lepilemur feeding observations from 

Northern Madagascar 

Andrew J. Lowin 

Society for Environmental Exploration / Frontier, 50-52 

Rivington Street, London EC2A 3QP, United Kingdom, 

research@frontier.ac.uk 

Lepilemur ankaranensis is the most northerly distributed 

member of the genus Lepilemur, with a range that extends 

south from Montagne d’Ambre National Park (Mittermeier 

et al.,2008).The behaviour and ecology of Lepilemur is poorly 

understood (Ratsirarson et al.,1987);this report summarises 

some preliminary observations of L. ankaranensis. 

Observations took place in a forest fragment (09°23.6E, 

46°07.3’S) 70km south of Antsiranana (Diego Suarez), near 

the town of Anivorano, west of the Route Nationale 6. The 

site is situated approximately 4 km south of the Mt.d’Ambre 

Park limit (Fig. 1). The area is heavily degraded, with only 

pockets of secondary dry deciduous forest remaining. 

Fig. 1: Study site in northern Madagascar. 

Casual feeding observations of L. ankaranensis took place 

from August 2009 to March 2010 (excluding the month of 

December). Observations were made in the first hour after 

sun set, with animals located using a flashlight. They were 

then followed,and any feeding bouts were recorded,with the 

plant species and food item noted. During this time, 32 % of 

all observations were of L. ankaranensis feeding on fruits, 

whereas all other observations were of leaf feeding. Five 

plant families were utilized for their fruits during the study: 

Moraceae, Verbenaceae, Rubiaceae, Pittosporaceae, and one 

that was not identified. 

As Lepilemur are thought to be predominantly folivorous 

(Ganzhorn et al., 2004; Thalmann and Ganzhorn, 2003), this 

proportion of fruit consumption seems to be high as compared 

to other Lepilemur species. For example, Thalmann 

(2001) found that during their study of L.edwardsi,only 0.3 % 

of 229 feeding bouts were feeding on fruits. 

Also during this study, L. ankaranensis was observed feeding 

on fruits with both Eulemur coronatus and Eulemur sanfordi in 

the same tree,also feeding on the same fruit,with no signs of 

aggression shown between any of the animals. A second 

study took place in June 2010, six dusk-till-dawn follows 

were carried out on consecutive nights, for a total of 64.3 

hours. Again, animals were located at dusk with a flashlight 

and followed until they returned to their sleeping sites in the


morning.During this time,175 feeding observations were recorded,and 

no fruit was consumed.During this second study, 

a focal animal was observed to be chased out of a feeding 

tree by a female E.coronatus.The female E.coronatus then began 

eating the unripe fruits of the tree.It thus seems that during 

times of fruit abundance L. ankaranensis utilize fruits as a 

food resource along with several other lemur species occurring 

in the area. However, when food resources were not 

abundant in the dry season, only leaves were eaten and 

interspecific competition appears to be higher. 

On several occasions during this second study, leaf stems 

were snapped from trees and white tree exudates were consumed. 

Latex exudates are thought to be a toxic defence 

mechanism and therefore usually avoided by primates (Glander,1994),but 

latex feeding by Colobus spp.has also been observed 

(Mckey, 1978). Other lemur species, such as Phaner 

furcifer (Petter et al., 1975;Petter,1978;Thalmann,2006) and 

Mirza coquereli (Hladik,1979),are also known to feed on tree 

exudates. A review of the literature on exudate feeding in 

primates by Coimbra-Filho and Mittermeier (1977) suggested 

that tree exudates, in addition to simple sugars, protein, 

and minerals,may also provide a source of calcium.However, 

the latter authors also suggested that for most primates 

exudate feeding was rare and of little nutritional importance. 

This short report highlighted some behaviors of Lepilemur 

ankaranensis,a relatively poorly studied member of the Lepilemur 

genus. Further field work is required to examine in detail 

the previously discussed observations and to improve 

our knowledge of this species. 

References 

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exudate-eating and the "short-tusked" condition in Callithrix 

and Cebuella. Pp. 105-115. In: D.G. Kleiman (ed.). The 

Biology and Conservation of the Callitrichidae. Smithsonian 

Institution Press, Washington, D. C. 

Ganzhorn,J.U.;Pietsch,T.;Fietz,J.;Gross,S.;Schmid,J;Steiner, 

N. 2004. Selection of food and ranging behavior in a sexually 

monomorphic folivorous lemur: Lepilemur ruficaudatus. 

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five sympatric nocturnal lemurs to seasonal variations in 

food production. Pp. 211-223. In: D.J. Chivers; J. Herbert 

(eds.). Recent Advances in Primatology, Vol. 1: Behavior. 

Academic Press, New York and London. 

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Press, New York. 

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mustelinus in Northern Madagascar. Primates 28: 

119-122. 

Thalmann, U. 2001. Food resource characteristics in two 

nocturnal lemurs with different social behavior: Avahi 

occidentalis and Lepilemur edwardsi.International Journal of 

Primatology 22: 287-324. 

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History of Madagascar. The University of Chicago Press, 

Chicago. 

Thalmann, U. 2006. Lemurs - Ambassadors for Madagascar. 

Madagascar Conservation and Development 1: 4-8. 

Hypotheses on ecological interactions 

between the aye-aye (Daubentonia madagascariensis) 

and microhylid frogs of the 

genus Platypelis in Tsaratanana bamboo 

forest 

Andolalao Rakotoarison 1*, Solohery A. Rasamison 1, 

Emile Rajeriarison 2,David R.Vieites 3,Miguel Vences 4 

1Département de Biologie Animale, Université d’Antananarivo, 

BP 906, Antananarivo 101, Madagascar 

2Research assistant, Ranomafana National Park, BP 2, Fivondronana, 

Ifanadiana, Ranomafana 312, Madagascar 

3Museo Nacional de Ciencias Naturales-CSIC, C/José Gutiérrez 

Abascal 2, 28006 Madrid, Spain 

4Zoological Institute, Technische Universität Braunschweig, 

Spielmannstr. 8, 38106 Braunschweig, Germany 

*Corresponding author: andomailaka@gmail.com 

The aye-aye (Daubentonia madagascariensis) is the most distinctive 

of all lemurs.It is the only known living species of the 

Daubentoniidae (Simon and Meyer, 2001). The hands of the 

aye-aye are highly specialised,with long and slender third fingers 

that are used for precise grooming, mainly at face level, 

to get food into the mouth with rapid movements,and to tap 

on the bark of tree trunks to detect insect larvae or other 

arthropods (Goix,1993).When an aye-aye locates a cavity,it 

will anchor the upper incisors into the wood and then gnaw 

away at the wood with the lower incisors to make a pit 

(Erickson, 1995a, 1994). This unique manner of foraging for 

arthropods leaves traces of biting on the wood cover which 

are often used to ascertain the presence of the species even 

without an actual sighting (Duckworth,1993 and own observations 

of one of us, ER). During a recent herpetological inventory 

on the Tsaratanana massif in northern Madagascar, 

we noticed bamboo holes that were possibly caused or enlarged 

by foraging aye-aye,and we observed frogs living inside 

these cavities. Here we report these observations and posit 

a number of hypotheses on the possible ecological interactions 

among these species, with the goal of stimulating further 

studies. 

During a herpetological inventory in Tsaratanana (the highest 

mountain massif of Madagascar,which rises up to 2876 m 

above sea level) one of us (AR) carried out an ecological 

study on frogs of the genus Platypelis (Mycrohylidae: Cophylinae), 

from the 9th to the 22nd of June 2010. Specifically, we 

worked in a mountain forest bordering the temporary pond 

locally called Matsabory Maiky (S 14°09’04.09"- E 48°57’ 

26.06" – 2,066 m elevation) - corresponding to campsite 2 on 

the trail from Mangindrano to the Maromokotro peak. The 

observed Platypelis occupy a specific microhabitat: the species 

live and breed inside the bamboo internodes which contain 

water and are accessible through small external holes. 

These frogs have endotrophic development: their non-feeding 

tadpoles develop inside the water retained in the tree 

holes and bamboo internodes. Based on a comparison with 

type material and DNA barcoding, we ascertained that the 

encountered Platypelis belong to two species described from


the Tsaratanana massif: P. tsaratananaensis (most common) 

and the much larger P. alticola (more rare). Detailed data on 

the ecology and reproductive biology of these frogs will be 

published elsewhere. Approximately 754 bamboo trunks, at 

five different study sites, were inspected around the campsite. 

These sites each had four plots of 10 x 10 m areas. Out 

of the 754 trunks,we discovered in 162 of them,a total number 

of 204 internode segments; small rounded holes that 

were most probably made by insects like Dinoderus minutus 

(Delobel and Tran, 1993). According to these authors, 

Dinoderus minutus deposit eggs in bamboo internodes in 

which their larvae develop (Fig. 1d). 

At one of the sites (ca. 600 m east of the pond), we discovered 

some bamboo stems with remarkably different kinds of 

holes which allowed access to the hollow cavity of the 

internodes. Parts of the bamboo had been damaged in an irregular 

way. This appeared similar to what has already been 

described as typical damage caused by the gnawing activity of 

the aye-aye, whereby a freshly ripped-back piece is still attached 

and solid (Duckworth, 1993) (Fig. 1a-b). 

On 20 of the 281 bamboo trunks at this study site,we found 

similar damages,with a total of 71 holes which were more or 

less oval and measured 5.2-29.7 mm vertically and 1.5- 

8.2 mm horizontally. The diameter measurements of the 

non-damaged bamboo trunks were 5.3-54.2 mm, and those 

with holes were 5.3-48.9 mm.On several bamboo trunks we 

observed such holes in various internodes (1-6 m above the 

Fig.1:Traces of animals on the trunks of bamboo at the study 

site: (a) bamboo internode segment with an upper and a 

lower node attributed to the aye-aye; (b) segments of two 

bamboo trunks with a hole attributed to the aye-aye on the 

right and bite traces on the left; (c) traces attributed to 

aye-aye upper and lower incisors on a "virgin" bamboo trunk 

segment; (d) typical regular-shaped hole in a bamboo segment 

attributed to insects. 

Fig. 2: Animals observed within bamboo segments: (a) frogs: 

various specimens of Platypelis tsaratananaensis in one segment; 

(b) spider; (c) myriapod; (d) insect (cockroach). 

ground),and some internodes had an upper and a lower hole. 

Most importantly for the hypotheses drawn below, on some 

of the trunks without holes, we observed clear traces of 

gnawing that probably represent the upper and lower incisors 

of the aye-aye (Fig. 1b). According to our observations, 

80 % of all the holes found in the study site were caused by 

the activity of insects, and 20 % by the aye-aye. 

Bamboo internodes accessible by both kinds of holes were 

populated by Platypelis frogs as well as a variety of insects,spiders 

and centipedes (Fig. 2). At the study site where the bite 

traces ascribed to the aye-aye were discovered, the altogether 

282 holes (putatively made by insects) contained: 61 

Platypelis distributed in 24 different holes, 12 insects in 8 different 

holes, and 2 myriapods in 2 holes. In the 71 holes 

ascribed to the activity of the aye-aye, we observed 30 

Platypelis in 11 holes, 4 insects in 3 holes, and 0 centipedes. 

Based on these observations, we posit the following (partly 

alternative) hypotheses which require verification and further 

study: 

(1) We are confident that the observed marks at one of our 

study sites, similar to those noted by Duckworth (1993),are 

indeed caused by the activity of the aye-aye. Fresh bamboo 

stems are externally smooth and very strong, and it seems 

unlikely that any other mammal or even a bird could cause 

such damage. However, the possibility that these holes may 

be made by rats (such as Rattus rattus (which we collected at 

Matsobory Maiky), or Brachytarsomys) needs to be excluded 

by direct observations. 

(2) We assume that the aye-aye will typically search for bamboo 

internodes which already have small holes made by insects.This 

is because in such internodes there is a high likelihood 

of finding prey.In addition to insect larvae and other arthropods, 

tree-hole breeding frogs like Platypelis may also be 

consumed. In areas with high bamboo density, these frogs 

may constitute an import part of the aye-aye diet. If proven, 

this fact - that aye-ayes may eat frogs in addition to invertebrates 

- would be an interesting discovery in terms of 

Primatology. 

(3) Alternatively, the aye-aye may also gnaw holes into previously 

untouched bamboo segments. The bite traces we en-


countered in such "virgin" internodes support this hypothesis.Reasons 

for this might either be a search for drinking water, 

or the search and detection of insect larvae which develop 

inside these internodes and which have not yet made a 

hole to emerge. 

(4) As a fourth and highly speculative hypothesis, the aye-aye 

may gnaw holes into "virgin" bamboo segments (or increase 

the size of pre-existing holes) as part of a long-term feeding 

strategy in which such holes are produced to make the bamboo 

segment suitable for colonization by arthropods and 

frogs. This would enable the aye-aye to "harvest" its food 

during a subsequent visit to the site several days later. Obviously, 

such a foresighted feeding strategy in a basal primate 

would be of extreme interest,but we are aware that alternative 

and more probable explanations exist. 

Detailed testing of these hypotheses will require long-term 

observations in an area of dense growth of large bamboo, 

probably including the deployment of a large number of camera 

traps and possibly hair traps to obtain evidence of aye-aye 

activity. Carrying out such studies at the site in Matsabory 

Maiky is difficult. It should be noted that the Tsaratanana 

massif is difficult to access. However, alternative sites, e.g. at 

Marojejy (Duckworth, 1993) might contain a large population 

of Platypelis (albeit other species) as well, and could be 

surveyed more systematically. 


A warm thanks to the Directorate of Waters and Forests 

and the Head of the Offices of Madagascar National Parks at 

Mangindrano and Ambanja for the research permits on 

Tsaratanana. We are also indebted to the many people who 

have logistically assisted in our expedition, especially to the 

local guides (Faly and Levaovao) from Mangindrano. 

References 

Delobel, A.; Tran, M. 1993. Les coléoptères des denrées alimentaires 

entreposées dans les regions chaudes. Faune 

tropical XXXII: 98-99. 

Duckworth,J.W.1992.Feeding damage left in bamboos,probably 

by aye-ayes (Daubentonia madagascariensis).International 

Journal of Primatology 14: 927-931. 

Erickson, C.J. 1994. Tap-scanning and extractive foraging in 

aye-ayes, Daubentonia madagascariensis. Folia Primatologica 

62: 125-135. 

Erickson, C.J. 1995a. Feeding sites for extractive foraging by 

the aye-aye,Daubentonia madagascariensis.American Journal 

of Primatology 35: 235-240. 

Goix,E.1993.L’utilisation de la main chez le aye-aye en captivité 

(Daubentonia madagascariensis) (Prosimiens, Daubentoniidés). 

Mammalia 57: 171-188. 

Milliken,G.W.;Ward,J.P.;Erickson,C.J.1991.Independent digit 

control in foraging by the aye-aye (Daubentonia madagascariensis). 


Simon,E.L.;Meyer,D.,2001.Folklore and beliefs about the aye 

aye (Daubentonia madagascarienis) Lemur News 6:11-16. 

Discovery of crowned sifaka (Propithecus 

coronatus) in Dabolava, Miandrivazo, Menabe 

Region 

Josia Razafindramanana 1*, Rija Rasamimanana 2 

1Groupe d’Etude et de Recherche sur les Primates de Madagascar 

(GERP), Lot 34 Cité des Professeurs Fort Duchesne, 

Ankatso, Antananarivo 101, Madagascar. 

2Pan African Mining Madagascar (PAMM), Lot 137 II AN 

Analamahitsy, Antananarivo 101, Madagascar. 

*Corresponding author: r_josia@hotmail.com 

Key words: Propithecus coronatus, Dabolava, distribution, 

lemurs, Indridae 

The crowned sifaka Propithecus coronatus was until recently 

regarded as one of four subspecies of P. verreauxi, family 

Indridae, which occur throughout western and southern 

Madagascar (Muller et al., 2000; Mittermeier et al., 1994; Tattersall, 

1986; Wilmé and Callmander, 2006). Recent taxonomic 

revisions (Mittermeier et al., 2008) have promoted all 

four subspecies to full species status (Mittermeier et al., 

2006). However, there is considerable debate about the validity 

of P. coronatus, and especially its relationship with P. 

deckeni (Mittermeier et al.,2008),due to the physical similarities 

and close geographical distributions of these taxa,including 

apparent sympatry at some sites (e.g. Tattersall, 1986; 

Curtis et al.,1998;Muller et al.,2000;Groves,2001;Thalmann 

et al., 2002). 

P. coronatus was previously assigned to the IUCN conservation 

rating "Critically Endangered",but has since been moved 

into the "Endangered" category; nevertheless, the distribution 

range and the ecology of this species are not yet well 

understood (IUCN, 2008). Crowned sifakas are diurnal, and 

their habitat is characterised by dry deciduous forests and 

mangroves (Petter and Andriatsarafara, 1987). They live in 

groups of two to eight individuals, with home ranges from 

1.2–1.5 ha.They feed mainly on buds,green fruits and mature 

leaves (Muller,1997).It is known that they reproduce seasonally, 

with females giving birth every 2-3 years (Curtis et al., 

1998; Mittermeier et al., 2006). Compared to other lemurs, 

their reproduction rate is very slow, making recovery of 

small populations even more problematic. 

The newly discovered crowned sifaka population is situated 

at Amboloando (UTM WGS 84,N 7822351 E 580189) in the 

Commune of Dabolava in central Madagascar,and is the most 

southerly record of the species.Amboloando lies about 4 km 

from Dabolava village, and 40 km to the southeast of Miandrivazo.Amboloando 

comprises 7 ha of dry semi-deciduous, 

secondary forest that exhibits the characteristics of riverine 

forests, consisting of deciduous as well as evergreen trees 

such as Acacia sp.,Nastus sp.and Macaranga sp.The altitude is 

about 600 m above sea level,and the area is characterized by 

a clearly defined wet and dry season.The sifaka population is 

composed of a single group,which constituted six adults and 

one juvenile when first discovered in June 2009 (Razafindramanana, 

2009). One of the adult males disappeared later in 

the year, presumed dead, leaving six individuals remaining. 

The animals appear to be classic P.coronatus (Fig.1),but some 

individuals show pelage colour variation, with dark fur on 

their back and arms (Fig.2).Behavioural studies of the group 

are underway,and a preliminary community-based conservation 

program has been established at the site, involving several 

organisations including GERP, The Aspinall Foundation, 

SAHA and Pan-African Mining Madagascar.Forests in Amboloando 

and the surrounding area are heavily degraded. Different 

factors threaten the survival of this species in Madagascar:in 

contrast to the other sites such as Anjamena (Muller,2000),hunting 

does not occur in Amboloando,partly due 

to the sifaka being regarded as holy by the local people. 

Therefore,other threats such as habitat destruction through 

slash-and-burn agriculture to make way for pasture for livestock,charcoal 

production,and mining exploitation affect the 

sifaka group. 

Surveys in the vicinity of Dabolava suggest that this is the 

only group of P.coronatus remaining in that area,despite local 

people claiming that other groups were present between 5 

and 10 years ago. Therefore, it appears that habitat destruc-


Fig.1:Crowned sifaka in the Amboloando 

Forest (top). 

Fig. 2: Crowned sifaka with dark colour 

on its arms (right). 

Fig. 3: Map showing the area of discovery and survey in Dabolava. 

tion for local livelihoods has resulted in the almost complete 

extirpation of crowned sifaka in the area, probably due to a 

combination of habitat loss and food scarcity. 

Some studies describe the range of P.coronatus as broadly restricted 

to the region between the rivers Betsiboka and 

Mahavavy (Muller et al., 2000; Wilmé and Callmander, 2006), 

with a population density of 173 individuals/km² at Anjamena 

(Muller, 2000). In an analysis of the distribution of lemurs in 

central western Madagascar, Thalman and Rakotoarison 

(1994) suggested that the faunal region bounded by the 

Betsiboka,the central highlands,the river Tsiribihina and the 

Mozambique Channel can be divided into four sub regions. 

These sub regions are separated by the three rivers: Mahavavy, 

Manambaho and Manambolo, but are interconnected 

with the Bongolava Massif.The discovery of a crowned sifaka 

population in Dabolava, which is located in the south of the 

central highlands sub region, confirms the hypothesis that 

the historical range of this species might spread along the 

central highlands of Madagascar. The record of a group of 

crowned sifaka in Andranotonga, slightly north of the Mahajilo 

River, was cited by Tattersall (1986). The present report 

appears to be the first location of P.coronatus to the south of 

this river. 

The Mahajilo is a tributary of the Tsiribihina 

River, which is considered to 

represent the north-western limit of P. 

verreauxi (Mittermeier et al., 2006). P. 

coronatus is therefore unlikely to be 

found much further south or southwest 

than Dabolava. Only 80 km 

south-west of Dabolava, a population 

of P. verreauxi is known from Ambatolahy 

(SAHA,2009),which lies within 

the Ambararata/Londa protected area 

complex (Fig. 3). Another tributary of 

the Mahajilo, the Mania River, lies between 

Ambatolahy and Dabolava and 

may therefore represent the distributional 

limit of P. coronatus and P. verreauxi 

in the south of Madagascar. A 

conservation programme and restoration 

of the remaining habitat with 

the local people are needed to save 

this population of P. coronatus. 

Acknowledgments 

I thank GERP – Groupe d’Etude et de 

Recherche sur les Primates de Madagascar, 

The Aspinall Foundation, Cotswold 

Wildlife Park, SECAS, Belfast 

Zoo, Besancon Museum and Parc 

Zoologique de Paris for funding this 

research. Many thanks to Dr Jonah 

Ratsimbazafy and Tony King for discussions 

about the project implementation.I 

am grateful for permission and 

assistance in the field from the Direction 

Régionale de l’Environnement et 

des Forêts and the Commune of 

Dabolava.I also thank Pan African Mining 

Madagascar for providing accommodation 

in their lovely camp site. 

References 

Curtis, D.J.; Velo, E.-O.; Raheliarisoa; Zaramody, A.; Muller, P. 

1998.Surveys on Propithecus verreauxi deckeni,a melanistic 

variant, and P.v. coronatus in Northwest Madagascar. Oryx 

32: 157-163. Groves, C.P. 2001. Primate taxonomy. Smithsonian 

Institution Press, Washington, D.C. 

Hawkins,A.F.A.;Durbin,J.C.;Reid,D.B.1998.The primates of 

the Baly Bay area,north-western Madagascar.Folia Primatologica 

69: 337-345. 

IUCN. 2008. 2008 IUCN Red list of threatned species. 

www.iucnredlist.org. 

Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.; 

Mast,R.1994.Lemurs of Madagascar.1st ed.Conservation 

International, Washington, D.C. 

Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis E.E.; 

Langrand, O.; Ratsimbazafy, J.; Rasoloarison, R.; Ganzhorn, 


Lemurs of Madagascar. 2nd ed. Conservation International, 


Mittermeier,R.A.;Ganzhorn,J.U.;Konstant,W.R.;Glander,K.; 


J.; Mayor, M.I.; Louis Jr., E.E.; Rumpler, Y.; Schwitzer, 

C.; Rasoloarison, R.M. 2008. Lemur diversity in Madagascar. 

International Journal of Primatology 29 (6): 1607- 

1656. 

Muller,P.1997.The behaviour and ecology of the crowned sifaka 

(Propithecus verreauxi coronatus) in north west Madagascar. 

Unpublished Ph.D. thesis, University of Zurich.


Muller,P.A.;Velo,E.-O.;Raheliarisoa;Zaramody A.;Curtis.D.J. 

2000. Surveys of five sympatric lemurs at Anjamena, 

northwest Madagascar. African Journal of Ecology 38: 

248-257. 

Petter, J.-J.; Andriatsarafara. F. 1987. Conservation Status and 

distribution of lemurs in the west and northwest of Madagascar. 

Primate Conservation 8: 169-171. 

Razafindramanana, J. 2009. Propithecus coronatus on the verge 

of extinction:Help to save them:GERP,Antananarivo,Madagascar, 

6. 

SAHA. 2009. Crèation de Nouvelle Aire Protegee pour le 

complexe Ambararata/Londa. 

Tattersall,A.I.1986.Notes on the distribution and taxonomic 

status of some species of Propithecus in Madagascar. Folia 

Primatologica 46: 51-63. 

Thalmann, U.; Rakotoarison, N. 1994. Distribution of lemurs 

in central western Madagascar, with a regional distribution 

hypothesis. Folia Primatologica 63: 156-161. 

Thalmann, U.; Kümmerli, R.; Zaramody, A. 2002. Why Propithecus 

verreauxi deckeni and P. v. coronatus are valid taxa – 

quantitative and qualitative arguments. Lemur News 7: 

11-16. 

Wilme, L.; Callmander M.W. 2006. Les populations reliques 

de primates: les Propithèques. Lemur News 11: 24-31. 

Inferences about the distant past in Madagascar 

Elwyn L. Simons 

Duke Lemur Center, Division of Fossil Primates, Dept. Evol. 

Anthropology, Duke University, 3705 Erwin Road, Durham, 

NC 27705, USA, esimons@duke.edu 

From Etienne de Flacourt (1658), the following is an English 

translation, from the original French, of an entry in his book 

"Histoire de la Grande Isle Madagascar:" 

"Tretretretre or Tratratratra. It is a large animal like a calf of 

two years old, with a round head and the face of a man: the 

fore feet are like those of a monkey (or ape),and the hind feet 

also.It has curly (or frizzy) hair,a short tail and ears like those 

of a man. It resembles the "Tanacht" described by Ambroise 

Paré. It can be seen near the pond of the Lipomami [tribe] 

and in that region is where it can be found. It is a highly 

solitary animal,the people of that country have a great fear of 

it and flee from it as it also does from them." From this context,it 

is not clear whether Flacourt actually had seen this animal. 

It is well known that in Madagascar village people tend to 

name lemurs after the sounds they make, following a sort of 

onomatopoeic pattern for animal names such as occurs in 

the case of the cuckoo bird in English. For instance, the 

ground predator alarm call of species of genus Propithecus is a 

loud "si-i-fak!" cry and so the name of this animal in the Malagasy 

language is "Sifaka".The mouse lemur makes a chittering 

alarm call and has the name "T’sit-sihy".The Avahi,a nocturnal 

lemur, has one call that sounds like the word "avahi!" In the 

case of Flacourt’s animal the name Tretretretre or Tratratratra 

is definitely onomatopoeic and sounds like an alarm 

bark–it is not unlike the bark alarm call of the southeastern 

Madagascan lemur Propithecus edwardsi,or that of another lemur 

related to it, Indri indri, or even the alarm bark of the 

chimpanzee. 

Hence, I have often considered this term, or name, to be a 

"fossil" sound and it seems likely that it would have been a 

replication, by members of the Lipomami tribe of the alarm 

call of this giant lemur when they told Flacourt about the animal. 

The location of the Lipomami region in southeastern 

Madagascar is known today (Tattersall,1982).Thus Flacourt’s 

name for the animal may be the only known "fossil" sound. 

There are frequent references by various scientists all agreeing 

that Flacourt must have been describing,in the above passage, 

one of the giant extinct lemurs; but which one? All lemurs 

have hands and feet like those of monkeys, but wavy 

hair is more restricted–mainly to members of the Indriid 

group or taxonomic family [this family includes only species 

of the extant genera Propithecus,Indri,and Avahi] and incidentally 

they all have rounded ears–like those of a man. One 

large giant extinct lemur of the south and southwest of the 

island is known as Palaeopropithecus ingens and taxonomists 

generally agree that genus Palaeopropithecus is related to the 

family Indriidae (Orlando et al., 2008), where curly or frizzy 

hair occurs. A number of scientists have speculated that 

Flacourt’s Tretretretre lemur belonged to the genus Megaladapis 

(Tattersall, 1982; Mittermeier et al., 1994), which also 

occurs as fossils from the southern part of the island, but 

species of this genus have a large snout and could never be 

described as having a round head. Moreover, the distal ends 

of the nasal bones in species of this genus are elongated and 

expanded and, in life, there must have been an expanded or 

bulbous nose or even a trunk. Hence, one could never say 

that the creature had "a face of a man." Differing from Megaladapis,Palaeopropithecus 

ingens does have a rounded humanlike 

head with forward directed eyes and a small face.In addition 

to all these other features,the living species Indri indri or 

babacoot,is the only lemur that has a short tail.In addition to 

this,the most complete skeleton of Palaeopropithecus,recovered 

by a Duke expedition in 1983, and the only associated 

skeleton of this animal ever found includes a sacrum that diminishes 

posteriorly and could only hold a very small and 

short tail.In opposition to all these conclusions,it can be said 

that Palaeopropithecus ingens could not possibly be construed 

to have been the size of a calf of two years in age–nor,in fact, 

would any of the extinct giant lemurs have been that large. 

Nevertheless, it is well known that exaggeration surrounds 

stories about such little known animals, and also Malagasy 

cattle tend to be small. For these reasons it would appear 

that the Tretretretre was a Palaeopropithecus species–a conclusion 

also implied by Godfrey and Jungers (2002). A year 

later these authors reconfirm the same position (Godfrey 

and Jungers, 2003). 

Between 1994 and the year 2003 teams from the Duke 

Lemur Center excavated fossils at two caves called Akomaka 

and Ankilitelo in southwestern Madagascar on the Mikoboka 

Plateau north of Tulear. This region is Madagascar’s most 

extensive and stratigraphically thickest calcareous plateau. 

Discoveries made at both of these caves show that the 

southwestern part of the island was inhabited comparatively 

recently by several giant lemurs including Palaeopropithecus 

ingens and Megaladapis edwardsi. More importantly, these 

species lived relatively recently (Godfrey and Jungers, 2002), 

as evidenced by radiocarbon dating based on specimens 

from Ankilitelo (Megaladapis at 630 ± 50 years B.P. and 

Palaeopropithecus, 510 ± 80 years B.P. The latter of these 

dates (calculated in 1996) ranged from 1406 to 1566 years 

AD and falls into historic times. These are relatively recent 

ages,not so far from the date of Flacourt’s observation of the 

Tretretretre which could have been at any time after he was 

named Governor of Ft.Dauphin Madagascar in 1648;approximately 

350 years ago. A more recent date determined in 

2008 on a Cryptoprocta bone from Ankilitelo gives a similar 

age to that of the Palaeopropithecus, estimated as between 

1408 and 1488 (Simons,1997;Muldoon et al.,2009;Muldoon 

and Simons, 2007). This suggests that the small mammals 

accumulated more or less contemporaneously with the giant 

lemurs. [The cave name, Ankilitelo, means "at the three kili 

(tamarind) trees" but no such trees grow there now. Mala-


Fig. 1: Harpagophytum grandidieri burr or "hitchhiker" from 

southern Madagascar.These barbs are approximately 9.5 cm 

across. 

gasy caves can be named for a nearby settlement or village.At 

present there is no nearby hamlet with such a name but in 

the past there could have been. Kili trees often grow in 

villages there.] 

At this point another speculation can be introduced.Pastoral 

grass burning has reduced present day forests on the Mahafaly 

Plateau, north of Tulear, to the ridges of hills and many 

present day plant species occurring in them must be the 

same as those of 500-1000 years ago. The remaining forests 

near Ankilitelo represent both spiny thicket and succulent 

woodland regions and examples of the dry deciduous forest 

are nearby (Simons, 1997; Muldoon et al., 2009). In general, 

grass burning in that region, near the Ankilitelo cave, is arrested 

from spreading by outcrops of limestone on the 

slopes of hills. In addition, it would appear that surviving forests 

on hilltops of the Ankilitelo region remain as they were a 

few hundred years ago.The surface finds of small mammals in 

Ankilitelo (34 species) do reflect those of a few hundred 

years ago–when the giant lemurs existed and when the cave 

was serving as a natural trap–these environmental conditions 

were similar to those of the present.The fauna suggests 

that, perhaps, the region then was slightly more humid, and 

definitely the forests of the region have undergone fragmentation 

(Simons, 1997; Muldoon and Simons, 2007). Each such 

natural trap cave serves as a window in time because, as the 

solution cavities open mainly from below,the fauna that fall in 

will only begin to collect when there is a surface opening. 

More dates are being determined for fossils from Ankilitelo 

but the window concerned here did not open long enough 

ago that the 34 small mammal species are different from 

those now extant in the region or relatively nearby.The commonest 

giant lemur at Ankilitelo is the awkwardly constructed 

Palaeopropithecus that presumably could not support 

itself on all fours on the ground. Once having fallen to 

earth it would have been restricted to swimming, sloth-like 

motions and this perhaps explains why so many P. ingens fell 

into the pit. Also, Godfrey and Jungers (2003) report a Malagasy 

tradition (p. 258) that an "ogre with the body of an animal 

but the face of a human" could be made helpless on 

smooth rock surfaces. Such clumsiness of the sloth-like 

Palaeopropithecus would account both for its abundance in 

Ankilitelo and its presumed inabilities on the ground. Whatever 

the pelage of this animal was like,or indeed that of any of 

the giant lemurs, their fur could not have resisted picking up 

burrs and other hitchhikers from the southwestern forests 

as has been noted with modern lemurs (personal observa- 

tions of Michelle Sauther).A present day student of behavior, 

Sauther,has observed individual Lemur catta that had become 

entangled with the large seeds of what is often called Uncarina 

grandidieri–but more properly,because of an earlier date 

of description, this species should be assigned to genus Harpagophytum–meaning 

"snatcher plant". This plant of southwestern 

Madagascar has amazing "hitchhikers" about 2.5 to 3 

inches across each of which has 30/35 protruding spines 

approximately 1.5 inches long (see Fig.l).Each of these spines 

is, in turn, tipped by 4 recurving fishhook-like projections. 

This huge seed pod is something that it seems would only 

have evolved to be transported by a much larger animal than 

any now extant on the Island–presumably a giant lemur like 

species of such southern genera as Palaeopropithecus, Megaladapis, 

or Archaeolemur or even transportation by the elephant 

bird (see below). 

Modern botanists report that the pasty pollen of this plant is 

spread by pollen-eating beetles, who, after feeding on pollen 

from the anthers, get it all over themselves and when covered 

by pollen fly from flower to flower where pollen is 

transmitted from them to the stigma.This sort of pollination 

may be the principal fertilization process, but lemur transport 

of these seed burrs does occur today (personal observations 

of Michelle Sauther), and must have also done so in 

the past. Working at the Beza Mahfaly Special Reserve, in 

southwestern Madagascar,Sauther has seen Lemur catta individuals 

with Harphagophytum burrs stuck on the face, feet, 

and tail. However, she has not seen them attached to Propithecus 

verreauxi; a second larger, diurnal lemur species which 

occurs at Beza but is more arboreal than L. catta. These dry 

seed pods would naturally attach to the skin, not necessarily 

fur,of any passing animals and be carried while attached until 

its spines were broken enough for the seed to drop off.It was 

recently suggested that dispersal of these seed pods might 

have been carried out primarily by the extinct elephant birds 

of Madagascar (Midgley and Illing, 2009). The authors presenting 

this view argue that the mature fruit more often accumulate 

on the ground as "trample burrs" and so are more 

likely to stick to the feet of these extinct giant ratites than to 

fur of arboreal animals. I suspect, however, that the giant lemurs 

did not always stay high up in trees but were often on 

or near to the ground. The mature terminal hooks of the 

Harpagophytum (Uncarina) burr have evolved so as to attach 

to any extremity,not necessarily fur.Also it is of interest that 

these plants are often called the "Mouse-trap tree" or "Grapple 

tree". These species belong in the sesame family (Pedaliaceae) 

and typically constitute shrubs or small trees. It is 

told that Malagasy people sometimes collect and put together 

bunches of these seeds and place cheese or other attractants 

at the center of the bunch. They then use this device 

to trap rats and mice: Hence, the origin of the common 

name. 

References 

Etienne de Flacourt.1658.Histoire de la grande Isle Madagascar, 

2 vols. Chez G. de Lvyne, Paris. 

Godfrey, L. R.; Jungers, W. L. 2002. Quaternary fossil lemurs. 

Pp. 1-530. In W.C. Hartwig (ed.). The Primate Fossil 

Record:Cambridge Studies in Biological and Evolutionary 

Anthropology No.33.Cambridge Univ.Press,Cambridge, 

UK. 

Godfrey,L.R.;Jungers,W.L.2003.The Extinct Sloth Lemurs of 

Madagascar. Evol. Anth. 12: 252-263. 

Midgley, J. J; Illing, N. 2009. Were Malagasy Uncarina fruits dispersed 

by the extinct elephant bird? So. Af. J. Sci. 105 

(11/12): 467-499. 

Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.; 

Mast, R.B. 1994. Lemurs of Madagascar. Conservation 

International, Washington, D.C., USA.


Muldoon, K.M.; de Blieux, D.D.; Simons, E.L.; Chatrath, P.S. 

2009.The subfossil occurrence and paleoecological significance 

of small mammals at Ankilitelo cave,southwestern 

Madagascar. Journ. Mammology 90 (5): 1111-1131. 

Muldoon,K.M.;Simons,E.L.2007.Ecogeographic Size Variation 

in Small-Bodied Subfossil Primates From Ankilitelo, 

Southwestern Madagascar.Am.J.Phys.Anth.134:152-161. 

Orlando, L.; Calvignac, S.; Schnebelen, C.; Douady, C.J.; Godfrey, 

L.R.; Hänni, C. 2008. DNA from extinct giant lemurs 

links archaeolemurids to extant indriids.BMC Evolutionary 

Biology 8: 121. 

Simons,E.L.1997.Lemurs:Old and New.In S.M.Goodman;B. 

D.Patterson (eds.). Natural Change and Human Impact in 

Madagascar. Smithsonian Inst. Press, Washington, D.C. 

Tattersall, I. M. 1982. The Primates of Madagascar. Columbia 

Univ. Press. 

Husbandry guidelines for mouse lemurs 

at Paris Zoo 

Delphine Roullet 

Parc Zoologique de Paris, MNHN, 53 avenue de Saint Maurice, 

75012 Paris, France, roullet@mnhn.fr 

There are two species of mouse lemur in captivity in Europe: 

the grey mouse lemur, Microcebus murinus, and the Goodman’s 

mouse lemur, Microcebus lehilahytsara (Pes, 2009). The 

European captive population of grey mouse lemurs was 

established at the end of the 1960s and is now composed of 

165 individuals (778.77.10; Pes, 2009),distributed in 29 institutions. 

The population of this species is of unknown origin. 

According to recent morphological measurements (Pes, 

2009) and preliminary results of mtDNA studies (Roos, 

2008, in Pes, 2009), this population can be divided into two 

pure lineages:one composed of pure breed animals from the 

region of Vohimena (SW Madagascar),and the other of pure 

breed animals from the region of Mandena (SE Madagascar). 

A third lineage is composed of hybrids between the two pure 

lineages (Pes, 2009). 

The European captive population of Goodman’s mouse lemurs 

was established in 2005 and is currently composed of 

62 individuals (33.29; Pes, 2009) distributed in two institutions. 

The first animals were imported from the area of 

Andasibé, Madagascar. They were recognized as a new species 

when they arrived in Europe (Rübel, pers. comm.). 

The Parc Zoologique de Paris has a success story with the 

grey mouse lemurs. The most important group arrived in 

December 1990 and was composed of 79 individuals. The 

origin of the animals that arrived in the 1980s is unknown. 

According to the analysis of the European captive population 

(conducted by Tomas Pes for the ESB), the animals coming 

from Paris appear to be hybrids between the two pure lineages 

described above (Pes, pers. comm.). 

The first births occurred in 1991 only a few months after the 

arrival of the first animals.During the period of 1991 to 2004, 

when the last grey mouse lemurs eventually left the zoo after 

the closing of the nocturnal area, there had been a total of 

224 successful births (young surviving longer than two 

months), with an 86.5 % birth success rate. 1994 was the 

most prolific year with 56 successful births. The colony of 

grey mouse lemurs in Paris occasionally reached more than 

150 individuals. 

Before 2001, we didn’t know much about the animals, especially 

the composition of the groups.Eleven females were the 

founders of the colony in Paris. Since the identity of the fathers 

was not recorded, the filiations were only built from 

the females. Potentially,28 males could have been the founders 

of the colony. 

Before 2002, most of the females lived alone in small cages 

and were introduced to males (of various group sizes) only 

during the few days of oestrus. The females were kept isolated 

again afterwards.The young were separated from their 

mothers just after their weaning to join a young animals 

group. 

In 2002 we decided to implement some changes in the management 

of the colony in order to improve the wellbeing of 

the animals: to increase the space available to them, to rearrange 

the enclosures according to the wild habitat of the animals, 

to carry out enclosure enrichment, and to re-constitute 

the groups to make them more similar to the ones observed 

in the wild. The following husbandry guidelines were 

established according to the new management of the colony 

set up in 2002. 

Only single sex groups could be seen by the public.The breeding 

groups were kept in a separated building. Moreover, 

from 2002 onwards,we limited the number of births (around 

10 per year) in order to be able to keep all the animals in 

good conditions (and no longer in small cages as had been 

done in the past). 

Facility standards 

1. Enclosure 

Size: The enclosure should have a minimum total floor size 

of 4 m² with a minimum height of 2 m for both male and female 

groups.For a mother with her young,the enclosure can 

be smaller during the first month.After this period,the young 

start to explore their environment and need more space. 

Temperature: 20°C (18-22° C). Not below 18° C. Below 

this temperature, the animals enter torpor. Torpor can also 

be provoked by intense stress such as prolonged capture of 

an animal. 

Inside Humidity: 50-70 %. 

Lighting and photoperiod:Similar to that found in Madagascar 

or Europe but the photoperiod must vary during the 

year for breeding. 

Furniture: Dense environment with thin branches and 

leaves. The animals need to have many places to hide from 

people and also from each other when they live in groups,especially 

when the animals are unrelated. 

Nest box (see Fig. 1): 

Size: 12x12x12 cm 

Entry diameter: 5 cm 

It’s very important to provide one nest box per animal,in 

different places, even if they sleep together. This allows 

them to be alone if they want to be. 

Fig.1:Grey mouse lemurs (Microcebus murinus) in nest box at 

Paris Zoo. (Photo: F.-G. Grandin, MNHN)


2. Feeding and watering 

Two feeding spots are needed if there are more than two 

adults in the group.Additional feeding spots are required according 

to the size of the group.If there are not enough nest 

boxes or feeding spots,the animals can be very aggressive towards 

one another, sometimes causing serious wounds, 

mostly to the tail. One water bottle per enclosure is sufficient. 

Diet for 1 animal/day: 

Monday: 1 tea-spoon of gruel* +3folivorous pellets 

Tuesday: 1/8 of apple + 2 slices of carrot + 3 folivorous pellets 

Wednesday:1/8 of apple + 1/8 of pear + 3 folivorous pellets 

Thursday: 1 tea-spoon of gruel* +3folivorous pellets 

Friday:1/8 of apple + 2 slices of carrot + 3 folivorous pellets 

Saturday: 1 tea-spoon of gruel* +3folivorous pellets 

Sunday: 1 slice of banana + 1/8 of mango + 5 mealworms + 3 

folivorous pellets 

*Gruel composition: folivorous pellet powder + milk 

powder + baby cereals + yolks + cottage-cheese + juice of 

squeezed oranges + vitamins (every Monday). 

For overweight animals (weight >100 g):from Tuesday to Saturday 

we provide only 2 slices of carrot and 3 folivorous pellets. 

No change for Sunday and Monday. 

For underweight animals (weight


females may be more interested in food during the breeding 

season than the males which,in turn,may be more interested 

in the females (Roullet, 1998). 

The enrichment made with a branch covered in fruit juice 

was the most used by the animals:This is a branch (diameter 

of around 5 cm, length of 30-40 cm) in which two trenches 

are dug along its length. The branch is fixed onto the mesh 

roof. We apply the fruit juice in the trenches with a brush 

(Roullet,1998).The animals spend their time licking the juice 

out of the branch as they would do with sap in wild (Martin, 

1973). 

Mixed-species exhibits 

Paris Zoo experienced 2 successful combinations:with ayesayes 

(Daubentonia madagascariensis) and greater tenrecs 

(Tenrec ecaudatus).We tried to put the mouse lemurs in with 

slow lorises (Nycticebus coucang), but without success: 

wounds were observed on the mouse lemurs’ tails, so they 

were removed from the exhibit. 

References 

Martin, R.D. 1973. A review of the behaviour and ecology of 

the lesser mouse lemur (Microcebus murinus). Pp. 1-68. In: 

R.P. Michael; J.H. Crook (eds.). Comparative ecology and 

behaviour of primates. Academic Press, London, UK. 

Pes, T. 2009. The European Studbook of Grey Mouse Lemur 

(Microcebus murinus). Zoo and Botanical Garden Plzen. 

Roullet, D. 1998. Effet d’un enrichissement physique sur les 

comportements agonistiques et exploratoires de plusieurs 

groupes de microcèbes murins (Microcebus murinus) 

en captivité. DESS d’Ethologie Appliquée et de Chronobiologie 

du Comportement. Université Paris XI. 

Articles 

Diurnal lemur density in the national 

park parcel Ivontaka Nord, UNESCO 

Biosphere Reserve of Mananara-Nord 

Marta Polasky Lyons 

School for International Training, Fort Dauphin, Madagascar; 

and Carleton College, Northfield, Minnesota, USA, 

marta.lyons@gmail.com 

Abstract 

Here I present a recent diurnal lemur density study performed 

in the Biosphere Reserve of Mananara Nord, conducted 

between the dates of November 9 and 22, 2008 in 

Ivontaka Nord; part of Mananara-Nord National Park. Densities 

were calculated using transect walks, and other information 

was gathered through interviews with local people 

and national park staff. The density of Eulemur fulvus albifrons 

appears to be over twice that recorded in other areas 

(Table 1), while the density of Varecia variegata variegata 

appears to be low,perhaps due to the latter’s preference for 

undisturbed habitat and past problems with overhunting. In 

addition, Eulemur rubriventer was found to inhabit the park, 

though its range was previously thought not to extend east 

into the biosphere. Having been, in the past, under strong 

pressures from local inhabitants,Ivontaka Nord represents a 

disturbed low altitude rainforest;however this report shows 

that the lemur populations within the parcel may be recovering. 

Introduction 

Background 

The Biosphere Reserve of Mananara-Nord was created in 

1989. It is one of 533 UNESCO (the United Nations Educational, 

Scientific, and Cultural Organization) Biospheres 

around the world.The Biosphere Reserve of Mananara Nord 

covers 144,000 ha,with 23,000 ha being devoted to a terrestrial 

park,and another 1,000 ha to a marine park.The terrestrial 

national park is split into three separate parcels. Going 

from north to south these parcels are: Ivontaka Nord, 

Ivontaka Sud, and Verezanantsoro. The parcel of Ivontaka 

Nord covers an area of only 827 ha.This is small compared to 

the other two parcels; Ivontaka Sud and Verezanantsoro, 

which are 1,300 ha and 20,685 ha respectively (Fig. 1). These 

parcels are connected by forest sections that are not protected 

by the national park system (ANGAP, 2005). 

Fig.1:National Park of Mananara-Nord.Three parcels in dark 

shade,going from north to south:Ivontaka Nord (where this 

survey was carried out), Ivontaka Sud, Verezanantsoro. 

Source: MNP. 

Threats 

It is estimated that approximately 1.9 to 2.2 % of the primary 

forest within the biosphere is cleared every year, usually for 

rice cultivation through tavy (slash and burn agriculture) 

(ANGAP, 2005). Besides just destroying necessary habitat, 

this deforestation further splits already extremely fragmented 

sections of primary forest.Another concern is illegal 

(and legal) selective extraction. The population within the 

biosphere relies on wood for constructing their houses and 

fuel for cooking.In the villages most cooking is done over an 

open flame using collected wood.The wood does not simply 

go to the villages on the periphery of a forest,but is also collected 

for sale in urban centers and areas farther away from 

the forest.


Another large concern in the biosphere has to do with 

poaching. Until recently, hunting was thought to not be of 

great concern in Madagascar, being far below the menace 

caused by deforestation (Goodman et al., 2003); however, 

that is quickly proving incorrect. Unlike in other areas of the 

country where eating lemurs is considered fady (taboo), the 

Betsimisaraka ethnic group that makes up the majority of inhabitants 

within the biosphere reserve is only known to have 

fady related to the eating of Indri indri,and even this taboo has 

been shown not to be universal (Mittermeier et al., 2006). In 

the area people have classically used laly (traditional lemur 

traps) and firearms in order to hunt lemurs for consumption 

(ANGAP, 2005). Lemurs within the reserve have been a traditional 

source of protein for villagers whose diets are based 

on the staple rice. 

Site 

The field camp was situated within the town of Ambodivoandrozana 

approximately 17 km south of Mananara Nord. The 

village is inaccessible by vehicle but rests less than 2 km from 

the national park parcel of Ivontaka Nord.Between the town 

and the national park lies another forest managed by the 

COBA (communaute de base) for conservation and local subsistence 

needs. This forest is split into three zones; Beantohiravina 

(21.7 ha), Betsingiala (11.52 ha), and Ambahinkarabo 

(41.06 ha). These zones are separated by areas of 

agricultural land and secondary forest. A Gestion Contractualisée 

des forêts (GCF) in 2005 transferred certain management 

and use rights to the local community. It is mostly 

low altitude primary growth rainforest, intermixed with 

zones of savoka (secondary growth). The parcel of Ivontaka 

Nord is made up of both primary forest and disturbed habitat.From 

past species inventories,it was believed that two diurnal 

lemur species (Eulemur albifrons and Varecia v.variegata) 

and multiple nocturnal species, including Microcebus rufus, 

Lepilemur mustelinus, Avahi laniger, and Daubentonia madagascariensis 

among others inhabited this national park. 

Methods 

A survey of diurnal lemur species was conducted in the low 

altitude (between 250 and 300 m above sea level) rainforest 

in the national park of Ivotaka Nord and adjoining community-managed 

forest (GCF).The survey was conducted from 

November 9-22,2008.Within the national park parcel,a preestablished 

2 km transect set up by ANGAP was followed,in 

addition to other transects along preexisting paths throughout 

the two forests. Additional permanent transects were 

not established because of time constraints and the desire to 

limit the impact on the habitat. Instead, distance was measured 

by walking at a constant pace along preexisting paths 

(approximately 900 m per hour). This pace was calculated 

using the 2 km transect, and variation in speed was used to 

create a range for densities. The total of 17 transects were 

walked, ranging from 600-2,500 m in length, with a total distance 

walked of approximately 30 km. 

For each transect the date, start and end time, weather, and 

location were recorded.When a lemur was spotted,the species 

was noted along with the group size,time of day,distance 

on transect, distance from path, habitat type, and the GPS 

coordinates. Density was then calculated by the number of 

individual lemurs/area. Area was based on the total length of 

transects walked multiplied by double the average perpendicular 

distance of lemur from the path (Whitesides et al., 

1988; Norscia et al., 2006). Common methods for this type 

of primate study include using a 50 % criterion for falloff 

distance based on histograms (Whitesides et al., 1988; John- 

son and Overdorff,1999;Erhart and Overdorff,2008) or the 

program DISTANCE (Quemere et al., 2010). However, given 

time constraints it was not possible to collect an adequate 

amount of data to perform these tests.Based on earlier studies 

on similar species in similar habitats we used a falloff distance 

of 20 m (Irwin, 2001). In addition to transect walks, 

interviews were conducted with employees of the ANGAP 

office of Mananara Nord and local people of the town of Ambodivoandrozana. 

Within the village, the interviews were 

done with prominent members of the community including 

members of Slow Food (an agriculture movement within the 

biosphere) and the COBA. 

Results 

The observed population density for Eulemur albifrons was 

46.13 ± 2.32 individuals/km 2 within the park (Tab.1).The average 

group size observed was 7 individuals. Multiple groups 

were observed with females carrying babies on their back, 

and overall this species was found in a variety of different 

habitats, both dense and sparse primary forest, and even on 

the edge of secondary growth. Through a combination of 

data collected in the forest and interviews with local people, 

we found that though Eulemur albifrons frequents multiple 

habitats,even leaving the forest to eat crops,they mainly rest 

within the National Park. 

Two other diurnal lemur species were observed within the 

parcel, Varecia variegata variegata and Eulemur rubriventer. V. v. 

variegata was observed to have a density of 1.06 ± 0.02 individuals/km 

2 (Tab. 1). This species was only observed on one 

occasion and only one individual was seen. According to the 

local guide and vice president of the COBA,eight individuals 

of this species exist within the parcel, a group of five and a 

group of three (F. Frejes, personal communication). This implies 

a density of 0.97 individuals/km 2.From traces observed 

on the ground (eaten fruit) and calls heard,it seems clear that 

this lemur spends the majority of its time within the limits of 

the parcel as it prefers deep valleys with tall trees; a habitat 

not found in the more disrupted community managed forest. 

Tab.1:Density of diurnal lemurs within the parcel of Ivontaka 

Nord. Densities were calculated of the three species observed 

within the parcel, Eulemur albifrons, Eulemur rubriventer,and 

Varecia v.variegata.A 50 % falloff distance was used 

for perpendicular distances over 20 m (Whitesides et al., 

1988; Irwin et al., 2000) Error bars represent the error in 

pace of transect walked and error in observation of animals 

from path. 

Species Number of 

individuals 

encountered 

Eulemur 

albifrons 

Eulemur 

rubriventer 

Varecia v. 

variegata 

Number of 

groups 

encountered 

Calculated density 

(individuals/km2) 

31 5 46.13 ± 2.32 

3 1 6.40 ± 0.38 

1 1 1.06 ± 0.02 

Eulemur rubriventer, being a species believed to live west of 

the biosphere in higher altitude rainforest, was thus not on 

the initial list of lemurs to be found in the area.From the one 

sighting of three individuals in a dense part of the primary 

forest, the calculated density is 6.40 ± 0.38 individuals/km 2 

for the parcel (Tab. 1). No sightings were made of this individual 

outside of the parcel,and as its eating habits closely mirror 

that of E.albifrons,it was not possible to tell the difference 

between traces found on the ground.


Discussion 

The parcel of Ivontaka Nord is considered to be an example 

of disturbed primary forest.Anthropogenic effects are much 

stronger in Ivontaka Nord than in other parcels,because it is 

the smallest of the three, and the closest to the town of 

Mananara. Before the creation of the national park, the area 

of Ivontaka Nord was frequently used as a place to grow 

crops through tavy,harvest wood for construction and cooking,and 

hunt lemurs.For these reasons,the lemur population 

of the parcel was at one time diminished to the point of localized 

extinction of the critically endangered Varecia v.variegata, 

in addition to the localized extinctions of Indri indri and Hapalemur 

griseus (J. Betsiahilika, personal communication). Yet, 

because it is connected through a corridor of community 

managed forest to the less disturbed parcel of Ivontaka Sud 

(Fig.1),in 2002,V.v.variegata migrated into the GCF corridor 

between Ivontaka Sud and Ivontaka Nord, and in 2005 these 

lemurs could again be found within the parcel. However, the 

species of Indri indri and Hapalemur griseus are not believed to 

currently inhabit the parcel, though they can be found in the 

other two parcels within the biosphere (J. Betsiahilika, personal 

communication). 

Within the parcel, there was convincing evidence to suggest 

that the lemurs might be recovering even more than previously 

known. During the two weeks of transects, Eulemur 

rubriventer was observed within the parcel, even though its 

recorded range ends west of the biosphere reserve. In addition,traces 

of the distinct eating habits (shredding the stems 

of tall plants whilst stripping off foliage) of Hapalemur griseus 

were found within the savoka. Additionally, there were reports 

of Indri indri spending time in the corridor between 

Ivontaka Sud and Ivontaka Nord (J. Betsiahilika, personal 

communication). 

Through field observations, E. albifrons was found to have a 

population twice that found in other areas such as Masoala 

(Mittermeier et al.,2006).They frequent both the parcel and 

community forest. During the season in which this study 

took place,when most of the fruit within the parcel is not yet 

ripe, many Eulemur albifrons were reported by local villagers 

to be exiting the parcel in search of other cultivated fruits 

like lychee and banana. V. v. variegata on the other hand 

appeared to not leave the protected parcel.It is possible that 

they adapt their feeding habits based on the season and thus 

do not eat cultivated fruits (Ratsimbazafy, 2002). 

The observed density of V. v. variegata was not congruent 

with the report of the local guide (F.Frejes,personal communication). 

This species, like Eulemur rubriventer, was observed 

on only one occasion. Because of time constraints, surveys 

had to be conducted over a two week period and with only 

the use of one field team. For more complete and definitive 

findings, a survey needs to be done over a longer period of 

time, possibly across different seasons. Also, it would be 

beneficial to conduct the same type of study after dark. 

Finally, it would be helpful to determine lemur densities in 

the other two parcels, and the community managed forests 

in-between. 

Conclusion 

It is currently of the utmost importance to make sure that 

the integrity of forested sections outside the realm of the 

parcel remains protected.These areas represent a buffer between 

the fragile low altitude rainforest and an ever expanding 

human population. The corridor formed by these community-managed 

forests, between the three national park 

parcels, is a priority area. It has already been shown to provide 

a bridge to facilitate migration between the three parcels,which 

is critical in a country where forest fragmentation 

is proving detrimental to the gene flow of lemurs (Louis et al., 

2006). 

Through the work of the biosphere reserve,significant steps 

have already been taken to encourage conservation. Both 

government officials and villagers appear to be working together 

to promote a healthy ecosystem. Already with the 

promotion of crops such as vanilla,cloves,and coffee,as an alternative 

to other more environmentally negative livelihoods, 

villagers say they have seen an improvement both in 

their lives and the forest health (Desana and Berger,personal 

communication). However, people have needs; the agricultural 

inhabitants of these rural areas need both to grow their 

food and to have wood and other materials for performing 

everyday tasks. It is possible to improve both the lives of the 

people in the area and decrease their negative impact on 

their environment through simple strategies such as rice intensification 

and promoting more efficient cooking methods. 

Maybe then populations of Indri indri, Hapalemur griseus, and 

Propithecus diadema,all species that at one time inhabited the 

area, will return to the parcel of Ivontaka Nord. 


This study was carried out under a Memorandum of Understanding 

between SIT and ANGAP Mananara for an internship 

for the author in November 2008.SIT thanks the Ministry 

of Higher Education and Scientific Research and the University 

of Antananarivo for the ongoing collaboration under 

which SIT Study abroad operates. This study could not have 

been possible without the help of Barry Ferguson and Jim 

Hansen of SIT in addition to the staff in the ANGAP office of 

Mananara, including Willy Mora, Jocelyn Bezara, Justin Besiahilika, 

and Jean Cristophe Josoa. 

References 

ANGAP. 2005. Plan de Gestion de la Conservation-Parc 

National Mananara Nord. 

COBA FMAA. 2005. Fifanekena Famindra-Pitantanana ny 

Atiala sy ny Harena Voajanahary azo Havaozina ao Ambodivoandrozana. 

Erhart, E.M.; Overdorff, D.J. 2008. Population demography 

and social structure changes in Eulemur fulvus rufus from 

1988 to 2003.American Journal of Physical Anthropology 

136: 183-193. 

Goodman, S.; Raselimanana, A. 2003. Hunting of wild animals 

by Sakalava of the Menabe region: a field report from 

Kirindy-Mite. Lemur News 8: 4-6. 

Irwin, M.T.; Samonds, K.E.; Raharison, J. 2001. A biological inventory 

of the lemur community of Réserve Spéciale de 

Kalambatritra, south-central Madagascar. Lemur News 6: 

24-28. 

Johnson, S.E.; Overdorff, D.J. 1999. Census of brown lemurs 

(Eulemur fulvus sspp.) in southeastern Madagascar: 

Methods-testing and conservation implications. American 


Louis, E.E. jr.; Coles, M.S.; Andriantompohavana, R.; Sommer, 

J.A.; Engberg, S.E.; Zaonarivelo, J.R.; Mayor, M.I.; Brenneman, 

R.A. 2006. Revision of the mouse lemurs (Microcebus) 

of Eastern Madagascar. International Journal of 

Primatology 27: 347-389. 

Mittermeier, R.; Tattersall, I.; Konstant, W.R.; Nash, S.D. 2006. 

Lemurs of Madagascar. Conservation International, Washington 

D.C. 

Norscia,I.;Rahanitriniaina,O.G.;Jolly,A.;Donati G.2006.Preliminary 

survey of lemur density in the semimontane rainforest 

of Anka,Fort-Dauphin region.Lemur News 11:14- 

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Quemere, E.; Champeau, J.; Besolo, A.; Rasolondraibe, E.; 

Rabarivola,C.;Crouau-Roy,B.;Chikhi,L.2010.Spatial Variation 

in Density and Total Size Estimates in Fragmented 

Primate Populations: The Golden-Crowned Sifaka (Propithecus 

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Distribution of Prolemur simus north of 

the Mangoro-Nosivolo River – how far 

north do we really have to look? 

Rainer Dolch 1*, Erik R. Patel 2, Jonah H. Ratsimbazafy 

3,4, Christopher D. Golden 5, Tianasoa Ratolojanahary 

1, Jean Rafalimandimby 1, Jonathan L. Fiely 1 

1Association MITSINJO, Lot 104 A, Andasibe 514, Madagascar 

2Cornell University, 211 Uris Hall, Ithaca, NY 14850, USA 

3Durrell Wildlife Conservation Trust,BP 8511,Antananarivo 

101, Madagascar 

4GERP, 34 Cité des Professeurs, Fort Duchesne, Antananarivo 


5University of California, Department of Environmental 

Science,Policy and Management,Mulford Hall #3114,Berkeley, 

CA 94720, USA 

*Corresponding author: rdolch@gmx.de 

Introduction 

While the Mangoro-Nosivolo river system is a recognized 

biogeographical divide for several lemur species (Goodman 

and Ganzhorn, 2004), this pattern does not hold for the 

Greater Bamboo Lemur (Prolemur simus). Findings from 

numerous subfossil sites indicate that the historical distribution 

of P. simus, now one of the rarest Malagasy primates 

(Wright et al.,2008,2009),once encompassed most of Madagascar 

(Godfrey and Vuillaume-Randriamanantena, 1986). 

During the last 150 years, documented sightings of the species 

became more and more scarce and by the middle of the 

last century it was already feared extinct (e.g. Napier and 

Napier, 1967). 

More recent discoveries came solely from southeastern 

Madagascar (Petter et al., 1977; Meier and Rumpler, 1987), 

which led to the unspoken assumption that P.simus had been 

extirpated from the rest of the island. Despite the fact that 

the last collected specimen of P. simus had come from 

Mananara in 1876 (Godfrey and Vuillaume-Randriamanantena, 

1986), not a single individual had been found north of 

the Mangoro river for more than 130 years, before Dolch et 

al.(2004,2008) rediscovered the species in Torotorofotsy,in 

the commune of Andasibe. 

In order to investigate further into the distribution and abundance 

of P. simus north of the Mangoro, several extensive 

surveys have recently been, and are currently being, conducted 

(King and Chamberlan, 2010). As preliminary results 

of these surveys are trickling in, accounts of P. simus from 

inhabitants of these regions also multiply rapidly and await 

verification. 

We do not aspire to anticipate survey results,but we believe 

that summarizing our current knowledge of P.simus north of 

the Mangoro is crucial for the planning of future surveys and 

conservation strategies. 

Methods 

We gathered and compiled information on P. simus north of 

the Mangoro deduced from our own research,from reports 

that villagers brought to our attention, and from anecdotal 

evidence in existing literature.The ten localities from where 

information was available included (from south to north) the 

Marolambo area, the Brickaville area, the western parts of 

the Ankeniheny-Zahamena Corridor (CAZ), Zahamena, the 

Soanierana-Ivongo area, Ambatovaky, Marotandrano, Mananara, 

Makira, and Marojejy (Fig. 1). 

Fig. 1: Localities from where information on P. simus was collected. 

Results 

Results of our compilation are summarized in Tab.1.Of all localities 

north of the Mangoro examined,two (Brickaville and 

western CAZ) have steadfast records of P.simus based on independently 

confirmed sightings. One locality (Soanierana- 

Ivongo) has a record based on a single observation, whereas 

for four others (Marolambo, Zahamena, Ambatovaky, Makira) 

evidence is only based on reports of villagers. The final 

three (Marotandrano, Mananara, Marojejy) do not have any 

records. Details are given below. 

Tab. 1: Potential localities for P. simus north of the Nosivolo- 

Mangoro. 

Region evidence based on 

Marolambo area reports by villagers 

Brickaville area sightings, confirmed 

Western CAZ sightings, confirmed 

Zahamena reports by villagers 

Soanierana-Ivongo area sightings, unconfirmed 

Ambatovaky reports by villagers 

Marotandrano (no evidence) 

Mananara (mo evidence) 

Makira reports by villagers 

Marojejy (no evidence) 

Marolambo area. A hotspot of endemic fish species richness, 

the Nosivolo and lower Mangoro rivers have recently 

received increased attention by researchers and conservationists 

alike.While working in this area,we received several 

accounts of villagers on P. simus from 2006-2009. Reports 

claiming the occurrence of P. simus come from 4 communes 

along the Nosivolo-Mangoro river, and focus (from west to 

east) on Ambohimilanja, Betampona, Marolambo and Ambinanidilana. 

In order to verify these accounts, a preliminary 

survey is currently being conducted within The Aspinall 

Foundation’s "Saving Prolemur simus" project (Ratsimbazafy, 

2010).


Brickaville area. A similar survey has already been conducted 

for the Brickaville area. A total of 6 sites in isolated 

fragments in the communes (from west to east) of Fanasana, 

Anivorano and Fetraomby have been found containing P. 

simus. Details are given by Ravaloharimanitra et al. (2010). 

Western CAZ.In the course of the same survey,12 sites in 

the western parts of the Ankeniheny-Zahamena corridor 

have been found containing P. simus (Ravaloharimanitra et al., 

2010). These findings follow earlier reports from villagers 

that had claimed its presence (e.g.Schmid and Alonso,2005). 

The communes where P. simus has been confirmed include 

(from south to north) Andasibe, Morarano-Gare, Fierenana, 

and Didy. 

Zahamena. Ganzhorn (2004) states that a report of the 

presence of P. simus in the PN Zahamena was brought to his 

attention in 1995, but that it "was questioned and eventually 

withdrawn".Information on a possible occurrence of P.simus 

in Zahamena, obviously derived from that report, is mentioned 

in Godfrey et al. (1997). 

Soanierana-Ivongo area. In their little noticed bulletin, 

the Association de Défense de la forêt d’Ambodiriana report 

that trainee Coralie Ebert, while studying the woolly 

lemurs (Avahi laniger) of this forest, claims to have observed 

an individual of Prolemur simus (ADEFA, 2009). An earlier 

lemur survey of the area (Beaucent and Fayolle, 2008) has 

not yielded evidence of P.simus.The forêt d’Ambodiriana lies 

just 30 km to the north of Soanierana-Ivongo,a region where, 

according to Mittermeier et al. (2006), halogodro and bokombolobe 

are still used as local names for P. simus. 

Ambatovaky.Ambatovaky is an area that has received only 

little attention due to its difficult accessibility. In the early 

1990s, a lemur survey was conducted by Evans et al. (1993- 

1994). They did not find tangible evidence for P. simus, but 

state that "local people indicated that there existed until recently 

a lemur which fed on giant bamboo along the Sandrangato 

and/or Marimbona rivers,known as alakoto or halokoto". 

Marotandrano.Lying to the northwest of Ambatovaky,Marotandrano 

has even received less attention than the former. 

A lemur survey by Ralison (2006) did not indicate presence 

of P. simus. 

Mananara. The last specimen of P. simus to be collected 

from north of the Nosivolo-Mangoro river came from an 

area close to Mananara (Godfrey and Vuillaume-Randriamanantena,1986).Although 

the exact collection locality can not 

be traced (due to unsuited transcription of its name by the 

collector J.P.Audebert),the assumption that P.simus may still 

occur in the forests around Mananara was still put forward 

by Nicoll and Langrand (1989). No evidence for P. simus in 

Mananara has been produced since. 

Makira. Being Madagascar’s largest continuous rainforest 

(317,000 ha), lemur surveys in Makira are not easy to conduct. 

Two years of intensive surveys by Rasolofoson et al. 

(2007) and Ratelolahy and Raivoarisoa (2007) have not uncovered 

any evidence of P.simus.Similarly,during seven years 

of relying on trusting relationships with hunters, Golden 

(2009) has not come across P. simus among the 23 mammal 

species hunted for consumption throughout southern,western, 

northern, and eastern Makira. However, villagers living 

adjacent to the newly discovered Antohaka Lava forest at the 

edge of northeastern Makira (20 km south of Andrakata on 

Marojejy’s southeastern border) have reported recent sightings 

of a large bamboo lemur with ear tufts known locally as 

bokombolobe.Unfortunately,several months of systematic lemur 

surveying of the Antohaka Lava forest between August 

and December 2009 did not confirm these reports, despite 

an exceptional primate diversity documented inthat area 

(Patel, 2009). 

Marojejy. A lemur survey by Sterling and McFadden (2000) 

found no evidence of P. simus. Alleged observations of bamboo 

lemurs other than Hapalemur griseus by tourists may be 

attributed to the possible presence of H. occidentalis, rather 

than P. simus (R. Mittermeier, pers. comm.). Moreover, during 

nine years of research on Propithecus candidus in Marojejy,no 

local reports or sightings of P.simus have been received (Patel, 

2009). 

Discussion 

Despite the scarcity of information, growing evidence supports 

that P. simus may still be widespread in Madagascar 

north of the Mangoro river. Since the species occupies large 

home ranges (Dolch et al., unpubl. data), appears to travel at 

night due to possible cathemerality (Santini-Palka, 1994), is 

cryptic, and often silent when unhabituated, it is conceivable 

that it has been overlooked in the past.However,because the 

P.simus vocal repertoire is distinct and extensive (Bergey and 

Patel,2008) and its feeding traces on giant bamboo (Cathariostachys 

madagascariensis) are unmistakable,attention to such 

indirect evidence of P. simus presence should be focused 

upon in all surveys. 

The report from Makira, if confirmed, is especially interesting, 

since the northernmost former record for the species 

(other than from subfossils) comes from Antongil Bay 

(Schwarz, 1931). 

Our experience shows that accounts of villagers are mostly 

reliable, and that people usually have a good sense of what 

animal species do or do not occur in their vicinity.Therefore, 

integrating local people is crucial for further studies into 

Prolemur distribution. Logically, collaboration with local people 

is one conservation recommendation given by the Prolemur 

Conservation Working Group (Madagascar Fauna 

Group, 2010). 

Based on information compiled,we tentatively conclude that 

P. simus is still more widespread than previously thought. 

Without sufficient data, given persisting threats to the habitats 

in which it occurs and our incomplete understanding of 

habitat requirements for the species,we do not dare say that 

a larger distribution area contributes in any way to relieving 

the species from extinction pressure. Unfortunately, P. simus 

still has to be considered one of the most threatened primates 

in the world. 


We thank Tokiniaina Hobinjatovo for helping with literature 

research and Coralie Ebert for additional information. We 

thank all individuals that have shared their observations and 

made available the information presented here. We would 

also like to thank the Margot Marsh Biodiversity Fund, and 

the National Geographic Society Conservation Trust Award 

#C135-08. 

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Enquête préliminaire de la distribution 

des lémuriens de bambou dans et autour 

du Corridor forestier Fandriana-Vondrozo, 

Madagascar 

Andry Rajaonson 1,2, Maherisoa Ratolojanahary 2, 

Jonah Ratsimbazafy 1, Anna Feistner 3, Tony King 2* 

1Groupe d’Etude sur les Primates de Madagascar (GERP), 

Lot 34 Cité des Professeurs, Fort Duchesne, Ankatso, Antananarivo 


2The Aspinall Foundation, BP 7170 Andravoahangy, Antananarivo 


3Centre ValBio, BP 33 Ranomafana, Ifanadiana 312, Madagascar 

(present address: anna@feistner.com) 

*Corresponding author: tonyk@aspinallfoundation.org 

Mots-clés: Prolemur simus, Hapalemur aureus, Varecia variegata 

editorum, bambou 

Introduction 

Le Grand Hapalémur (Prolemur simus) mangeur de bambou, 

est classé comme étant "gravement menacé CR" sur la liste 

rouge de l’Union Internationale pour la Conservation de la 

Nature (UICN,2009).Il est aussi l’une des quatre espèces de 

Madagascar faisant partie des 25 primates considérés 

comme les plus menacés au monde (Mittermeier et al.,2007, 

2009). Des individus capturés dans les années 1800 venaient 

d’une région plus étendue, ce qui laisse supposer que son 

habitat a diminué.Les sites de subfossiles avec des squelettes 

identiques à celui de P. simus sont nombreux, impliquant une 

distribution encore plus vaste à une époque très ancienne 

(Godfrey et Vuillaume-Randriamanantena, 1986; Godfrey et 

al., 2004). 

Actuellement, la distribution géographique de l’espèce semble 

très étroite.On pense que P.simus est seulement présent 

dans quelques fragments de forêt tropicale humide près de la 

côte Est de Madagascar (Mittermeier et al.,2006 ;Dolch et al., 

2008;Wright et al.,2008).Wright et al.(2008) résume la crise


actuelle de l’espèce: sur les 70 localités étudiées, la présence 

de P.simus est confirmée seulement sur 11 d’entre elles,à une 

altitude comprise entre 121 et 1600 m. 

La plupart des sites connus abritant P.simus se trouvent dans 

ou autour du Corridor Fandriana-Vondrozo, dans le Sud-Est 

du pays (Andriaholinirina et al.,2003;Meier et Rumpler,1987; 

Sterling et Ramaroson, 1996; Wright et al., 1987, 2008; Mittermeier 

et al., 2009). Deux autres espèces de lémuriens de 

bambou vivant dans le corridor, Hapalemur aureus et H. 

griseus, sont aussi menacées (Mittermeier et al., 2006; IUCN, 

2009).Notons que P. simus est le plus grand des lémuriens de 

bambou,avec un pelage gris brun (comme H. griseus).Il se distingue 

facilement des deux autres espèces par des touffes de 

poils blancs sur les oreilles. Sa face est aussi plus allongée, et 

on le trouve souvent au sol, alors que les autres espèces y 

sont rarement (Wright et al., 1987). Selon l’étude de Tan 

(1999) à Ranomafana, le régime alimentaire de P. simus est 

constitué à 95 % d’une seule espèce de bambou Cathariostachys 

sp. (ou volohosy dans le dialecte local malgache), 3 % 

d’autres espèces de bambous et de graminées,0,5 % de fruits 

et 1,5 % d’autres éléments (principalement de la terre et des 

champignons). 

La présente étude a été organisée dans le cadre du Projet 

Varibolomavo proposé par The Aspinall Foundation (TAF). 

Ce projet veut mettre en place des actions rapides, efficaces 

et collaboratives pour sauver Prolemur simus. Plus précisément, 

le deuxième objectif du projet est d’organiser une 

étude de la distribution et de l’abondance de P. simus (TAF, 

2008, 2009; King and Chamberlan, 2010). Par conséquent, le 

but de cette étude était de contribuer à réaliser ces objectifs, 

dans et autour du corridor Fandriana-Vondrozo, par a) la 

récolte des connaissances indigènes locales sur les distributions 

des lémuriens; et b) la recherche des signes de présence 

des lémuriens de bambou. Nous présentons ici un 

résumé des résultats de l’étude, exposés de façon plus 

détaillée par Ratolojanahary et al. (2009). 

Méthodes 

Entre les 27 novembre 2008 et 25 mai 2009, nous avons enquêté 

dans 14 zones situées dans et autour du Corridor 

Fandriana-Vondrozo (Tab. 1, Fig. 1). Pour chaque commune, 

des entretiens avec les autorités locales ont eu lieu.Des collaborations 

avec ces personnes ont permis d’organiser les 

réunions villageoises pour mener les enquêtes participatives, 

à l’aide de photos des espèces de lémuriens supposées 

coexister dans ce couloir forestier (Prolemur simus, Hapalemur 

aureus, H. griseus, Eulemur rufus, E. rubriventer, Propithecus 

edwardsi, Varecia variegata editorum, Microcebus rufus, 

Cheirogaleus major, Avahi laniger, Lepilemur microdon, Daubentonia 

madagascariensis). Les appellations locales des différentes 

espèces connues par les communautés villageoises 

ont été relevées lors de chaque enquête.De plus,nous avons 

utilisé la méthode de cartographie participative (Jones et al., 

2005) durant la réunion dans les communes de Mahazoarivo, 

Iandraina, Sahamadio et Evato. Suite aux résultats des enquêtes 

villageoises, nous avons visité des forêts et sites 

intéressants dans la région, toujours accompagnés par un 

guide local et des agents de recherche du Centre ValBio de 

Ranomafana. Nous nous arrêtions tous les 25 mètres pour 

relever la localisation des bambous et des espèces de lémuriens, 

ainsi que les signes de présence de ces dernières. 

La présence des espèces de lémuriens était révélée soit par 

l’observation directe (animal vu), soit par l’observation indirecte 

(signes de nourrissage,excréments ou vocalisation).La 

recherche des signes de nourrissage des lémuriens de bambou 

était faite dans les zones de bambous, et les signes 

Tab. 1: Les zones visitées pendant l’enquête. 

Zone Sites visités Dates 

Zones situées dans le corridor forestier 

Ambendrana 1 27-28 nov 2008 

Amindrabe 1 29 nov - 1 déc 2008 

Ambodiara 1 11-13 déc 2008 

Antarehimamy 1 14-16 déc 2008 

Antaranjaha 4 

29 jan - 6 fév 2009 (dont Tsianivoho 

et Ambolomadinika) 

Manambolo 1 9-11 mai 2009 

Zones situées autour du corridor forestier 

Mananjary 4 14-16 jan 2009 

Sahalanona 9 17-23 jan 2009 

Manakara 1 25 jan 2009 

Mahazoarivo 2 18-19 mai 2009 (dont Ifasy) 

Iandraina 1 21 mai 2009 

Sahamadio 1 22-23 mai 2009 

Evato 0 24-25 mai 2009 

Mahafasa 0 25 mai 2009 

14 zones 27 sites 27 nov 2008 - 25 mai 2009 

Fig. 1: Les sites visités lors de l’enquête dans et autour du 

Corridor forestier Fandriana-Vondrozo. 

étaient examinés précautionneusement afin d’identifier l’espèce 

qui en était responsable.Prolemur simus préfère surtout 

les bambous de grand diamètre, et les parties de bambou 

privilégiées varient avec les saisons.Entre juillet et novembre, 

P. simus consomme principalement la moelle tendre de 

bambou géant (Tan,1999),après avoir ouvert la tige en deux 

et l’avoir déchirée en petits morceaux (Wright et al., 1987). 

Ainsi, l’échantillon à rechercher devrait être des tiges déchirées 

sans ou avec peu de moelle.Par contre,entre novembre 

et avril, il se concentre sur les jeunes pousses (Tan, 1999), 

donc les échantillons devraient être des bouts de jeunes 

pousses de bambou géant. Avec de l’expérience, il est égale-


ment possible de distinguer les signes laissés sur les feuilles 

de bambou. P. simus se nourrit des jeunes feuilles matures et 

ne consomme pas la partie tranchante de la feuille. Par 

contre, Hapalemur aureus et H. griseus consomment uniquement 

la base de la feuille, H. aureus déchirant la gaine de 

chaume à l’aide de ses dents avant de manger les branches. 

Résultats et Interprétations 

Espèces de lémuriens recensées 

Les populations locales ont souvent des noms vernaculaires 

distincts pour les différentes espèces de lémuriens.En outre, 

nous avons remarqué qu’ils ne pouvaient pas toujours identifier 

les espèces sur les photos, alors qu’ils pouvaient les reconnaître 

dans la nature.Les informations récoltées lors des 

enquêtes villageoises doivent donc être utilisées avec précaution 

et sont toujours à vérifier sur le terrain. Durant les 

vérifications, nous n’avons trouvé qu’un seul site présentant 

des signes de nourrissage de Prolemur simus (Tab. 2). Par 

contre, nous avons effectué 39 observations (directes et 

indirectes) de Hapalemur griseus, dans 10 des 12 zones visitées 

(Tab. 2), et huit observations (indirectes) de H. aureus 

(Tab.3).Toutes ces observations ont été faites dans les zones 

situées au sein du corridor forestier, mais aucune dans ses 

alentours (Tab. 2). Cette espèce a laissé des signes de nour- 

Tab. 2: Espèces de lémuriens rencontrées dans chaque zone. 

Zone Hapalemur Hapalemur Prolemur Autres espèces 

griseus aureus simus 

Zones situées dans le corridor forestier 

Ambendrana vu & signes signe 

E rubriventer (vu) 

Microcebus sp. (nid) 

Amindrabe vu & signes signe 

signe P. edwardsi (vu et entendu) 

(environ 1 an) D. madagascariensis (signes) 

Ambodiara signes signes V. variegata (entendu) 

Antarehimamy signes signes V. variegata (entendu) 

Antaranjaha signes 

signes & 

entendus 

V. variegata (entendu) 

E. rufus (vu) 

Manambolo signes E. rufus (vu) E. rubriventer (vu) 

Zones situées autour du corridor forestier 

Sahalanona vus & signes M. rufus et A. laniger à vendre 

Mananjary vus & signes 

Microcebus sp. (nid) 

Cheirogaleus sp. (signes) 

Manakara E. rufus (vu) A. laniger (vu) 

Mahazoarivo vus 

Iandraina signes 

Sahamadio signes 

Tab. 3: Observations de Hapalemur aureus faites pendant l’étude. 

Zone Remarque Latitude Longitude Altitude (m) 

Ambendrana signe de nourrissage S 21º 22’ 22.7" E 047º 20’ 46.5" 1182 

Amindrabe signe de nourrissage S 21º 24’ 13.1" E 047º 22’ 47.7" 1070 

Ambodiara signe de nourrissage S 21º 53’ 27.0" E 047º 21’ 18.9" 825 

Antarehimamy signe de nourrissage S 21º 54’ 47.5" E 047º 20’ 38.4" 1074 

Antaranjaha signe de nourrissage S 21º 58’ 20.3" E 047º 20’ 16.7" 828 

Antaranjaha entendu des cris S 21º 58’ 25.3" E 047º 20’ 17.7" 783 

Antaranjaha signe de nourrissage S 21º 58’ 39.1" E 047º 19’ 43.8" 786 

Manambolo signes de nourrissage S 22º 04’ 06.2" E 046º 59’ 27.5" 1238 

Tab. 4: Observations de Varecia variegata faites pendant l’étude. 

Zone Remarque Latitude Longitude Altitude (m) 

Ambodiara entendu des cris S 21º 53’ 17.4" E 047º 21’ 42.3" 500 

Ambodiara entendu des cris S 21º 53’ 17.7" E 047º 21’ 34.5" 825 

Antarehimamy entendu des cris S 21º 55’ 00.4" E 047º 22’ 10.3" 489 

Antaranjaha entendu des cris S 21º 58’ 23.6" E 047º 20’ 15.4" 828 

Antaranjaha entendu des cris S 21º 58’ 31.5" E 047º 20’ 13.6" 743 

rissage et émis des cris. La présence de huit autres espèces 

de lémuriens a également été constatée (Tab. 2). L’une d’entre 

elles, Varecia variegata editorum, est une sous-espèce 

gravement menacée selon l’UICN (2009), et les détails de 

toutes les observations de cette espèce sont présentés dans 

le Tab. 4. 

Résultats par zone située dans le corridor forestier de Fandriana- 

Vondrozo 

Ambendrana: Le village d’Ambendrana (S21°22’44.9" E 047° 

18’31.0",altitude 1121 m) est placé sous l’autorité de la commune 

rurale d’Androy et situé à une vingtaine de kilomètres 

au sud du Parc National de Ranomafana. Ce village est entouré 

de rizières localisées tout autour du corridor.La forêt 

d’Ambendrana a une superficie de 1.496 hectares et est 

gérée par la communauté de base depuis 2003. Au cours de 

l’enquête, les villageois n’ont reconnu que 3 espèces de 

lémuriens des 12 présentées sur les photos, notamment 

l’espèce Hapalemur griseus. D’après nos observations, la 

forêt d’Ambendrana est perturbée. Cependant, nous avons 

pu localiser quelques groupes de lémuriens, dont un groupe 

de H. griseus, et des signes de nourrissage. 

Amindrabe: La forêt d’Amindrabe a une superficie de 5.800 

hectares et est également gérée par la communauté de base 

depuis 2003.Cette forêt est située à 5,7 km du village d’Ambendrana. 

Le Fokontany Amindrabe (S21° 

23’14.8" E047°21’ 46.4", altitude 1096 m) 

fait également partie de la commune ru- 

rale d’Androy et comprend plusieurs villages. 

Durant l’enquête, les villageois ont 

reconnu 5 espèces de lémuriens, dont H. 

griseus. Pendant l’expédition dans le site 

d’Amindrabe, deux anciens signes de 

nourrissage (vieux d’environ un an d’après 

nos constatations) de P.simus ont été trouvés 

sur le tronc d’une espèce de bambou 

localement appelé Volotsangana (S21º24’ 

22.5", E047º23’07.2", altitude 1055 m). 

Nous avons également trouvé deux 

groupes de H. griseus, des Propithecus edwardsi 

et des signes d’alimentation de Daubentonia 

madagascariensis, attestant de la 

grande diversité de ce site en espèces de 

lémuriens. 

Ambodiara: Le Fokontany d’Ambodiara 

(S21°54’41.3", E047°23’29.2, altitude 

346 m) existe depuis 1910 et est composé 

de 8 villages. Le village d’Ambodiara est 

situé à 5,9 km, c’est-à-dire à environ 3 

heures de marche à l’ouest d’Ikongo. Au 

cours de l’enquête, les villageois ont reconnu 

9 espèces de lémuriens,dont Hapalemur 

aureus, H. griseus et également Prolemur 

simus. D’après nos observations, la 

forêt d’Ambodiara est perturbée. Nous 

n’avons pas trouvé P.simus sur ce site,mais 

nous avons constaté la présence de Cathariostachys 

sp. Par contre, Varecia variegata 

editorum abonde dans cette localité, et 

nous avons trouvé des signes de nourrissage 

de H. aureus et H. griseus. 

Antarehimamy: Situé dans le district 

d’Ikongo, le village d’Antarehimamy (S21° 

55’59.2",E047°22’17.6",altitude 410 m) se 

situe à 3,16 km au Nord-Est d’Ambodiara 

et à 9,01 km à l’ouest de la commune 

rurale d’Ikongo.Lors de l’enquête,les villa-


geois ont reconnu 7 espèces de lémuriens,mais aucun lémurien 

de bambou.Néanmoins,nous avons trouvé des signes de 

nourrissage de Hapalemur aureus et H. griseus. Nous avons 

également remarqué l’abondance de Varecia variegata editorum 

sur le site. 

Antaranjaha/Ambolomadinika/Tsianivoho:Située dans le district 

d’Ikongo, la commune d’Ambolomadinika gère 12 

Fokontany, dont Antaranjaha et Tsianivoho. Le Fokontany 

Antaranjaha (S21°59’42.3",E047°25’40.2") est situé à 3,7 km 

au Sud-Ouest d’Ambolomadinika. Dans ce site, les villageois 

n’ont pas reconnu de lémuriens de bambou. Cependant, des 

cris de Hapalemur aureus ont été entendus dans la forêt à 

150 m environ de notre campement,c’est-à-dire à Ankazondrano.A 

Marofototra,situé à 30 minutes du Fokontany d’Antaranjaha, 

toutes les jeunes pousses de bambou Cathariostachys 

sp.étaient coupées.Ce sont des signes de nourrissage de 

H. griseus. De plus, un villageois a confirmé avoir trouvé un 

groupe de H. griseus comprenant 12 individus à cet endroit. 

Nous avons également entendu des cris de Varecia variegata 

editorum, vu un groupe de Eulemur rufus, et trouvé un piège à 

lémuriens dans la forêt. Dans la commune rurale d’Ambolomadinika, 

on remarque beaucoup de zones agricoles déboisées. 

Malheureusement, les lémuriens de bambou sont menacés 

à cause de la coupe massive de bambous dans ces 

zones et la chasse pratiquée par les habitants de la Commune. 

Manambolo:Située dans la région de Fianarantsoa,la forêt de 

Manambolo (S22°04’06.2",E046°59’27.5",1238 m) se trouve 

dans le Fokontany de Morafeno, commune rurale de Sendrisoa. 

La gestion de la forêt est assurée par le FI.TE.MA 

(FIkambanan’ny TEraky MAnambolo) et concerne cinq villages: 

Mandamako, Mahavita, Ambinda, Ankazobe, Ampidira. 

La survie de la population locale dépend largement de l’agriculture, 

l’élevage et la production du rhum traditionnel. 

Cependant, la culture sur brûlis est encore pratiquée sur la 

lisière forestière. Lors de l’enquête, les villageois ont reconnu 

4 espèces de lémuriens, dont Hapalemur griseus et H. 

aureus. La vérification en forêt nous a révélé des signes de 

nourrissage de H.aureus, et nous avons vu directement Eulemur 

rufus et E. rubriventer. 

Résultats par zone située autour du corridor forestier de Fandriana-Vondrozo 

Mananjary: Notre campement à Tsararivotra (S21°10’41.8", 

E048°13’19.6", altitude 39 m) était situé à 23 km au nordouest 

de la ville de Mananjary. Le site de Tsararivotra est 

inclus dans le Fokontany de Volomborona Asakatara et fait 

partie de la commune de Morafeno Mananjary.Nous n’avons 

trouvé que Hapalemur griseus, Cheirogaleus major et Microcebus 

rufus dans cette zone. 

Sahalanona: La commune de Sahalanona fait partie du 

District d’Ikongo et inclut 9 Fokontany (Sahalanona, Mahaly, 

etc.). La population est composée d’agriculteurs, d’éleveurs 

et de pêcheurs. Le village de Sahalanona (S22°03’19.2" 

E047°37’ 12.2", altitude 129 m) existe depuis environ 300 

ans. Malgré l’abondante présence de bambous, dont le 

bambou géant Cathariostachys sp., nous n’avons trouvé que 

Hapalemur griseus dans cette zone. Cette espèce est menacée 

par la chasse que pratiquent les villageois. D’autres 

espèces de lémuriens sont également en danger car elles 

sont aussi chassées et vendues par les villageois, notamment 

Avahi laniger (chassé pour l’alimentation et l’usage domestique) 

et Microcebus rufus (dont le prix est de 5.000 Ariary 

par individu). 

Manakara: Le village d’Ambila se trouve à 17 km au nord de 

Manakara. Le Fokontany Ambila fait partie de la commune 

d’Ambila (S22°00’11.6", E047°58’19.9") de la région de Manakara.Notre 

observation a été effectuée directement dans 

la forêt de Tsiazombazaha située à 10 km du village d’Ambila. 

L’enquête n’a pas eu lieu dans ce site car il n’y avait plus de 

village (principalement notre cible) autour de la forêt. Cette 

forêt est gérée par la communauté de base du Fokontany 

d’Ambila.A cet endroit,nous n’avons pas trouvé de bambou, 

et avons trouvé seulement deux espèces de lémuriens, Avahi 

laniger et Eulemur rufus, après vérification dans la forêt. 

Mahazoarivo/Ifasy: Située dans la région de Farafangana, la 

commune de Mahazoarivo (S22°39’49.0", E047°18’42.4", 

222 m) fait partie du corridor forestier, et la population pratique 

l’agriculture et l’élevage. L’exploitation des ressources 

minières, surtout des pierres précieuses, représente une 

source de revenus importante pour la population. Lors de 

l’enquête, les villageois n’ont reconnu que deux espèces de 

lémuriens,Hapalemur griseus et Microcebus rufus.Nous avons 

visité deux forêts dans cette commune,à Mitimboto (Fokontany 

de Mahazoarivo) et Ifasy ou Mahafasy (Fokontany Mahatsara) 

où deux groupes de H. griseus ont été vus sur chaque 

site. A Ifasy (S22°39’13.0", E47°14’56.1"), des individus de H. 

griseus de très grande taille ont été localisés, similaires à 

Prolemur simus, mais l’absence des touffes de poils blancs sur 

les oreilles nous a permis de faire la distinction. Nous avons 

également remarqué que le nom local de H. griseus était 

différent à Mitimbato et Ifasy,respectivement Varibolo madinika 

et Varibolo vaventy. 

Iandraina: Le Fokontany d’Iandraina fait partie de la commune 

Rurale de Vohimasy. Il se situe à 15 km au nord-ouest 

de Farafangana. La forêt de Befoza et celle d’Ambolosy 

(S22°46’07.0", E047°41’07.0", 53 m) se trouvent dans ce 

fokontany. Les populations sont constituées principalement 

d’agriculteurs et d’éleveurs. La pratique des cultures vivrières 

constitue l’activité principale. Contrairement aux 

autres sites que nous avons visités dans le sud de la zone 

d’étude, nous avons trouvé une population de Cathariostachys 

sp. à Ambolosy. 

Sahamadio: Située dans la région de Farafangana, cette zone 

est plus ou moins enclavée (absence d’infrastructure routière) 

et même la circulation et le transport de produits 

locaux s’effectuent toujours par pirogue.La commune rurale 

de Sahamadio (S22°31’13.4", E047°35’02.8", altitude 27 m) 

dépend beaucoup de l’agriculture. L’existence de signes de 

nourrissage dans la forêt de Sahamadio nous a permis d’établir 

que Hapalemur griseus,localement appelé "varibolo" y est 

présent.L’enquête effectuée au niveau de la commune rurale 

d’Ambalatany a également confirmé la présence de lémuriens 

de bambou de grande taille dans la forêt d’Ambalakazaha. 

Evato:Dans la commune d’Evato (S 22°36’42",E 047°41’20"), 

dans la région de Farafangana, le développement des différentes 

infrastructures est remarquable,citons comme exemple 

les hôpitaux, écoles, marchés et routes en bon état. Un 

bloc de forêt primaire se trouve à Iaboloha dans cette 

commune. Notre enquête nous a donné des informations 

sur la présence de plusieurs espèces de lémuriens dans cette 

forêt. Ce site mérite donc d`être visité pour une prochaine 

vérification. 

Mahafasa: Dans cette zone située également dans la région 

de Farafangana, ce qui reste de forêt primaire est en général 

la forêt de bambou,un endroit où se trouvent des tombeaux. 

Etant donné la situation actuelle de sécurité, nous n’avons 

pas obtenu la permission de visiter cette forêt sacrée de 

bambou. Cette dernière recouvre une grande surface, environ 

3 km de longueur et jusqu’à 100 m de largeur,et pourrait 

être importante en tant qu’habitat de lémuriens.


Discussion 

La série d’expéditions menée le long du Corridor forestier 

Fandriana - Vondrozo nous a permis d’évaluer provisoirement 

la répartition des lémuriens de bambou. Concernant 

Prolemur simus,un seul signe de nourrissage a été identifié,et 

ce signe remontait à un an, confirmant les résultats des 

études précédentes qui indiquent que l’espèce a une distribution 

fragmentée dans la région (Wright et al., 2008). Pour 

Hapalemur aureus (espèce menacée EN), la découverte de 

l’évidence de sa présence sur six zones,toutes dans le corridor 

forestier, est encourageante car cela implique une large 

distribution dans celui-ci, bien que l’espèce ne semble pas 

exister en-dehors.Hapalemur griseus (espèce vulnérable VU) 

a été trouvé dans presque toutes les zones visitées, dans le 

corridor forestier mais également dans des zones éloignées 

de ce dernier. 

La menace principale pour les espèces de bambou dans la 

région du corridor Fandriana-Vondrozo est la destruction 

des habitats naturels et leur conversion en champs de culture.Cette 

technique est appelée "agriculture sur brûlis".Par 

conséquent, cette pression entraîne la raréfaction et même 

la disparition des espèces autochtones de bambous. Malgré 

la présence de bambous à l’intérieur du corridor, la persistance 

de la pratique du tavy,les coupes de bambous en permanence 

et surtout la chasse aux lémuriens mettent en péril 

la survie des espèces de lémuriens. En outre, la taille de la 

forêt du Corridor Fandriana-Vondrozo est petite par rapport 

aux autres corridors forestiers du pays. Sa largeur est 

très réduite surtout dans sa partie sud, et sur la photo 

aérienne, la voûte forestière apparaît très ouverte. Tous ces 

facteurs menacent la viabilité des populations de lémuriens 

vivant dans le corridor, et tout particulièrement les espèces 

présentant une distribution fragmentée telles que P. simus. 

Actuellement, beaucoup de lémuriens sont chassés et vendus 

par les villageois (exemple:Sahalanona,Antaranjaha).Les 

forêts de bambous sont fragmentées et isolées les unes des 

autres, ce qui laisse à penser que ces lémuriens de bambou 

sont réellement en danger. En outre, les utilisations des 

bambous dans la région sont nombreuses. La population locale 

utilise les différentes espèces de bambou suivant leur 

taille pour la construction des maisons, particulièrement 

pour les toitures, les murs, et des clôtures. Les bambous 

servent également à fabriquer du matériel pour les usages 

quotidiens, parmi lesquels les paniers à fruits, volailles, écrevisses 

et anguilles. Enfin, ils permettent de transporter des 

bagages. Aussi, les espèces de bambous de plus grand diamètre 

sont utilisées comme récipient pour transporter de 

l’eau. La conséquence négative de l’utilisation des bambous 

est minime par rapport à la destruction des habitats. Exemple:le 

Corridor d’Ampitsinjovabe (site d’Antarehimamy) est 

une bonne localité pour trouver H. griseus, H. aureus et V. 

variegata editorum, mais ces trois espèces sont menacées à 

cause de la chasse et des coupes sélectives de bois pratiquées 

par les habitants résidant autour du corridor. 

Pour la conservation de Prolemur simus,il faudrait accroître la 

taille des aires protégées en y incluant les forêts de bambous, 

et restaurer les fragments d’habitats isolés au sein d’un 

paysage agricole déboisé, afin d’équilibrer la valence écologique, 

c’est-à-dire la zone supportable pour l’espèce (en 

pratiquant une reforestation de bambou). Cependant, d’une 

façon générale, il y a un besoin immédiat de sensibilisation, 

pour conscientiser la population aux problèmes de coupe de 

bambous,de tavy et de chasse des lémuriens,afin d’assurer la 

survie d’espèces de lémuriens dans et autour du Corridor 

Fandriana-Vondrozo. Finalement, les sites d’Ambodiara, Mahazoarivo 

(Alafady, Ranomena), Ambalakazaha et Mahafasa 

sont recommandés pour une nouvelle vérification de la présence 

ou non des lémuriens de bambou. En effet, la population 

locale semble être convaincue d’avoir trouvé P. simus à 

ces endroits. 

Remerciements 

Nos vifs remerciements vont: au Ministère de l’Environnement, 

des forêts et du Tourisme, à la Direction Générale de 

l’Environnement et des forêts, et à la Direction du Système 

des Aires Protégées, Madagascar, pour leur accord et la 

délivrance de l’autorisation de recherche (permis n°279/08/ 

MEFT/SG/DGEF/DSAP/SSE); à The Aspinall Foundation, GB, 

pour le financement de l’enquête dans le cadre du Projet 

"Sauver Prolemur simus";au Groupe d’Etude et de Recherche 

sur les Primates de Madagascar (G.E.R.P) et son personnel 

administratif; au Centre International de Formation pour la 

Valorisation de la Biodiversité (Centre ValBio) et son personnel 

administratif; à l’ICTE et Conservation International, 

Antananarivo,pour leurs conseils et entire collaboration;aux 

communes, Fokontany, et COBAS des zone visitées pour 

leurs amabilité et collaboration; et enfin, aux assistants de 

recherche du Centre ValBio à Ranomafana, Justin Rakotonjatovo, 

Dominique Razafindraibe, Jean-Guy Razafindraibe, 

Aime-Victor Tombotiana et Telo Albert, et au chauffeur de 

The Aspinall Foundation,Mohamad Mbaraka,pour leur assistance 

sur le terrain. 

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8: 84-88 

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Lehman, S.; Louis, E.E. Jr.; Arrigo-Nelson, S.J.; Raharison, 

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simus). Primate Conservation 23: 5-17. 

Effect of red ruffed lemur gut passage on 

the germination of native rainforest plant 

species 

Onja H. Razafindratsima 1,2*, Emilienne Razafimahatratra 

1 

1Department of Animal Biology, University of Antananarivo, 

Madagascar 

2Department of Ecology and Evolutionary Biology, Rice University 

– MS 170, 6100 Main St., Houston, TX 77005, USA 

(current affiliation) 

*Corresponding author: ohr1@rice.edu, onjhar@hotmail.com 

Key words: seed dispersal, germination success, Varecia 

rubra, primate, corridor restoration, Masoala 

Abstract 

Like much of Madagascar’s remaining rainforest,the forest of 

Masoala National Park is facing severe threats from deforestation 

and fragmentation. The remaining fragmented areas 

are connected by degraded corridors which are important 

for biological exchange. Frugivorous animals such as lemurs 

may have an important role in the restoration of such degraded 

areas through seed dispersal. Unfortunately,no studies 

have been carried out before concerning the role lemurs 

play in the restoration of the largest corridor in Masoala, 

Ambatoledama. This study explores the effect of seed passage 

inside the gut of the frugivorous red-ruffed lemur 

(Varecia rubra) on the germination of some native tropical 

plants with the aim to understand the capacity of V. rubra to 

help in the restoration of the Ambatoledama corridor. We 

planted seeds of nine plant species that we collected from V. 

rubra’s fresh feces in a nursery to compare with seeds that 

we extracted manually from corresponding fruits. The germination 

of seeds was monitored each month after planting 

them. Results showed that defecated seeds had overall a significantly 

higher germination rate than non-passed seeds. 

Thus, lemur ingestion of seeds has the capacity to improve 

seed germination of several species and some plants require 

the physiological treatment inside the gut to germinate. Results 

suggested that restoration projects in the area including 

the Ambatoledama corridor should take into account the 

important role Varecia rubra plays in the regeneration of the 

forest and corridor.Management actions that increase movement 

and protection of animals moving into and out of the 

corridor will be important for the long term success of the 

project. 

Introduction 

The rainforest of the Masoala Peninsula suffers greatly from 

loss and fragmentation caused by the human population 

living around the area.The forest is subdivided into different 

fragments, connected by corridors of degraded habitat 

which are Ambatoledama,Analambolo and Ilampy (Holloway, 

1997). Corridors are vital for enabling gene flow and dispersal 

of wildlife among habitat fragments (Mech and Hallett, 

2001). The largest of these is the Ambatoledama corridor, 

which connects two large parcels of the forest (Fig. 1). The 

restoration of this corridor is critical for safeguarding wildlife 

populations in the fragments and for preserving gene flow 

between fragments (Mech and Hallett, 2001; Haddad et al., 

2003). To restore this degraded corridor, it is necessary to 

plant native trees or to encourage zoochory (biological dispersal 

of seeds through animal defecation) (Duncan and 

Chapman, 2002; Neilan et al., 2006). Since 1997, Madagascar 

National Parks (MNP) and the Wildlife Conservation Society 

(WCS) have established a restoration project in the Ambatoledama 

corridor by planting native fruiting trees (Holloway,1997) 

with the aim of attracting frugivorous vertebrates 

which will in turn carry seeds into the degraded parts of the 

forest and into forest clearings. Unfortunately, no studies 

have previously been carried out to shed light on the importance 

of frugivorous animals,especially lemurs,in the reforestation 

of the Ambatoledama corridor.Unlike the majority of 

tropical forests,the diversity of the frugivorous bird community 

in Madagascar is impoverished, and therefore primates 

are the principal dispersers of its tropical trees (Goodman, 

1997; Dew and Wright, 1998; Ganzhorn et al., 1999; Bleher 

and Böhning-Gaese,2001).Ten lemur species are indentified 

as living in the Masoala Forest (Mittermeier et al., 2006); one 

of which (Varecia rubra) is endemic to this region and has Endangered 

status (IUCN, 2008), and can be found in both the 

corridor habitat and adjacent forest fragments (Razakamaharavo 

et al., 2010). Previous studies demonstrated that 

Varecia variegata is an effective disperser in the southeastern 

rainforests (Dew and Wright,1998).However,we know very 

little about the potential role of V.rubra for regeneration and 

restoration of the corridor habitat in Ambatoledama. 

In this study, we explored the germination success of seeds 

defecated by Varecia rubra in order to understand their capacity 

for seed dispersal and potential impact on the restoration 

of the degraded rainforest corridor at Ambatoledama. 

Our objective was to shed light on the role of this species in 

forest regeneration. Understanding their influence on tree 

germination is particularly important given the threatened 

status of this species. This paper tested the hypothesis that 

gut passage of seeds by Varecia rubra facilitates seed germination. 

Our prediction was that lemur-gut-passed seeds have a 

higher germination rate than non-passed seeds because of 

the physiological treatment affecting the seed coat inside the 

gut.


Materials and methods 

Field site 

This study was carried out at the Ambatoledama corridor 

(S15°27’ E050°01’) on the north-eastern part of the Masoala 

Peninsula. Ambatoledama connects Masoala National Park 

with Makira National Forest to the West. Its forest has undergone 

significant deforestation but restoration projects 

have augmented Ambatoledama such that it now forms a 1 

km wide corridor of secondary forest (Hekkala et al., 2007; 

Razakamaharavo et al.,2010).It consists of a dense evergreen 

rainforest with an altitude ranging from 300 to 700 m. The 

forest is mostly characterized by the presence of tree species 

of the Pandanacea, Ebenaceae, Clusiaceae, Euphorbiaceae, 

Sapotaceae and Rubiaceae families (Martinez, unpublished). 

Study species 

Varecia rubra belongs to the family Lemuridae (Gray, 1821) 

and is one of two species recognized within the genus (Mittermeier 

et al., 2006). V. rubra is only found on the Masoala 

peninsula and it is classified by the World Conservation Union 

(IUCN) as Endangered (IUCN, 2008). V. rubra is a largesized 

diurnal species with a body length ranging from 43 to 

57 cm (Vasey, 2003) and has a typically frugivorous diet 

(Rigamonti, 1993; Vasey, 1997). They currently inhabit both 

the corridor habitat and the adjacent protected areas (Razakamaharavo 

et al., 2010) and are thus potentially important 

for regeneration of the corridor habitat. 

Field experiment 

Focal animals were followed for 3-5 days per week from 

dawn to dusk (from 0600 hours to 1800 hours) to collect 

fresh fecal samples (Dew and Wright,1998;Kaplin and Moermond,1998;Stevenson,2000;Poulsen 

et al.,2001;Link and Di 

Fiore, 2006). Each fecal sample was washed and filtered 

through a 1-mm sieve (Stevenson,2000).Seeds were extracted 

and then identified with the help of local research guides 

and an expert local botanist familiar with the Masoala flora. 

We planted gut-passed seeds and control seeds that were 

extracted manually from fruits in an outdoor nursery adjacent 

to the corridor at Ambatoledama. The nursery consisted 

of two "flower beds" of 11.2 m 2: one for defecated 

Fig. 1: Location of the Ambatoledama corridor. 

seeds and the other one for non-passed seeds. Following 

methods used by the conservation agents of MNP in Ambatoledama, 

a sunshade of 80 cm height, composed of Longoza 

leaves (Afromomum angustifolium) was placed above each 

flower bed to imitate the closed canopy of the forest. Also, 

the soil of the nursery was mixed with fertile soil from cultivated 

field. Seeds were placed in the soil mixture and 

covered by 1 mm-thick river sand to keep a constant temperature. 

An equal number of seeds were planted within each species 

per treatment. However, the numbers varied between species 

depending on how many seeds were collected from 

lemur feces. The germination of seeds was assessed each 

month after planting. 

Data analysis 

We performed a paired t-test to test for differences between 

the germination rate of lemur-gut-passed and nonpassed 

seeds, an ANOVA analysis to test if the two factors 

(seed species and treatment) had effects on the germination 

rate of the seeds and to determine whether there was interaction 

between these factors. We analyzed the germination 

of each species in order to assess the influence by lemur gut 

passage, with Pearson test using contingency tables, which 

was adjusted with Bonferroni correction for multiple comparisons 

(Sokal and Rohlf, 1995). 

Results 

In total, 268 fresh fecal samples from three individuals of 

red-ruffed lemur were collected during 58 days of observation. 

The fecal samples contained fleshy fruit parts, stalks, 

leaves, soil and fecal liquid. 95.52 % of these contained seeds, 

to some of which fleshy fruit parts were still attached. 906 

seeds of more than 1mm size were extracted. A majority of 

them were intact with minor scarification.They represented 

34 different plant species that belong to 15 Families.Based on 

our collected sample, the most common seed species found 

in lemur defecations were the nine species we chose to 

study here (Tab. 1). In the nursery, we planted a total of 390 

defecated seeds and compared them with 398 non-passed 

seeds. 

Lemur-gut-passed seeds had significantly higher germination 

rates overall than non-passed seeds (t=3.284,df=8,p=0.011). 

Passed seeds had a germination rate 

of 64.61 %, whereas non-passed 

seeds had a rate of 39.69 %.For each 

species, seeds that had been defecated 

had a higher germination rate 

than non-passed seeds, except for 

Tsilaitra (Tab. 1). This pattern was 

driven primarily by four species, including 

Antaivaratra,Matahobaratra, 

Tsilaitra, and Vongobe species. 

In a two factor analysis of variance 

for seed germination, there was a 

significant interaction between the 

species of seeds and their treatment 

(passed or non-passed) (F= 4.2004, 

p


Tab. 1: List of species studied with their germination rate. Sample sizes are represented in 

brackets. The star on p-values corresponds to their significance (Pearson test) after a 

Bonferroni correction for multiple tests. 

# Malagasy Scientific Family Germination rate Pearson test 

name name 

gut-passed non-passed ChiP- seeds seeds squarevalue 1 Antaivaratra Potameia sp. Lauraceae 41.67 (n = 48) 17.86 (n = 56) 7.139 0.0075* 

2 Hazondronono Sideroxylon Sapotaceae 80.00 (n = 10) 60.00 (n = 10) 0.952 0.3291 

3 Karaka Pandanus Pandanaceae 40.00 (n = 20) 15.00 (n = 20) 3.135 0.0766 

4 Matahobaratra Garcinia sp. Clusiaceae 58.06 (n = 31) 00.00 (n = 31) 25.364


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Feeding ecology of the crowned sifaka 

(Propithecus coronatus) in a coastal dry 

forest in northwest Madagascar (SFUM, 

Antrema) 

Claire Pichon 1*, Rivo Ramanamisata 2, Laurent Tarnaud 

1, Françoise Bayart 1, Annette Hladik 1, Claude 

Marcel Hladik 1, Bruno Simmen 1 

1UMR 7206, Eco-anthropologie et Ethnobiologie, Centre 

National de la Recherche Scientifique,and Museum National 

d’Histoire Naturelle, 4 avenue du Petit Château, 91800 Brunoy, 

France 

2Département de Biologie Animale,Faculté des Sciences,B.P. 

906, Université d’Antananarivo, Antananarivo (101), Madagascar 

*Corresponding author: cpichon@mnhn.fr 

Key words: diet, primate, activity budget, forest composition 

The crowned sifaka (Propithecus coronatus; Milne-Edwards, 

1871) inhabits dry forests, riparian forests and mangroves of 

northwest Madagascar. Originally believed to occur in a restricted 

area between the Mahavavy River in the southwest 

(where it overlaps with P.deckenii) and the Betsiboka River in 

the northeast (which separates it from P. coquereli), sightings 

west of the Mahavavy River and along the Bongolava Massif 

suggest that the distribution of this medium-sized species is 

wider (Tattersall, 1986; Thalmann et al., 2002). The distribution 

and the taxonomic status of crowned sifakas have long 

been debated, but the combination of morphological and 

biogeographic evidence supports considering it as a valid 

species (Thalmann et al., 2002; Mittermeier et al., 2006; 

Groves and Helgen, 2007; Mittermeier et al., 2008). Considered 

as Endangered (A2 c,d) by the IUCN (2010), populations 

of crowned sifakas were estimated not to exceed 1,000 

individuals in the wild.However,recently discovered populations 

in restricted fragmented forests extend the species’


distribution range farther towards the Southwest. A "Biocultural 

Project" was therefore initiated at Antrema (a site 

located in the Mahajanga region) in 2000 to promote sustainable 

management (Gauthier et al., 2001).The project aims at 

preserving a coastal environment in which crowned sifakas 

occur in high densities while allowing villagers, mainly fishermen, 

to use natural products of the environment with parsimony 

and to benefit from technical and economical help.The 

project also aims at promoting local socio-cultural rules and 

a way of life that tends to respect the forest environment,including 

useful plants and several sympatric lemur species 

(Propithecus coronatus, Eulemur fulvus, Eulemur mongoz, Lepilemur 

sp., Microcebus murinus). The site contains three of the 

Northwest’s typical ecosystems (dry semi-deciduous forest, 

mangrove swamp, savanna), which suffer moderate anthropogenic 

pressure (Gauthier et al., 2001). Owing to local 

beliefs, especially the sacred (“masina”) nature of sifakas, the 

Sakalava community plays a central role in this conservation 

process (Harpet et al.,2000,2008).In this context,a few studies 

started investigating the behavior in relation to habitat 

and food supply of the lemur species of Antrema (Gauthier et 

al., 1999; Razafimahefa, 2001; Ramanikirahina, 2004). However,a 

detailed analysis of the feeding ecology and population 

densities of P. coronatus is still lacking. We present here preliminary 

data on the plant species composition of the whitesand 

coastal forest inhabited by a dense population of sifakas 

(among other prosimian species) and on the feeding ecology 

of sifaka groups censused since 2008. 

Methods 

Study site 

The Antrema station is a coastal area of 12,300 ha located on 

the left riverside of the Betsiboka estuary, northwestern 

Madagascar (15°42’-15°50’S, 46°-46°15’E; Gauthier et al., 

2001). The region undergoes a distinct dry season of 7 

months from April to October.The mean annual rainfall (n = 

9 years) in the Mahajanga region is 1,410 mm (with a peak in 

January-February), with irregular rainfall during the dry season.With 

an annual mean of 27° C,temperature is highest in 

October and lowest between June and August (Airport of 

Mahajanga, 2000-2009). 

Although the Antrema area has been traditionally protected 

by the local Sakalava beliefs, forest areas where studies are 

conducted are fragmented. After two first surveys in November 

2007 and April-May 2008, we decided to establish 

the study site at Badrala (15°45.665’S, 46°12.300’E). With 

about 24 ha just behind the littoral dune,this non-sacred forest 

site offers suitable conditions to study the socioecology 

of sifakas and the dynamics of a dry forest in Madagascar.The 

forest there is partly split by a sandy open dune that sifaka 

groups can cross easily. Tree logging occurs at low intensity 

(with few selected species for defined use, e.g. for boats or 

coffins) and small trees are sporadically cut for fences and 

house building. We studied floristic composition by inventorying 

trees along four North/South-oriented line transects, 

10 m-wide each, that were roughly perpendicular to the sea 

front.Within this 0.73 ha,we tagged each tree > 10 cm diameter 

at breast height (DBH) with plastic labels, recorded 

their DBH, the number of stems and their vernacular name. 

Likewise, we counted woody lianas and herbaceous vines 

> 1 m high within eight 10 x 10 m (800 m 2) plots regularly 

spaced along the transects. Plant species were sampled and 

dried for later botanical identification. 

Sifaka population density 

In order to locate and identify groups of P. coronatus in 

Badrala, we initially mapped groups encountered during 

repeated transect walks. We drew individuals’ facial masks 

for each of the followed groups, noted their sex from visual 

inspection of the genitalia and other external characteristics 

(cysts, scars, damaged ears, fur colour), and took pictures. 

Knowledge gained progressively of groups and individuals 

allowed us to provide a preliminary estimate of population 

density for the Badrala site. 

Behavioral data collection 

We collected behavioral data during 4 periods (06 to 21 July 

2008; 11 November to 12 December 2008; 05 April to 06 

June 2009;17 October to 22 November 2009). Most groups 

were already accustomed to the sporadic presence of local 

people. Once we could observe animals at close distances, 

we followed each group successively over 2 to 5-day periods, 

from 06:30 to 18:30 hours. 

We used the instantaneous scan-sampling method (Altmann, 

1974) to study group activity budget.Every 5 minutes,we recorded 

the individuals’ activity using one of the following categories:resting 

(immobile,with eyes open or closed),moving 

(more than 0.5 m),foraging (searching for a food item),feeding 

(processing or chewing a food item), social activity (displaying 

agonistic and affiliation behaviors with other individuals) 

and other miscellaneous behaviors. We noted the plant 

part and species eaten by individuals. 

Besides recording activity budgets,we determined diet from 

mouthful counts converted into weight of ingested matter 

(Hladik, 1977) for 2 periods: April-June 2009 and October- 

November 2009.We estimated food intake in focal individuals 

that were followed continuously for 30 minutes each. 

Observations were alternated across males and females (excluding 

juveniles) within groups. 

Results 

Forest composition 

Plant families occurring at Badrala are presented separately 

for trees and lianas/vines in Fig.1.To date,91 tree and liana or 

vine species have been identified at least at the family level, 

and taxonomic identification of 15 more putative species is 

still in progress.The 5 richest families in terms of the number 

of species are Fabaceae, Sapindaceae, Ebenaceae, Euphorbiaceae 

and Apocynaceae.The most dominant tree species are 

Strychnos decussata, Vitex beraviensis, Mimusops occidentalis, 

Baudouinia fluggeiformis and Macphersonia gracilis that represent 

almost one third of total basal area and tagged trees. 

Combretum coccineum, Hypoestes sp., Landolphia perrieri and 

Reissantia sp.accounted for more than one third of the lianas 

and vines. 

Density of trees inventoried on the 4 transects (n=486) corresponds 

to 666 inds. ha 1 with a total basal area of 14.5 m² 

ha -1.We found a high density of woody lianas and herbaceous 

vines in the 800m² plots (n=373). 

Sifaka population density and group composition 

Groups at Badrala have 1-3 breeding adult males, 1-4 breeding 

adult females, and 1-4 immature offspring. We encountered 

between 15 and 20 groups at this site. Based on current 

recognition of individuals within these groups, a minimum 

estimate of 300 inds. km 2 was calculated. Mean size of 

focal groups was 4.3 ± 1.8 individuals (n=16). 

Diet and activity pattern 

Sifakas consumed at least 60 plant species from 32 families. 

Tab. 1 lists major food species eaten. During the dry season, 

14 plant species represented 75 % of the diet whereas only 7 

species were the main food resource in the wet season.


Tab. 1: Food species accounting for 50 and 75 % of the diet of Propithecus coronatus during the dry season and the wet season. 

Eaten plant species are listed in decreasing order and their abundance in transects and plots (see text) is indicated. 

Family Species Vernacular name Items Abundance (%) 

Dry season 

Lamiaceae Vitex beraviensis Mojiro yl 10,7 

Fabaceae Baudouinia fluggeiformis Manjakabentany yl ml 4.7 

Sapotaceae Mimusops occidentalis Natofotsy yl stems 3.9 

Anacardiaceae Operculicarya gummifera Atokonjo ml buds 3.5 

Sapindaceae Majidea zanguebarica Tsipopoka yl ml fl 2.3 

Oleaceae Noronhia boinensis Tsilaitra beravina yl 1.9 

Moraceae Trilepisium occidentalis Kililo ml 1.2 

Sphaerosepalaceae Rhopalocarpus lucidus Hazondringitra yl fr 1.0 

Melastomataceae Warneckea sp. Voatrotrokoala yl 0.6 75 % 

Burseraceae Commiphora sp. Arofy fr buds 0.4 

Fabaceae Bussea perrieri Morango ml 0.2 

Olacaceae Olax dissitiflora Ambiotsy ml 0.2 

Moraceae Ficus pyrifolia Nonika fr 

Unidentified - RR80 ml 

Wet season 

Anacardiaceae Abrahamia deflexa Motsovavy yl fl 3,1 

Anacardiaceae Abrahamia sp. Manavodrevo buds yl fl 1.2 

Fabaceae Chadsia flammea Fanamohazo buds yl fl 0.8 50 % 

Sapotaceae Capurodendron gracilifolium Natoboay buds yl fr 0.2 

Apocynaceae Landolphia perrieri Vahipira yl 6.8 75 % 

Combretaceae Terminalia sp. Taly buds yl 3.1 

Anacardiaceae Operculicarya gummifera Atokonjo yl 3.5 

yl: young leaves; ml: mature leaves; fl: flowers; fr: fuits 

Fig.1:Abundance of plant families plotted in 

decreasing number of individuals among a) 

trees with DBH>10cm (based on transects; 

0.73 ha) and b) woody lianas and herbaceous 

vines >1 m height (based on plots; 

0.08 ha).Striped bars refer to the plant families 

with the highest number of species.


58% 

(a) Dry season (b) Wet season 

9% 

2% 

1% 

30% 

25% 

11% 

6% 

1% 

Sifakas were highly folivorous during both seasons, supplementing 

their diet with flowers, fruits, vegetative buds, and 

sometimes young stems (Fig. 2). They consumed more mature 

leaves and fruits in the dry season and more flowers in 

the wet season. 

The activity budget of the sifakas is presented in Fig. 3. Although 

‘resting’ predominated throughout the study, 

individuals rested more in the dry season than in the wet season.Inversely,they 

travelled less and engaged in feeding activities 

more often during the wet season. 

Discussion 

The sifaka density was found to be high in the dry forest of 

Antrema, with a minimum estimate far above the 173 inds. 

km 2 found in the riparian forest of Anjamena (Muller et al., 

2000) or for other sifaka species in dry or wet forests 

(O’Connor, 1988; Ganzhorn, 1992). This high density might 

be related to some peculiar characteristics of the forest in 

terms of food quantity and/or food quality available to this 

prosimian species. However,tree basal area was not particularly 

high compared with other dry forests in Madagascar 

and Mayotte (Hladik, 1980; Simmen et al., 2005). It is not yet 

clear also whether high density is related to a putative low 

predation pressure.To our knowledge,no sightings or traces 

of viverrid carnivores have been reported; large raptors and 

boas would be the only predators that could affect the 

demography of the Antrema sifaka population (Garbutt, 

2007; Sinclair and Legrand, 2008). 

As regards their feeding behavior, crowned sifakas fed primarily 

on leaves from a few tree, liana and vine species, and 

supplemented their diet with a wide range of secondary 

items as commonly occurs in other Propithecus species 

(Meyers and Wright, 1993; Simmen et al., 2003; Lehman and 

Mayor, 2004; Irwin, 2008). Although this species remained 

57% Young leaves 

Mature leaves 

Flowers 

Fruits 

Other 

Fig.2:Food categories in the diet of Propithecus coronatus during the dry season 

(a) and the wet season (b). 

29,4% 

(a) Dry season 

0,8% 5,9% 

50,0% 

(b) Wet season 

3,9% 

35,8% 

5,3% 

46,6% 

Resting 

Locomotion 

Foraging 

Feeding 

6,8% 

7,1% 

3,6% 4,8% 

Social 

Other 

Fig. 3: Activity budget of Propithecus coronatus during the dry season (a) and the 

wet season (b). 

folivorous during our study,its diet changed 

with seasons. Young leaves were the preferred 

food type in the early wet season, 

while mature leaves were the dominant one 

in the beginning of the dry season. In addition, 

P. coronatus ate a higher proportion of 

liana and vine parts during the wet season. 

Crowned sifakas also followed the typical 

activity pattern of other sifaka species 

(Norscia et al.,2006;Patel,2006;Charrier et 

al., 2007), spending most of their time resting 

and devoting a substantial amount of 

time to feeding activities and locomotion. 

Activity budget nevertheless changed with 

seasons. It is generally suggested that the 

cool dry season represents a period of food 

scarcity for animals,which they compensate 

for by reducing their energy expenditure, 

travelling less and resting more. In a recent 

joint research project, the content of litter 

traps regularly distributed along the transects 

was collected and weighted every two 

weeks throughout one year. It was found 

that plant species could be grouped according 

to their temporal pattern of leaf loss 

(Ranaivoson et al., 2010; see also Razakanirina, 

2010). Several trees, lianas and vines 

lost their leaves more or less regularly 

throughout the dry season while others 

were characterized by delayed leaf loss or 

on the contrary by precocious leaf fall.One 

consequence is that leaves are available 

throughout the year, although as different sets of species 

varying in quantity,diversity,and presumably,nutritional quality.This 

at least could explain why sifakas are able to increase 

the diversity of consumed plants (and adopt a more opportunistic 

strategy) during the dry season, a period normally 

described by the scarcity of food resources. 

Future work on seasonal variations in the diet’s nutritional 

and chemical content will allow us to examine the role of 

qualitative aspects in food choices (Moss,1991;Dearing et al., 

2000) and further examine potential differences between 

genders with regard to the importance of energy conservation 

for female sifakas (Wright, 1999; Charrier et al., 2007). 

Conclusion 

Better knowledge of the ecology and the villagers’ social perception 

of this flagship species may contribute to conservation 

of other diurnal lemurs, by incorporating the villagers’ 

symbolic perception of their natural environment. Investigating 

the interactions between this species and plants of the 

coastal dry forest ecosystem will undoubtedly result in 

better conservation decisions for Antrema. From an evolutionary 

ecology standpoint, the studies we have planned for 

the next years at Antrema will also contribute to better 

identify the selective pressures that have been driving the 

evolution of prosimian typical life-history traits such as 

reproductive synchronization or dominance-based feeding 

priority of females over males in gregarious species (Wright, 

1999; Dewar and Richard, 2007). 


We thank the Malagasy Institutions that authorized to collect 

and export the plant samples,the Ministère de l’Environnement, 

des Eaux et forêts et du Tourisme. We also thank 

Antrema’s project staff for assistance with the field work as


well as Master degree students S.Razakanirina,T.Ranaivoson, 

V. Randriantoposon and L. Razafindramahatra for helping 

with data collection. Special thanks to the specialists of the 

Dept of Phanerogamy,J.N.Labat,P.Phillipson and Pete Lowry, 

for helping with plant identification. Finally, we thank C.A. 

Gauthier, E. Roger, D. Rakotondravony and H. Razafindraibe 

for logistic support and collaborative work, and E.G. Leigh 

for helpful comments on an earlier draft. This study was 

funded by the UMR 7206 - CNRS,and was conducted under 

the "Convention cadre de cooperation" between the Université 

d’Antananarivo and the Museum National d’Histoire 

Naturelle, Paris. 

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Effet de la dégradation de l’habitat sur la 

consommation alimentaire d’Eulemur 

rubriventer dans deux sites: Talatakely et 

Vatoharanana,du Parc National de Ranomafana 

Laingoniaina H.Rakotonirina 1,2*,Germain J.Spiral 1,2, 

Jonah H. Ratsimbazafy 1,2, Soanorolalao Ravelonjanahary 

1,Raharizelina Ralaiarison 1,2,Stacey Tecot 3,Alex 

Hall 4, Tricia Calhoon 4, Gisèle R. Randria 1,2 

1Département de Paléontologie et d’Anthropologie Biologique, 

Faculté des Sciences, B.P. 906, Université d’Antananarivo, 

Madagascar 

2Groupe d’Etudes et de Recherche sur les Primates de Madagascar 

(GERP) 

3Department of Anthropology, University of Stony Brook, 

USA 

4Voluntary field assistant 

*Contact de l’auteur principal: laingoniaina2000@yahoo.fr 

Mots clés:Eulemur rubriventer,dégradation,habitat,consommation 

alimentaire, Ranomafana, Madagascar 

Introduction 

La grande île est potentiellement riche en matière de biodiversité 

et est par conséquent renommée pour sa remarquable 

richesse écologique, biologique et génétique (Ganzhorn 

et al.,2001).Cette richesse qui est gravement menacée 

par la diminution et la destruction immuables des habitats 

naturels de nombreuses espèces fait de Madagascar un des 

huit "hotspots" les plus considérés de notre planète (Ganzhorn 

et al., 2001). 

A l’échelle mondiale, la menace la plus grave pour la population 

des primates est la destruction et la dégradation de leur 

habitat, notamment les forêts tropicales qui hébergent aujourd’hui 

environ 90 % des primates non humains du monde 

(Mittermeier et al., 2006, 2010). Les lémuriens malgaches ne 

font pas exception à cette constatation. La dégradation des 

forêts affecte la biologie générale des lémuriens car non 

seulement elles leur fournissent des abris et de la nourriture, 

mais aussi elles servent de supports à la locomotion et aux 

différentes activités de ces animaux (Razafimahazo, 2001). 

Selon Randriatahina en 2001, la fragmentation de l’habitat 

affecte en premier lieu la distribution et la dispersion de la 

nourriture.Certains facteurs influencent le rythme d’activité 

et le budget-temps des primates:il s’agit surtout des facteurs 

écologiques majeurs tels que la structure de l’habitat,le type 

d’alimentation (Zaonarivelo, 1999). Par ailleurs, Dunbar 

(1988) affirme que les primates pourraient augmenter leur 

déplacement journalier pour trouver de la nourriture ou 

inversement en vue d’économiser leur énergie. 

Notre présent travail est axé sur la corrélation entre l’habitat 

et l’alimentation des lémuriens.Les lémuriformes montrent 

un degré de variabilité en ce qui concerne la spécialisation 

aux régimes alimentaires.La plupart d’entre-eux (les 

Lémuridés,les Mégaladapidés,les Indridés) se spécialisent au 

régime végétarien. Cependant, la proportion de feuilles, de 

fleurs, et de fruits consommés varie suivant les espèces et 

sous-espèces,d’une région à une autre,et de saison en saison 

(Richard, 1978). Selon Zaonarivelo (1999), des facteurs écologiques 

influencent les comportements des lémuriens et la 

perturbation de leur habitat affecte leur organisation sociale 

et l’exploitation des ressources alimentaires. 

En tenant compte de toutes ces observations, nous avons 

effectué une étude concernant l’effet de la dégradation de 

l’habitat sur la consommation alimentaire d’Eulemur rubriventer 

dans deux sites: Talatakely et Vatoharanana du Parc National 

de Ranomafana, dans la province de Fianarantsoa. 

Il y a lieu de souligner qu’Eulemur rubriventer, une espèce 

hautement frugivore (Overdorff, 1993), dispose d’une haute 

importance écologique car elle participe activement à la dispersion 

des graines dans la région du Sud-Est de Madagascar, 

en particulier dans le Parc National de Ranomafana. A cet 

égard, bien que l’animal soit encore classé dans la catégorie 

vulnérable selon la liste rouge de l’UICN (Mittermeier et al., 

1994,2006,2010),il a besoin d’ une action de conservation. 

C’est la raison pour laquelle le parc national de Ranomafana a 

été choisi comme notre station de recherche car par rapport 

aux autres régions de l’île, les lémurs à ventre roux y 

sont les plus répandus (Mittermeier et al.,2006);et leur habitat 

présente un degré variable de dégradation. 

Compte tenu de cette variation du degré de dégradation et 

de perturbation du milieu de vie d’Eulemur rubriventer dans le 

Parc National de Ranomafana, nous pouvons avancer une 

hypothèse selon laquelle la consommation alimentaire n’est 

pas statistiquement différente entre celle de Vatoharanana 

et celle de Talatakely. 

Ce projet a été réalisé dans le cadre de la collaboration interdépartementale 

entre l’Université d’Antananarivo, l’ICTE/ 

MICET,le MNP et l’Université de Texas.Ainsi,le présent travail 

qui vise en la conservation des lémurs à ventre roux a 

comme objectifs d’inventorier les différentes espèces de 

plantes consommées par Eulemur rubriventer,de comparer le 

régime alimentaire adopté dans chaque site d’étude, de 

déterminer les caractéristiques des plantes consommées 

dans les deux sites d’études à dégradations différentes. 

Site d’études 

Le parc National de Ranomafana se trouve dans le Sud Est de 

Madagascar.Sa superficie est de 41.600 ha.Ce parc se localise 

au Nord-Est de Fianarantsoa, à 70 km à l’Ouest de l’Océan 

Indien et à 400 km d’Antananarivo.Il est situé entre 47°18’ à 

47°37’ Est de longitude et 21°2’ à 21°25’ Sud de latitude. La 

température moyenne annuelle est de l’ordre de 21°C selon 

Turk en 1995. Quant à la pluviosité, Overdorff a affirmé en 

1996 que la pluie y est saisonnière avec une précipitation 

moyenne de 2000 mm. Deux sites ont été choisis pour 

effectuer notre travail de recherche. Il s’agit de: 

Talatakely:milieu perturbé et plus dégradé.Il est situé à environ 

10 mn de marche de la poste de garde et de contrôle 

du Parc d’Ambodiamontana. Ce site de 1020 m d’altitude 

(Brady et al., 1996), est caractérisé par une visite fréquente 

de touristes. Notons également que, à cause des abattages 

intensifs des arbres par les bûcherons (Kremen, 1992), la 

forêt de Talatakely se trouve fortement dégradée. 

Vatoharanana: un milieu moins perturbé et moins dégradé. 

Cet endroit est à 1090 m d’altitude et se trouve à 2 

heures de marche de Talatakely. Ce champ de forêt était exploité 

par les bûcherons il y a 25 ans (Brady et al.,1996).Mais, 

la dégradation était moins intense que celle de Talatakely 

(Kremen, 1992). La visite des touristes dans cette station 

d’études est également moins fréquente. Vatoharanana est 

donc un site moins perturbé par rapport à Talatakely. 

Les deux stations de recherche sont représentées dans la 

figure 1. 

Espèces étudiées 

Afin de répondre à toutes nos questions, une espèce hautement 

frugivore, qui participe activement à la dispersion des 

graines (Overdorff,1993),a fait l’objet de notre étude.Il s’agit 

du Lémur à ventre roux ou Eulemur rubriventer. Généralement,cette 

espèce de taille moyenne vit en petit groupe de 2


Fig. 1: Localisation des deux sites d’études dans le Parc 

National de Ranomafana. 

à 5 individus. Elle consomme également des feuilles, des 

fleurs, de la terre, des mille-pattes. Selon Overdorff (1996), 

l’animal mange beaucoup plus de fruits et moins de feuilles. 

Eulemur rubriventer présente un dimorphisme sexuel au 

niveau de la morphologie. En effet, le ventre des femelles est 

clair;tandis que la poitrine et la partie inférieure du corps du 

mâle sont visiblement colorées en marron roux. Le mâle 

diffère également de la femelle par la présence de tâche 

blanche sur le coin interne des yeux (Dague et Petter,1988). 

En ce qui concerne notre étude, nous avons suivi cinq 

groupes de lémurs à ventre roux dont trois à Talatakely et 

deux à Vatoharanana. 

Suivi écologique 

Le suivi écologique proprement dit était précédé de la familiarisation 

de tous les groupes d’études. A ce propos, durant 

cinq mois d’études sur terrain,du mois de décembre 2003 au 

mois d’avril 2004, la fréquence d’observation était de cinq 

jours par semaine. L’observation s’étale de sept à douze 

heures dans la matinée et de treize à quinze heures dans 

l’après midi. Au total, l’équipe a suivi cinq groupes pour les 

sites de Talatakely et de Vatoharanana, pendant 588 heures 

45 minutes et 54 secondes. 

La méthode d’enregistrement continu de données a été 

adoptée.Elle nous donne des informations plus fiables et plus 

pratiques par rapport à une méthode d’enregistrement instantané 

(Martin et Bateson, 1986). En outre, nous avons 

collecté les données sur l’alimentation en suivant la méthode 

de "focal animal sampling" (Altmann, 1974). Durant ces observations,nous 

avons enregistré les informations suivantes: 

Site d’études: Talatakely ou Vatoharanana; 

Groupe d’étude; 

Focal animal; 

Date de l’observation; 

Heure d’observation (Début et fin); 

Temps dépensé (Début et fin) à la consommation alimentaire 

et aux autres activités; 

Nom de chaque espèce consommée (plantes ou autres); 

Parties consommées des plantes: fruit, feuille, fleur; 

Etat des parties comsommées (fruit immature ou mûr, 

jeune feuille, feuille mature, fleur ouverte ou fermée); 

Piste la plus proche. 

Analyses statistiques 

Test de similarité entre deux échantillons 

Ce test sert à vérifier la similarité entre le régime alimentaire 

adopté par les lémurs à ventre roux de Talatakely et celui de 

Vatoharanana.Il se base sur la valeur du coefficient de Jaccard 

(Brower et al.,1990).Ce coefficient est donné par la formule 

suivante: 

CC: Coefficient de Jaccard 

S1: Effectif d’espèces végétales, animales et autres dans le 

régime de l’espèce de Talatakely (ET); 

S2: Effectif d’espèces végétales, animales et autres dans le 

régime d’Eulemur rubriventer de Vatoharanana (EV); 

C:Effectif d’espèces végétales,animales et autres communes 

(ET et EV) 

Par souci de conformité, nous avons adopté les échelles 

suivantes: 

0-40 %: faible similarité entre les deux régimes; 

40-60 %: similarité moyenne entre les deux régimes; 

60-80 %: grande similarité entre les deux régimes; 

80-100 %: forte similarité entre les deux régimes. 

Test de Chi-deux: 

Cette méthode sert à comparer la durée moyenne journalière 

(en minute) consacrée à la consommation alimentaire 

des lémurs à ventre roux de Talatakely et de Vatoharanana.A 

cet effet, les variables utilisées sont les durées moyennes 

journalières dépensées à la consommation des fruits, des 

feuilles, des fleurs et autres (sol, eau, champignon, insectes,…); 

et ce dans des intervalles de temps bien déterminés; 

c’-est-à-dire, entre 7 et 8h, 8 et 9h, 9 et 10h, 10 et 11h, 11 et 

12h, 13 et 14h et finalement entre 14 et 15h. Plus précisément, 

elle nous permet de vérifier si la différence entre la 

consommation de ces aliments est statistiquement significative 

ou non dans les deux milieux. 

Résultats 

Temps dédié à l’activité alimentaire 

Fig. 2 montre l’allure générale de la durée moyenne journalière 

en ce qui concerne la consommation alimentaire 

générale des lémurs à ventre roux du milieu plus dégradé de 

Talatakely et celle du site moins dégradé de Vatoharanana. 

Selon cette figure,la prise de nourriture débute entre 7 et 8h. 

Concernant le site de Talatakely,elle dessine un pic entre 9 et 

10h. Cette activité diminue jusqu’à 12h, puis remonte pour 

atteindre le maximum vers 14 à 15h. Quant à l’Eulemur rubriventer 

de Vatoharanana,le pic de l’alimentation se situe entre 

8 et 9h. La courbe diminue jusqu’à 12h environ. Ensuite, une 

légère remontée de l’activité est constatée jusqu’à 15h envi-


ron. L’hypothèse nulle est acceptée car l’analyse statistique 

( 2= 9,95 ;avec ddl=6,et p>0,05) indique une différence non 

significative concernant la prise de la nourriture entre 

chaque intervalle de temps. 

20 

18 

16 

14 

12 

10 

8 

6 

4 

2 

0 

7-8h 8-9h 9-10h 10-11h 11-12h 13-14h 14-15h 

Intervalle de temps 

Talatakely Vatoharanana 

Fig. 2: Temps consacré à la consommation alimentaire pour 

Eulemur rubriventer dans les deux sites d’étude: Talatakely et 

Vatoharanana. 

Durée en minute 

Similarité entre les régimes alimentaires d’Eulemur rubriventer du 

site dégradé de Talatakely et de celui de la station moins dégradée 

de Vatoharanana 

L’inventaire des espèces consommées par les lémurs à 

ventre roux dans les deux sites nous a permis de calculer le 

coefficient de Jaccard afin de tester s’il existe ou non une 

similarité de régimes dans les deux milieux. Nous avons 

recensé 52 espèces de plantes qui sont utilisées comme 

source de leur nourriture pour le site de Talatakely. En 

revanche, 60 espèces sont inventoriées dans le site de 

Vatoharanana.Il est à noter que 35 d’entre elles sont à la fois 

consommées à Talatakely et à Vatoharanana. La liste des 

espèces végétales utilisées par ces animaux comme source 

de leur nourriture (avec la durée de consommation correspondante) 

est résumée dans le tableau récapitulatif suivant. 

Tab. 1: Liste des espèces végétales consommées (61) par 

Eulemur rubriventer, avec la durée de consommation correspondante, 

dans les deux sites d’étude. 

Nom 

malagasy 

Ramy 

Kalafambakaka 

Durée (en mn) 

Genre Famille Fr Fe Fl Tala Vato 

Canarium 

madagascariensis 

Burseraceae + + 

Oncostemum sp. Myrsinaceae + + + 23,57 242,55 

Vakoana Pandanus sp. Pandanaceae + + + 21,40 

Nonoka Ficus sp. Moraceae + 24,02 269,75 

Vahimberana 

Strongylodon c 

raveniae 

Fabaceae + 2,03 189,72 

Voara Ficus sp. Moraceae + 70,08 39,45 

Voara rano Ficus botryoides Moraceae + + 23,13 36,15 

Sandramy Protorhus sp. 

Anacardiaceae 

+ 17,08 

Voandavenona 

+ 12,70 87,63 

Famakilela Ficus sp. Moraceae + + 3,65 7,55 

Tavolo malady Cryptocarya 

acuminata 

Lauraceae + 98,32 

Vahitamboro Danais sp. Rubiaceae + + 19,10 12,97 

Mahanoro 

Streblus 

dimepate 

Moraceae + 62,58 

Tsirika Pandanus sp. Pandanaceae + 38,00 1,67 

Rotra Syzygium sp. Myrtaceae + 27,28 

Apana Ficus sp. Moraceae + 8,070 135,15 

Sira Neodypsis sp. Arecaceae + 36,30 21,47 

Rotra mena Syzygium sp. Myrtaceae + 9,020 17,32 

Fandramanana Aphloia 

theaeformis 

Flacourtiaceae 

+ 127,82 

Velatra spécial Ruellia sp. Acanthaceae + 68,68 27,05 

Durée (en mn) 

Nom 

malagasy 

Genre Famille Fr Fe Fl Tala Vato 

Faritraty Memecylon sp. 

Melastomataceae 

+ 16,83 101,57 

Maranitratoraka 

Vernonia sp. Asteraceae + 68,40 

Vahivoraka 

Mendoncia 

avani 

Mendonciaceae 

+ + 169,48 

Fatsikahitra Alberta humblotii Rubiaceae + + 9,03 425,63 

Albizia Albizia chinensis Leguminosae + 29,67 

Rotra fotsy Syzygium sp. Myrtaceae + 30,97 

Tongoalahy Bakerella sp. 

Loranthaceae 

+ + 3,25 25,68 

Kalafana 

spécial 

Oncostemum 

botryoides 

Myrsinaceae + + + 7,67 130,45 

Hafipotsy Grewia sp. Tiliaceae + 35,92 

Fanalamangidy + 

Vavaporetaka Melanophylla 

crenata 

Melanophyllaceae 

+ 26,05 16,58 

Kaboka Voacanga sp. Apocynaceae + 86,73 66,30 

Sandramy fot- 

Protorhus sp. 

syAnacardiaceae 

+ 20,32 

Apaliala Treculia africana Moraceae + 14,85 

Ramandriona Dilobeia thouarsii Proteaceae + 7,15 15,28 

Andriambo. 

lamena 

Menispermaceae 

+ 138,73 

Bararata Gaertnera sp. Rubiaceae + 26,75 11,62 

Nato jabo 

Mammea 

vatoensis 

Clusiaceae + 69,60 

Tavilona Vernonia sp. Asteraceae + 4,07 

Malanimanta 

Apodytes 

thouvenotii 

Icacinaceae + 3,52 

Voantsosoka + 45,08 

Lambinanala Nuxia sp. Loganiaceae + 17,88 

Rahiaka 

Chrysophyllum 

boivinianum 

Sapotaceae + 6,30 663,50 

Amontana Ficus lutea Moraceae + 22,42 

Goavy 

Psidium 

cattleianum 

Myrtaceae + 589,02 

Fanorafa 

Euphorbiaceae 

+ 173,10 

Kimba spécial Symphonia sp. Clusiaceae + 3,23 

Veso 

Terminalia tetranoCombretaraceae + 2,02 

Vahirano Cissus sp. Vitaceae + 15,73 104,17 

Fohaninasity Psychotria sp. Rubiaceae + 260,93 

Rohindambo Smilax anceps Smilacaceae + 5,67 

Sary 

Potameia 

chartacea 

Lauraceae + 36,60 

Kalamasina 

Embelia madagascariensis 

Myrsinaceae + 10,43 

Ambora 

Tambourissa 

thouvenotii 

Monimiaceae + 10,85 

Harongana 

Harungana mada- 

Clusiaceae 

gascariensis 

+ 21,45 

Nato spécial Sideroxylon sp. Sapotaceae + 1,82 

Holatra Champignon + 

Amboralahy 

Decarydendron he- 

Monimiaceae 

lenae 

+ 10,33 

Sandramy 

Mena 

Protorhus sp. 

Anacardiaceae 

+ 1,50 

Inconnue 1,37 8,82 

Champignon 6,60 

Tala = site de Talatakely; Vato = site de Vatoharanana; Fr = fruits; Fe = feuilles ; 

Fl = fleurs.; *: consommé 

Le calcul du coefficient de Jaccard offre une valeur de 0,45; 

soit 45 %. Il en découle que le régime alimentaire d’Eulemur 

rubriventer des deux sites présente une similarité moyenne. 

Aussi, la différence entre le régime d’Eulemur rubriventer des 

deux milieux peut être révélée par la constatation de la 

durée consacrée à la consommation de chaque catégorie 

alimentaire telle que les fruits, les fleurs, les feuilles et bien 

d’autres (Cf. Tab. 2 et Fig. 2). 

En se basant sur ce tableau récapitulatif (Tab.2) et sur la Fig.2, 

nous constatons que dans le milieu dégradé de Talatakely, la 

consommation des fruits s’avère très importante par rapport 

à celle du milieu moins dégradé de Vatoharanana. De 

plus en comparant avec l’espèce du site de Talatakely,celle de


la station de Vatoharanana comble beaucoup plus sa nourriture 

avec des fleurs, des feuilles, et d’autres types d’aliments. 

Tab. 2: Comparaison de la consommation journalière de 

chaque catégorie alimentaire d’Eulemur rubriventer dans les 

deux sites durant la période d’observation (en minute). 

Sites Fruit Fleur Feuille Autres 

Talatakely 80,80 0,52 4,54 0,12 

Vatoharanana 72,58 8,32 11,16 1,63 

Durée de la consommation 

(en minute) 

90 

80 

70 

60 

50 

40 

30 

20 

10 

0 

Fruit Fleur Feuille Autres 

Catégorie alimentaire 

Talatakely Vatoharanana 

Fig.3:Allure de la consommation journalière de chaque catégorie 

alimentaire d’Eulemur rubriventer dans les deux sites 

durant la période d’observation. 

Discussion 

Eulemur rubriventer est un lémurien hautement frugivore. 

Mais la proportion des fruits qu’il consomme varie suivant le 

degré de perturbation,de dégradation,de l’altitude du milieu, 

ainsi que de la saison. Notons également que la consommation 

des différentes catégories alimentaires dépend de la 

disponibilité des ressources alimentaires (Rasolofonirina, 

2001). Lors de notre étude, nous constatons que dans le 

milieu dégradé de Talatakely, la consommation des fruits 

s’avère très importante que dans le milieu moins dégradé de 

Vatoharanana.De plus,en comparant avec l’espèce du site de 

Talatakely, celle de la station de Vatoharanana comble beaucoup 

plus sa nourriture avec des fleurs, des feuilles, et d’autres 

types d’aliments. Ceci est dû certainement à la saison 

cyclonique durant laquelle le vent est violent.Comme le site 

de Vatoharanana est beaucoup plus élevé par rapport à Talatakely,il 

s’avère logique qu’il est beaucoup plus affecté par ce 

vent violent.En effet,les fruits deviennent rares car beaucoup 

d’entre eux tombent par terre. Selon Zaonarivelo (1999), 

pendant la période de crise, Varecia variegata variegata augmente 

le taux de folivorie même si les feuilles sont des aliments 

de compensation.Cette stratégie adoptée par l’animal 

lors de la période de crise est également observée chez 

d’autres espèces de lémuriens de la forêt dense humide 

(Ganzhorn, 1988) entre-autres l’Eulemur rubriventer,leEulemur 

mongoz qui se nourrissent de fleurs en plus des fruits et 

des feuilles durant la période de floraison (Sussman, 1975). 

Par ailleurs,Garbutt (1999) argumente que les fruits constituent 

la majeure partie de l’alimentation d’Eulemur rubriventer. 

Mais quand ils ne sont pas disponibles,les feuilles et les fleurs 

sont aussi consommées (Rasolofonirina, 2001). Notons à la 

même occasion que les primates adoptent différentes stratégies 

pour affronter le manque de nourriture de base (le 

fruit pour notre cas) soit en augmentant le temps de la 

recherche de nourriture en se déplaçant beaucoup, soit en 

acceptant de consommer des aliments de basse qualité 

(Zaonarivelo, 1999). 

Par rapport au milieu dégradé,l’abondance des fruits dans le 

milieu moins dégradé est évidente.Mais en tenant compte de 

l’étude phénologique mensuelle des plantes recensées, ce 

n’est pas toujours le cas; car plusieurs facteurs pourraient 

agir sur ce milieu.Ainsi le climat,en particulier la pluviosité et 

le cyclone, influence la qualité et la quantité de la nourriture 

disponible (Dajoz, 1985). En effet, durant la période cyclonique, 

l’altitude de la station joue un rôle important sur les 

arbres à semences. Autrement dit, plus l’altitude d’un milieu 

est élevée,plus le vent agit directement sur les arbres et plus 

les fruits tombent. Par conséquent, les arbres portent moins 

de fruits. L’animal est obligé de se rabattre sur d’autres 

catégories alimentaires comme les feuilles, les fleurs, les 

champignons, les insectes pour pouvoir combler l’insuffisance 

de nourriture de base.Ce cas se rencontre dans le site 

de Vatoharanana qui est considéré comme milieu moins 

dégradé et qui est situé à une altitude plus élevée (1090 m) 

par rapport au site dégradé de Talatakely qui se trouve à une 

altitude de 1020 m (Brady et al, 1996). Concernant le site de 

Talatakely, la dispersion des fruits dans l’espace est insuffisante 

à cause de la dégradation de ce milieu.Ainsi,au lieu de 

combler sa nourriture par d’autres types d’aliments, il est 

contraint à se déplacer loin afin de consommer de la nourriture 

de haute qualité qui lui apporte beaucoup plus d’énergie 

comme le glucide, le lipide et les protéines (Zaonarivelo, 

1999). 

Finalement, en se basant sur les données phénologiques, 

nous avons constaté que la consommation de chaque catégorie 

alimentaire varie suivant leur disponibilité mensuelle 

dans chaque mileu. Au mois de décembre, par exemple, 

l’Eulemur rubriventer de Vatoharanana consomme beaucoup 

plus de fleurs que de fruits par rapport à celui de Talatakely 

car pendant ce mois, les fleurs y sont disponibles. Remarquons 

également que l’espèce de la station de Talatakely ne 

consomme que des fruits durant la fructification de l’espèce 

introduite de goyave (Psidium cattleianum). Ce type de fruit 

est très apprécié par l’animal. 

Le régime alimentaire d’Eulemur rubriventer est moyennement 

similaire dans les deux milieux étudiés. 

Cependant, quelques espèces de plantes consommées par 

l’animal sont propres à chaque site. L’espèce introduite de 

goyave Psidium cattleianum se rencontre uniquement à Talatakely. 

Notons que l’introduction de cette espèce marque la 

dégradation de cette station. Par contre, l’animal de Vatoharanana 

a l’opportunité de consommer, par exemple, l’espèce 

Mammae vatoensis qui est endémique à ce site. Cette 

similarité moyenne entre les deux régimes implique que la 

dégradation du site de Talatakely n’est pas encore poussée à 

l’extrême (Randriamahaleo,2005).En effet,il est classé parmi 

les sites moyennement dégradés. Selon Tam-Alkis (1997), le 

site de Talatakely est séparé de Vatoharanana par une barrière 

biogéographique (rivière Fompohonona). Il paraît que 

la dispersion des graines de part et d’autre de cette rivière 

est empêchée. Voilà pourquoi certaines espèces de plantes, 

utilisées comme source alimentaire, caractérisent uniquement 

l’un de ces sites. Par conséquent son régime varie 

suivant le site. 

Conclusion 

Cette étude nous a permis de fournir plus d’informations sur 

la relation entre la dégradation de l’habitat d’Eulemur rubriventer 

et sa consommation alimentaire.L’investigation révèle 

une similarité moyenne entre le régime alimentaire adopté 

par l’animal de Talatakely et celui de Vatoharanana. L’espèce 

semble être principalement frugivore quelque soit le site. 

Cependant, la proportion de consommation varie suivant le 

milieu. A Talatakely, par exemple, ce lémur à ventre roux se


nourrit beaucoup plus de fruits par rapport à celui de Vatoharanana 

où il comble sa nourriture avec des fleurs, des 

feuilles, et bien d’autres catégories alimentaires. Ceci est dû 

sans doute au passage des deux cyclones (Elita et Gafilo) 

dans la région durant la période d’étude. En fait, Vatoharanana 

se trouve à une altitude très élevée par rapport à Talatakely.En 

effet,les vents violents agissent directement sur les 

arbres fruitiers, conduisant ainsi l’insuffisance des fruits. 

Aussi, la consommation de Psidium cattleianum (Myrtaceae), 

qui est une espèce introduite propre à Talatakely semble être 

très importante dans ce milieu. Par contre, Chrysophillum 

boivinianum s’avère être la plus appréciée par l’animal de 

Vatoharanana. Toutes ces constatations nous conduisent à 

dire que la consommation alimentaire d’Eulemur rubriventer 

paraît être conditionnée par la disponibilité de la nourriture 

dans chacun des sites visités et par l’état de l’habitat. 

Remerciements 

Nous tenons à remercier le Centre Valbio, le MICET représentés 

respectivement par le Professeur Patricia Wright et le 

Docteur Benjamin Andriamihaja pour leur soutien et leur 

collaboration durant la longue haleine de travail dans le Parc 

National de Ranomafana. Nos vifs remerciements s’adressent 

également à tous les guides de recherche pour leur 

assistance (Telo Albert, Victor, Koto, Nirina) et à tout ce qui 

contribue, de près ou de loin à la réalisation de ce projet. 

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des Sciences, Université d’Antananarivo. 

Observations of terrestrial latrine behaviour 

by the southern gentle lemur Hapalemur 

meridionalis in the Mandena littoral 

forest, southeast Madagascar 

Timothy M. Eppley* and Giuseppe Donati 

Nocturnal Primate Research Group,Department of Anthropology 

and Geography, Oxford Brookes University, Gipsy 

Lane, OX3 0BP, Oxford, UK 

*Corresponding author: eppleyti@gmail.com 

Key words: southern gentle lemur, Hapalemur meridionalis, 

defecation, latrines, Mandena 

Latrine behaviour is defined as the non-random selection of 

a specific defecation site,and although it is rarely described in 

primates it is well known among other mammalian species as 

a form of olfactory communication (Irwin et al.,2004).Olfactory 

compounds, which convey chemical signals, are transmitted 

via scent-producing skin gland secretions, saliva, and/ 

or waste products (Epple, 1986). Olfaction signals may be 

advantageous as they are not limited spatially and temporally, 

allowing individuals of a predominately visual communication


to receive signals when distant from the source (Schilling, 

1979;Irwin et al.,2004).Olfactory signals may transmit information 

pertaining to environmental familiarisation (Schilling, 

1979), reproductive behaviour and condition (Epple, 1986), 

territorial demarcation (Mertl-Milhollen,1979),and inter- or 

intra-group spacing (Schilling, 1979; Epple, 1986). 

Previous literature has discussed latrines by varying in location 

(arboreal, terrestrial, or subterranean), and being further 

analysed by volume of faeces and spatial distribution 

(Schilling, 1980; Boonstra et al., 1996; Irwin et al., 2004; Pouvelle 

et al., 2009). This behaviour appears to be well studied 

within other mammals but for primates, especially strepsirhines, 

latrine use is only sparsely mentioned within much 

broader scope research (Irwin et al.,2004).In this report we 

present our observations of this peculiar behaviour exhibited 

by the southern gentle lemur, Hapalemur meridionalis,a 

threatened primate that occurs in southeast Madagascar.We 

will use our observations to review the hypotheses offered 

thus far to explain latrine use,i.e.advertisement of sexual cycle,predation 

avoidance,intra- group and inter-group spacing 

in the context of the fragmented littoral forest. 

Methods 

This research was conducted from May to July 2008, on the 

southern gentle lemur within the Mandena littoral forest 

(24°95’S 46°99’E), a coastal forest in southeast Madagascar. 

The littoral forest is among the most endangered ecosystems 

in Madagascar (Bollen and Donati, 2006) and Mandena 

is a protected conservation zone encompassing 230 hectares 

of fragmented and partially degraded littoral forest 

interspersed with marsh and swamp. Three groups of H. 

meridionalis (mean = 5.7 ind/group) were habituated and 

followed daily from dawn to dusk with 62 hours of observation 

recorded (Eppley and Donati, in press). These lemurs 

are of particular interest, as they do not subsist exclusively 

on bamboo like the majority of their congeners. Rather, the 

southern gentle lemur exhibits a dietary predilection for 

terrestrial (turf) grasses while displaying a unique grazing 

behaviour (Eppley and Donati, in press). Although the main 

research being conducted focused on the feeding ecology of 

these animals, opportunistic observations of latrine behaviour 

were collected ad libitum. In addition to habitat characteristics 

and GPS waypoints taken at the feeding and resting 

sites of three groups, latrine locations were also recorded. 

Spatial analyses were carried out with ArcMap version 9.3, 

with the outermost feeding and resting site waypoints for 

each group being used to create the minimum polygon for 

their respective ranging areas. 

Results 

On three separate occasions an entire group of foraging H. 

meridionalis were witnessed descending to the ground and 

defecating in succession either near or under a high-rooted 

tree of different species.Latrine bouts were recorded ad libitum 

on a single occasion for Group B (four individuals) and 

twice for Group C (seven individuals). Observations took 

place within the forest fragment, and were never witnessed 

in open canopy areas. The three occurrences of this terrestrial 

latrine behaviour were observed shortly after individuals 

had awoken from a midday resting bout (between 11:00 

and 14:00) at distances approx. 20 m from the resting site. 

The three locations where the latrine behaviour was exhibited 

are shown in Figure 1.Little overlap between the ranging 

areas of the three groups was observed with 0.92 ha (7.74 %) 

calculated to exist between groups A and B,0.18 ha (1.68 %) 

between groups A and C, and 0.06 ha (0.73 %) between 

groups B and C. After an individual had defecated, they 

ascended three to four meters and continued feeding.Subsequently, 

the group followed an order of sequential defecation, 

where by one conspecific would defecate and the next 

would rapidly follow.The individuals remaining at an elevated 

height always appeared vigilant, scanning the surrounding 

area and sometimes continuing to forage, while the conspecific 

was on the ground. Upon inspection of two of the 

defecation sites, accumulations of hardened faecal matter 

were identified. Dissimilarly, the third site consisted of only 

fresh faeces with no sign of old faecal matter. 

Fig. 1: Ranging areas of the three observed H. meridionalis 

groups within the northeast corner of the Mandena littoral 

forest. The three latrine sites were observed within the territorial 

boundary buffer zones of Groups A & C and B & C. 

Discussion 

Among primates,latrine behaviour has been recorded (Table 

1) in Alouatta seniculus (Gilbert, 1997; Feeley, 2005; Neves et 

al., 2009; Pouvelle et al., 2009), Ateles geoffroyi (Notman et al., 

2009), as well as the strepsirhines: Cheirogaleus major, C. 

medius, Lepilemur leucopus (Charles-Dominique and Hladik, 

1971; Russell, 1977), L. microdon, L. ruficaudatus, Hapalemur 

aureus, H. griseus, Prolemur simus, and Lemur catta (Irwin et al., 

2004). With the exception of Neotropical primates, however,minimal 

research has been conducted to understand its 

function within primates and latrine utilisation is often a matter 

of debate. In Ranomafana National Park, for example, H. 

griseus have been observed to occasionally use the same 

resting/sleeping sites and since they often defecate very soon 

after they wake up (Tan,pers.comm.),an accumulation of faecal 

material under these trees may appear inadvertently 

latrine-like (Notman et al., 2009; Pouvelle et al., 2009). Thus, 

latrine use by H.griseus may be anecdotal within this population. 

Irwin et al. (2004) presented four non-mutually exclusive 

hypotheses for the possible adaptive function of primate 

latrines:1) advertisement of sexual cycle,2) predation avoidance, 

3) intra-group spacing, 4) and inter-group spacing. 

The conveyance of scent-marks is well known to advertise 

sexual activity and receptivity (Asa, 2008). If this holds true 

for H. meridionalis, latrines should be used more frequently 

during the breeding seasons. It has been reported that H. 

griseus mate between June and July, experiencing a gestation 

length of approximately 137 days (Tan, 2006). Though our 

research potentially overlapped with the mating season, we 

made no observations of mating during the study period.


Thus, we had no possibilities to compare the mating season 

with pre- or post-mating periods, as the hypothesis testing 

would require. However, since two of the latrines at Mandena 

appeared to be utilized long-term, similar to the findings 

of Irwin et al.(2004),we conclude that the advertisement 

of sexual receptivity is unlikely the sole function of lemur 

latrines. 

Tab. 1: Observed primates that have exhibited terrestrial 

latrine behaviour. Adapted from Irwin et al. (2004). 

Species Localities References 

Hapalemur 

meridionalis 

Mandena Conservation 

Zone 

This study 

Hapalemur 

griseus 

Analamazaotra 

Special Reserve 

Irwin et al. (2004) 

Prolemur 

simus 

Ranomafana 

National Park 

P. Wright (in Irwin et al., 2004) 

Lemur catta Isalo National Park 

J. Jernvall and P. Wright 

(in Irwin et al., 2004) 

Lepilemur 

leucopus 

Beza Mahafaly 


L. Nash (in Irwin et al., 2004) 

Lepilemur Manombo Special J. Ratsimbazafy (in Irwin et al., 

microdon Reserve 

2004) 

Lepilemur sp. 

(?microdon) 

Kalambatritra 


Irwin et al. (2004) 

Lepilemur 

ruficaudatus 

Kirindy Forest 

J.U. Ganzhorn (in Irwin et al., 

2004) 

Ateles 

geoffroyi 

Runaway Creek Nature 

Preserve, Belize 

Notman et al. (2009) 

Alouatta 

seniculus 

Nouragues Reserve, 

French Guiana 

Pouvelle et al. (2009) 

Gentle lemurs have a particularly effective predator avoidance 

strategy including camouflage from cryptic pelage, rapid 

flight behaviour,and potential cathemeral activity pattern 

(Mutschler et al., 1999; Curtis et al., 2006; Tan, 2006). Several 

potential predators of Hapalemur exist in the littoral forest. 

There have been documented cases of fossa Cryptoprocta 

ferox preying on H. griseus (Goodman and Pidgeon, 1999; 

Sterling and McFadden,2000).The Madagascar tree boa Sanzinia 

madagascariensis (= Boa manditra) also prey on Hapalemur 

spp.(Goodman et al.,1993;Rakotandravany et al.,1998), 

and several aerial predators (Madagascar harrier hawk Polyboroides 

radiatus, Frances’s sparrowhawk Accipiter francesii, 

Henst’s goshawk Accipiter henstii,common barn owl Tyto alba, 

and the Madagascar long-eared owl Asio madagascariensis) 

represent a threat to medium-sized lemurs (Goodman et al., 

1993; Wright, 1997; Karpanty and Goodman 1999). In fact, 

the concealment of Hapalemur faeces under large high-rooted 

trees may theoretically act as a safeguard against predation 

by impairing the ability of a predator to detect the prey 

population (Boonstra et al.,1996;Irwin et al.,2004).Although 

these observations are in accord with the anti-predator idea, 

single faecal deposits were also detected at indiscriminate 

locations. Thus, more data are necessary to test the hypothesis 

of latrine behaviour as an anti-predator strategy. 

Intra-group spacing has also been suggested to advertise 

proximal resource use and assist in inter-individual spacing 

(Kruuk,1992).In accord with Irwin et al.(2004),however,it is 

unlikely that Hapalemur latrine behaviour is used for intragroup 

spacing, as they live in cohesive family units. 

The territorial demarcation hypothesis suggests that scentmarks 

are placed around home range boundaries to act as a 

delineation of the territory, i.e. inter-group spacing (Mertl- 

Millhollen, 1979; Lewis, 2005). In fact, it is evident from our 

observations that H.meridionalis chose defecation sites in the 

narrow areas of overlap with neighbouring conspecifics 

groups (Fig. 1). If this adaptive function holds true, latrine 

behaviour might be even more common in areas of dense 

population (Irwin et al.,2004),such as the forest fragments of 

Mandena (Eppley and Donati, in press). 

Although the exhibition of preferred, non-random defecation 

sites is most likely multifactorial,latrines in Mandena appear 

to best fulfil the function of inter-group spacing. Therefore, 

latrines may be a low-energy behavioural response to 

the ecological challenge of defending resources with minimal 

rates of agonism (Irwin et al.,2004).In the future,more quantitative 

studies should focus on seasonal and spatial exhibition 

of latrine use to verify whether this behaviour is intrinsically 

linked to territorial delineation and resource defence in 

lemurs. 


We would like to thank the Commission Tripartite of the 

Malagasy government, Ministère de l’Environnement, des 

Eaux et forêts of the Malagasy government,the University of 

Antananarivo, and CAFF/CORE for permission to conduct 

research, as well as the Malagasy Institute for the Conservation 

of Tropical Environments (MICET) for all of their logistical 

assistance. Financial support was provided partly by the 

Chester Zoo (NEZS) and QMM.We would also like to thank 

the QMM Environmental Team, most especially Manon Vincelette, 

Jean-Baptiste Ramanamanjato, Johny Rabenantoandry, 

Faly Randriatafika, and Christophe Rambolamanana for 

all of their advice and logistical help. We are grateful to Jörg 

Ganzhorn for all of his continuous support and scientific advice.Thank 

you to the entire staff of the Oxford Brookes Primate 

Conservation MSc program, especially Simon Bearder, 

Anna Nekaris, and Vincent Nijman. We greatly appreciate 

the GIS assistance of Maureen Mullen. My sincere gratitude 

goes to my field guide Robertin "Tintin" Ravelomanantsoa 

and research assistant Abi Coleman for their companionship 

and tireless help in the marecage. 

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Conservation des lémuriens via la protection 

de leurs habitats et le développement 

communautaire dans les corridors 

de Betaolana et Tsaratanana-Betaolana, 

région de SAVA 

Lala Razafy Fara 1*, Iarilanto Andriamarosolo 2 

1WWF Madagascar & Western Indian Ocean PO, BP 738 

Antananarivo 101, Madagascar 

2WWF Andapa, BP 28, Andapa 205, Madagascar 

*Corresponding author: frazafy@wwf.mg 

Contextes 

Ecologique: 

Madagascar est connu pour sa haute valeur en biodiversité. 

Sa flore et sa faune ont une valeur d’endémicité très élevée 

(au dessus de 80 %) due entre autres à son insularité. Madagascar 

est donc plus riche en espèces endémiques comparées 

à d’autres continents du monde.Le microclimat de l’île a 

permis que d’une région à une autre, la flore et la faune constituent 

une richesse spectaculaire à part. 

Pour la partie Nord de Madagascar, la flore luxuriante, avec 

des espèces endémiques du genre Dalbergia, est encore à 

découvrir.Le WWF a travaillé dans la région de SAVA pour la 

mise en place de deux Aires Protégées (AP),Parc National de 

Marojejy et Réserve Spéciale d’Anjanaharibe Sud, actuellement 

sous gestion du Madagascar National Parks. Pour le 

WWF,la conservation du flux génétique implique le maintien 

et la restauration de la connectivité écologique.Ce maintien 

peut être le plus important paramètre le long des pentes 

altitudinales car cette connectivité est actuellement très 

rare au sein de l’écorégion de l’Est.De plus,il peut constituer 

un refuge pour la biodiversité vu les changements du climat 

(Erdmann et al., 2005). 

Eu égard aux efforts déjà investis dans la protection des deux 

AP (Parc de Marojejy et de la Réserve Spéciale d’Anjanaharibe 

Sud), le WWF a continué ses efforts de conservations 

dans les deux corridors forestiers (Betaolana et Tsaratanana-Betaolana). 

Dans ces localités, les espèces endémiques 

sont aussi très remarquables comme le palmier Marojejya 

insignis et la fougère Asplenium marojyense. La forêt dense et 

humide de cette partie de l’île abrite une multitude de faune 

à découvrir et à protéger. 

La forêt occupe encore 35,60 % de la région de SAVA (données 

images satellites de 2000),ce taux est élevé par rapport 

à d’autres régions de Madagascar.La richesse en biodiversité 

de cette zone est démontrée par de nombreuses études,sur 

la flore et sur la faune, conduites dans cette zone. 

Pour ces deux corridors Betaolana et Tsaratanana-Betaolana, 

les inventaires effectués par Rajaonson et Rakotonirina 

(2007) pour le WWF ont montré l’existence de 10 espèces 

de lémuriens pour le corridor Betaolana, et de sept espèces 

pour le corridor Tsaratanana Betaolana, 84 espèces d’oiseaux 

pour l’ensemble des deux corridors. En comparaison 

avec les inventaires des lémuriens effectués par Goodman et 

al. (2003), la région d’Andapa, incluant les AP de Marojejy et 

d’Anjanaharibe-Sud et le corridor forestier de Betaolana, 

possèdent une richesse spécifique en communautés de lémuriens 

et tient une place importante en biodiversité à Madagascar.La 

synthèse des études déjà effectuées dans la zone a 

recensé 12 espèces de lémuriens dont: Microcebus rufus, 

Cheirogaleus major, Allocebus trichotis, Phaner furcifer, Avahi 

laniger, Lepilemur mustelinus, Daubentonia madagascariensis, 

Hapalemur griseus griseus, Eulemur rubriventer, Eulemur fulvus


albifrons, Propithecus candidus, Indri indri. Parmi ces espèces 

citées, celles non rencontrées dans les deux corridors sont: 

Indri indri et Phaner furcifer (pour les deux corridors Betaolana 

et Tsaratanana-Betaolana) et Allocebus trichotis pour le 

corridor Tsaratanana Betaolana). 

Parmi les espèces de lémuriens, le Simpona (Propithecus candidus) 

est une des 25 espèces de primates les plus menacées 

dans le monde (Mittermeier et al., 2005), d’où l’intérêt de se 

focaliser sur sa conservation. Certains auteurs (Petter et al., 

1979; Tattersall, 1982) pensent que l’habitat du Simpona 

pourrait éventuellement s’étendre vers le Sud jusqu’aux 

forêts de la péninsule de Masoala. Il est par contre prouvé 

que le versant Ouest de Anjanaharibe-Sud, actuellement 

sans statut de protection, héberge une population importante 

de Simpona. La population de Simpona de la région 

Andapa et donc mondiale est estimée à une valeur comprise 

entre 100-1000 individus (Mittermeier et al., 1994). Des 

inventaires effectués par le WWF vers la fin 2006 ont relevé 

que le Simpona se rencontre aussi vers le Nord du Corridor 

de Betaolana en allant vers le Nord Ouest du côté du massif 

de Tsaratanana. 

Social et économique: 

Les deux corridors forestiers, Betaolana et Tsaratanana 

Betaolana, couvrent approximativement une superficie respective 

de 16 500 ha et de 130 900 ha et sont très riches en 

forêt.En se référant aux délimitations sur la carte 1,les deux 

corridors renferment encore jusqu’à 90 % de forêts. Situées 

sur des sols ferralitiques, les forêts des deux corridors sont 

localisées entre les altitudes 850 et 1600 m pour Betaolana 

et 800 à 2280 m d’altitude pour Tsaratanana-Betaolana. Les 

deux corridors jouissent encore du climat humide de la 

région de SAVA. 

Les deux corridors appartiennent aux districts d’Andapa 

(pour le Corridor de Betaolana) et de Bealanana (pour le 

corridor de Tsaratanana-Betaolana). Les populations des 

communes locales vivant à la périphérie de ces corridors 

sont estimées, en 2009, à 72 521 pour le corridor Betaolana 

et à 41 333 pour le corridor Tsaratanana-Betaolana (à partir 

des données d’INSTAT 1993). 

Dans la région de SAVA, les forêts dans les deux corridors 

Betaolana et Tsaratanana Betaolana font partie des domaines 

forestiers de l’Etat. Ils n’ont de ce fait pas de statuts 

particuliers (Garreau et Manantsara, 2003). Toutefois, le 

WWF, connaissant l’importance de la flore et la faune dans 

cette zone, a mis en œuvre de 2005 à 2008 deux projets 

spécifiques pour la conservation des lémuriens et de leurs 

habitats dans la région de SAVA. Dans le cadre des projets 

mis en œuvre par le WWF qui se sont succédés dans la région 

de SAVA, celui-ci a travaillé avec les populations locales 

pour mettre ces deux corridors sous statuts de Nouvelles 

Aires Protégées. Pour le Corridor Betaolana, huit communautés 

de bases ayant reçu des contrats de transferts de gestion 

des lots de forêts ont déjà déposé leur manifestation 

d’intérêt dans ce sens en 2008.Les différentes étapes et procédures 

à respecter suivent actuellement normalement leur 

cours. 

Cet article, eu égard à tous les acquis dans le cadre des 

projets du WWF, est une capitalisation des expériences de 

WWF Madagascar pour les projets de conservation des 

lémuriens et de leurs habitats dans le Nord de Madagascar. 

L’un des deux projets est connu sous son appellation courte, 

projet Simpona. Son but est en effet de mettre en exergue 

cette espèce phare bien qu’elle ne soit pas la seule à être 

protégée dans le cadre des projets du WWF. 

Approche de conservation 

Les sites des deux projets 

La région de SAVA se trouve dans l’ex Province d’Antsiranana 

au Nord de Madagascar. Les deux projets du WWF 

dans cette localité ont été conduits dans deux corridors 

forestiers entre les Aires Protégées Tsaratanana (Réserve 

Naturelle Intégrale), Marojejy (Parc National) et Anjanaharibe 

Sud (Réserve Spéciale).La carte suivante montre la situation 

générale de ces localités. 

Les objectifs de conservation 

A partir de la connaissance des richesses de cette localité, 

des pressions et menaces sur les espèces cibles,la finalité des 

deux projets était de freiner l’utilisation irrationnelle de la 

forêt tout en construisant un environnement où les populations 

locales pourront vivre en harmonie avec la nature. 

Découlant de cette finalité,les objectifs des deux projets ont 

été de: 1) Mieux connaître le niveau de menace sur les 

lémuriens et sur leurs habitats; 2) Elaborer des scénario de 

zonages et de gestion; 3) Conscientiser et éduquer les 

communautés locales concernant les menaces et la protection 

des lémuriens; 4) Initier de nouveaux transferts de 

gestion des forêts auprès des communautés locales de base 

(sur la partie Ouest du Corridor de Betaolana); 5) Protéger 

et/ou restaurer les habitats des lémuriens; 6) Renforcer la 

protection des lémuriens par l’extension et la création de 

nouvelles aires protégées; 7) Vulgariser et promouvoir les 

produits de substitution des principaux produits forestiers; 

8) Procéder au suivi d’évaluation des méthodes et procédures 

de gestion internes des associations des forêts; 9) 

Restaurer les terrains défrichés de la périphérie du corridor. 

En bref,les deux projets essaient de renforcer les conditions 

requises pour la conservation à long terme de la biodiversité 

via la conservation des fonctions écologiques des deux corridors 

(Betaolana et Tsaratanana-Betaolana) de manière participative. 

Diagnostic participatif: Mieux connaitre pour mieux 

protéger 

Cartographie 

A partir des connaissances issues de la littérature, une cartographie 

simplifiée basée sur des cartes topographiques 

(échelles 1/100 000) et des interprétations des images satellites 

2000 a été élaborée. L’objectif est de partir des limites 

des forêts à partir de ces images pour délimiter des zones où 

les inventaires forestiers vont être réalisés.De même,de par 

cette technique,des zonages forestiers et des cartes d’occupation 

des sols sont élaborés.Le zonage forestier se base sur 

les états de dégradations des forêts. Les unités semblables 

ont été groupées dans une même catégorie définie pour un 

objectif spécifique (conservation, enrichissement / restauration, 

droit d’usage). 

Inventaire des lémuriens par des primatologues 

Pour mieux intégrer les populations locales dans l’importance 

de la conservation des lémuriens,elles ont été invitées 

à participer aux inventaires des lémuriens.Leur connaissance 

de base, combinée aux connaissances scientifiques de deux 

primatologues recrutés ont été pour renforcer les acquis 

dans le cadre de la littérature.Cette approche a aussi permis 

d’élaborer des documents de base pour le suivi des lémuriens 

et de choisir des espèces pour la restauration 

forestière.


Fig. 1: Carte de localisation générale des sites. 

Etudes des pressions et menaces sur les lémuriens et leurs 

habitats 

Les études sur les pressions et menaces ont été conduites 

selon des enquêtes informelles et formelles et selon des observations 

directes.Les enquêtes informelles servent à la fois 

à gagner la confiance des populations locales et à recueillir le 

maximum d’informations. Les enquêtes formelles sont utilisées 

par les consultants primatologues pour une meilleure 

représentativité des réponses. Les membres de l’équipe de 

WWF,à mesure de l’avancement de la conduite des activités 

dans le cadre des deux projets, ne cessent de faire des 

recoupements entre toutes les réponses obtenues. 

La participation des membres des communautés de base est 

indispensable dans les deux types d’enquêtes pour qu’ils se 

sentent responsables et pour confronter les réponses au fur 

et à mesure que des biais sont observés. Les méthodes 

classiques de recherches participatives utilisées sont du 

genre MARP: Méthode Accélérée de Recherche Participative 

et PALM: Participatory Learning Methods ou Méthode 

d’apprentissage Participatives. Les membres de l’équipe du 

WWF ont reçu auparavant des formations sur ces méthodes. 

Dans le cadre de leur travail, ces méthodes sont 

conduites lors des premières approches au sein d’une nouvelle 

communauté demandant à gérer une ressource naturelle 

dans leur terroir. 

Responsabilisation sous forme de transfert de 

gestion 

Inventaire forestier et zonage 

Dans chaque localité de travail avec une communauté, des 

inventaires forestiers ont été conduits.La finalité est de connaitre 

les potentialités des différents types de forêts en 

produits forestiers ligneux et non ligneux. Une uniformisation 

des méthodes de relevés a été réalisée. A partir des 

cartes de zonages élaborées, les inventaires ont été concentrés 

dans les zones de conservations et des droits 

d’usage.Des parcelles imbriquées sont définies.Elles ont des 

surfaces de 100 x 100 m pour des diamètres de 1,30 m (dhp) 

supérieur ou égal à 30 cm;deux parcelles de 25 x 25 m pour 

les arbres de dhp compris entre 10 cm à 30 cm; et deux 

parcelles de 5x5m.Lesdimensions de ces parcelles peuvent 

varier en fonction du relief (si le terrain est trop accidenté,la 

réduction des dimensions est préconisée) et leur nombre 

varié.Mais quelques soient les variantes,une moyenne de un 

hectare par type de forêt est requise pour les gros diamètres. 

Elaboration de plan d’aménagement 

En vue de l’obtention des contrats de transfert de gestion 

des forêts auprès du Ministère des forêts, chaque communauté 

devrait établir des plans d’aménagement des forêts.


Les plans d’aménagement suivent les directives proposées 

par la Direction Générale des forêts. Le WWF,pour l’élaboration 

de ces plans,a surtout veillé à ce que les prélèvements 

respectent le principe de la durabilité à savoir: ne prélever 

que le volume correspondant aux accroissements moyens 

annuels. Pour un meilleur équilibre entre les besoins des 

populations locales, des reboisements en espèces introduites 

sont aussi proposés dans ces plans d’aménagement, 

mais dans des parcelles en dehors des forêts naturelles. 

Appropriation sous forme de contrôle 

L’approche d’appropriation sous forme de contrôle a aussi 

été développée pour atteindre les objectifs des deux projets. 

Elle consiste à encourager les membres des communautés à 

l’établissement de parcelles d’observation en forêt en se 

basant sur les pistes existantes et sur les signes visibles de 

pressions (coupe, pièges). Des représentants dynamiques au 

sein de chaque communauté sont ainsi élus et font des visites 

(contrôles) régulières, parfois inattendues, en forêts. Ces 

visites sont organisées pour observer la biodiversité 

(floraison des plantes,augmentation du nombre des groupes 

de lémuriens observés, passage d’un oiseau ou autre type 

d’animal faisant la particularité de leur forêt etc.) et pour 

constater si des pressions sur les lémuriens et la forêt sont 

encore présentes. Ces membres notent leurs observations 

dans des cahiers réservés pour ces observations et tout 

autre évènement qu’ils jugent importants (rencontre avec 

d’autres personnes, etc.). 

Mesures d’accompagnement 

Le WWF défini les mesures d’accompagnement de toute 

activité développée pour contribuer à l’amélioration des 

conditions de vie des populations locales. Ces activités doivent 

à la fois réduire les pressions sur la biodiversité en générale 

et sur les cibles de conservation en particulier (forêt 

naturelle et lémurien) et compenser les efforts de conservations 

entrepris. L’approche consiste d’abord à analyser les 

résultats de toutes les études préalables aux transferts de 

gestion des forêts. A partir de ces analyses, les potentialités 

de chaque terroir (à partir des cartes d’occupation des sols 

et des rendements obtenus pour chaque spéculation engagée) 

dans lesquels vivent les communautés sont dégagées. 

Les mesures à développer cherchent ainsi à augmenter les 

sources de revenus des paysans et à améliorer leur alimentation 

pour une meilleure santé. 

Résultats et discussions 

Inventaires forestiers et zonages 

Au total, 14 inventaires forestiers (correspondants au nombre 

de transfert de gestion des forêts) ont été effectués dans 

la région de SAVA. La potentialité des forêts est variable. Le 

nombre des arbres de dhp > 30 varie de 200 à 400/ha. Au 

total, 200 espèces de plantes (ligneuses et non ligneuses) 

sont inventoriées. D’un site à l’autre, le nombre des espèces 

ligneuses inventoriées varie de 45 à 87. 

Les espèces les plus fréquemment rencontrées sont: Tambourissa 

religiosa,Weinmannia rutembergii,Zantoxyllon mananarense, 

Chrysophyllum boivinianum, Canarium madagascariensis, 

Symphonia fasciculata, Diospyros aff.ambilensis, Macaranga 

decaryana, Erythroxylum sphaeranthum, Brachylaena merana, 

Syzygium emirnense, Uapacca densiflora, Ocotea cymosa… 

En fonction de l’éloignement des forêts par rapport aux villages, 

leur état de dégradation diffère. Cette observation a 

permis de faire trois grande classifications: forêt naturelle 

plus ou moins intacte,classée à protéger ou à conserver (for- 

mant donc le noyau dur);forêt partiellement dégradée,localisée 

encore en plein cœur de la forêt,classée comme forêt à 

restaurer;forêt naturelle à faible potentialité, due à un degré 

d’écrémage localisée à la périphérie des lisières, classée 

comme zone de cantonnement de droit d’usage. 

Transfert de gestion des forêts 

Le WWF a pu mettre en place dans le cadre de ces deux projets 

14 transferts de gestion. Pour chaque communauté, la 

surface totale des forêts dans ces transferts varie de 300 à 

5000 ha (dépendant du taux de couverture forestière dans le 

territoire de chaque village d’appartenance de la communauté). 

Les zonages des forêts sont décrits précédemment. 

En principe, les contrats sont établis pour trois ans. Ensuite 

une évaluation devra se faire par le Service Forestier en partenariat 

avec les communes d’appartenance des communautés 

gestionnaires des forêts.Etablis à partir de 2007,certains 

contrats nécessitent ainsi une évaluation à partir de cette 

année ou au plus tard en début de l’année prochaine. 

Au total,les 14 transferts de gestion des forêts ont permis de 

sécuriser sous la gestion des COBA,27 000 ha de forêts.Ces 

forêts incluent tous les types de forêts à différents usages définis 

auparavant (conservation, restauration, droit d’usage). 

Les valeurs de la restauration pour le WWF sont présentées 

dans son document de la vision de la biodiversité (Erdmann 

et al., 2005) qui a défini 40 aires comme Aires Prioritaires de 

conservation de par leur valeur en biodiversité. Les études 

sur leur état de dégradation ont relevé aussi que 23 de ces aires 

auront probablement besoin d’importante restauration 

(< 20 % de forêt),si elles doivent entièrement concourir à la 

conservation de la biodiversité. Dans tous les sites de transferts 

de gestion des forêts, le WWF met ainsi l’accent sur 

l’importance de la restauration. Les communautés avec l’encadrement 

du WWF, des observations sur terrain et des 

inventaires effectués définissent ainsi des zones de restauration 

dont la superficie varie d’une communauté à une 

autre.Les espèces utilisées pour ces restaurations des forêts 

dégradées sont essentiellement des essences autochtones. 

Leur choix est justifié par leur emplacement (héliophile pour 

les zones très ouverts et/ou périphérie de la forêt; nomade 

pour les zones sous couvert des espèces héliophiles) ou par 

leur utilisation (construction, nourriture des lémuriens). 

Suivis et contrôles 

Le WWF, pour faciliter l’uniformisation des suivis et contrôle 

en matière des lémuriens, a élaboré un livret sur les 

lémuriens avec la photo des espèces et une description 

sommaire des espèces.L’idée de départ était d’exploiter ces 

livrets peu avant la fin des projets pour les analyser. Il a 

pourtant été constaté que les représentants des communautés 

n’ont pas utilisé les livrets,mais les a bien classés dans 

leur valise. Leur explication est que le livret (en couleur) est 

trop beau pour être amené et abimé en forêt à cause de 

l’humidité. Néanmoins, les suivis et contrôles ont été effectivement 

effectués par les communautés. De plus, les données 

ont été stockées soit dans des cahiers à part, soit dans 

leur tête. 

Dans le cadre des suivis et contrôles instaurés de manière 

participative,des suivis et contrôles à part doivent aussi être 

faits par les organismes techniques d’appuis des communautés. 

En effet, à travers les expériences de l’élaboration du 

livret, il a été constaté que les instructions et formations 

n’ont pas été suivies correctement.Les agents des deux projets, 

ne se sont donc rendus compte qu’un peu tardivement 

de la nécessité de faire aussi des suivis et contrôles rapprochés.


Table 1: Caractéristiques des localités d’observation des lémuriens. 

Localités Andrakengy Andasipiro Ambodivoara Ambodimandresy 

Période 20 Novembre - 6 Décembre - 19 Novembre - 5 Décembre - 

d’observation 3 Décembre 2006 17 Décembre 2006 3 Décembre 2006 17 Décembre 2006 

Coordonnées S 14° 18’ 53.4’’ S 14° 12’ 10.0’’ S 14° 32’ 05.0’’ S 14° 32’ 05.0’’ 

géographiques E 049< 16’ 38.3’’ E 049< 22’ 34.8’’ E 049< 26’ 42.1’’ E 049< 30’ 21.1’’ 

Les suivis et contrôles effectués par les paysans ont quand 

même permis de localiser des sites d’observations de l’espèce 

Propithecus candidus (Simpona).Patel, en échangeant les 

données avec l’équipe des projets du WWF a publié en 2009 

que l’espèce Propithecus candidus est aussi rencontrée dans le 

Corridor de Betaolana et de Tsaratanana-Betaolana. Quelques 

caractéristiques des localités où les inventaires ont été 

effectués et où cette espèce a été observée sont synthétisées 

dans le tableau 1. 

Les alternatives aux pressions et menaces 

Les pressions sont définies comme étant les activités causant 

des impacts négatifs aussi bien sur les forêts que sur les 

lémuriens. Ces activités peuvent être légales ou illégales. Par 

contre, les menaces sont des activités pouvant apparaître 

dans le futur et pouvant avoir des impacts négatifs sur les 

cibles (dans le cadre de ce projet les cibles sont les lémuriens 

et leur habitat). 

Les analyses,effectuées dans le cadre de ces deux projets ont 

montré que les pressions et les menaces sur les cibles sont 

principalement constituées de: la déforestation causée essentiellement 

par les cultures sur brûlis, la dégradation des 

forêts engendrée par les prélvements diverses (bois de construction 

ou autre matériels pour la construction tels que les 

lianes et les bambous, cueillette de miel), la chasse moderne 

et le piège traditionnel. La déforestation tue à la fois les 

lémuriens et détruit leur habitat. Il en est de même des 

pièges traditionnels mais à un degré moindre. 

Pour arrêter les pressions sur les forêts, les deux projets du 

WWF ont donc analysé les activités pouvant remplacer 

celles formant une pression et constituant une menace dans 

le futur.En bref les activités développées sont:l’amélioration 

de l’exploitation de l’espace par l’agroforesterie; la promotion 

des cultures maraichères,l’amélioration des cultures de 

riz sur les bas fonds étroits par les Systèmes de Riziculture 

Améliorés (SRA),la promotion des briques pour la construction 

des maisons et le reboisement des espèces à croissance 

rapide; l’apiculture, la pisciculture et l’amélioration de l’élevage 

des volailles. 

Elaboration d’une stratégie régionale pour la conservation des 

lémuriens 

Les principales cibles de conservations des deux projets sont 

la forêt humide et les lémuriens. A part les transferts de 

gestion,les deux projets ont aussi été conduits pour éduquer, 

informer et sensibiliser les populations locales (partant des 

élèves dans les écoles primaires aux écoles secondaires mais 

aussi l’ensemble des populations concernées dans les communes). 

Pour la sauvegarde des lémuriens, les engagements des parties 

prenantes sont recherchés à travers l’élaboration et la 

mise en œuvre d’une stratégie régionale pour la conservation 

des lémuriens. Dans cette stratégie, les activités de 

chaque partie sont définies de manière à minimiser les 

dépenses monétaires afin de les rendre réalisables. 

Pour une meilleure intégration ou engagement de chaque 

entité, une Association des Amis des lémuriens a été créée. 

Cette association est mise en réseau via un site web aux 

amoureux des lémuriens dans le monde. La vision a été 

développée pour le long terme. Toutefois, 

le WWF s’est rendu comte 

qu’avec l’isolement de chaque localité, 

les réponses instantanées et directes 

des membres aux intéressés via le site 

web sont très limités et précaires. En 

effet,pour se connecter,les élèves doivent 

aller à Andapa; et même si cer- 

tains élèves d’Andapa sont concernés, le reflexe avec cette 

haute technologie nécessite encore un encadrement rapproché 

pour être efficace. 

Pour mieux unir les efforts,une Union Régionale des Associations 

de gestion des forêts, appelée aussi "Gestion Unie du 

Corridor de Betaolana" a aussi été créé. Malgré les efforts 

investis pour créer cette Union, elle est restée au stade de 

constitution (dépôt de dossier) au moment où les phases des 

deux projets sont terminées. Toutefois, elle est engagée et 

sera reprise dans les autres projets du WWF dans cette 

localité. Il est donc souligné ici l’importance des encadrements 

par des organismes promoteurs dans le long terme ou 

du moins à moyen terme pour obtenir de meilleurs impacts 

dans les phases du projet. 

Les parties prenantes définies dans ce document 

sont composées par les partenaires techniques et les autres 

organismes et/ou associations travaillant dans les domaines 

de l’environnement, les communautés de base, les organes 

de décentralisation et de déconcentration de l’Etat (les 

régions, les communes, les districts et les Fokontany). 

Conclusions 

La capitalisation des deux projets du WWF dans le Nord a 

permis de comprendre les efforts encore à fournir dans le 

cadre de la conservation des lémuriens et de leur habitat.Les 

deux projets conduits dans la région de SAVA sont complémentaires, 

ceci a permis une uniformisation des approches 

techniques et scientifiques. Les deux projets ont enrichis les 

données sur la diversité de la zone en flore et en faune.Ils ont 

pu se réaliser de manière participative. En effet, ils ont été 

très bien accueillis par les populations locales du fait que ces 

projets étaient majoritairement axés sur les activités de 

développement pour atteindre leurs objectifs de conservation. 

En conclusion, les défis à lancer doivent se concentrer sur 

deux aspects. 

Aspect technique: des études plus poussées sur la flore et la 

faune sont à conduire. En effet, les découvertes à faire sont 

encore immenses vu l’étendue du massif forestier. 

Aspect développement: des activités sont engagées et n’en 

sont qu’au début de leur mise en œuvre. Il serait nécessaire 

dans le futur de faire des évaluations de ces projets en 

termes d’impact sur la conservation des habitats et des 

lémuriens ainsi que sur l’amélioration des conditions de vie 

de la population locale. 

Ces deux projets ne sont pas des projets de recherches. 

Toutefois, ils ont pu être enrichis par les interventions des 

scientifiques consultants d’une part et des équipes du WWF 

d’autre part. 

Remerciements 

Nos vifs remerciements s’adressent aux bailleurs qui ont 

financés les projets du WWF Madagascar dans la région de 

SAVA. Ils ont permis de conduire à la fois des études scientifiques 

et des activités de développement.Ces bailleurs sont: 

WWF Allemagne, WWF Suède et la Conservation Internationale. 

Bien qu’à faible volume en terme de fonds, nous 

tenons aussi à remercier le WWF Danemark pour des fonds 

spécifiquement alloués à l’élaboration des support de com-


munication (poster,teeshirt,banderoles etc.) pour une meilleure 

compréhension des aspects de la conservation auprès 

des paysans en particulier et des parties prenantes en 

général. 

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of the Parc National de Marojejy and the Reserve 

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Lémuriens.Pp. 

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d’inventaires biologiques faisant référence à l’altitude 

dans la région des massifs montagneux de Marojejy 

et d’Anjanaharibe-Sud. Recherches pour le Développement, 

série sciences biologiques no. 19. Centre d’Information 

et de Documentation Scientifique et Technique, 

Antananarivo, Madagascar. 

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Mast,R.1994.Lemurs of Madagascar.Conservation International, 

Washington, D.C., USA. 

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A.A.; Butynski, T.M.; Ganzhorn, J.U.; Kormos, R.; Aiguiar, J. 

M.; Walker, S. 2005. The World’s 25 Most Endangered 

Primates 2004-2005. Lemur News 10: 3-6. 

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in prosimians. Pp. 247-305. In: G.A. Doyle and R.D. Martin 

(eds.).The Study of Prosimian Behavior.Academic Press. 

Rajaoson,A.;Rakotonirina,L.H.2007.Rapport pour le WWF; 

Inventaire des lémuriens et des forêts dans le Corridor de 

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University Press, New York, USA. 

Genetic diversity in ten Indri (Indri indri) 

populations compared to other lemur 

species 

John Zaonarivelo 1,Rick Brenneman 2*,Rambinintsoa 

Andriantompohavana 3, Edward E. Louis, Jr. 2,3 

1University of North Madagascar, Antsiranana, Madagascar 

2Center for Conservation and Research, Omaha’s Henry 

Doorly Zoo,3701 South 10th Street,Omaha,NE 68107,USA 

3Madagascar Biodiversity Partnership, Antananarivo, 101 

Madagascar. 

*Corresponding author: rabr@omahazoo.com 

Genetic diversity is considered by most to be the key to long 

term survival and the maintenance of the evolutionary trajectory 

of a species.Genetic variation at loci under selection 

gives the species as a whole the mechanisms with which to 

respond or adapt to environmental changes.Lemurs in general 

are poorly studied with respect to establishing baseline 

parameter estimates for genetic diversity.Only limited popu- 

lation genetics studies exist on the genera Propithecus, Avahi, 

Varecia, Eulemur, Microcebus, and Mirza (Tab. 1). To date, the 

genus Indri is depauperate of population genetic data that 

would help better understand the genetic diversity harbored 

in its populations. 

Tab.1:Lemur studies using multilocus microsatellite suites to 

estimate genetic diversity as observed (HO) and expected 

(HE) heterozygosity levels. 

Species PopuSam- Loci HO HE Reference 

lationsples Indri indri 2 20 20 0.654 0.766 Zaonarivelo 

et al., 2007b 

Propithecus 

deckeni 

2 20 14* 0.790 0.851 Lei et al., 2008a 

P. deckeni 1 10 18* 0.776 0.776 Lei et al., 2008b 

P. coquereli 1 25 20 0.635 0.771 Rakotoarisoa 

et al., 2006a 

P. candidus 2 18 17* 0.648 0.614 McGuire et al., 2009 

P. coronatus 1 10 18* 0.771 0.774 Lei et al., 2008b 

P. diadema 2 20 13* 0.818 0.814 Ramarokoto 

et al., 2008 

P. edwardsi 2 20 12* 0.681 0.618 Bailey et al., 2009 

P. verreauxi 3 30 13 0.670 0.712 Rakotoarisoa 

et al., 2006b 

P. tattersalli 2 20 16* 0.673 0.683 Razafindrakoto 

et al., 2008 

P. tattersalli **3 75 13 

Quéméré et al., 

0.699 0.682 

2009 

P. tattersalli 9 224 13 

Quéméré et al., 

0.690 0.660 

2010 

Avahi laniger 5 37 22 0.640 0.838 Andriantompohavana 

et al., 2004 

A. occidentalis 

1 7 22 0.514 0.586 Andriantompohavana 

et al., 2004 

Varecia 

rubra 

2 32 15 0.616 0.618 Razakamaharavo 

et al., 2010 

V. variegata 

variegata 

4 35 25 0.337 0.506 Louis et al., 2005 

Eulemur 

cinereiceps 

2 21 16* 0.598 0.641 

Tokiniaina et al., 

2009 

E. collaris 4 40 10* 0.617 0.576 Ranaivoarisoa 

et al., 2010 

E. sanfordi 5 54 11* 0.562 0.567 Ramanamahefa 

et al., 2010a 

E. coronatus 6 80 11* 0.636 0.673 Ramanamahefa 

et al., 2010b 

E. rubriventer 

2 12 20 0.531 0.643 Andriantompohavana 

et al., 2007 

Lemur catta 1 24 7* 

Zaonarivelo et al., 

0.837 0.838 

2006 

Microcebus 

ravelobensis 

8 205 7 

Olivieri et al., 

0.615 0.605 

2007 

M. ravelobensis 

12 187 8 

Radespiel et al., 

0.708 0.734 

2008 

M. bongolensis 

3 45 8 

Olivieri et al., 

0.557 0.565 

2008 

M. danfossi 7 78 8 0.628 0.662 Olivieri et al., 2008 

Mirza 

coquereli 

***1 69 7 

Markolf et al., 

0.712 0.799 

2008 

* Heterozygosity averages calculated using loci with null allele 

frequency estimates less than 0.10. 

** Estimated over genetic clusters, not actual populations 

*** Samples taken from 1993-2006. 

The Indri (Indri indri, Gmelin, 1788), or Babakoto as it is 

known in most of eastern Madagascar,is the largest extant lemur 

(Powzyk and Thalmann, 2003). Indri are primarily midlevel 

forest folivores preferentially feeding on immature 

leaves and somewhat also on mature leaf matter, flowers, 

fruits,seeds and even bark when necessary (Britt et al.,2002). 

The Babakoto is currently threatened by the rapid reduction


of forest cover and fragmentation of suitable habitat (Harper 

et al., 2007) which limits the species’ density and range 

(Glessner and Britt, 2005). Although protected, the Babakoto 

is also threatened by subsistence hunting pressure and 

bush meat trade (Golden, 2005). Designated as Endangered 

according to the IUCN Red List of Threatened Species 

(IUCN, 2008), I. indri is currently split into two subspecies, 

Indri indri indri (Gmelin, 1788) and Indri i. variegatus (Gray, 

1872). Here, we present population genetic parameter estimates 

from populations along the entire range of the species 

analyzed from nuclear microsatellite multilocus genotypes. 

Methods 

Samples were collected from 106 Indri from 10 sites across 

the geographical range of the species (Fig. 1). From north to 

south,the forests represented in the collection were Anjanaharibe-Sud 

Special Reserve, Marotandrano Special Reserve, 

Ambatovaky Special Reserve,Zahamena Special Reserve and 

National Park, Betampona Nature Reserve, Anjozorobe 

Regional Forest Reserve, Mantadia National Park, Analamazoatra 

Special Reserve (Andasibe),Maromizaha Classified 

Forest, and Anosibe an’ala Classified Forest. The elevations 

of the sampling sites ranged from lowland forests (Anosibe 

An’ala, 125 m asl) to highland forests (Anjozorobe, 1358 m 

asl). 

Fig. 1: Map of Madagascar indicating the study areas. 

Immobilization and collection 

All lemurs investigated in this study were free-ranging and 

were immobilized with a CO2 powered DAN-INJECT (Brrkop, 

Denmark) Model JM rifle propelling Pneu-Darts (Williamsport, 

PA) loaded with 10 mg/kg estimated body weight 

of Telazol ® (Fort Dodge).We recorded the location (within 6 

m accuracy) of all of the immobilized lemurs using a global 

positioning system (GPS) device. Each individual was transported 

back to the base camp where complete morphometric 

data were taken (Zaonarivelo et al., 2007a). Whole 

blood (1.0 cc per kilogram) from the femoral artery and 

2.0 mm skin biopsies from the ear pinnae were collected 

from each sedated lemur (Junge and Louis, 2002). A Home 

Again ® (Home Again Pet Recovery Service, East Syracuse, 

NY) microchip was placed subcutaneously between the scapulae 

of each lemur to positively identify individuals re-captured 

during any future immobilizations. Following data and 

sample collection, an injection of lactated Ringer’s solution 

was administered subcutaneously to support maintenance 

requirements and to dissipate the effect of the Telazol ®.Animals 

were monitored for three hours post recovery then 

released according to the capture GPS coordinates. 

Data generation 

Ear punches were dissected into quarters and DNA was 

extracted using standard PCI/Chloroform procedures (Sambrook 

et al., 1989). Approximately 50 ng of genomic DNA 

was used for each PCR reaction. Multilocus genotypes were 

generated from a suite of 20 Indri-specific microsatellite loci 

as described in Zaonarivelo et al. (2007b). The genotype file 

was checked for typographical errors,scoring errors,stutter 

bands and allele dropout with Micro-Checker (van Oosterhout 

et al., 2004) and Microsatellite Analyser (MSA; Dieringer 

and Schlötterer, 2002). We used CERVUS (version 2.0, 

Marshall et al.,1998;Slate et al.,2000) to identify loci with excessive 

null allele frequency estimates (nf > 0.10) and to estimate 

polymorphic information content of the loci.Moderate 

(0.05 < nf < 0.20) and high (0.20 < nf) null allele frequencies 

can have significant effects on population genetics parameter 

estimates (Chapuis and Estoup, 2007). The process of redesigning 

primer pairs is both costly and time consuming; 

therefore,we opted to delete problematic loci from the data 

set. We deleted eight loci with moderate null allele frequencies 

(nf > 0.1) to reduce the bias from misclassification of null 

heterozygotes as homozygotes (Callen et al., 1993; Hoffman 

and Amos, 2005) and to control the variance of parameter 

estimates (Chapuis and Estoup, 2007). The accepted loci 

were verified for independence of linkage disequilibrium 

(with Bonferroni-adjusted P-values) in FSTAT (Goudet,1995, 

2001). 

Hardy-Weinberg exact tests (Guo and Thompson, 1992) 

were performed by locus and population in Genepop (version 

4.0, Raymond and Rousset, 1995). Initially, we used the 

default settings for the MCMC estimation of HWE then 

increased the batch size from 100 to 250 to reduce the standard 

error of the P-value to below 0.01. Genetic diversity 

was measured as observed heterozygosity (HO) and expected 

heterozygosity (HE). In addition, the number of effective 

migrants was estimated globally and pair-wise using the private 

allele method. We used FSTAT to calculate the total 

number of alleles (k), mean number of alleles (MNA), and 

rarefacted allelic richness (AR; Leberg, 2002) by locus and 

population.Allelic richness estimates the allelic diversity in a 

data set based on the population with the fewest number of 

individuals contributing genotypes by locus. This is an unbiased 

comparison of allelic diversity since populations with 

more contributors provide a greater opportunity to capture 

more alleles from lower frequency occurrences. Wright’s Fstatistics 

were estimated in FSTAT for within population 

similarity (FIS) and between population differences (FST) 

according to Weir and Cockerham (1984). 

The effective population sizes were estimated with the linkage 

disequilibrium (LD) option in NeEstimator (Peel et al., 

2004; Hill, 1981; Waples, 1991). We tested all populations 

having met the minimum statistical threshold required (n = 

20 genes or 10 individuals) for the presence of bottleneck 

events using Bottleneck (version 2.0, Cornuet and Luikart,


1996; Luikart et al., 1998; Piry et al., 1999) under the Infinite 

Alleles Model (IAM; Kimura and Crow, 1964), the Stepwise 

Mutation Model (SMM;Ohta and Kimura,1973),and the Two 

Phase Model (TPM;di Rienzo et al.,1995).We varied the proportion 

of the single step contribution to the TPM to identify 

the P < 0.05 threshold of significance.The program identifies 

populations with an excess of heterozygosity relative to 

mutation-drift equilibrium which is indicative of a reduction 

in the effective population size (Maruyama and Fuerst,1985). 

An estimate of relationships among all individuals sampled at 

each forest was done in SPAGeDi (Hardy and Vekemans, 

2002),then compared to a simulation of known pedigreed individuals. 

The analysis was performed to calculate the relationship 

coefficients described in Queller and Goodnight 

(1989) in the absence of spatial data. 

Results 

Genetic diversity as mean number of alleles ranged from 

6.08-8.92 per population. Using the rarefacted allelic richness, 

the range lowered to 5.87-7.67. The expected heterozygosity 

ranged from 0.77-0.86 (P > 0.05;Fig.2) with an average 

of 0.81, while the observed heterozygosity ranged from 

0.65-0.84 (P < 0.05; Fig. 3) with an average of 0.74. The number 

of effective breeders in the sampled populations averaged 

between 12.6 and 39.6 per population (Tab. 2). 

Results from Bottleneck showed that none of the 10 populations 

deviated from a mutation-drift equilibrium under the 

SMM.Three populations,Anjanaharibe Sud,Ambatovaky and 

Anjozorobe did not show evidence of population bottleneck 

under the IAM either.The rest of the populations were significant 

for bottleneck events under the IAM and varying proportions 

of single step contributions under the TPM. 

The frequencies of the relationship coefficients estimated 

using SPAGeDi were overlaid upon a simulation generated 

from known pedigreed data so that each of the population’s 

relative distribution of relationships could be compared with 

parent offspring, full sibling, half sibling and unrelated relationship 

coefficient distributions (Fig. 4). The data indicated 

that the sampling was from individuals that were somewhat 

related more than the unrelated individuals in the reference 

simulation. Inbreeding can also be due to background relatedness 

where an increased allelic identity by descent is a result 

from bottleneck events in the population’s history.Relationship 

coefficient distributions sup- 

port the assumption that the individuals 

sampled were often from family 

groups. All of these sources may potentially 

be due to the effects of habitat 

fragmentation which is certainly 

the case in the Anosibe an’ala population 

where the habitat is so fragmented 

that although multiple family 

groups were encountered, they were 

found in isolated forest fragments. 

Discussion 

Of the 10 Indri populations sampled, 

six (Anjanaharibe Sud, Ambatovaky, 

Zahamena, Betampona, Mantadia, and 

Anosibe an’ala) deviated from HWE 

with an excess of homozygotes.Considering 

inbreeding as potential cause, 

five of the populations (Anjanaharibe 

Sud, Ambatovaky, Betampona, Mantadia, 

and Andasibe) had relatively 

high FIS and one (Anosibe an’ala) had 

Y 

.90 

.80 

.70 

.60 

55DD 

A B C D E F G H I J 

X 

Fig. 2: Ranges of expected heterozygosities with 95 % confidence 

intervals: A) Anjanaharibe Sud; B) Marotandrano; C) 

Ambatovaky; D) Zahamena; E) Betampona; F) Anjozorobe; 

G) Mantadia;H) Andasibe;I) Maromizaha; J) Anosibe an’ala. 

4DF4 

Y 

.90 

.85 

.80 

.75 

A B C D E F G H I J 

X 

Fig. 3: Ranges of observed heterozygosities with 95 % confidence 

intervals: A) Anjanaharibe Sud; B) Marotandrano; C) 

Ambatovaky; D) Zahamena; E) Betampona; F) Anjozorobe; 

G) Mantadia;H) Andasibe;I) Maromizaha; J) Anosibe an’ala. 

Tab. 2: Population genetic parameter estimates for 10 populations comprised of n 

samples each derived from 12 microsatellite loci for number of alleles (k), the mean 

number of alleles (MNA), allelic richness (AR), probability of satisfying Hardy-Weinberg 

Equilibrium (HWE), observed (HO) and expected (HE) heterozygosities, inbreeding 

estimate (FIS), the number of effective breeders (Neb) estimated with the 

linkage disequilibrium method and 95 % confidence interval,and results from the Bottleneck 

test under the infinite allele model (IAM) and the two phased model (TPM) 

with proportion of multistep mutations contributing to the P < 0.05 significance level. 

n k MNA AR HWE HO HE FIS Neb 95% CI IAM TPM 

ANJ 10 80 6.67 6.41 * 0.67 0.77 0.135 23.0 16.2-37.8 NS NS 

TAND 10 73 6.08 5.87 NS 0.75 0.77 0.024 21.9 15.1-36.9 ** 70 

VAK 11 76 6.33 5.92 NS 0.69 0.77 0.121 20.1 14.7-30.4 NS NS 

ZAH 14 107 8.92 7.65 NS 0.81 0.86 0.060 39.6 28.8-61.3 ** 10 

BET 10 79 6.58 6.37 ** 0.73 0.80 0.093 18.3 13.5-27.1 ** 20 

ANJZ 10 83 6.92 6.70 NS 0.84 0.79 -0.066 20.9 15.1-32.5 NS NS 

TAD 10 96 8.00 7.67 NS 0.75 0.85 0.120 20.1 15.3-28.3 ** 70 

DASI 11 81 6.75 6.40 NS 0.73 0.81 0.095 12.6 10.2-16.2 ** 5 

MIZA 10 89 7.42 7.17 NS 0.83 0.84 0.021 15.9 12.4-21.3 ** 5 

ANOSIBE 10 92 7.67 7.38 * 0.65 0.85 *0.236 28.8 19.9-49.3 ** NS 

* P < 0.05, ** P < 0.001; Anjanaharibe-Sud(ANJ), Marotandrano (TAND), Ambatovaky (VAK), 

Zahamena (ZAH), Betampona (BET), Anjozorobe (ANJZ), Mantadia (TAD), Andasibe (DASI). 

Maromizaha (MIZA), Anosibe an’ala (ANOSIBE).


Frequency 

Frequency 

Frequency 

Frequency 

Frequency 

Frequency 

Frequency 

0,4 

0,3 

0,2 

0,1 

0 

0,4 

0,3 

0,2 

0,1 

0 

0,4 

0,3 

0,2 

0,1 

0 

0,4 

0,3 

0,2 

0,1 

0 

0,4 

0,3 

0,2 

0,1 

0 

0,4 

0,3 

0,2 

0,1 

0 

0,4 

0,3 

0,2 

0,1 

0 

Anjanaharibe-Sud 

-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9 

Relationship Coefficients 

Morontandrano 

-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9 


Ambatovaky 

-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9 


Zahamena 

-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9 


Betampona 

-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9 


Anjozorobe 

-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9 


Mantadia 

-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9 


Frequency 

Frequency 

Frequency 

0,4 

0,3 

0,2 

0,1 

0 

0,4 

0,3 

0,2 

0,1 

0 

0,4 

0,3 

0,2 

0,1 

0 

Andasibe 

-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9 


Maromizaha 

-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9 


Anosibe an'ala 

-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9 


Parent-Offspring Full Sibling Half Sibling 

Unrelated All Populations 

Fig. 4: Distributions of relationship coefficients estimated in 

each population overlaid on a simulation of 10,000 known 

genotypes and pedigreed relationships (Queller and Goodnight, 

1989). 

a significantly high FIS estimate. Considering relationship 

among the samples as one manner of capturing inbreeding,all 

populations show some degree of relationship above what 

would be expected if the sampled individuals were unrelated. 

This is not surprising since samples were collected as found 

and this could include individuals that are members of a family 

group which is supported by the relationship coefficient 

distributions in Fig. 2. 

All populations demonstrated some degree of recent reduction 

in the effective population sizes.The bottlenecks did not 

appear to have been a single global event as different populations 

showed differing degrees to which the bottlenecks 

were detected. Using Lawler’s (2008) 4*Neb*generations, 

the bottlenecks detected would have had an expected window 

of occurrence of up to 250 to 800 years ago, within the 

timeframe of human encroachment. Hence, we postulate 

that among other things, anthropogenic disturbances, 

whether habitat destruction or subsistence hunting, may 

have influenced the demographic reduction that we detected 

in the bottleneck test. These baseline values could be useful 

in long-term or future studies to determine genetic health 

trends over time under various forest or habitat conditions. 

Genetic diversity is considered to be the most important 

factor in determining the genetic health of a species or population.Among 

the lemurs,there is little information in the literature 

that addresses the genetic diversity estimations for 

multiple populations of a given species. Ranaivoarisoa et al. 

(2010) found the observed heterozygosity in three of four 

Eulemur collaris populations to be higher than the expected 

heterozygosity. In E. coronatus, Ramanamahefa et al. (2010a)


found the expected heterozygosity to be higher in five out of 

six populations,one comparison was significantly higher (P < 

0.01).In E.sanfordi,Ramanamahefa et al.(2010b) found that in 

all populations sampled,the observed heterozygosities were 

higher than the expected heterozygosities. Lastly, Quéméré 

et al. (2009), found that their study estimated the expected 

heterozygosity level for several populations of P. tattersalli to 

be higher than the estimate found by Razafindrakoto et al. 

(2008) using a different marker suite in different populations. 

Other studies (included in Tab. 2) have estimated heterozygosity 

levels for one or two populations of various lemur 

species in recent years.While these estimates provide some 

general sense of genetic diversity, they are not standardized 

(e.g. not the same markers were used in each study) so the 

estimates are contingent on the polymorphic information 

content and amplification quality in each independent study. 

What we do see is a relative trend that the observed 

heterozygosities are in general lower, but usually not significantly, 

than the expected heterozygosities under the assumptions 

of HWE. 

In this study on I. indri, we found the differences among the 

HO estimates to differ but not significantly and the differences 

among the HE estimates to differ with low significance 

(P < 0.05). The average estimates for heterozygosity, thus 

genetic diversity, were in the upper range of those found in 

limited population studies on other lemur species. These 

estimate trends provide the basis for future and integrative 

studies where multiple species might be considered in sympatric 

zones to investigate the overall genetic health of the 

biodiversity and to better understand the effects that humans 

may be having on the evolutionary potential of lemurs. 


We acknowledge the Ministry of Environment and Eaux & 

Forets, Madagascar National Parks, U.S. Fish & Wildlife Service, 

Professor Gisele and the Department of Paleontology 

and Anthropology, University of Antananarivo for their help. 

This project would not have been possible without the support 

of the staff, guides, and drivers of Henry Doorly Zoo 

and the Madagascar Biodiversity Partnership. We also wish 

to thank the Theodore F.and Claire M.Hubbard Family Foundation, 

Bill and Berniece Grewcock, the Ahmanson Foundation, 

the James Family Foundation and the Hawks Family 

Foundation for their support of this project. 

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Zaonarivelo, J.R.; Andriantompohavana, R.; Shore G.D.; Engberg, 

S.E.; McGuire, S.M.; Louis, E.E. Jr.; Brenneman, R.A. 

2006.Characterization of 21 microsatellite marker loci in 

the ring-tailed lemur (Lemur catta). Conserv. Genet. 8(5): 

1209-1212. 

Verreaux’s sifaka fur condition in the 

spiny forest of southern Androy 

Ivan Norscia 1*, Jean Lambotsimihampy 2, Elisabetta 

Palagi 1,3 

1Natural History Museum, UniversitB di Pisa, Via Roma, 79, 

56011, Calci (PI), Italy 

2Berenty Village, Antandroy, Madagascar 

3Unit of Cognitive Primatology and Primate Center,Institute 

of Cognitive Sciences and Technologies, The National Research 

Council (CNR), Rome, Italy 

*Corresponding author: norscia@hotmail.com 

Résumé 

Les conditions du pelage des animaux peuvent représenter 

un moyen fiable et un indicateur non invasif pour comprendre 

l’état de santé d’une population et distinguer des segments 

différents de la même population.En 2008 nous avons 

effectué un recensement de sifaka (Propithecus verreauxi) 

dans les forêts riveraines de la réserve de Berenty (foret 

galerie et de transition de Malaza et forêt secondaire aux


espèces allochtones de Ankoba) et dans six portions de 

forêt épineuse qui sont inclues dans le domaine privé de 

Berenty. Nous avons relevé l’état du pelage des animaux 

selon trois conditions: fourrure intègre (niveau 1), fourrure 

faiblement endommagée (ponctuée par des petites zones, 

sans pelage, couvrant moins de 30 % du corps; niveau 2), 

fourrure fortement endommagée (pelage manquant sur une 

surface supérieure à 30 %;niveau 3).Nous avons aperçu seulement 

quatre sifaka au niveau de pelage 3 et, par conséquence, 

nous avons pu évaluer statistiquement les différences 

seulement entre les niveaux de pelage 1 et 2, en comparant 

forêt épineuse et les forêts riveraines, soi au niveau 

des groups d’animaux (n= 41) soi au niveau des zones recensée 

(n= 9). Même si avons détecté un nombre significativement 

plus haut de sifaka avec la fourrure faiblement 

endommagée dans le domaine épineuse,la nature et surtout 

l’entité du dommage indiquent que les conditions du pelage 

n’arrivent pas vraiment à différentier des segments distincts 

dans la population de sifaka de Berenty. 

Introduction 

An index of coat condition can be a non-invasive tool for 

tracking health and stress at the population level (Jolly, 

2009a). In fact, pelage growth can be directly influenced by 

the proximate stimulus of light (acting through neuro-endocrine 

pathways), by the nutritional status, and indirectly by 

temperature and behavior (Ling, 1970). Two main functions 

of fur are a) insulation, which allows conservation of body 

heat, thus reducing energy expenditure and food requirements; 

and b) shielding, which protects day-active mammals 

from excessive heat load from solar radiation (Scott et al., 

2001; Kenagy and Pearson, 2000). 

Here, we considered coat condition of Propithecus verreauxi 

(Verreaux’s sifaka) as a possible indicator of the "health status" 

of animals in different habitats and investigated whether 

it could provide information on possible population stress in 

the poorly investigated spiny forest of south Madagascar. 

The dry spiny forest of southern Madagascar is a thorny environment,both 

metaphorically and literally speaking.Listed as 

one of the 200 most important ecological regions in the 

world,it harbors the highest level of plant endemism in Madagascar 

(Elmqvist et al., 2007). In spite of its importance, the 

spiny forest is underrepresented in terms of protection and 

conservation programs (Fenn, 2003; Ganzhorn et al., 2003; 

Seddon et al.,2000).To fill,at least in part,this gap,we investigated 

sifaka fur condition in different spiny forest parcels 

inside the Berenty Estate (Androy region,south Madagascar) 

and compared it with sifaka inhabiting the riverine forest 

areas inside the Berenty Reserve, a habitat much richer in 

staple food for lemurs. 

Methods 

Study site, survey technique, and fur condition evaluation 

In March-April 2008, a comprehensive sifaka survey was conducted 

in the Berenty Estate, covering 134 ha of spiny forest 

and 60 ha of non-spiny forest areas. The Berenty Estate is located 

in the semi-arid Androy Region (rainfall averages less 

than 500 mm per year). The spiny forest is usually 3 to 6 m in 

height with dwarf and xerophyte plants,and emerging trees of 

the Family Didieraceae that may reach more than 10 m in 

height,such as the keystone species Allouadia procera (Elmqvist 

et al., 2007). We performed the survey in all accessible spiny 

forest parcels (sacred areas, used as a cemetery, cannot be 

accessed by anyone except for local family clans) and in three 

riverine areas of the Berenty Reserve (on the Mandrare river), 

comprising a northern section (occupied by the 40 ha secondary 

forest of Ankoba dominated by the exotic legume species 

Pithecellobium dulce;S 24.99°;E 46.29°) and a southern section 

(Malaza: S 25.01°; E 46,31°) (Fig. 1 shows study locations). Inside 

Malaza we considered the 7 ha gallery forest (dominated 

by tamarinds;Tamarindus indica) and the front-transitional forest 

(13 ha) (Jolly et al.,2006).In all the areas considered in this 

study,logging and hunting are prohibited,and the fossa (Cryptoprocta 

ferox) isabsent. 

Fig.1:Study site location:Berenty reserve (solid outline;white 

area:scrub;diagonal lines:Ankoba and Malaza riverine forests) 

and spiny forest fragments (black areas): 1 = Spiny Malaza; 2 = 

Spiny Reserve 1;3 = Spiny Reserve 2;4 = West Rapily;5 = Fragment 

X; 6 = Anjapolo, about 13 km north-west of Berenty. 

Dashed outlines include degraded spiny and/or scrub areas. 

The rest of the territory (white) is covered by pasture and 

sisal fields. (Map based on Google Earth satellite view). 

We performed the survey via walking, at a speed of about 1 

km/h,along preexisting trails and through forest paths chosen 

ad hoc to have visibility of at least 50 m to the right and left (to 

avoid pseudoreplication we followed Norscia and Palagi, 

2008). 

During the census we evaluated the fur condition of each individual 

lemur.We scored coat condition on a 3-point scale:coat 

undamaged, with fur fully covering the body (level 1); ruffled 

coat,with fur punctuated by small areas of reduced/missing fur 

(on less than 30 % of the body, especially on elbows and/or 

knees;level 2);patchy coat,usually with black skin areas clearly 

visible due to reduced/missing fur (on more than 30 % of the 

body, especially on elbows/knees, external sides of forearms 

and thighs, fingers and toes; level 3) (Fig. 3). 

Statistics 

We performed the analyses at group or at forest site level. 

Owing to the small sample size (n


in the spiny and two in the non-spiny forest areas). Thus, we 

considered only level 1 (undamaged coat) and 2 (moderately 

missing fur) for the analyses and found that the proportion of 

individuals with level-1 fur was significantly higher in the nonspiny 

than in the spiny areas both in the analysis per forest 

site (Exact Mann-Whitney U test, nnon-spiny=3, nspiny=6, Z= 

-2.35, p=0.024) (Fig. 2) and in the analysis per group (Mann- 

Whitney U test, nnon-spiny=31, nspiny=11, Z=-3.26, p=0.001). 

FUR FUR CONDITION 

CONDITION 

0.90 

0.80 

0.70 

0.60 

0.50 

0.40 

0.30 

0.20 

SPINY FOREST RIVERINE 

AREAS FOREST AREAS 

Fig. 2: Difference in fur condition (proportion of individuals 

showing level-1 fur) between the spiny and riverine forest areas. 

The difference is significant (p


Fenn, M.D. 2003. The spiny forest ecoregion. Pp. 1525-1530. 

In: S.M. Goodman; J.P. Benstead (eds.). The Natural History 

of Madagascar. University of Chicago Press, Chicago. 

Ganzhorn, J.U.; Goodman, S.M.; Dehgan, A. 2003. Effects of 

forest fragmentation on small mammals and lemurs. Pp. 

1228-1234. In: S.M. Goodman; J.P. Benstead (eds.). The 

Natural History of Madagascar. University of Chicago 

Press, Chicago. 

Jolly, A. 2009a. Coat conditions of ringtailed lemurs, Lemur 

catta, at Berenty Reserve, Madagascar: I. Differences by 

age, sex, density and tourism, 1996-2006. Am. J. Primatol. 

71: 191-198. 

Jolly, A. 2009b. Coat conditions of ringtailed lemurs, Lemur 

catta, at Berenty Reserve, Madagascar: II. Coat and tail 

alopecia associated with Leucaena leucocephala. Am. J. 

Primatol. 71: 199-205. 

Jolly, A; Koyama, N; Rasamimanana, H; Crowley, H; Williams, 

G.2006.Berenty Reserve:a research site in southern Madagascar.Pp.32-42.In:A.Jolly,R.W.Sussman,N.Koyama,H. 

Rasamimanana (eds.). Ringtailed Lemur Biology: Lemur 

catta in Madagascar. Springer Verlag, New York. 

Kenagy, G.J.; Pearson, O.P. 2000. Life with fur and without: 

experimental field energetics and survival of naked meadow 

voles. Oecologia 122: 220-224. 

Ling, J.K. 1970. Pelage and molting in wild mammals with 

special reference to aquatic forms.Q.Rev.Biol.45:16-54. 

Mundry, R; Fischer, J. 1998. Use of statistical programs for 

nonparametric tests of small samples often leads to 

incorrect P values: examples from Animal Behaviour. 

Anim. Behav. 56: 256-259. 

Norscia, I.; Palagi, E. 2010. Fragment quality and distribution 

of the arboreal primate Propithecus verreauxi in the spiny 

forest of south Madagascar. J. Trop. Ecol. DOI: 

10.1017/S0266467410000519. 

Norscia,I.;Palagi E.2008.Berenty 2006:census of Propithecus 

verreauxi and possible evidence of population stress. Int. J. 

Primatol. 29: 1099-1115. 

Scott, D.W.; Miller, W.H.; Griffin, C.E. 2001. Structure and 

function of skin.Muller and Kirk’s Small Animal Dermatology, 

6th ed. W.B. Saunders, Philadelphia. 

Seddon, N.; Tobias, J; Yount, J; Ramanampamonjy, J.M.; 

Butchart,S;Randrianizahana H.2000.Conservation issues 

and priorities in the Mikea forest of south-western Madagascar. 

Oryx 34: 287-304. 

Siegel,S.;Castellan,N.J.Jr.1988.Nonparametric Statistics for 

the Behavioral Sciences, Second edition. MacGraw-Hill, 

New York. 

Rediscovery of Sibree’s dwarf lemur in 

the fragmented forests of Tsinjoarivo, 

central-eastern Madagascar 

Marina B. Blanco 

Department of Anthropology, University of Massachusetts, 

240 Hicks Way, Amherst, MA 01003, USA, 

mbblanco@anthro.umass.edu 

The recent genetic confirmation of a rare dwarf lemur 

species, C. sibreei, at Tsinjoarivo is bitter-sweet. The excitement 

of reporting the first known living population of this 

species is tainted by conservation concerns, as the forest 

fragment in which Sibree’s dwarf lemurs were captured is 

highly disturbed and targeted for illicit logging. This species, 

like many others inhabiting rapidly degrading forests, faces 

the serious threat of extinction. 

Taxonomic background of the genus, first field discovery, 

and subsequent recognition of C. sibreei 

During the 19th century,the small nocturnal lemurs of Madagascar 

were clumped in a chaotic array of species and genera. 

For most of the 20th century,however,dwarf lemurs (Cheirogaleus) 

were classified in only two species: the eastern C. 

major and the western C.medius (Schwarz,1931).Around the 

turn of the century, Groves (2000) conducted a taxonomic 

revision of the genus on the basis of morphological analysis 

of museum specimens and increased the species number to 

seven: C.medius,C.adipicaudatus,C.major,C.ravus,C.crossleyi, 

C. minusculus and C. sibreei. This last species, in fact, had been 

originally described by the Swiss naturalist Forsyth Major in 

1896 during one of his expeditions to Madagascar (Forsyth 

Major, 1896). He had named it Chirogale sibreei in honor of 

fellow naturalist James Sibree,who had spent more than fifty 

years in Madagascar and had written extensively about its 

people, fauna, flora and geology. Forsyth Major published 

measurements of an individual "obtained from the neighbourhood 

of Ankeramadinika," a locality vaguely described 

by its discoverer as "one day’s journey to the east of Antananarivo",but 

in fact a well-known village at the time,located in 

the central high plateau on the road that connected Antananarivo 

to Mahatsara on the east coast (Capitaine "X", 

1901). In his taxonomic revision, Groves (2000) included as 

Cheirogaleus sibreei not only the holotype from Ankeramadinika 

(currently housed at the Natural History Museum in 

London) but also three additional specimens (3 skins and 1 

skull), two of which came from Ampasindava, northwestern 

Madagascar, and one from an unclear provenance (Imerina, 

which refers to a region of the central highlands around 

Antananarivo). 

The taxonomic shrinkage of Cheirogaleus 

The increase in the number of species within the genus 

Cheirogaleus was not surprising because dwarf lemurs occupy 

a wide variety of habitats in Madagascar,and their close relatives,the 

mouse lemurs (Microcebus), had undergone a taxonomic 

explosion of their own with more than 10 species 

described during the past 15 years (Louis et al.,2008;Olivieri 

et al., 2007; Radespiel et al., 2008). However, Groves’ 2000 

revision of dwarf lemur taxonomy did not escape criticism, 

not least of which had to do with the criteria that he used to 

define species, the lack of reliable locality information from 

museum specimens, and the absence of on-the-ground surveys 

to assess geographic boundaries and variation among 

species (Blanco et al., 2009; Tattersall, 2007). A recent and 

more comprehensive revision of dwarf lemur taxonomy was 

carried out by Groeneveld and colleagues, who compiled 

genetic and morphometric data from field as well as museum 

specimens from a variety of localities across Madagascar, 

including some of the specimens studied by Groves (Groeneveld 

et al., 2009; 2011). This research showed overall consistency 

between morphological and genetic data in recognizing 

only three Cheirogaleus species: C. medius, C. major 

and C. crossleyi. Individuals that previously had been assigned 

to C. adipicaudatus fell within the C. medius clade, and those 

named as C.ravus grouped with C.major.Results were inconclusive 

for C.minusculus and C.sibreei because holotype specimens 

were not available for sampling and their genetic affiliation 

could not be determined. Genetic data from one of the 

C. sibreei museum specimens from Ampasindava linked this 

specimen to C. medius. Nevertheless, the C. sibreei holotype 

from Ankeramadinika was larger and did not group morphologically 

with other C. medius. This suggested that the individuals 

from Ampasindava may have been misclassified by 

Groves as C. sibreei (Groeneveld et al., 2010). The status of 

this species remained equivocal. 

Second field discovery of C. sibreei, at last 

The story of a dwarf lemur named "May" told by Mitchell 

Irwin (2002) turned out to be rather prophetic. Irwin’s 

research team rescued this dwarf lemur badly burned in a


human-induced fire nearby Andasivodihazo,one of the forest 

fragments at Tsinjoarivo (Fig. 1). At the time, "May" was believed 

to be, as were all eastern rainforest dwarf lemurs, C. 

major. Unfortunately, this female could not fully recover and 

died soon after the salvage, but her skeletal remains were 

carefully preserved and stored at the University of Antananarivo 

by Irwin’s team. Seven years later this specimen came 

to play a key role in our morphological analysis of dwarf 

lemurs at Tsinjoarivo. 

Fig.1: Map showing Tsinjoarivo and other localities associated 

with Sibree’s dwarf lemurs; see text for details. 

In 2006, with the logistic help of Mitchell Irwin and Jean-Luc 

Raharison, I began a survey of nocturnal lemurs at Tsinjoarivo. 

My assistants and I successfully trapped dwarf lemurs 

at two study sites: in one of the forest fragments (Andasivodihazo, 

19º41’15"S, 47º46’25"E, 1660 m) and within continuous 

forest (Vatateza, 19º43’15"S, 47º51’25"E, 1396 m) 

(Blanco et al., 2009). Even to an inexperienced eye, fragment 

dwarf lemurs looked different from continuous forest individuals, 

in that they were overall smaller, with grayer fur, 

marked eye rings and significantly larger female genitalia (Fig. 

2). Our morphological and dental analyses determined that 

of all the species described by Groves,C.sibreei was the most 

similar to forest fragment dwarf lemurs (Blanco et al., 2009). 

(Hopefully, sampling of C.sibreei’s holotype will be allowed in 

the near future to definitely determine whether or not there 

is a genetic match between this specimen and fragment 

dwarf lemurs from Tsinjoarivo.) Recent genetic analyses 

have confirmed not only that dwarf lemurs from Andasivodihazo 

constitute a different clade (and therefore an independent 

phylogenetic lineage), but also that the fragment 

dwarf lemur species had branched off first and was ancestral 

to the other dwarf lemur species (Groeneveld et al., 2010). 

To date, no other living population of C. sibreei has been 

reported in the wild and more intensive surveys around the 

Tsinjoarivo area (including possibly remaining forests nearby 

Ankeramadinika, ~100 km from Tsinjoarivo) are warranted 

to assess geographic boundaries and population density. 

Fig.2: Sibree’s dwarf lemur captured at Andasivodihazo, one 

of the forest fragments at Tsinjoarivo. 

Conservation concerns 

The genetic confirmation of three clades corresponding to C. 

medius, C. major and C. crossleyi, each of which has broad 

geographic distributions, implied that dwarf lemurs might be 

less threatened than previously thought (Groeneveld et al., 

2009). However, the situation for C. sibreei is radically different. 

Tsinjoarivo’s unique geographic setting, continuous 

with the central plateau on the west and the steep escarpment 

of rainforest in the east, may harbor a unique array of 

animal communities. To date, C. sibreei has been captured in 

sympatry with C. crossleyi at one forest fragment, Andasivodihazo,and 

at one intermediate location,Ankadivory.Both of 

these areas are subjected to illicit logging and heavy deforestation 

(Fig.3).Furthermore,these forest sites are located towards 

Tsinjoarivo’s western boundary which reaches some 

of the highest altitudes (up to ~1650 m) known in eastern 

Fig. 3: Example of logging near Ankadivory, one of the forest 

sites where Sibree’s dwarf lemurs were captured.


rainforests in Madagascar.So far,only C.crossleyi has been captured 

at Vatateza,a lower altitude site located within continuous 

and less disturbed forest.Virtually everything has to be 

learned about the "ancestral" C.sibreei,including how to prevent 

its extinction through habitat loss.Concerted efforts by 

organizations such as Sadabe (www.sadabe.org) are instrumental 

in raising awareness of endangered species and promoting 

research and educational programs in the Tsinjoarivo 

area. 


I am particularly thankful to the research team in Madagascar: 

Malagasy students Vololonirina Rahalinarivo and Mamihasimbola 

Rakotondratsima, and local assistants Noel Rakotoniaina, 

Edmond and Nirina Razanadrakoto. Additional 

thanks to Jean-Luc Raharison and Mitchell Irwin for providing 

logistical support and assistance in the field. I am also 

grateful to Laurie Godfrey, Stacy Gebo and Christoph 

Schwitzer for their comments. This research was supported 

by funds from the Rufford Foundation, MMBF/Conservation 

International Primate Action Fund and Primate Conservation 

Inc. I would like to thank the Ministère de l’Environnement 

et des forêts of the Malagasy government and the University 

of Antananarivo for permission to conduct this research.The 

project in Madagascar was facilitated by the Institute 

for the Conservation of Tropical Environments (ICTE, 

Patricia C. Wright) and the Madagascar Institute pour la 

Conservation des Ecosystèmes Tropicaux (MICET), especially 

Benjamin Andriamihaja. 

References 

"X." 1901. Voyage du Général Gallieni: cinq mois autour de 

Madagascar, progrès de l’agriculture, développement 

commercial, ressources industrielles, moyens de colonisation. 

Hachette, Paris. 

Blanco, M. B.; Godfrey, L.R.; Rakotondratsima, M.; Rahalinarivo, 

V.; Samonds, K. E.; Raharison, J-L.; Irwin, M. T. 2009. 

Discovery of sympatric dwarf lemur species in the high 

altitude forest of Tsinjoarivo, eastern Madagascar: implications 

for biogeography and conservation.Folia Primatol. 

80: 1-17. 

Forsyth Major, C. I. 1896. Diagnosis of new mammals from 

Madagascar. Ann. Mag. Nat. Hist. 6: 318-325. 

Groeneveld, L. F.; Weisrock, D. W.; Rasoloarison, R. M.; Yoder, 

A. D.; Kappeler, P. M. 2009. Species delimitation in lemurs: 

multiple genetic loci reveal low levels of species diversity 

in the genus Cheirogaleus. BMC Evol. Biol. 9: 30. 

Groeneveld, L. F.; Blanco, M. B.; Raharison. J-L.; Rahalinarivo, V.; 

Kappeler, P. M.; Godfrey, L. R.; Irwin, M. T. 2010. MtDNA and 

nDNA corroborate existence of sympatric dwarf lemur species 

at Tsinjoarivo,eastern Madagascar.Mol.Phylogenet.Evol. 

55: 833-845. 

Groeneveld, L. F.; Rasoloarison, R. M.; Kappeler, P. M. 2011 

Morphometrics confirm taxonomic deflation in dwarf 

lemurs (Primates: Cheirogaleidae) as suggested by genetics. 

Zool. J. Linn. Soc. 161: 229-244. 

Groves,C.P.2000.The genus Cheirogaleus:unrecognized biodiversity 

in dwarf lemurs. Int. J. Primatol. 21: 943-962. 

Irwin,M.T.2002.The story of May,a courageous dwarf lemur. 

IPPL News April: 12-14. 

Louis, E. E.; Engberg, S. E.; McGuire, S. M.; McCormick, M. J.; 

Randriamampionona, R.; Ranaivoarisoa, J. F.; Bailey, C. A.; 

Mittermeier, R. A.; Lei, R. 2008. Revision of the mouse 

lemurs,Microcebus (Primates,Lemuriformes),of northern 

and northwestern Madagascar with descriptions of two 

new species at Montagne d’Ambre National Park and 

Antafondro Classified Forest. Primate Conservation 23: 

19- 38. 

Mullens, J. 1875. Twelve Months in Madagascar. James Nisbet 

& Co., London. 

Olivieri,G.;Zimmermann,E.;Randrianambinina,B.;Rasoloharijaona, 

S.; Rakotondravony, D.; Guschanski, K.; Radespiel, 

U. 2007. The ever-increasing diversity in mouse lemurs: 

Three new species in north and northwestern Madagascar. 

Mol. Phylogenet. Evol. 43: 309-327. 

Radespiel, U.; Olivieri, G.; Rasolofoson, D. W.; Rakotondratsimba, 

G.; Rakotonirainy, O.; Rasoloharijaona, S.; Randrianambinina, 

B.; Ratsimbazafy, J. H.; Ratelolahy, F.; Randriamboavonjy, 

T.; Rasolofoharivelo, T.; Craul, M.; Rakotozafy, L.; 

Randrianarison,R.M.2008.Exceptional diversity of mouse 

lemurs (Microcebus spp.) in the Makira region with the 

description of one new species. Am. J. Primatol. 70: 1033- 

1046. 

Schwarz,E.1931.A revision of the genera and species of Madagascar 

Lemuridae.Proc.Zool.Soc.Lond.1931:399-426. 

Tattersall, I. 2007. Madagascar’s lemurs: Cryptic diversity or 

taxonomic inflation? Evol. Anthropol. 16: 12-23. 

Funding and Training 

AEECL Small Grants 

Since 2009,AEECL awards two small grants of up to € 1,000 

each year to graduate students, qualified conservationists 

and/or researchers to study lemurs in their natural habitat. 

Priority is given to proposals covering conservation-relevant 

research on those species red-listed as Vulnerable, Endangered, 

Critically Endangered or Data Deficient by the IUCN. 

We support original research 

that helps with establishing 

conservation action plans for 

the studied species. Grants 

are normally given to recent 

graduates from Malagasy universities to help building local 

capacity. 

We may also, in special circumstances, support studies on 

Malagasy species other than lemurs if the proposal provides 

satisfactory information as to how lemurs or the respective 

habitat/ecosystem as a whole will benefit from the research. 

All proposals will be assessed by the Board of Directors of 

AEECL and/or by external referees.The deadline for applications 

is February 15th of each year. Successful applicants will 

be notified by June 1st.More information can be found on the 

AEECL website, www.aeecl.org. 

The Mohamed bin Zayed Species Conservation 

Fund 

Announced at the World Conservation Congress in Barcelona 

in 2008,The Mohamed bin Zayed Species Conservation 

Fund is a significant philanthropic endowment established to 

do the following: 

Provide targeted grants to individual species conservation 

initiatives; 

Recognize leaders in the field of species conservation;and Elevate 

the importance of species in the broader conservation 

debate.


The fund’s reach is truly global,and its species interest is nondiscriminatory. 

It is open to applications for funding support 

from conservationists based in all parts of the world,and will 

potentially support projects focused on any and all kinds of 

plant and animal species, subject to the approval of an independent 

evaluation committee. 

Details on this important new source for species conservation 

initiatives and research can be found at 

www.mbzspeciesconservation.org 

CI Primate Action Fund 

The principal objective of Conservation International’s Primate 

Action Fund is to contribute to global biodiversity 

conservation by providing strategically targeted, catalytic 

support for the conservation of endangered nonhuman 

primates and their natural habitats. 

Projects submitted to the foundation should have one or 

more of the following characteristics: 

A focus on critically endangered and endangered nonhuman 

primates (and most especially those included in the 

biennial listing of the World’s 25 Most Endangered Primates) 

living in their natural habitats; 

Location in areas of high overall biodiversity and under great 

threat (e.g., "threatened hotspots", "megadiversity" countries) 

- to ensure maximum multiplier effect for each project; 

Direction and management by nationals from the tropical 

countries,to help increase local capacity for implementing 

biodiversity conservation; 

The ability to strengthen international networks of fieldbased 

primate specialists and enhance their capacity to be 

successful conservationists; and 

Projects that result in publication of information on endangered 

primate species in a format that is useful both to 

experts and the general public. 

Applications for support are considered throughout the year 

with no deadlines for submittal. Proposals should be sent by 

electronic mail to: 

Anthony B. Rylands, Primate Action Fund, Conservation International, 

2011 Crystal Drive, Suite 500, Arlington, VA 

22202, USA, a.rylands@conservation.org 

Recent Publications 

Lemurs of Madagascar, 3rd edition, by Russell A. Mittermeier, 

Edward E. Louis Jr., Matthew Richardson, Christoph 

Schwitzer,Olivier Langrand,Anthony B.Rylands,Frank Hawkins, 

Serge Rajaobelina, Jonah Ratsimbazafy, Rodin Rasoloarison, 

Christian Roos, Peter M. Kappeler and James Mackinnon.Illustrated 

by Stephen D.Nash.Conservation International, 

Tropical Field Guide Series, Arlington, VA, 2010. 762 

pp. ISBN: 978-1-934151-23-5. US$55.00. 

In 2006,Madagascar was making significant progress towards 

conservation by expanding the protection of its natural treasures.At 

the same time,the second edition of Conservation 

International’s Tropical Field Guide Series, Lemurs of Madagascar,had 

just come off the press,a full twelve years after its 

much-celebrated predecessor. A lot has changed in four 

years. Political and economic instability has imperiled both 

the Malagasy people and their unique wildlife. Conservation 

has taken drastic steps backwards as the desperation of the 

masses and greed of a few elites and international profiteers 

has exacerbated the conflict between the domains of humans 

and wildlife. CI has answered the call to action by releasing 

a new third edition of its lemur field guide, dwarfing 

previous editions in both size and its depth of research and 

detail. With nearly 1,100 references to support it–up from 

approximately 500 references in the second edition–the 

third edition stands as more than just a complete compendium 

of our knowledge about lemurs, but the perfect guide 

for appreciating the history, diversity, uniqueness, and pure 

beauty of our strepsirrhine cousins. 

According to CI’s Jill Lucena,from early 2009,the 13 authors 

and dozens of contributors have worked tirelessly on the 

third edition,dedicating thousands of hours towards its production. 

Authors Matthew 

Richardson and Anthony B. 

Rylands, as well as illustrator 

Stephen D. Nash and 

graphic designer Paula K. 

Rylands,labored exclusively 

on the project for nine 

months. The end result is a 

field guide that will leave 

other academic fields envious! 

This new volume is 247 

pages longer than the previous 

edition, with 767 pages 

of carefully organized maps, 

photos, and colorful illustrations, 

in addition to all 

the details lemur enthusiasts 

and researchers have 

come to expect from this 

book.The content is so rich 

that the book’s dimensions 

have increased from 7.5" x 

4.5" x 1" to slightly more than 9.25" x 6.25" x 1.25" just to accommodate 

everything.And herein lies what may be the only 

problem with this new edition. Reviews of the previous two 

editions had lavished praise for not only the content and 

scope, but also the portability of the books. Although it will 

fit comfortably in a backpack, it will add more weight and 

consume more space than its predecessors. But given the 

content, that may be a small price to pay. 

The layout of the book has not changed much since the last 

edition. An enthralling chapter on Madagascar’s ancient geological 

history has been added, providing tantalizing details 

about the mysteries of Madagascar’s ancient past,while a few 

familiar chapters and appendices have been reordered. Each 

lemur family has now been assigned its own chapter.The section 

entitled "How to Use This Field Guide" still walks new 

readers through the layout of the book. The "Quick Visual 

Reference" and colored tabs facilitate speedy navigation and 

help to satiate an ecotourists’ spontaneous hunger for specific 

information. The "Lemur Life-list" returns in a more 

readable table format to help ecotourists record their first 

sightings of the numerous lemur species. Even the maps of 

the island have been revamped and are easier to read in this 

larger format. 

Introductory chapters discuss ancient geology,lemur origins, 

the extinct subfossil (giant) lemurs, the history of lemur re-


search, and lemur conservation. Each chapter contains significant 

updates from the second edition. The ancient geology 

of Madagascar is covered in meticulous detail, while the theories 

of lemur origins are explored in depth, leaving both 

reader and researcher alike desperate for more definitive 

answers.The chapter on the extinct subfossil lemurs is beautifully 

illustrated with new peer-reviewed artwork from 

award-winning illustrator Stephen D. Nash, and has new details 

about their biology and extinction.The history of lemur 

research and discovery expands greatly upon the work from 

the last half century–a topic greatly underrepresented in the 

previous edition. Additionally, the chapter is loaded with 

newly added artwork from the 1700s and 1800s. The lemur 

conservation chapter provides a critical update on the newest 

emerging threats faced by lemurs and their habitat, 

namely the logging of precious hardwoods and bushmeat 

hunting. Additional detail is also provided about other important 

threats that received little mention previously, such 

as invasive species, cattle-raising, and mining. 

The bulk of the book details the description, geographic 

range,natural history,conservation status,and best locations 

to observe each of the 101 species and subspecies of lemur. 

(This total is up from the 71 taxa detailed in the second edition,yet 

the authors note that upcoming research may reveal 

as many as 110 to 125 lemur taxa!) Each species section 

sports a portrait photo, detailed range map, and other photos 

to enrich the lavish textual content.Once again,the third 

edition sets itself apart from the previous editions with its 

encyclopedic coverage of details from the lemur research literature.Species,such 

as the silky sifaka (Propithecus candidus), 

are discussed in significantly greater detail,bringing everyone 

from bright-eyed ecotourists to veteran lemur researcher 

up to speed on the latest findings. 

The second edition of Lemurs of Madagascar sold out quickly, 

leaving shelves empty as early as February 2008. The third 

edition is poised to do the same. With a print run of 10,000 

copies,nearly two-thirds are already spoken for according to 

Jill Lucena.To help promote conservation education in Madagascar, 

CI is generously donating 3,000 copies to its partner 

in the field,NGO Fanamby.CI hopes that the remaining copies 

will spark the public’s interest in Madagascar’s ecological 

gems and spur a new wave of ecotourism to bolster conservation 

efforts. 

Once again, CI has provided an invaluable tool for a diverse 

audience, which includes ecotourists, Malagasy tour guides, 

students,lemur researchers,and lemur enthusiasts.Although 

larger and not quite as portable as its predecessors, the increased 

size of the third edition hosts a wealth of enhanced 

encyclopedic detail,new and stunning artwork by Stephen D. 

Nash, and additional color photos and illustrations. With a 

copy of this printing in hand, the only things missing are a 

backpack full of supplies and an airline ticket to Madagascar. 

So what are you waiting for? 

Alex Dunkel 

Theses Completed 

Blanco,M.B.2010.Reproductive biology of mouse and dwarf 

lemurs of eastern Madagascar,with an emphasis on brown 

mouse lemurs (Microcebus rufus) at Ranomafana National 

Park,a southeastern rainforest.PhD Dissertation.University 

of Massachusetts, Amherst. 

This dissertation investigates reproductive schedules of 

brown mouse lemurs at Ranomafana,using intensive trap- 

ping techniques. The reproductive condition of female 

mouse lemurs was recorded on the basis of vaginal morphology,vaginal 

smears,body mass gain profiles and nipple 

development. Testis size was measured in males throughout 

the reproductive season. The timing of the first seasonal 

estrus was determined in frequently captured females 

over multiple years and it showed individual periodicities 

close to 365 days,consistent with endogenous regulation 

and entrainment by photoperiod. The timing of 

estrus did not correlate with female age or body mass. 

Males showed testicular regression during the rainy season, 

although there was high inter-individual variation in 

testes size at any given point during the reproductive season. 

Furthermore, some individuals completed testicular 

regression earlier than others. Implications for polyestry 

are discussed. 

For comparative purposes,mouse lemurs were also trapped 

at two study sites in the Tsinjoarivo area: one in a forest 

fragment and the other within continuous forest. 

These forests are higher in altitude than the main study 

area at Ranomafana. Trapping success for mouse lemurs 

was lower at Tsinjoarivo than Ranomafana.Albeit preliminary, 

data from Tsinjoarivo suggest that females have lower 

reproductive success than do females at Ranomafana. 

Nevertheless, mouse lemurs in the Tsinjoarivo forest 

fragment did not appear to be in "poorer" condition than 

those in the continuous forest.It had been reported in the 

literature that western gray mouse lemurs captured in secondary 

forests have lower body masses and lower recapture 

rates than those captured in primary forest; in fact, 

the opposite was true of the mouse lemurs at Tsinjoarivo. 

I additionally collected data on a larger member of the 

family Cheirogaleidae, the dwarf lemurs (Cheirogaleus), 

which live in sympatry with Microcebus at Ranomafana and 

Tsinjoarivo. I analyzed the patterns of growth, development 

and reproduction in Cheirogaleus and Microcebus and 

compared dwarf and mouse lemurs to other similarlysized 

prosimians which do not undergo torpor or hibernation.These 

comparisons draw attention to the unusual 

reproductive and metabolic strategies employed by cheirogaleids 

to cope with Madagascar’s unpredictable environments, 

which ultimately define their very unique life 

histories. 

Key words: Cheirogaleus, Madagascar, Microcebus, Mouse 

lemurs, Rainforest, Ranomafana, Reproduction, Tsinjoarivo. 

Bonaventure, R.T.A.R. 2010. Ecologie et comportement de 

Propithecus verrreauxi dans les zones d’extension de la 

Réserve Spéciale de Bezà Mahafaly.Engineer in agronomy, 

option Eaux et forêts.Eaux et forêts,Ecole Supérieure des 

Sciences Agronomiques de l’Université d’Antananarivo 

(ESSA), Madagascar. 

The population of Propithecus verreauxi in the special 

reserve of Bezà Mahafaly is one of the conservation targets 

of the site which is the subject of a long-term follow. 

Natural destruction of their habitat and the pressure of 

hunting which is exerted on this species outside of the 

current reserve are the origin of the sifakas’ decline.With 

the park extension project under way, furthering the 

knowledge on behavior and ecology of Propithecus verreauxi 

in disturbed areas outside the current reserve is essential 

for decision-making regarding conservation measures 

for this species. Thus,a study of behavior and ecology 

of the sifakas was carried out in the extension area of the 

special reserve of Bezà Mahafaly in gallery and transition 

forest at the end of the dry period. The study was centered 

on 3 focal groups including one in the gallery forest 

and two in the transition forest. The method of focal animal 

sampling was chosen to study their behavior.A floristic 

inventory according to the Gentry method, which 

includes transects of 2 x 50 m, was carried out to study 

the habitat. On the whole, 120h of observations of sifaka 

behavior were carried out and 12 transects were walked. 

In disturbed areas, the sifakas still consumed preferred 

plants.This resulted in a high intake of 2 or 3 easily digestible 

plant species while at the same time a large variety of 

other species was consumed.Thus,the disturbance of the 

sites did not influence food intake of the sifakas.Moreover,


it was the group of sifakas in the gallery forest which was 

the most affected by resource scarcity in the dry period, 

necessitating a priorisation of the sifaka groups in this forest 

formation in the extension of the reserve.In addition, 

the availability of nutritional resources influenced spacial 

dispersion of group members as well as group size. With 

the current park extension project different access restrictions 

will be implemented, having as an objective the 

conservation of the sifaka groups as well as other conservation 

targets outside of the current reserve,while introducing 

a range of alternative income-generating activities 

and sustainable resource management practices for the 

local communities. 

Key words: Lemur, Propithecus verreauxi,Ecology, Behavior, 

Conservation, Special reserve, BezB Mahafaly, Madagascar. 

Delmore,K.2009.Maintenance of stability in the Andringitra 

brown lemur hybrid zone.M.A.thesis,University of Calgary, 

Calgary, Alberta, Canada. 

Two models of hybrid zone stability have been proposed: 

1) the tension zone model, which predicts that intrinsic 

selection acts against hybrids but is counteracted by dispersal 

of parentals into the zone and 2) the bounded superiority 

model,which predicts that exogenous selection 

favours hybrids within transitional habitats and parentals 

outside the zone.I used morphological,genetic and ecological 

data to evaluate these models in a stable hybrid zone 

between Eulemur rufifrons and E.cinereiceps in southeastern 

Madagascar. This zone appears to conform to the 

bounded superiority model: it was relatively wide and 

composed mostly of hybrids that were equally as fit as 

parentals.Gene flow between parental and hybrid populations 

was also limited, clines in multiple characters were 

non-coincident and significant ecological correlates were 

identified. Results suggest that hybridization can serve as 

an important evolutionary force and need not always be 

considered a conservation risk for endangered taxa. 

Hobinjatovo, T. 2009. Etude morphométrique et génétique 

de conservation d’Eulemur cinereiceps (Milne-Edwards et 

Grandidier, 1890) dans les forêts de Mahabo, de Manombo 

et de Vevembe, Madagascar. Mémoire de DEA en Biologie,Ecologie 

et Conservation Animale,Département de 

Biologie Animale,Faculté des Sciences,Université d’Antananarivo. 

Cette étude a pour but, d’une part, de connaître et de 

comparer la morphométrie d'’Eulemur cinereiceps de la forêt 

de Mahabo,de Manombo et de Vevembe - Madagascar, 

ainsi donc de voir le degré du dimorphisme sexuel et 

d’autre part, d’établir des données de bases génétiques 

pour avoir de plus amples informations en vue de la conservation 

de cette espèce en danger critique.La descente 

sur le terrain et la collecte des données ont été effectuées 

pendant deux semaines du mois de mai 2006 et le travail 

au laboratoire pour l’étude génétique a été fait du mois de 

Février au mois de Mai 2007.Ce lémurien pése en moyenne 

2,04 kg avec une longueur moyenne de la couronne de 

la tête de 10,84 cm,celle du corps de 33,31 cm et est doté 

d’une queue de 50,44 cm. La comparaison de la morphométrie 

des individus des sites d’étude, par le test U de 

Mann-Whitney montre que le Lémurien à Collier Blanc 

de Vevembe est de plus grande taille que ceux des 2 autres 

sites.La différence de taille entre le mâle et la femelle 

n’est pas significative. Chez cette espèce, le dimorphisme 

sexuel est très marqué concernant la couleur du pelage et 

la dimension de la canine supérieure,considérable chez le 

mâle.Le testicule droit du mâle est plus long,large et volumineux 

que le gauche.14 marqueurs génétiques polymorphiques 

ont été sélectionnés pour effectuer le génotypage 

par l’utilisation de la technologie moléculaire et à l’aide 

de la réaction en chaîne polymérasique. Les programmes 

de Cervus, GenePop, Fstat, Structure et Bottleneck ont 

été utilisés pour déterminer la structure et la nature génétique 

de la population d’Eulemur cinereiceps. Ces analyses 

ont permis de déterminer que la valeur de la diversité 

des gènes des populations est pareille et modérée, oscillant 

autour de 57 % ;la divergence génétique est comprise 

entre 0,05 à 0,15 et est qualifiée de modérée; la richesse 

allélique varie de 2,928 à 3,632;une migration entre la population 

de Mahabo et de Vevembe,d’une part et celle de 

Manombo et de Vevembe d’autre part,a été identifiée;un 

certain degré de consanguinité a été constaté à Mahabo. 

La population de Manombo subit un goulot démographique 

et aucune structure distincte ni sous - structure n’a 

été observée au sein des populations de ces 3 sites.Même 

si cette espèce est en danger critique, sa santé génétique 

est modérée.Elle pourrait être bonne si les solutions adéquates 

sur la conservation génétique étaient appliquées. 

Dans le cas contraire,elle deviendrait désastreuse.La prise 

immédiate de mesures de conservation efficientes est 

donc nécessaire afin de préserver les populations pures 

d’E. cinereiceps et de protéger ses habitats. 

Mots-clés: Eulemur cinericeps, Mahabo, Manombo, Vevembe, 

Madagascar, Morphométrie, Mensuration, Génétique, 

Population, Conservation. 

Ingraldi, C. 2010. Forest fragmentation and edge effects on 

eight sympatric lemur species in southeast Madagascar. 

M.A. thesis, University of Calgary, Calgary, Alberta, Canada. 

Extensive slash-and-burn agriculture in southeastern Madagascar 

has led to the fragmentation of forests in this region, 

creating a constricted available habitat area and increasing 

the proportion of forest edge. I investigated the 

response to forest fragmentation and edge effects in eight 

lemur species through comparisons of species density 

and diversity between fragments, as well as correlation 

analyses including population distribution patterns,ecological 

variables,and distance from forest edge.I also include 

a more detailed focus on the behavioural response of Eulemur 

cinereiceps. Results were highly varied, with no species 

showing strong aversion to forest edge but with higher 

overall densities in larger,more connected fragments. 

Eulemur cinereiceps spent significantly more time near the 

forest edge while resting, but edge did not affect feeding 

patterns or food availability. These results suggest that 

conservation management should focus on maintaining 

large, complex fragments and improving connectivity 

through forest corridors. 

Mihaminekena,T.H.2010. Etude de la relation entre la dégradation 

de l’habitat et les activités de Propithecus edwardsi 

du Parc National de Ranomafana Ifanadiana, Madagascar. 

Mémoire de DEA en Paléontologie et évolution biologique, 

Biologie Evolutive, Primatologie, Département de 

Paléontologie et d’Anthropologie biologique, Faculté des 


Une étude sur le comportement et l’habitat de Propithecus 

edwardsi a été réalisée dans le Parc National de Ranomafana.Elle 

a été réalisée dans trois sites ayant chacun un 

degré de perturbation inégal: Talatakely (fortement perturbé); 

Sakaroa (moyennement perturbé) et Valohoaka 

(non perturbé). Sa finalité est d’analyser le type de comportement 

biologique adopté par l’espèce en réponse à la 

dégradation de son habitat et de le comparer entre les 

trois sites. La méthode adoptée est celle décrite par Altmann 

en 1974 qui consiste à déterminer l’activité du focal 

animal toutes les dix minutes et celle de tous les groupes 

toutes les cinq minutes. Pour toutes les activités, sauf le 

déplacement, la différence est toujours significative pour 

les trois sites. Le repos est plus élevé dans le site intact 

(36.6%) par rapport à l’alimentation.Inversement le repos 

est moins fréquent (28.9 %) que l’alimentation (53.6 %) 

dans le site fortement perturbé. La fréquence des activités 

de l’espèce dans le site moyennement perturbé est 

toujours comprise entre les deux sites perturbés et non 

perturbés. Pour ses activités, l’espèce utilise certains niveaux 

de strates, spécialement ceux compris entre 10 et 

15 et 15 et 20 m. Néanmoins, l’espèce habitant le site intact 

se place à un niveau plus haut que celle de la forêt perturbée. 

Les grands arbres sont plus abondants à Valohoaka 

qu’à Sakaroa et à Talatakely:respectivement la hauteur 

varie de 10.80, 9.53 et 9.47 m; celui du DHP est de 13.59 

cm;12.17 et 11.09 cm.Les épaisseurs de la couronne sont 

respectivement 4.42,4.28 et 3.91 m.Les parties de plantes


consommées pour chaque site sont significativement très 

différentes. La corrélation entre la consommation de jeunes 

feuilles et les activités exercées est toujours positive et significative 

quel que soit le site. Les femelles choisissent un niveau 

plus haut des arbres que les mâles à Valohoaka et à Talatakely 

par contre à Sakaroa c’est l’inverse. La cohésion du 

groupe est plus observée dans le site intact par rapport à celui 

dégradé.Bref,la perturbation influe les activités générales 

et la structure de l’habitat de Propithecus edwardsi. 

Mots-clés:Propithecus edwardsi, Lémuriens, Degré de perturbation, 

Habitat, Activités, Parc National de Ranomafana, 

Madagascar. 

Polowinsky, S.Y. 2009. Nutrition of captive Sclater’s lemurs 

(Eulemur macaco flavifrons GRAY, 1867) and crowned 

lemurs (Eulemur coronatus GRAY,1842),with special emphasis 

on the problem of obesity. PhD dissertation, Biology 

and Geography, University of Duisburg-Essen, Germany. 

This study was concerned with the obesity problem of blueeyed 

black lemurs and crowned lemurs in captivity. Its aims 

were to optimize the species’ diet in captivity by combining 

data obtained from individuals kept at different European 

zoos as well as from wild blue-eyed black lemurs to gain a 

better understanding of the ecological and nutritional needs 

of Eulemur macaco flavifrons in order to prevent individuals 

from becoming obese and to assist planned conservation 

measures. 

The captive part of the study was conducted in two European 

zoos:Cologne Zoo (Germany) and Parc Zoologique 

et Botanique de Mulhouse, Sud-Alsace (France). A longterm 

study with one group of blue-eyed black lemurs 

(1.3) and one group of crowned lemurs (1.2) was carried 

out at Cologne Zoo.In addition,three groups of blue-eyed 

black lemurs (2.1; 1.1; 1.1) and three groups of crowed 

lemurs (2.2; 2.1; 3.2) were studied at Mulhouse Zoo. The 

body weight development of captive individuals was registered 

and compared to body weight data of wild individuals. 

The obesity rate in captivity was recorded. An obese 

animal was identified as one weighing more than two standard 

deviations over the mean wild weight. Moreover, 

nutrient and energy intake of Eulemur macaco flavifrons 

and Eulemur coronatus at Cologne Zoo and Mulhouse Zoo 

were registered. In addition, digestibility trials were conducted. 

Samples of feeds and faeces were analyzed using 

Weende analysis and detergent analysis. In Madagascar, 

four groups of Eulemur macaco flavifrons in two forest fragments,one 

mainly primary forest and the other predominantly 

secondary forest,were observed.Samples of plants 

utilized by free-ranging blue-eyed black lemurs were collected.They 

were botanically classified and analyzed using 

Weende analysis and detergent analysis. 

The mean body weights of Eulemur macaco flavifrons as well 

as Eulemur coronatus in captivity were significantly higher 

than the mean body weight of free-ranging individuals.100 

% of the Eulemur macaco flavifrons sample and 33.3 % of 

the Eulemur coronatus sample were obese.Significant body 

weight differences were found between the groups studied 

at Cologne Zoo and Mulhouse Zoo, which could be 

explained by different feeding regimes. Comparing the 

diet of free-ranging blue-eyed black lemurs to the zoo 

diets that were based mainly on fruits and vegetables at 

Cologne Zoo and Mulhouse Zoo, considerable differences 

were found with respect to NDF,ADF,ADL and crude 

protein content, whereas ash and crude lipid content varied 

only slightly.The NFC and energy content in the zoo 

diets were almost twice as high as those in the diet of wild 

blue-eyed black lemurs. The high NFC,crude protein and 

metabolizable energy content and low fibre content of 

the zoo diets as compared to the wild diet,combined with 

a relatively high apparent digestibility of ~80 % for Eulemur 

macaco flavifrons and ~84 % for Eulemur coronatus, respectively, 

and in combination with lemurs’ typically low basal 

metabolic rates, all clearly contribute to the obesity problem 

of captive Eulemur macaco flavifrons. 

The presented data of food consumed by Eulemur macaco 

flavifrons in captivity and in the wild reveals elementary 

differences concerning nutrient and energy composition. 

Although a bright variety of fruits and vegetables could 

protect animals in captivity from stereotypic behaviour, a 

systematic reassessment of the zoo diet is suggested: 

increasing fibre content and decreasing energy density by 

feeding vegetables,and whenever possible,fresh plant material 

in appropriate quantities instead of energy-rich 

fruits, gruel or commercial feeds. Although the utilization 

of the food fibre content by a generalist frugivore like Eulemur 

macaco flavifrons or Eulemur coronatus is limited, fibre 

content plays an important role in the maintenance of 

physiological health. A zoo diet corresponding to the natural 

requirements of lemurs guarantees an optimization 

of breeding programmes and presents a valuable and necessary 

contribution to the preservation of these highly 

endangered species. 

Key words: Eulemur macaco flavifrons, Eulemur coronatus, 

Nutrition, Digestibility, Obesity, Captivity, Energy intake. 

Rafaliarison R.R. 2010. Activité générale du Prolemur simus: 

transition saison sèche - saison de pluies et activité de la 

femelle avant et après mise bas dans le Parc National 

Ranomafana.Département de Paléontologie et d’Anthropologie 

Biologique, Université d’Antananarivo, Madagascar. 

Cette étude a été réalisée dans la parcelle 3 du parc national 

Ranomafana qui abrite le seul groupe du parc.Elle nous 

aidera à mieux comprendre les variations de l’activité générale 

du Prolemur simus pendant la transition de la saison 

sèche à la saison de pluie ainsi que la variation de l’activité 

de la femelle avant et après mise bas. Les résultats ont 

montré que le Prolemur simus a dépensé la moitié de leur 

temps à l’alimentation suivi du repos. Les variations de la 

fréquence de l’activité sont en relation avec la partie consommée 

(tige, moelle ou jeunes pousses), la disponibilité 

alimentaire,la température et la pluie ainsi que la disponibilité 

en eau.La strate la plus utilisée est comprise entre 0 

à 5 m. La présence d’un nouveau né a une influence sur 

l’activité et la proximité des individus du groupe. Pour la 

femelle, il y a une diminution de la fréquence de l’alimentation 

après la mise bas.Il y a aussi une augmentation très 

marquée de la fréquence du repos après la mise bas.Le juvénile 

s’éloigne de la femelle après mise bas tandis que le 

mâle reste toujours près de la femelle. 

Mots clés: Prolemur simus, Activités, Mise bas, Parc National 

Ranomafana, Madagascar. 

This study was carried out in Parcel 3 of Ranomafana National 

Park,where the only group of Prolemur simus within 

the park is present.It concerns the variation in the general 

activities of P.simus during the transition from the dry season 

to the rainy season as well as the activity of the female 

before and after giving birth.The results showed that P. simus 

spent half of their time feeding, followed by resting. 

The variation in frequency of activities was related to the 

consumed plant parts (trunk,culm pith or bamboo shoot), 

availability of food,temperature,rain and the availability of 

water. The most frequently used forest stratum was between 

0 and 5 m of height. The presence of the new-born 

had an influence on the activity and the spacing of the individuals 

in the group.For the female,there was a reduction 

of the frequency of feeding after giving birth. There was 

also a very marked increase in the frequency of resting after 

birth. The juvenile stayed away from the female after 

she had given birth,but the male always remained close to 

the female. 

Key words:Prolemur simus,Activity,New born,Ranomafana 

National Park, Madagascar. 

Raharivololona,B.M.2010.Intestinal parasite infection of the 

gray mouse lemur (Microcebus murinus, J.F. Miller, 1777) in 

the south-eastern littoral forest of Madagascar. PhD Dissertation, 

Hamburg University, Hamburg, Germany. 

Madagascar’s plants and animals belong to one of the most 

unique and threatened biotas of the world. Lemurs are 

the flagship species associated with the biological crisis of 

the island and notably vulnerable to habitat degradation. 

While most studies on the effect of habitat destruction 

on species survival have focused on population reduction 

and forest degradation, indirect effects, such as altered 

parasite loads have received little attention. Parasitologi-


cal studies have concentrated on large primates, such as 

apes and monkeys.This is probably due to epidemiological 

interest in apes, which are genetically closer to humans 

and are known to be a reservoir of certain pests and 

diseases fatal to humans. Prosimians’ gastrointestinal parasites 

are less studied. 

The goal of this project was to assess and describe the 

gastrointestinal parasites of the lemur species Microcebus 

murinus (Family Cheirogaleidae), also known as the gray 

mouse lemur, from the littoral forest fragments of Mandena 

in extreme southeastern Madagascar. In addition, I 

wanted to evaluate the utility of determining gastrointestinal 

parasite loads based on fecal samples. From April 

2003 to October 2005, 427 fecal samples obtained 

from169 different individuals of M. murinus from five forest 

fragments were analyzed to assess the parasite species 

richness of this animal based on parasite egg morphology.Three 

individuals of M.murinus were also sacrified in 

order to look for adult worms for identification and confirmation 

of parasite species, and to localize their gastrointestinal 

parasites in the digestive tract. Screening all 

fecal samples by using the modified technique of the 

McMaster flotation, I noted that M. murinus harbored at 

least nine different intestinal parasites, which included 1) 

six Nemathelminthes:a member of the family Ascarididae, 

one species of the family Subuluridae represented by the 

genus Subulura,an unidentified Strongylida,a species of the 

genus Trichuris (Trichuridae), two species of the family 

Oxyuridae: the first belongs to the genus Lemuricola and 

the second is still unidentified;2) two Plathelminthes:two 

cestodes of the genus Hymenolepis (Hymenolepididae);3) 

one Protozoa: belonging to the order Coccidia. 

These gastrointesinal parasites of M.murinus from Mandena 

have not been previously described from this primate. 

The cestode infection deserves special attention,as these 

parasites have not been previously reprted from lemurs. 

Adult worms of Trichuris species were found in the caecum, 

as well as Lemuricola worms in the caecum and large 

intestine.Subulura worms were more abundant in the caecum 

than in the small and large intestine. A large number 

of Subulura larvae were observed in the caecum.As exemplified 

by the data on Subulura sp. worms in the digestive 

tract of M.murinus,the number of nematode parasite eggs 

and larvae found in the feces are correlated with the intensity 

of infection in the digestive tract. 

To assess effects of forest fragmentation and degradation,fecal 

samples from the first captureof 169 individuals of Microcebus 

murinus living in five littoral forest fragments were 

analyzed for gastrointestinal parasites. The fragments differed 

in size and forest quality. In good quality forest 

blocks, lemurs from a smaller fragment had higher prevalences 

and intensities of infection of gastrointestinal nematodes 

and protozoans than animals from a larger forest 

fragment. In larger forest blocks, excretion of eggs from 

Ascarididae and tapeworms was higher in a degraded forest 

fragment than in a better quality forest fragment.This 

situation was reversed in small forest fragments with fewer 

eggs of Subulura nematodes and protozoans shed by 

lemurs in the degraded fragment than by lemurs from the 

good quality fragment. The analyses are hampered by the 

fact that only one forest fragment was available per type 

of treatment. Keeping this limitation in mind, the results 

are consistent with other studies and indicate that forest 

degradation and fragmentation have marked effects on 

the level of parasitism of Madagascar’s lemurs. 

To assess seasonal effects on the excretion of gastrointetinal 

parasites I screened fecal samples from M. murinus 

caught during monthly trapping sessions for eggs and 

larvae of intestinal parasites. Parasite excretions changed 

seasonally when analyzed at the level of individual hosts. 

The number of parasite species and the abundance of parasite 

eggs and larvae in Microcebus feces were higher during 

the hot season than the cold season.Reduced parasite 

excretion during the cold season could be due to environmental 

factors or due to the ability of M. murinus to 

enter torpor and hibernation during the cold season 

which might lead to reduced metabolism of intestinal parasites 

and results in reduced shedding of parasite eggs. 

No such seasonal variation was found on the level of the 

lemur population when the analyses were based on samples 

of unknown origin. 

The study revealed noticeable effects of forest fragmentation 

on parasite loads as measured via the excretion of 

parasites. The disadvantageous consequences of increased 

parasite infections on the health of these animals is due 

to changes in habitat conditions and is a factor that needs 

to receive more attention when developing conservation 

plans. 

Key words: Microcebus murinus, Parasites, Habitat degradation, 

Fragmentation, Forest quality, Hymenolepis. 

Randrianarimanana, H.L. 2009. Etude comparative de l’alimentation 

et du comportement des deux espèces sympatriques 

d’Indriidés :Propitecus diadema et Indri indri dans 

le Réserve Naturelle Intégrale n°1 de Betampona (Tamatave).Mémoire 

de DEA en Paléontologie et évolution biologique,Biologie 

Evolutive,Primatologie,Département de 

Paléontologie et d’Anthropologie biologique, Faculté des 


Des études comportementales et nutritionnelles des 

deux espèces sympatriques d’Indriidés (Propithecus diadema 

et Indri indri) ont été réalisées pendant les mois de 

mars,avril,juin et juillet 2008 dans la Réserve Naturelle Intégrale 

numéro un de Betampona (Tamatave). Les données 

comportementales et nutritionnelles,la hauteur fréquentée 

et la nature des supports et des coordonnées 

géographiques ont été enregistrés toutes les 10 minutes. 

Des analyses statistiques ont été réalisées pour étudier 

comment ces deux plus grands lémuriens partagent leurs 

nourritures et habitats. Même si les deux espèces ont la 

même fréquence d’alimentation et sont toutes deux considérées 

folivores, Propithecus diadema consomme un peu 

plus de fruit qu’Indri indri (respectivement 33,6 et 9,4 %) et 

utilise beaucoup plus d’espèces végétales comme source 

de nourriture. Propithecus diadema fréquente des hauteurs 

beaucoup plus basses qu’Indri indri durant ses activités 

(8,3071et 10,208 m). De plus, cette dernière espèce 

utilise beaucoup de petits supports (respectivement 3,80 

et 5,38 cm) et peu inclinés (40,32° et 47,79°). Propithecus 

diadema se déplace beaucoup tandis qu’Indri indri se repose 

davantage.Malgré le chevauchement de leur territoire, 

ces deux espèces montrent une séparation de leur niche 

écologique. 

Mots-clés: Propithecus diadema, Indri indri, Alimentation, 

Comportement, Comparaison. 

Razafindratsima, O.H. 2009. Rôle écologique de Varecia rubra 

et d’Eulemur albifrons dans le Corridor Ambatolaidama du 

Parc National Masoala.Mémoire de DEA en Biologie,Ecologie 

et Conservation Animale, Département de Biologie 

Animale, Faculté des Sciences, Université d’Antananarivo. 

Une étude a été effectuée sur deux espèces de lémuriens 

sympatriques dans les forêts tropicales humides du corridor 

Ambatolaidama du Parc National Masoala - Varecia 

rubra (Geoffroy, 1812) et Eulemur albifrons (Geoffroy, 

1796). Le but est d’étudier les rôles écologiques de ces 

espèces dans la reforestation du corridor en tant que disséminatrices 

de graines. Ceci afin de mettre en évidence 

leur importance au niveau de ce site et par conséquent, 

d’élaborer une stratégie de conservation. Un groupe d’E. 

albifrons et trois groupes de V.rubra ont été suivis.Trois femelles 

de V.rubra,choisies comme animaux focaux,ont été 

munies de colliers à radio émetteur. Ces espèces ont fait 

l’objet de suivis quotidiens, de novembre 2006 à janvier 

2007, afin de collecter des informations sur leur régime 

alimentaire et leur défécation. Pour E. albifrons, aucune 

donnée sur son alimentation n’a été obtenue de part la 

difficulté de son suivi dû à l’absence de collier. Les fèces 

collectées sont analysées afin d’en extraire des graines qui 

ont été, ensuite, inventoriées, mesurées et identifiées. La 

viabilité de ces graines a été étudiée par un test d’immersion 

dans l’eau,puis par la mise en terre dans une pépinière 

des graines déféquées comparées avec celles extraites 

manuellement des fruits. Afin de comprendre le devenir 

de ces graines après leur dépôt, une étude de l’habitat où


les fèces ont été déposées a été réalisée. Les résultats de 

cette étude ont démontré que V.rubra a une frugivore élevée 

(86.1 %). Aussi, ces deux espèces participent activement 

à la dissémination des graines de la majorité des 

espèces végétales du corridor, avec 16 graines par jour 

disséminées par V. rubra représentées par 34 espèces végétales 

appartenant à 15 familles, et 10 graines par jour 

pour E. albifrons réparties dans 8 familles avec 11 espèces. 

De plus, après leur passage au niveau du tube digestif de 

ces animaux,les graines sont viables (x2 = 107,283,ddl = 2, 

p = 0,0001) et ont une germination plus élevée que les 

graines témoin (x2 = 55,680, ddl = 1, p = 0,0001). Très peu 

d’entre elles ont subi des dommages (seulement de 3,8 % 

et 0,7 % respectivement pour les graines déféquées par V. 

rubra et E. albifons). L’étude de l’habitat démontre une 

réussite de germination malgré une forte pente.Ces deux 

espèces de lémuriens sont donc d’importantes disséminatrices 

de graines de la forêt humide du corridor Ambatolaidama.Elles 

jouent un rôle important dans le maintien 

de l’équilibre écologique et contribuent à la reforestation 

du corridor. 

Mots-clés: Varecia rubra, Eulemur albifrons, Primates, Régime 

alimentaire, Dispersion des graines, Parc National 

Masoala, Corridor, Ambatolaidama, Madagascar. 

Razakanirina H. 2010. Suivi phénologique global et statut de 

conservation de 4 espèces végétales (Strychnos decussata, 

Diospyros ferrea,Gardenia decaryi et Capurodendron gracilifolium) 

consommées par Propithecus verreauxi coronatus dans 

la forêt de Badrala (Antrema - Région Boeny). Mémoire 

de DEA en Biologie et Ecologie Végétale,option Ecologie 

végétale, Département de Biologie et Ecologie Végétale, 

Faculté des Sciences, Université d’Antananarivo. 

La Station Forestière à Usage Multiple d’Antrema constitue 

un des habitats de Propithecus verreauxi coronatus.Dans 

cette région, ces Lémuriens sont vénérés comme étant 

les représentants des ancêtres des "Sakalava". Cette culture 

leur offre donc un haut niveau de protection mais 

est- ce que la forêt peut leur fournir la nourriture dont ils 

ont besoin? 

Des études sur la phénologie et une évaluation du statut 

de conservation des quelques espèces consommées par 

ce lémurien ont été réalisées dans la forêt sèche sur dune 

de Badrala (partie Nord de la station),afin de faire ressortir 

les différents types phénologiques et la saisonnalité des 

différentes phénophases des arbres de cette forêt. Trois 

(3) plots de 2000 m2 ont été montés dans la forêt et dans 

chaque plot,tous les arbres à DHP > $10 cm sont numérotés 

et des paniers collecteurs de litières sont installés 

suivant un transect de 200 m. La phénologie des arbres a 

été suivie pendant une année grâce à des observations directes 

de chaque individu et à l’analyse des litières qui 

sont collectées tous les 15 jours. La période de floraison 

maximale des espèces se produit au début de la saison humide, 

avant l’apparition des feuilles et la défeuillaison est 

assez importante au milieu de la saison sèche. Strychnos 

decussata, Diospyros ferrea, Gardenia decaryi et Capurodendron 

gracilifolium sont classées en danger d’extinction 

(EN). Ainsi, des mesures de conservation sont à entreprendre 

afin de protéger ces espèces et les habitats de ce 

lémurien. 

Mots-clés: Suivi phénologique, Statut de conservation, 

Plantes consommées,Propithecus verreauxi coronatus,forêt 

sèche, Badrala, Station Forestière à Usage Multiple d’Antrema, 

Madagascar. 

Rued, A.C. 2009. Social structure and female foraging strategies 

in white-collared lemurs (Eulemur cinereiceps). M.A. 

thesis, University of Calgary, Calgary, Alberta, Canada. 

This thesis examines the nature of male-female affiliation 

in Eulemur cinereiceps, specifically whether it consists of 

special relationships or a central male social structure. A 

special relationship includes an unrelated male and female 

adult who preferentially associate and affiliate with each 

other over all other individuals within the group. I also 

examine the flexibility of female foraging strategies in 

response to changes in resource availability and energy 

requirements. I tested the resource defence hypothesis, 

which proposes that reproductive females form special 

relationships with males to improve foraging success and 

offset the energetic costs of reproduction.Data were collected 

on two small groups in Mahabo forest,on the southeastern 

coast of Madagascar.Analysis of social structure 

data suggested central male structure when resources 

were scarce and central female structure during the period 

of relative resource abundance.The resource defence 

hypothesis was not supported by foraging data. 

Solomon,S.K.2010.Living on the edge:a preliminary dry season 

study of crowned lemur (Eulemur coronatus, Gray 

1842) and Sanford’s lemur (E.sanfordi,Archbold 1932),responses 

to anthropogenic habitat changes in northern 

Madagascar. M.A., Anthropology (Environment & Sustainability),University 

of Western Ontario,London,Canada. 

Habitat fragmentation through anthropogenic disturbance 

is a significant threat to primates in all biogeographic 

areas. Recent research has shown that primates have 

non-patterned responses to this disturbance and that general 

models of changing primate behaviour are not effective 

conservation tools. Previous research on primates in 

fragments is concentrated in the Neotropics demonstrating 

a need to investigate species-specific responses in other 

areas of the world. This study examined two sympatric 

lemur species, the crowned lemur (Eulemur coronatus) 

and Sanford’s brown lemur (Eulemur sanfordi) and their responses 

to anthropogenic habitat fragmentation in northern 

Madagascar. Although habitat generalists, Sanford’s 

lemur was extirpated at the study site while crowned lemur 

density was low but viable; they were restricted to 

forest fragments on the periphery and top of limestone 

massifs. Conservation initiatives in these fragments are 

reliant on preserving fruit trees located in the remaining 

forest flatlands and the commitment of a community conservation 

group. 

Key words: Northern Madagascar, Crowned lemur, Sanford’s 

brown lemur,Forest fragmentation,Edge effects,Lemur 

density,Community conservation,Dry season,Deciduous 

forest.

Guidelines for authors 

Lemur News publishes manuscripts that deal largely or exclusively with lemurs and their habitats. The aims of Lemur News are: 1) to 

provide a forum for exchange of information about all aspects of lemur biology and conservation, and 2) to alert interested parties 

to particular threats to lemurs as they arise. Lemur News is distributed free of charge to all interested individuals and institutions. To 

the extent that donations are sufficient to meet production and distribution costs, the policy of free distribution will continue. Manuscripts 

should be sent to one of the editors electronically (see addresses for contributions on the inside front cover). Lemur News 

welcomes the results of original research, field surveys, advances in field and laboratory techniques, book reviews, and informal status 

reports from research, conservation, and management programs with lemurs in Madagascar and from around the world. In addition, 

notes on public awareness programs, the availability of new educational materials (include the name and address of distributor and 

cost, if applicable), and notification of newly published scientific papers, technical reports and academic theses are all appropriate 

contributions. Readers are also encouraged to alert Lemur News to pertinent campaigns and other activities which may need the 

support of the lemur research and conservation community. Finally, Lemur News serves as a conduit for debate and discussion and 

welcomes contributions on any aspect of the legal or scientific status of lemurs, or on conservation philosophy. 

Manuscripts should be in English or French, double spaced with generous margins, and should be submitted electronically in Word 

(*.doc or *.docx) or rich text format (*.rtf). They should generally be 1-8 pages long, including references and figures. Submissions to 

the “Articles” section should be divided into Introduction, Methods, Results and Discussion and should include a list of 4-6 key words. 

Short reports and other submissions do not need subheadings or key words. Ideally, English articles should include a French abstract 

and vice versa. Articles should include a map of the area discussed, including all major locations mentioned in the text. Macros, 

complex formatting (such as section breaks) and automatic numbering as provided by text processing software must be avoided. The 

corresponding author’s affiliation and full address must be provided, including e-mail and telephone number. For all other authors, 

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should be numbered using roman numerals. Please give all measurements in metric units. Please accent all foreign words 

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Maps should always be made as concise as possible and should include an inset showing the location of the area discussed in relation 

to the whole of Madagascar. 

Photographs: Black-and-white photographs are ideal. Color photographs are acceptable if they can be printed in greyscale without 

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name each file with the photographer credit and the number of the identifying caption (e.g. “Schwitzer_Fig.1”). We are always 

interested in receiving high quality photographs for our covers, especially those of little known and rarely photographed lemurs, even 

if they do not accompany an article. 

All figures should include concise captions. Captions should be listed on a separate sheet, or after the References section of the 

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should be sent electronically in any one of the following formats: BMP, CDR, EMF, WMF, XLS. Please name all files with the name of 

the first author of the manuscript to which they belong. Do not send figures embedded in the text of the manuscript. 

References: In the text, references should be cited consecutively with the author's surname and year of publication in brackets (e.g. 

Schwitzer et al., 2010; Kaumanns and Schwitzer, 2001). The reference list should be arranged alphabetically by first author's surname. 

Examples are given below. 

Journal article 

Ranaivoarisoa, J.F.; Ramanamahefa, R.; Louis, Jr., E.E.; Brenneman, R.A. 2006. Range extension of Perrier’s sifaka, Propithecus perrieri, in 

the Andrafiamena Classified Forest. Lemur News 11: 17-21. 

Book chapter 

Ganzhorn, J.U. 1994. Les lémuriens. Pp. 70-72. In: S.M. Goodman; O. Langrand (eds.). Inventaire biologique; Forêt de Zombitse. 

Recherches pour le Développement, Série Sciences Biologiques, n° Spécial. Centre d’Information et de Documentation Scientifique 

et Technique, Antananarivo, Madagascar. 

Book 

Mittermeier, R.A.; Konstant, W.R.; Hawkins, A.F.; Louis, E.E.; Langrand, O.; Ratsimbazafy, H.J.; Rasoloarison, M.R.; Ganzhorn, J.U.; 

Rajaobelina, S.; Tattersall, I.; Meyers, D.M. 2006. Lemurs of Madagascar. Second edition. Conservation International, Washington, DC, 

USA. 

Thesis 

Freed, B.Z. 1996. Co-occurrence among crowned lemurs 

(Lemur coronatus) and Sanford’s lemur (Lemur fulvus sanfordi) 

of Madagascar. Ph.D. thesis, Washington University, St. Louis, USA. 

Website 

IUCN. 2008. IUCN Red List of Threatened Species. 

. Downloaded on 21 April 2009. 

Call for voluntary contributions 

As most readers of Lemur News are certainly aware, 

fundraising has become more difficult. We will continue 

to distribute Lemur News free of charge to all interested 

individuals and institutions. However, we would like to ask 

subscribers for voluntary contributions to cover production 

costs. Please contact one of the editors for information on 

how to make contributions. 

Drawing by Stephen D. Nash

Editorial.................................................................................1 

Feature: Madagascar’s Environmental Crisis 

Madagascar’s illegal logging crisis: An update 

and discussion of possible solutions 

Erik R. Patel...............................................................................2 

Ongoing threats to lemurs and their habitat 

inside the Sahamalaza - Iles Radama National 

Park 

Melanie Seiler, Guy H. Randriatahina, 

Christoph Schwitzer.................................................................7 

News and Announcements..........................................9 

Short Communications 

Preliminary conservation status assessment for 

the Data Deficient northern giant mouse lemur 

Mirza zaza 

Eva Johanna Rode, K. Anne-Isola Nekaris, 

Christoph Schwitzer...............................................................11 

An observation of the hairy-eared dwarf lemur, 

Allocebus trichotis, in the Lakato region, eastern 

Madagascar 

Erwan Lagadec, Steven M. Goodman..................................12 

When big lemurs swallow up small ones: 

Coquerel’s dwarf lemur as a predator of grey 

mouse lemurs and endemic rodents 

Susanne Schliehe-Diecks, Matthias Markolf, 

Elise Huchard.........................................................................13 

Collective mobbing of a boa by a group of 

red-fronted lemurs (Eulemur fulvus rufus) 

Lennart Pyritz, Tianasoa Andrianjanahary..........................14 

Response of two nocturnal lemurs (Microcebus 

murinus and Lepilemur leucopus) to a potential 

boiidae (Sanzinia madagascariensis) predator 

Krista Fish...............................................................................16 

Effective predation defence in Cheirogaleus 

medius 

Kathrin H. Dausmann...........................................................18 

Lepilemur feeding observations from Northern 

Madagascar 

Andrew J. Lowin......................................................................20 

Hypotheses on ecological interactions between 

the aye-aye (Daubentonia madagascariensis) 

and microhylid frogs of the genus Platypelis in 

Tsaratanana bamboo forest 

Andolalao Rakotoarison, Solohery A. Rasamison, 

Emile Rajeriarison, David R. Vieites, Miguel Vences............21 

Discovery of crowned sifaka (Propithecus 

coronatus) in Dabolava, Miandrivazo, Menabe 

Region 

Josia Razafindramanana and Rija Rasamimanana............23 

Inferences about the distant past in Madagascar 

Elwyn L. Simons......................................................................25 

Husbandry guidelines for mouse lemurs at Paris 

Zoo 

Delphine Roullet....................................................................27 

Lemur News Vol. 15, 2010 

ISSN 1608-1439 

Table of contents 

Articles 

Diurnal lemur density in the national park 

parcel Ivontaka Nord, UNESCO Biosphere 

Reserve of Mananara-Nord 

Marta Polasky Lyons..............................................................29 

Distribution of Prolemur simus north of the 

Mangoro-Nosivolo River - how far north do we 

really have to look? 

Rainer Dolch, Erik R. Patel, Jonah H. Ratsimbazafy, 

Christopher D. Golden, Tianasoa Ratolojanahary, 

Jean Rafalimandimby, Jonathan L. Fiely................................32 

Enquête préliminaire de la distribution des 

lémuriens de bambou dans et autour du 

Corridor forestier Fandriana-Vondrozo, 

Madagascar 

Andry Rajaonson, Maherisoa Ratolojanahary, 

Jonah Ratsimbazafy, Anna Feistner, Tony King....................34 

Effect of red ruffed lemur gut passage on the 

germination of native rainforest plant species 

Onja H. Razafindratsima, Emilienne Razafimahatratra.....39 

Feeding ecology of the crowned sifaka 

(Propithecus coronatus) in a coastal dry forest 

in northwest Madagascar (SFUM, Antrema) 

Claire Pichon, Rivo Ramanamisata, Laurent Tarnaud, 

Françoise Bayart, Annette Hladik, Claude Marcel 

Hladik, Bruno Simmen.......................................................... 42 

Effet de la dégradation de l'habitat sur la 

consommation alimentaire d'Eulemur 

rubriventer dans deux sites: Talatakely et 

Vatoharanana, du Parc National de Ranomafana 

Laingoniaina H. Rakotonirina, Germain J. Spiral, 

Jonah H. Ratsimbazafy, Soanorolalao Ravelonjanahary, 

Raharizelina Ralaiarison, Stacey Tecot, Alex Hall, 

Tricia Calhoon, Gisèle R. Randria..........................................47 

Observations of terrestrial latrine behaviour by 

the southern gentle lemur Hapalemur 

meridionalis in the Mandena littoral forest, 

southeast Madagascar 

Timothy M. Eppley, Giuseppe Donati....................................51 

Conservation des lémuriens via la protection de 

leurs habitats et le développement communautaire 

dans les corridors de Betaolana et Tsaratanana- 

Betaolana, région de SAVA 

Lala Razafy Fara, Iarilanto Andriamarosolo.........................54 

Genetic diversity in ten Indri (Indri indri) 

populations compared to other lemur species 

John Zaonarivelo, Rick Brenneman, Rambinintsoa 

Andriantompohavana, Edward E. Louis, Jr............................59 

Verreaux’s sifaka fur condition in the spiny forest 

of southern Androy 

Ivan Norscia, Jean Lambotsimihampy, Elisabetta Palagi.....64 

Rediscovery of Sibree’s dwarf lemur in the 

fragmented forests of Tsinjoarivo, central-eastern 

Madagascar 

Marina B. Blanco....................................................................67 

Funding and Training.....................................................69 

Recent Publications........................................................70 

Theses Completed..........................................................71

Vol. 15 - Deutsches Primatenzentrum

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