Vol. 15 - Deutsches Primatenzentrum
Vol. 15 - Deutsches Primatenzentrum
Vol. 15 - Deutsches Primatenzentrum
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LEMUR NEWS<br />
The Newsletter of the Madagascar Section of the IUCN/SSC Primate Specialist Group<br />
<strong>Vol</strong>. <strong>15</strong>, 2010 ISSN 1608-1439<br />
Editors<br />
Christoph Schwitzer (Editor-in-chief)<br />
Bristol Conservation and Science Foundation, Bristol Zoo Gardens, UK; cschwitzer@bcsf.org.uk<br />
Claudia Fichtel<br />
German Primate Center, Göttingen, Germany; claudia.fichtel@gwdg.de<br />
Jörg U. Ganzhorn<br />
University of Hamburg, Germany; ganzhorn@biologie.uni-hamburg.de<br />
Rodin M. Rasoloarison<br />
German Primate Center, Göttingen, Germany; kirindy@simicro.mg<br />
Jonah Ratsimbazafy<br />
GERP, Antananarivo, Madagascar; gerp@wanadoo.mg<br />
Anne D. Yoder<br />
Duke University Lemur Center, Durham, USA; anne.yoder@duke.edu<br />
IUCN/SSC Primate Specialist Group<br />
Chairman Russell A. Mittermeier, Conservation International, Arlington, VA, USA<br />
Deputy Chair Anthony B. Rylands, Conservation International, Arlington, VA, USA<br />
Coordinator – Section on Great Apes Liz Williamson, Stirling University, Stirling, Scotland, UK<br />
Regional Coordinators – Neotropics<br />
Mesoamerica – Liliana Cortés-Ortiz, Museum of Zoology & Department of Ecology and Evolutionary<br />
Biology, University of Michigan, Ann Arbor, MI, USA<br />
Andean Countries – Erwin Palacios, Conservation International Colombia, Bogotá, Colombia and<br />
Eckhard W. Heymann, <strong>Deutsches</strong> <strong>Primatenzentrum</strong>, Göttingen, Germany<br />
Brazil and the Guianas – M. Cecília M. Kierulff, Instituto para a Conservação dos Carnívoros<br />
Neotropicais – Pró-Carnívoros, Atibaia, São Paulo, Brazil, Fabiano Rodrigues de Melo, Universidade<br />
Federal de Goiás, Jataí, Goiás, Brazil, and Maurício Talebi, Universidade Federal de São Paulo, Diadema,<br />
São Paulo, Brazil<br />
Regional Coordinators – Africa<br />
West Africa – W. Scott McGraw, The Ohio State University, Columbus, OH, USA<br />
Regional Coordinators – Madagascar<br />
Jörg U. Ganzhorn, Hamburg University, Hamburg, Germany, and Christoph Schwitzer, Bristol Conservation<br />
and Science Foundation, Bristol Zoo Gardens, Bristol, UK<br />
Regional Coordinators – Asia<br />
China – Long Yongcheng, The Nature Conservancy, China<br />
Southeast Asia – Jatna Supriatna, Conservation International Indonesia Program, Jakarta, Indonesia, and<br />
Christian Roos, <strong>Deutsches</strong> <strong>Primatenzentrum</strong>, Göttingen, Germany<br />
IndoBurma – Ben Rawson, Conservation International, Hanoi, Vietnam<br />
South Asia – Sally Walker, Zoo Outreach Organization, Coimbatore, Tamil Nadu, India, and Sanjay Molur,<br />
Wildlife Information Liaison Development, Coimbatore, Tamil Nadu, India<br />
Editorial assistants<br />
Nicola Davies, Rose Marie Randrianarison<br />
Layout<br />
Heike Klensang, Anna Francis<br />
Front cover: The Endangered golden-crowned sifaka (Propithecus tattersalli) at the edge of an area devastated<br />
by gold mining activities in the Daraina region of north-eastern Madagascar. © Pete Oxford/naturepl.com<br />
Addresses for contributions<br />
Christoph Schwitzer<br />
Bristol Conservation and Science Foundation<br />
Bristol Zoo Gardens<br />
Clifton, Bristol BS8 3HA<br />
United Kingdom<br />
Fax: +44 (0)117 973 6814<br />
Email: cschwitzer@bristolzoo.org.uk<br />
Lemur News online<br />
All <strong>15</strong> volumes are available online at www.primate-sg.org, www.aeecl.org and www.dpz.eu<br />
This volume of Lemur News was kindly supported by the Margot Marsh Biodiversity Foundation<br />
(through Conservation International’s Primate Action Fund) and by WWF Madagascar.<br />
Printed by Goltze GmbH & Co. KG, Göttingen, Germany<br />
Jonah Ratsimbazafy<br />
GERP<br />
34, Cité des Professeurs<br />
Antananarivo 101<br />
Madagascar<br />
Email: gerp@wanadoo.mg
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 1<br />
Editorial<br />
I am writing this Editorial only a couple of days after another<br />
attempted (and failed) Coup d’Etat in Madagascar,in which a<br />
faction of the army tried to topple the Transition Government.<br />
For nearly two years now, since the start of the political<br />
crisis in early 2009,the country has not seen a week without<br />
demonstrations, tensions between different political<br />
parties and attempts from international mediators to get<br />
power-sharing agreements signed by all sides. Most donors,<br />
governments and multinational organisations alike, have<br />
frozen all non-humanitarian aid for Madagascar,which has led<br />
to severe funding shortages in the environmental and conservation<br />
sector. The political crisis has thus quickly turned<br />
into a full-blown environmental crisis, with large-scale illegal<br />
logging taking place mainly in eastern Madagascar (Marojejy,<br />
Masoala, Makira), and unseen levels of lemur poaching all<br />
across the island.To keep people aware of the seriousness of<br />
the situation we have decided to run another feature on<br />
Madagascar’s environmental crisis in this issue of Lemur News,<br />
with an excellent update on illegal logging by Erik Patel as<br />
well as a case study of ongoing threats to lemurs and their<br />
habitat in Sahamalaza National Park by Melanie Seiler and<br />
colleagues.<br />
The conservation situation of lemurs has also been a big concern<br />
in several presentations given at the most recent 23rd<br />
Congress of the International Primatological Society in<br />
Kyoto, Japan. The talk that I remember best was by Lemur<br />
News co-editor Jonah Ratsimbazafy, who reminded the audience<br />
in a very emotional way that scientists and conservationists<br />
working in Madagascar had a moral responsibility to<br />
respond to the "cries of the lemurs", as otherwise these<br />
would remain unheard by the Malagasy and international<br />
community.In the biennial discussion session of "Primates in<br />
Peril", the list of the world’s top 25 most endangered primates,<br />
issued jointly by the IUCN/SSC Primate Specialist<br />
Group and IPS,lemurs remained a very high priority and will<br />
again make up 20% of the 25 listed species in the next biennium.Sadly,Madagascar<br />
thus retains its first place (along with<br />
Vietnam) as the country harbouring the highest number of<br />
the top 25. It can only be hoped that the political classes of<br />
Madagascar come to agree a way out of the current crisis<br />
sooner rather than later, as otherwise we run the very serious<br />
risk,during the UN Decade of Biodiversity 2011-2020,of<br />
losing a substantial proportion of the endemic biodiversity of<br />
this amazing megadiversity country.<br />
Alison Jolly with Russ Mittermeier at the IPS Lifetime Achievement<br />
Award 2010 ceremony in Kyoto. (Photo: R. Mittermeier)<br />
For a change,on a very positive note,I am thrilled to say that<br />
Alison Jolly was awarded the IPS Lifetime Achievement<br />
Award for her long-term commitment to lemur conservation<br />
and environmental education in Madagascar (see News<br />
and Announcements). My two daughters (now 4 and 2 years<br />
old) and I particularly enjoy reading Alison’s children’s book<br />
on Bitika,the mouse lemur,as,I am sure,do lots of children in<br />
Madagascar and elsewhere in the world.<br />
It is encouraging to see that this volume of Lemur News is<br />
again full of articles and short reports not only on lemur species<br />
red-listed in one of the Threatened categories (VU, EN<br />
or CR),but also on Data Deficient nocturnal species such as<br />
Mirza zaza,Lepilemur leucopus and the recently rediscovered<br />
Cheirogaleus sibreei (see the articles by Rode et al., Fish, and<br />
Blanco, respectively). As Johanna Rode and colleagues point<br />
out in their short report on Mirza zaza,Madagascar is in the<br />
unusual situation that 45 % of its primate species are redlisted<br />
as Data Deficient,which is a far higher percentage than<br />
in any other primate habitat country and mainly derived<br />
from the discovery of dozens of cryptic species in the genera<br />
Lepilemur and Microcebus over the last couple of years. Many<br />
of those species are only known from their type localities<br />
and may in fact be highly endangered. The more research is<br />
conducted and published on them, the easier it will become<br />
to assign them a conservation status and target them with<br />
conservation measures. It will require a concerted effort of<br />
the lemur research and conservation community over the<br />
next decade or so to to reduce the number of Data Deficient<br />
species to a level comparable to other regions (or, ideally, to<br />
zero).<br />
Another encouraging development is the frenzy of research<br />
and conservation activities now under way for Prolemur simus<br />
at various locations both south and north of the Mangoro<br />
River,reported by Dolch et al.as well as Rajaonson et al.in this<br />
volume. The greater bamboo lemur undoubtedly remains<br />
one of the most endangered of Madagascar’s lemurs. However,<br />
with several additional populations having been discovered<br />
over the last two years, workshops having been conducted<br />
that have led to a joint-up approach to this species’<br />
conservation,and the ex situ population having been included<br />
as an integral part of conservation efforts, I now think that<br />
we stand a real chance of saving Prolemur simus from extinction.<br />
As Jörg Ganzhorn announced in his editorial to Lemur News<br />
14, I have taken over the coordination of this newsletter<br />
from him after the 2009 volume, hence this is now the first<br />
volume that I have helped produce (which is my humble excuse<br />
for its slightly late publication). Jörg has been involved<br />
with Lemur News since its inception in 1993,first as a member<br />
of its Editorial Board and from volume 3 (1998) as its Editor.I<br />
am thus pleased to say that we will not lose his experience<br />
and backing,as he has kindly agreed to remain part of the editorial<br />
team. Likewise, Jonah Ratsimbazafy and Rodin Rasoloarison,who<br />
have been the newsletter’s Malagasy coordinators<br />
since 2006, and Anne Yoder, who represents the Duke<br />
Lemur Center, will carry on as editorial team members, for<br />
which I am grateful.I am indebted to Heike Klensang,who has<br />
been doing the layout for Lemur News now for more than a<br />
decade and is still not tired of it,and to Anna Francis,who has<br />
designed the beautiful new logo and front cover. Very many<br />
thanks also to Stephen D. Nash for the wonderful lemur silhouettes<br />
that we printed on the inside back cover.<br />
This volume of Lemur News was kindly supported by the<br />
Margot Marsh Biodiversity Foundation through Conservation<br />
International’s Primate Action Fund, and by the WWF<br />
Madagascar and West Indian Ocean Programme Office.
Page 2 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
I very much look forward to helping to take Lemur News into<br />
the UN Decade of Biodiversity together with the editorial<br />
team and with its base of loyal contributors and readers,and I<br />
will do my best to ensure that the newsletter will continue to<br />
help promote the conservation of lemurs as it has done for<br />
the last 17 years.<br />
Christoph Schwitzer<br />
Feature: Madagascar’s<br />
Environmental Crisis<br />
Madagascar’s illegal logging crisis: an update<br />
and discussion of possible solutions<br />
Erik R. Patel<br />
Cornell University, 211 Uris Hall, Ithaca, NY 14850, USA,<br />
patel.erik@gmail.com<br />
How sure are you that your favorite rosewood or ebony<br />
acoustic guitar was not made from rare,illegally logged trees<br />
in Madagascar;an exceptional biodiversity hotspot with desperately<br />
little original forest remaining? What is the origin of<br />
the wood in the expensive oriental-style rosewood furniture<br />
which is heavily advertised for sale on the internet? Unfinished<br />
rosewood boards from Madagascar are openly sold<br />
even in the United States (www.gilmerwood.com/boards_<br />
rosewood-exotic_unique.htm) and the United Kingdom (www.<br />
exotichardwoods.co.uk/Woods_List/Madagascar_Rosewood.asp).<br />
Can such vendors prove that the rosewood was legally (and<br />
ethically) obtained? The answer is usually "no".These can be<br />
difficult questions for consumers to answer, but purchasing<br />
these products can prolong the ongoing logging crisis in<br />
northeastern Madagascar in some of the most unique and biologically<br />
diverse forests in the world.<br />
Consumers should be suspicious since none of these rapidly<br />
disappearing Madagascan rosewood and ebony species are<br />
yet protected under CITES,the Convention on International<br />
Trade in Endangered Species. In November of last year, Gibson<br />
Guitars, one of the two largest U.S. stringed-instrument<br />
companies,came under federal investigation for violating the<br />
Lacey Act by allegedly using illegal rosewood from Madagascar<br />
which had first been shipped to Germany and then the<br />
United States (Michaels, 2009). Most of the illegally logged<br />
rosewood in Madagascar is used for the manufacture of furniture<br />
in China. Some of this is known to be sold in China as<br />
luxurious "Ming Dynasty style" furniture (Global Witness<br />
and Environmental Investigation Agency, 2009). Some may<br />
well be exported to western countries. China is the world’s<br />
leading exporter of furniture.According to the Office of the<br />
United States Trade Representative, the United States imported<br />
16 billion dollars of Chinese furniture in 2009,making<br />
it the USA’s fifth largest import from China.<br />
Illegal logging of rosewood (Dalbergia spp.) and ebony (Diospyros<br />
spp.) has emerged as the most severe threat to Madagascar’s<br />
dwindling northeastern rainforests. In 2009, a year<br />
of political upheaval in Madagascar due to an undemocratic<br />
change of power,approximately 100,000 of these trees were<br />
illegally cut in the UNESCO World Heritage Sites of Masoala<br />
National Park,Marojejy National Park,the Makira Conservation<br />
Site, and Mananara Biosphere Reserve (also a national<br />
park).Needless to say,the wood is extremely valuable.Rosewood<br />
can sell for US$5,000 per cubic meter,more than double<br />
the price of mahogany.Several hundred million dollars of<br />
these precious hardwoods were cut in 2009 in protected<br />
areas. The overwhelming majority of these profits are taken<br />
by a rosewood mafia of a few dozen organizing individuals,<br />
many of whose identities are well known.Few others benefit.<br />
Harvesting these extremely heavy hardwoods is a labor intensive<br />
activity requiring coordination between local residents<br />
who manually cut the trees, but receive little profit<br />
(about US$5/day), and a criminal network of exporters, domestic<br />
transporters, and corrupt officials who initiate the<br />
process and reap most of the enormous profits. This is a<br />
"tragedy with villains" unlike habitat disturbance from subsistence<br />
slash-and-burn agriculture which has been well described<br />
as a "tragedy without villains" (Barrett et al., 2010;<br />
Débois, 2009; Global Witness and Environmental Investigation<br />
Agency,2009;Patel,2007,2009;Randriamalala and Liu,in<br />
press;Schuurman and Lowry,2009;Schuurman,2009;Wilme<br />
et al., 2009; Wilme et al., in press).<br />
Globally, illegal logging results in an estimated US$10 billion<br />
lost per year to the economies of timber producing countries<br />
(Furones, 2006). In addition to depriving the government<br />
of Madagascar of millions of dollars of taxable revenue,<br />
illegal logging of this precious wood has decimated tourism<br />
in northeastern Madagascar, which had become a growing<br />
source of local income. Although selective logging results in<br />
less absolute forest loss than clear-cutting, it is often<br />
accompanied by substantial peripheral damage such as decreases<br />
in genetic diversity and increases in the susceptibility<br />
of the impacted areas to burning and bushmeat hunting.<br />
Documented long-term ecological consequences of selective<br />
logging in Madagascar include invasion of persistent,<br />
dominant non-native plant species, impaired faunal habitat,<br />
and a diminution of endemic mammalian species richness<br />
(Gillies, 1999; Cochrane and Schultze, 1998; Brown and<br />
Gurevitch, 2004; Stephenson, 1993). In actual practice, rosewood<br />
logging has turned out to be far less "selective" than<br />
originally believed. Often rafts made of a lighter species of<br />
wood (Dombeya spp.) are constructed to float the much<br />
more dense rosewood logs down rivers. Approximately five<br />
Dombeya trees are cut as "raft wood" for every one rosewood<br />
tree (Randriamalala and Liu,in press).Tall adult trees of<br />
a variety of species, that simply happen to be very close to<br />
rosewood trees, must often be cut simply to gain access to<br />
cut down a rosewood tree. This has been observed in<br />
Marojejy (pers. obs.).<br />
Red ruffed lemurs (Varecia rubra) are probably the most negatively<br />
impacted lemur since many were hunted by these loggers<br />
and this species is known to feed on ebony trees<br />
(Diospyros spp.) as well as pallisandre (Dalbergia spp.) in<br />
Masoala (Vasey, pers. comm.). Varecia rubra probably also<br />
feeds on the fruits and leaves of the logged "raft wood"<br />
Dombeya spp. trees like Varecia v. editorium in Manombo Forest<br />
in southeastern Madagascar (Ratsimbazafy, 2007). In<br />
Mantadia National Park, Indri indri and Propithecus diadema<br />
consume young leaves of one species of actual rosewood<br />
(Dalbergai baronii) which is also consumed by Milne-Edwards’<br />
sifakas (Propithecus edwardsi) in Ranomafana National Park<br />
(Powzyk and Mowry,2003;Arrigo-Nelson,2007).Propithecus<br />
diadema at Tsinjoarivo consume the unripe fruit of ebony<br />
trees (Irwin, 2006). In Marojejy, silky sifakas (Propithecus<br />
candidus) not uncommonly feed on the young leaves of pallisandre<br />
(Dalbergia chapelieri) which is also a preferred sleeping<br />
tree (pers. obs.).<br />
When discussing the impacts of precious wood logging, it is<br />
important not to forget how damaging all this has been to local<br />
communities as well.Local residents have suffered as foreign<br />
and domestic elites have corrupted the forest service,<br />
leading to losses of sustainable employment in tourism, re-
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 3<br />
search, and conservation. In some cases, community life has<br />
suddenly changed as gambling, prostitution, and crime have<br />
increased in rural communities. Moreover, the risks of local<br />
food shortages and nutritional deficiencies mount when<br />
farmers abandon subsistence agriculture for temporary,physically<br />
dangerous illegal logging work (Global Witness and<br />
Environmental Investigation Agency,2009;Patel,2007,2009).<br />
Moreover,illegal loggers trample on the beliefs and taboos of<br />
local people.In traditional Sakalava culture,ebony is a sacred<br />
wood only cut by priests who conduct traditional ceremonies<br />
with ebony staffs. The chief of Ankalontany, a Sakalava<br />
Malagasy village in the northeast, explained that "Some<br />
strangers from outside our village came here. They started<br />
cutting ebony and they clearly had no right. We asked for<br />
their authorization but they said they didn’t have to show us<br />
papers.They said they had police clearance and we can’t stop<br />
them." Laurent Tutu, president of the forest association of<br />
Ankalontany, remarked "It hurts us to see our trees cut like<br />
this. The forest loses its personality" (Cocks, 2005).<br />
Although illegal logging in Madagascar has received some media<br />
attention recently, confusion still remains regarding a<br />
number of key facts. The aim of this report is to provide an<br />
update (at the time of writing: May 25, 2010), dispel a few<br />
myths,discuss some of the possible solutions to this ongoing<br />
crisis, and present a comprehensive bibliography of articles,<br />
photos, films, and videos related to this topic.<br />
Four myths about illegal logging in Madagascar<br />
Myth #1: "Plenty of Madagascar rosewood is harvested legally…"<br />
says Bob Taylor, founder of Taylor Guitars. Quote<br />
from Gill, C. (2010). Log Jam. Guitar Aficionado. Spring Issue.<br />
P. 68<br />
This is simply not true. A vast amount of published evidence<br />
clearly shows that very very little,if any,of the rosewood logging<br />
in Madagascar is legal.The overwhelming majority of exported<br />
Madagascar rosewood is illegally logged within Masoala<br />
National Park and Marojejy National Park (which are part<br />
of a UNESCO World Heritage Site) as well as Mananara Biosphere<br />
Reserve (also a national park) and the vast Makria<br />
Conservation Site (Barrett et al., 2010; Débois, 2009; Global<br />
Witness and Environmental Investigation Agency,2009;Patel,<br />
2007, 2009; Randriamalala and Liu, in press; Schuurman and<br />
Lowry, 2009; Schuurman, 2009; Wilme et al., 2009; Wilme et<br />
al., in press).<br />
Myth #2: The current ban has stopped illegal logging.<br />
In late March, the government of Madagascar announced a<br />
new two to five year ban on export and cutting of ebony and<br />
rosewood. The decree #2010-141 officially passed on April<br />
14, 2010. Clearly this was an important and large step forward.<br />
However, the decree does not apparently include<br />
pallisandre, a precious hardwood in the same genus (Dalbergia)<br />
as rosewood. Illegal logging of pallisandre has heavily<br />
impacted some reserves such as Betampona Natural Reserve<br />
(Kett, 2005; Bollen, 2009). At the time of writing (May<br />
25, 2010), there have been no new exports since the recent<br />
ban.However,illegal rosewood and ebony logging still continues<br />
inside Mananara Biosphere Reserve and the Makira Conservation<br />
Site according to reliable anonymous informants.<br />
The clearest information has come from Mananara where at<br />
least several hundred, recently cut, rosewood logs were observed.<br />
Myth #3: Illegal logging was never a big problem in Madagascar<br />
until the recent political crisis.<br />
Illegal logging in Madagascar of rosewood (Dalbergia spp.) and<br />
ebony (Diospyros spp.) did not begin with the culmination of<br />
the political crisis in March 2009.A major illegal logging crisis<br />
in World Heritage Sites (Masoala National Park and Marojejy<br />
National Park) took place during 2004-2005,a time of political<br />
stability. The earliest documented case of rosewood logging<br />
in Madagascar and foreign export dates to 1902.Foreign<br />
exports of Madagascar rosewood occurred at "low" levels<br />
(1000 to 5000 tonnes) between 1998 and 2007. In 2008, exports<br />
jumped to 13,000 tonnes,and jumped again in 2009 to<br />
more than 35,000 tonnes (Botokely,1902;Randriamalala and<br />
Liu,in press;Global Witness and Environmental Investigation<br />
Agency, 2009).<br />
Myth #4: There are 43 species of rosewood trees in Madagascar.<br />
Some recent reports had mistakenly made this statement. It<br />
is not entirely clear exactly how many rosewood species are<br />
found in Madagascar. More botanical research is needed.<br />
However, currently, there are believed to be 10 species of<br />
rosewood in Madagascar in the genus Dalbergia which contains<br />
48 total species. The rosewood species are presumed<br />
to be Dalbergia baronii [VU], D. bathiei [EN], D. davidii [EN],D.<br />
louvelii [EN], D. mollis [NT], D. monticola [VU], D. normandii<br />
[EN], D. purpurascens [VU], D. tsiandalana [EN], and D. viguieri<br />
[VU] (Barrett et al., 2010).<br />
Rosewood stockpile solutions?<br />
Approximately 10,280 tonnes of illegally logged rosewood<br />
remain stockpiled in numerous locations in northeastern<br />
Madagascar, such as the ports of Vohemar and Antalaha as<br />
well as private residences in those cities and Sambava,<br />
Ampanifena,Ambohitralalana,and others.Each <strong>15</strong>0 kg log has<br />
an approximate market value of US$1,300 usd.As unfinished<br />
logs, the value of the current stockpile is therefore approximately<br />
US$90 million.Value increases dramatically,of course,<br />
after being constructed, for example, into high-end Ming<br />
Dynasty style furniture in China.A single armoire composed<br />
of only a few logs can sell for US$20,000 or more.It’s a horrid<br />
contrast to the annual income in Madagascar (about<br />
US$400) or the daily wage provided to loggers (US$5) for<br />
the dangerous and physically debilitating work (Randriamalala<br />
and Liu, in press; Global Witness and Environmental<br />
Investigation Agency, 2009; anonymous local informants).<br />
If the export ban holds (numerous other bans did not),what<br />
should be done with these stockpiles? Several ideas have<br />
been suggested.<br />
1. The "Forest Counterpart Fund" (Wilme et al., 2009) aims<br />
to create a conservation and charitable works fund to assist<br />
local communities and forests damaged by the illegal logging.<br />
The logs are not sold on the open market as in the second<br />
proposal below.Rather,philanthropists,conservation organizations,<br />
and international aid agencies pay to "adopt" a log.<br />
Each log can be "adopted" for its market value (about<br />
US$1,300). The logs themselves are given to (carefully selected)<br />
local residents who are victims of the illegal logging.<br />
The logs would then be carved,engraved,and customized for<br />
public display as symbols. If sufficient donors can be found,<br />
this proposal offers a win-win solution for Madagascar’s forests<br />
as well as people.<br />
2. The Moratorium-Conservation-Amnesty-Reforestation<br />
(MCAR) program (Butler, 2009). This is essentially a one-off<br />
actual sale with conservation benefits. Logs would be auctioned<br />
via a transparent market system in which the price<br />
and the log code would be recorded, publicly available, and<br />
digitally traceable.Funds generated would mainly go towards
Page 4 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
conservation programs such as reforestation and forest<br />
monitoring. Criminal traders would receive amnesty from<br />
prosecution as well as a very small percentage of the funds.<br />
An export moratorium would be required.There is always a<br />
danger that one-off sales can encourage further logging; a<br />
topic which has been extensively debated with respect to<br />
confiscated elephant ivory stockpiles. An impressive recent<br />
review paper in Science (Wasser et al., 2010):<br />
www.sciencemag.org/cgi/content/short/327/5971/1331)<br />
argued that no one-off ivory sales should be approved even if<br />
the funds go towards conservation.<br />
3. Destroy the stockpile. This was recently reiterated by<br />
Global Witness (GW) and Environmental Investigation<br />
Agency (EIA). Andrea Johnson, Director of Forest Campaigns<br />
at EIA explained that "To end the cycle of illegal harvest<br />
and corruption, the government should take the step of<br />
destroying all stocks that are not contained in the latest official<br />
inventories…Traders, who are currently stockpiling illegal<br />
timber, hoping for another ‘exceptional’ export authorization,must<br />
receive a clear signal that it will be impossible to<br />
profit from the illegal trade in the future." Numerous examples<br />
can be found from around the world of simple and effective<br />
destruction of stockpiles of contraband such as small<br />
arms,drugs,and ivory.Destruction also eliminates the not insignificant<br />
expense of storing and guarding the items.Burning<br />
the rosewood stockpiles would create a lot of pollution, it<br />
has been argued, and might be dangerous given the high volume.<br />
Other ways of destroying the wood are possible however.<br />
The wood could be hacked into tiny unusable pieces.<br />
This is already done sometimes by park rangers in Madagascar.<br />
This would take a very long time, but would be a fitting<br />
punishment of hard labor for members of the rich rosewood<br />
mafia! Of course, destruction of the wood, whatever the<br />
method,would contribute no money for any conservation or<br />
community development funds.<br />
Any of these possibilities are better than what has happened<br />
in the past:seized wood was auctioned off to the highest bidder.Foreign<br />
export remains a possibility too,despite the ban.<br />
French shipping company CMA-CGM Delmas exported<br />
rosewood from Madagascar several times in 2009 and 2010.<br />
Long-term solutions?<br />
Thinking long-term, what can be done to prevent another illegal<br />
logging crisis in Madagascar?<br />
Some may argue that so little rosewood and ebony remains,<br />
logging on this scale could never happen again.However,this<br />
had been claimed before 2009 too. More surveys are clearly<br />
needed. One hopes that some of the more impenetrable regions<br />
of mountainous Marojejy National Park may still have<br />
rosewood. But because rosewood tends to be harvested at<br />
lower elevations, near rivers (where the largest individuals<br />
are found), it is less protected by the physical challenges of<br />
the massif than some other tree species. It is encouraging<br />
that some Dalbergia and Diospyros species can form stump<br />
sprouts which can grow into a new tree over many many<br />
years. Unfortunately, some entire rosewood stumps are removed<br />
either to hide evidence of logging or for wood for<br />
small, locally made rosewood vases. Rosewood trees are<br />
known to be some of the oldest trees in the eastern Malagasy<br />
humid forests. They can live to be more than 400 years old,<br />
according to local guides. Traders explain that they can be<br />
harvested after 50 years (Patel 2007, 2009).<br />
1. CITES<br />
The surest way to reduce the likelihood of another illegal<br />
logging crisis in Madagascar, is to list all species in the genera<br />
of Dalbergia and Diospyros on CITES Appendix 1. Currently<br />
none of Madagascar’s ebony or rosewood species are protected<br />
under any appendices within the Convention on International<br />
Trade in Endangered Species (CITES). Globally,<br />
only one species of rosewood,Brazilian rosewood (Dalbergia<br />
nigra), is listed under CITES Appendix 1. This is the most<br />
stringent category,and prohibits all commercial trade of that<br />
wood from the date of listing. This has generally been effective.Guitars<br />
in the United States made of Brazilian rosewood<br />
are known to have risen in price and are harder to find since<br />
Appendix 1 listing. Similarly, Appendix 1 listing of Alerce<br />
(Fitzroya cupressoides),a heavily logged South American conifer,<br />
has significantly reduced logging and trade (Barrett et al.,<br />
2010; Keong, 2006).<br />
A few other Brazilian and Central American rosewood species<br />
are listed under CITES Appendix 2 and 3. These lower<br />
appendices aim to regulate trade,not prohibit it.Just this year,<br />
another species of Brazilian rosewood (Aniba rosaeodora),exported<br />
extensively as fragrant oil, was listed under CITES<br />
Appendix 2. Two additional species of Central American<br />
rosewood (D. retusa and D. stevensonii) are listed under Appendix<br />
3. Appendix 2, unlike Appendix 3, does require that<br />
the CITES authorities in the export nation determine that<br />
the species were legally obtained and that their export will<br />
not be detrimental to species survival.There seem to be few<br />
cases where Appendix 3 listing was sufficient, except as a<br />
means to Appendix 2 or higher listing. The well examined<br />
case-studies of big-leaf mahogany (Swietenia macrophylla) and<br />
ramin (Gonystylus spp.) both began as Appendix 3 species<br />
(which only requires unilateral listing by a habitat country)<br />
and were later voted in as Appendix 2 species by the CITES<br />
parties (Keong, 2006).<br />
To what degree can CITES regulations be implemented and<br />
enforced? The need for more officially trained import inspectors<br />
has been suggested numerous times. The agency<br />
chosen as the CITES management authority should be free<br />
of corruption and have experience in forest management.<br />
Insufficient trained staff has also hindered the ability of export<br />
authorities to determine whether an Appendix 2 species<br />
was legally obtained and non-detrimental to species survival.Range<br />
countries often require assistance in this respect.<br />
An unusually good example comes from Indonesia where biological<br />
data for ramin has been used in non-detriment findings<br />
to examine sustainability. Missing "certificates of origin"<br />
have been a problem for some Appendix 3 species. While<br />
ramin and big-leaf mahogany were listed on Appendix 3, the<br />
required ‘certificates of origin’ were not consistently issued<br />
by exporting nations;while importing countries were not always<br />
diligent about confirming that shipments arrived with<br />
such certificates (Blundell, 2007; Keong, 2006).<br />
2. Independent forest monitoring (IFM)<br />
In addition to CITES,actual improvements in forest monitoring<br />
on the ground are needed. A new system called independent<br />
forest monitoring (IFM) may be needed in order<br />
stop illegal logging, monitor implementation of REDD (Reducing<br />
Emissions from Deforestation and Forest Destruction)<br />
programs, restore the confidence of international donors,<br />
and ultimately to save Madagascar’s precious forests as<br />
well as attain social justice for Madagascar’s impoverished<br />
population. IFM has been defined as "the use of an independent<br />
third party that,by agreement with state authorities,provides<br />
an assessment of legal compliance, and observation of<br />
and guidance on official forest law enforcement systems"<br />
p. 18 (Global Witness,2005). IFM is similar in principle to unbiased<br />
international election observers. Local and international<br />
expertise is utilized, and monitoring teams operate
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 5<br />
independently but with the consent of the host government.<br />
Independent forest monitors are strictly observers, law enforcement<br />
remains the responsibility of local officials and<br />
governments.<br />
Of course other nations have been faced with similar forest<br />
monitoring problems.IFM has already been used successfully<br />
in several African and Central American nations seeking to<br />
improve the effectiveness of their forest monitoring. Since it<br />
was first introduced in 1999, IFM has been established in<br />
Cameroon, Cambodia, and Honduras. Smaller scale feasibility<br />
and pilot studies have been conducted in Ghana, Peru,<br />
Mozambique, Republic of Congo, Tanzania, and Democratic<br />
Republic of Congo. In Cambodia and Cameroon, donor<br />
countries have been the impetus behind IFM.Though in Honduras,the<br />
incentive for IFM was domestic,and hosted by the<br />
Honduran Commission for Human Rights (CONADEH).<br />
Furones (2006) and Young (2007) review the results of IFM in<br />
these nations, and consider them to be "broadly positive".<br />
Specific examples of the impact of IFM in these nations include:documentation<br />
of hundreds of forest crimes,cancellation<br />
of logging concessions,moratoriums on logging and timber<br />
transport,and creation of new "forest crimes monitoring<br />
units" in the forestry administrations. In some cases,IFM has<br />
earned money for these governments by identifying violations<br />
which led to large fines against logging companies and<br />
individuals breaching the law and forest management regulations.<br />
3. Update IUCN Red List assessments<br />
The approximately 10 Madagascar rosewood species listed<br />
above have not had their official conservation status evaluated<br />
by the IUCN since 1998. At that time, all were threatened<br />
except for D. mollis.Five of the ten were already classified<br />
as ‘endangered’ then. Given the extreme logging since<br />
that time, it is likely that their Red List categories should be<br />
reassessed (IUCN, 2010).<br />
4. UNESCO World Heritage Sites "in danger"<br />
The majority of the illegally logged rosewood in Madagascar<br />
comes from two UNESCO World Heritage Sites: Masoala<br />
National Park and Marojejy National Park.Why have Masoala<br />
and Marojejy not been placed on the World Heritage Sites<br />
"In Danger" List? After all,2010 is the United Nations "International<br />
Year of Biodiversity". Nine national parks and seven<br />
other protected natural areas are currently on this danger<br />
list, mainly for extensive anthropogenic disturbance such as<br />
poaching, logging, and war. The extent of the logging damage<br />
in Masoala National Park, in particular, over the past 5 years,<br />
must rival that of some of the other national parks "in danger".<br />
Placing a site on the UNESCO "danger list" is not utter<br />
de-listing. It is a reversible process meant to draw attention<br />
to and attract possible resources which can alleviate the crisis.There<br />
are specific funds that can become available if a site<br />
is placed on the danger list. One can only speculate that the<br />
reasons for no change in status may well be political and<br />
practical. Perhaps it complicates matters that eight national<br />
parks (which include these two) comprise the single Atsinanana<br />
World Heritage Site Complex. Perhaps there are<br />
fears of triggering an even greater loss of tourism.Whatever<br />
the reasons may be, it is odd that UNESCO has not been<br />
more vocal or active in its support of these two national<br />
parks which are the biodiversity jewels of the Atsinanana<br />
World Heritage Site Complex (IUCN, 2007).<br />
5. DNA fingerprinting<br />
DNA fingerprinting has recently been used on confiscated<br />
ivory to determine which populations of African elephants<br />
were slaughtered. Similar genetic techniques would be of<br />
great assistance in determining which populations of Madagascar<br />
rosewood are being logged the most,and in identifying<br />
species. DNA testing has already been used to track timber,<br />
but not yet in Madagascar.One of the biggest methodological<br />
challenges is extracting DNA from the heartwood of dead<br />
tree trunks (e.g., rosewood stockpiles), which consist of<br />
dead cells with partly degraded DNA. In living trees, it is a<br />
routine process to obtain DNA from the cambium just beneath<br />
the bark or leaves or buds. Nevertheless, several new<br />
techniques have successfully extracted DNA from dry wood<br />
of ramin (Gonystylus spp.) and other woods including 1000<br />
year old beech (Fagus spp.) (Nielson and Kjaer, 2008).<br />
References and rosewood logging resources<br />
Barrett, M.A.; Brown, J.L.; Morikawa, M.K.; Labat, J-N.; Yoder,<br />
A.D. In press. CITES designation for endangered rosewood<br />
in Madagascar. Science.<br />
Blundell,A.G.2007.Implementing CITES regulations for timber.<br />
Ecological Applications 17: 323-330.<br />
Bohannon, J. 2010. Madagascar’s forests get a reprieve – But<br />
for how long? Science 328: 23-25.<br />
Bollen, A. 2009. Eighth continent quarterly. The Newsletter<br />
of the Madagascar Fauna Group.Autumn Issue.<br />
Bosser, J.; Rabevohitra, R. 1996. Taxa et noms nouveaux dans<br />
le genre Dalbergia (Papilionaceae) à Madagascar et aux<br />
Comores. Bulletin du Museum national d'Histoire Naturelle,<br />
4e sér., 18: 171-212.<br />
Bosser,J.;Rabevohitra,R.2005.espèces nouvelles dans le genre<br />
Dalbergia (Fabaceae, Papilionoideae) à Madagascar.<br />
Adansonia, Sér. 3, 27, 2: 209-216.<br />
Botokely (Marc Clique).1902.Chronique commerciale,industrielle<br />
et agricole. Revue de Madagascar 4: 356-365.<br />
Braun, D. 2009. Lemurs, rare forests, threatened by Madagascar<br />
strife. NatGeo News Watch.<br />
blogs.nationalgeographic.com/blogs/news/chiefeditor/<br />
2009/03/lemurs-threatened-by-madagascar-strife.html.<br />
Downloaded on 23 March 2009.<br />
Braun, D. 2010. Conservationists applaud renewed ban on<br />
Madagascar rosewood. NatGeo News Watch.<br />
blogs.nationalgeographic.com/blogs/news/chiefeditor/2010/<br />
03/madagascar-rosewood-ban-reaction.html. Downloaded on<br />
31 March 2010.<br />
Brown,K.A.;Gurevitch,J.2004.Long-term impacts of logging<br />
on forest diversity in Madagascar. Proceedings of the National<br />
Academy of Sciences 101: 6045-6049.<br />
Butler, R. A. 2010. How to end Madagascar’s logging crisis.<br />
news.mongabay.com/2010/0211-madagascar.html. Downloaded<br />
on 10 February 2010.<br />
Cochrane,M.A.;Schulze,M.D.1998.Forest fires in the Brazilian<br />
Amazon. Conservation Biology 12: 948-950.<br />
Cocks, T. 2005. Loggers cut madagascan rainforest with impunity.<br />
Reuters. July 4.<br />
Débois, R.2009. La fièvre de l’or rouge saigne la forêt malgache.<br />
Univers Maoré 13: 8-<strong>15</strong>.<br />
Du Puy, D. J.; Labat, J.-N.; Rabevohitra, R.;Villiers, J.-F.;Bosser,<br />
J.; Moat, J. 2002. The Leguminosae of Madagascar. Royal<br />
Botanic Gardens, Kew, U.K.<br />
Furones, L. 2006. Independent forest monitoring: Improving<br />
forest governance and tackling illegal logging and corruption.<br />
Trócaire Development Review 135-148.<br />
Gerety, R.M. 2010. Major international banks, shipping companies,<br />
and consumers play key role in Madagascar’s logging<br />
crisis.<br />
news.mongabay.com/2009/12<strong>15</strong>-rowan_madagascar.html.<br />
Downloaded on 16 December 2010.<br />
Gill, C. 2010. Log Jam. Guitar Aficionado. Spring Issue.<br />
Gillies, A.C.M. 1999. Genetic diversity in Mesoamerican populations<br />
of mahogany (Swietenia macrophylla), assessed<br />
using RAPDs. Heredity 83: 722-732.<br />
Global Witness and Environmental Investigation Agency.<br />
2009.Investigation into the illegal felling,transport and export<br />
of precious wood in SAVA Region Madagascar. Unpublished<br />
report to the Government of Madagascar.
Page 6 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
www.illegal-logging.info/uploads/madagascarreportrevi<br />
sedfinalen.pdf. Downloaded on 20 November 2010.<br />
Irwin, M. T. 2006. Ecological impacts of forest fragmentation<br />
on diademed sifakas (Propithecus diadema) at Tsinjoarivo,<br />
Eastern Madagascar:Implications for conservation in fragmented<br />
landscapes. Ph.D. thesis, Stony Brook University,<br />
New York, USA.<br />
IUCN. 2007. World heritage nomination. IUCN technical<br />
evaluation. Rainforests of the Atsinanana (Madagascar).<br />
IUCN Evaluation Report. ID No. 1257.<br />
IUCN. 2010. IUCN Red List of Threatened Species. Version<br />
3.1. www.iucnredlist.org. Downloaded on 25 May 2010.<br />
Keong,C.H.2006.The role of CITES in combating illegal logging:<br />
Current and Potential. Traffic Online Report Series,<br />
No. 13. www.illegal-logging.info/item_single. php? it_id=<br />
504&it=document. Downloaded on 20 November 2010.<br />
Kett, G. 2005. Checking the reserve. Monthly from Madagascar.<br />
March. Madagascar Fauna Group.<br />
Labat, J.N.; Moat, J. 2003. Leguminosae (Fabaceae). Pp. 346-<br />
373. In: S.M. Goodman; J.P. Benstead (eds.) The Natural<br />
History of Madagascar. University of Chicago Press, Chicago,<br />
USA.<br />
Michaels, S. 2009. Gibson guitars raided for alleged use of<br />
smuggled wood. www.guardian.co.uk/music/2009/nov/<br />
20/gibson-guitars-raided. Downloaded on 20 November<br />
2009.<br />
Nielsen, L.R.; KjFr, E.D. 2008. Tracing timber from forest to<br />
consumer with DNA markers. Danish Ministry of the<br />
Environment, Forest and Nature Agency.<br />
www.skovognatur.dk/udgivelser. Electronic Publication.<br />
Office of the United States Trade Representative. 2010. US-<br />
China trade facts. www.ustr.gov/countries-regions/china.<br />
Downloaded on 23 May 23 2010.<br />
Patel, E.R. 2007. Logging of rare rosewood and pallisandre<br />
(Dalbergia spp.) within Marojejy National Park, Madagascar.<br />
Madagascar Conservation and Development 2(1):<br />
11-16.<br />
www.erikpatel.com/Logging_of_Rosewood_Patel_2007.pdf.<br />
Electronic Publication.<br />
Patel, E.R. In press. A tragedy with villains: Severe resurgence<br />
of selective rosewood logging in Marojejy National Park<br />
leads to temporary park closure. Lemur News.<br />
Patel,E.R.;Rasarely,E.;Tegtmeter,R.;Furones,N.;Fritz-Vietta,<br />
N.; Malan, S.; Waeber, P. In Prep. Beyond Ecological Monitoring:<br />
A proposal for "Independent Forest Monitoring"<br />
in Madagascar. Madagascar Conservation and Development.<br />
Randriamalala,H.;Liu,Z.In press.Bois de rose de Madagascar:<br />
Entre democratie et protection. Madagascar Conservation<br />
and Development.<br />
Ratsimbazafy, J. 2006. Diet composition, foraging, and feeding<br />
behavior in relation to habitat disturbance: Implications<br />
for the adaptability of ruffed lemurs (Varecia v.editorium)<br />
in Manombo forest, Madagascar. Pp. 403-422. In L. Gould;<br />
M.L.Sauther,(eds.) Lemurs:ecology and adaptation.Springer,<br />
New York.<br />
Rubel, A.; Hatchwell, M.; Mackinnon, J.; Ketterer, P. 2003. Masoala–L’oeil<br />
de la Forêt. Zoo Zurich.<br />
Schuurman, D.; Lowry, P.L. 2009. The Madagascar rosewood<br />
massacre. Madagascar Conservation and Development<br />
4(2): 98-102. www.mwc-info.net. Electronic Publication.<br />
Schuurman,D.2009.Illegal logging of rosewood in the rainforests<br />
of northeast Madagascar. TRAFFIC Bulletin 22(2):<br />
49.<br />
Stasse, A. 2002. La Filière Bois de Rose. Région d’Antalaha –<br />
Nord-est de Madagascar. Thèse de mastère non publiée,<br />
Université de Montpellier, France.<br />
Stephenson, P.J. 1993. The small mammal fauna of Reserve<br />
Speciale d’Analamazaotra, Madagascar: The effects of<br />
human disturbance on endemic species diversity. Biodiversity<br />
and Conservation 2: 603-6<strong>15</strong>.<br />
Wasser, S.; Poole, J.; Lee, P.; Lindsay, K.; Dobson, A.; Hart, J.;<br />
Douglas-Hamilton, I.; Wittemyer, G.; Granli, P.; Morgan, B.;<br />
Gunn,J.;Alberts,S.;Beyers,R.;Chiyo,P.;Croze,H.;Estes,R.;<br />
Gobush,K.;Joram,P.;Kikoti,A.;Kingdon,J.;King,L.;Macdonald,<br />
D.; Moss, C.; Mutayoba, B.; Njumbi, S.; Omondi, P.;<br />
Nowak, K. 2010. Elephants, ivory, and trade. Science 327<br />
(5971): 1331-1332.<br />
Wilmé,L.;Schuurman,D.;Lowry II,P.P.In Press.A forest counterpart<br />
fund:Madagascar’s wounded forests can erase the<br />
debt owed to them while securing their future, with support<br />
from the citizens of Madagascar. Lemur News.<br />
Wilmé, L.;Schuurman,D.; Lowry II, P.P.; Raven, P.H. 2009. Precious<br />
trees pay off – but who pays? Poster prepared for<br />
the World Forestry Congress in Buenos Aires,Argentina.<br />
www.mwc-info.net/en/services/Journal_PDF%27s/<br />
Issue4-2/MCD_2009_vol4_iss2_rosewood_massacre_<br />
Supplementary_Material.pdf. Downloaded on 23 October<br />
2009.<br />
Young, D. 2007. Independent forest monitoring: Seven years<br />
on. International Forestry Review 9(1): 563-574.<br />
Rosewood logging photos<br />
Photographer Toby Smith:<br />
www.telegraph.co.uk/culture/photography/7625511/<br />
Madagascar-undercover-slideshow.html<br />
Photographer Chris Maluszynsk:<br />
www.photoshelter.com/c/moment/gallery/ Rosewood-loggingin-Madagasar-by-Chris-<br />
Maluszynski/ G0000JWMAJa78LJ0/<br />
Rosewood logging films<br />
Dan Rather Reports:Treasure Island.Episode 437.A detailed<br />
investigation of the impact of the recent political crisis in<br />
Madagascar on the unique biodiversity of this island continent.<br />
Filmed in high-definition, active rosewood logging<br />
camps are shown. The impact of such habitat disturbance on<br />
the silky sifaka and the World Heritage Sites of Marojejy NP<br />
and Masoala NP are discussed. The debates surrounding the<br />
Ambatovy nickel mine adjacent to Andasibe-Mantadia NP<br />
are also discussed. The mine may be endangering one of the<br />
rarest animals on earth,the greater bamboo lemur (Prolemur<br />
simus) which is being protected there by the NGO Mitsinjo.<br />
Aired on HD-NET cable television November 2009. Purchasable<br />
and downloadable on I-Tunes in the United States.<br />
DVDs can be purchased online:<br />
hdnet-store.stores.yahoo.net/danrare437.html<br />
Sample Clip 1:<br />
www.facebook.com/video/video.php?v= 600388589544<br />
Sample Clip 2: www.youtube.com/watch?v= dEi-yRlJ-mk<br />
Carte Blanche: Madagascar (Part 1 and Part 2). Two short<br />
films examining illegal rosewood logging in Madagascar and<br />
the impact on the critically endangered silky sifaka. They<br />
were produced by Neil Shaw and commissioned and funded<br />
by Carte Blanche which is one of the most respected television<br />
news programs in the Southern Hemisphere. Aired on<br />
South African Television in April,2010,and streams freely online<br />
here:<br />
Carte Blanche: Madagascar Part 1:<br />
beta.mnet.co.za/carteblanche/Article.aspx?Id= 3919&ShowId=1<br />
Carte Blanche: Madagascar Part 2:<br />
beta.mnet.co.za/mnetvideo/browseVideo.aspx?vid=25570<br />
506: Bois de Rose. A Documentary Film by Joseph Areddy.<br />
2003. RSI, Comano/Signe, Genve/GAP, Antananarivo.<br />
Rosewood logging videos<br />
Madagascar Rainforest Massacre (English):<br />
www.youtube.com/watch?v=FzWNPHBRrAc<br />
Madagascar Rainforest Massacre (French):<br />
www.youtube.com/watch?v=KtjmFWpGNKs&feature=related<br />
Madagascar Rainforest Massacre (Malagasy):<br />
www.youtube.com/watch?v=rHYYhhLHeQw&feature=related
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 7<br />
Global Witness – Environmental Investigation Agency - Illegal<br />
logging in Madagascar – Part 1<br />
www.youtube.com/watch?v=T1hPviSbRcU<br />
Global Witness – Environmental Investigation Agency - Illegal<br />
logging in Madagascar – Part 2<br />
www.youtube.com/watch?v=LBtsNBpWW0E<br />
Global Witness – Environmental Investigation Agency - Illegal<br />
logging in Madagascar – Part 3<br />
www.youtube.com/watch?v=payUUJed0dc<br />
Global Witness – Environmental Investigation Agency - Illegal<br />
logging in Madagascar – Part 4<br />
www.youtube.com/watch?v=lm6a6Hrat3o<br />
Rosewood logging radio programs<br />
BBC World Service – Africa. September 17, 2009.<br />
www.bbc.co.uk/worldservice/africa/2009/09/090917_<br />
madge_rosewood2.shtml<br />
Ongoing threats to lemurs and their habitat<br />
inside the Sahamalaza - Iles Radama<br />
National Park<br />
Melanie Seiler 1,2, Guy H. Randriatahina 3, Christoph<br />
Schwitzer 1*<br />
1Bristol Conservation and Science Foundation, Bristol Zoo<br />
Gardens, Clifton, Bristol BS8 3HA, UK<br />
2University of Bristol, School of Biological Sciences, Woodland<br />
Road, Bristol BS8 1UG, UK<br />
3Association Européenne pour l’Etude et la Conservation<br />
des Lémuriens (AEECL), Lot: IVH 169 N Ambohimanandray,<br />
Ambohimanarina, Antananarivo 101, Madagascar<br />
*Corresponding author: cschwitzer@bcsf.org.uk<br />
The Sahamalaza - Iles Radama National Park,officially inaugurated<br />
in July 2007 and managed by Madagascar National<br />
Parks (MNP), includes both marine and terrestrial ecosystems<br />
and is the first park that was created under the<br />
"Programme Environnemental III" of the Malagasy government<br />
and the World Bank. In addition to the few remaining<br />
forest fragments of the Southern Sambirano ecoregion, the<br />
park is home to extensive mangrove forests, which harbour<br />
their own highly endangered fauna, and also includes offshore<br />
coral reefs. In 2003, researchers from the Cologne<br />
Zoo, funded by AEECL, undertook an expedition to Sahamalaza<br />
to explore the opportunities for the establishment of<br />
a permanent field station in order to study and protect the<br />
Critically Endangered blue-eyed black lemur (Eulemur flavifrons)<br />
and its habitat.In 2004 and 2005,the field station in the<br />
Ankarafa Forest became reality (Schwitzer et al.,2006),and it<br />
has since been used by both European and Malagasy scientists<br />
as a basis for research on E. flavifrons and other lemur<br />
species, especially the Sahamalaza sportive lemur (Lepilemur<br />
sahamalazensis) and the northern giant mouse lemur (Mirza<br />
zaza),occurring on the Sahamalaza Peninsula (Schwitzer and<br />
Randriatahina, 2009).<br />
Sahamalaza - Iles Radama National Park lies within a transition<br />
zone between the Sambirano region in the north and<br />
the western dry deciduous forest region in the south, harbouring<br />
semi-humid forests with tree heights of up to 30m<br />
(Schwitzer et al.,2006).The forests include a mixture of plant<br />
species typical of both domains (Birkinshaw, 2004), and the<br />
remaining primary and secondary forest fragments vary in<br />
their degree of degradation. There are no larger connected<br />
areas of intact primary forest left on the Sahamalaza Penin-<br />
sula, and the remaining fragments all show some degree of<br />
anthropogenic disturbance and/or edge effects (Schwitzer et<br />
al., 2007). The forests and forest fragments are separated by<br />
grass savannah and shrubs. Sahamalaza is the only protected<br />
area that harbours the blue-eyed black lemur,the Sahamalaza<br />
sportive lemur and the northern giant mouse lemur. Other<br />
lemur species in the park include the aye-aye (Daubentonia<br />
madagascariensis), the western bamboo lemur (Hapalemur<br />
occidentalis),and an as yet unidentified species of dwarf lemur<br />
(Cheirogaleus spec.).<br />
The remaining forest of the Sahamalaza Peninsula and its<br />
unique fauna are in grave danger of disappearing.The habitat<br />
is already extremely degraded, nonetheless bush fires and<br />
tree-felling are activities that are routinely pursued and accepted<br />
within the local society (Ruperti et al., 2008). During<br />
the first field season of a study on the impact of habitat degradation<br />
and fragmentation on the ecology and behaviour of<br />
the Sahamalaza Peninsula sportive lemur (Lepilemur sahamalazensis),<br />
conducted by MS in 2009, local people from the<br />
villages surrounding the protected area were found logging<br />
trees in the already small forest fragments almost on a daily<br />
basis. Logging activities mainly occurred in forest fragments<br />
where no researchers had been present in previous years.<br />
During walks through different forest fragments, in addition<br />
to large numbers of logged trees, two places where trees<br />
were processed for further use were found. Trees were<br />
felled mainly in the early morning hours,on the one hand because<br />
of the high temperatures later in the day, on the other<br />
hand probably because of the assumption that the researchers<br />
started observing animals later in the day and therefore<br />
would not realise the illegal logging activities. Nonetheless,<br />
trees were sometimes also felled in the afternoons. Because<br />
locals immediately fled when becoming aware of researchers’<br />
presence, we believe that the presence of researchers<br />
and/or field guides, park authorities or park rangers is a crucial<br />
factor in stopping illegal logging in the remaining fragments.For<br />
the next field season (2010) we therefore plan to<br />
expand the observations of Lepilemur to other, not yet used<br />
forest fragments to help prevent their destruction. Of<br />
course this cannot be a long-term solution to this problem.<br />
The presence of park rangers and further environmental education<br />
of the local people will thus be extremely important<br />
to save the Sahamalaza forests from further degradation.<br />
About five times between August and October 2009, fires<br />
occurred near the Ankarafa field station, three times in the<br />
savannah and twice in the forest itself. After having extin-<br />
Fig. 1: Lepilemur sahamalazensis<br />
poached<br />
and roasted by locals in<br />
Sahamalaza - Iles Radama<br />
National Park“.
Page 8 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
guished these fires it became obvious that they had all<br />
started right beside the fire breaks that are frequently used<br />
as paths by people on their way between villages. The Ankarafa<br />
field guides, all of them locals from the surrounding villages,<br />
assumed that the fires were set by villagers to show<br />
their dissatisfaction with the recently established national<br />
park that prohibits the use of the forests for collecting building<br />
material for their dwellings. As we followed the smoke<br />
that was coming from another fire, we found an area inside<br />
one of the core zones of the national park that was inhabited<br />
by a young couple. They harvested a rice field and regularly<br />
burned undergrowth around it. Additionally, they kept cattle<br />
and goats and had built 2 houses at this site,one for the cattle<br />
and one for themselves. As we talked to them, they claimed<br />
that they were allowed to stay on this site and that MNP had<br />
sold this part of the forest to them.They affirmed that,if they<br />
set fire on this site, they would keep an eye on it and would<br />
prevent the fire from expanding into the forest. Unfortunately<br />
this was not the case,however,as we later observed a<br />
fire around this site without anyone near it. Overall, it<br />
seemed that there were various people living inside the national<br />
park on permits given to them by what they claimed to<br />
have been MNP agents;we were told that there was a map of<br />
the park showing all the "excluded" areas available for housing<br />
and agriculture, which could be seen in the village of<br />
Marovato. If that was indeed the case (we did not have the<br />
opportunity to verify the information),it would be a massive<br />
problem for protecting Sahamalaza’s unique wildlife and forests.<br />
If people claiming to be MNP staff illegally sold permits<br />
for activities inside the national park, the destruction of the<br />
small forest fragments will continue rapidly.<br />
Another big problem comes with cattle; every day zebu cattle<br />
were observed in all forest fragments and on the savannah<br />
in Ankarafa, as people from nearby villages let their cattle<br />
roam freely.The abundance of zebu themselves and their excrements<br />
indicated that they frequently used the forest fragments<br />
as grazing grounds, especially those with remaining<br />
primary forest parts. When zebu were grazing in the forest<br />
rather than on the savannah, their movements were accompanied<br />
by crashing and breaking sounds;they were undoubtedly<br />
hindering the growth of many saplings,if not eating them.<br />
This is an additional threat to the forest fragments, and furthermore,<br />
the abundance of the excrements of local zebu<br />
has been found to negatively correlate with the density of L.<br />
sahamalazensis (Ruperti, 2007). Additionally, the introduced<br />
bush pig is responsible for considerable habitat destruction<br />
due to digging up large areas, thus hindering the growth of<br />
saplings. Unfortunately, the bush pig is reproducing wildly as<br />
it is regarded as fady (taboo) by the local people and therefore<br />
not hunted.<br />
Not only the activities of local people seem to be a threat to<br />
the endangered wildlife on the Sahamalaza Peninsula.One of<br />
the Ankarafa field guides encountered a foreigner,probably a<br />
resident living in Madagascar (since he spoke Malagasy fluently),<br />
with a 4x4 car and two local guides about 1 km from<br />
the researchers’ camp. These people had set up a tent and<br />
told the Ankarafa field guide that they were visiting all villages<br />
on the Sahamalaza Peninsula to look for fish. As we checked<br />
their camp site the next day, the three men were gone, but<br />
signs of a fire,logged branches and feathers of a harrier hawk<br />
were found,indicating that they had caught and killed this endangered<br />
bird of prey. We wrote a report about this event<br />
and handed it over, together with feathers of the bird, to<br />
MNP in Maromandia. However, as long as there are no signs,<br />
borders or fences indicating the national park area and its<br />
restrictions, these problems will continue.<br />
The ongoing political crisis is a further big concern that hinders<br />
the effective protection not only of the biodiversity of<br />
the Sahamalaza Peninsula, but of Madagascar and its national<br />
parks system as a whole. Only 10 years ago, Madagascar was<br />
notorious for its environmental degradation and deforestation,<br />
but that began to change in 2003 when then President<br />
Marc Ravalomanana, working with international conservation<br />
organizations and local groups, set aside 10 % of the<br />
country’s surface area as national parks and started supporting<br />
ecotourism, which slowed deforestation and helped to<br />
safeguard biodiversity.After the political events in early 2009<br />
that saw the ousting of the President and the installation of a<br />
transitional government, the majority of donor funds, which<br />
provided half the government’s annual budget, have been<br />
withdrawn, leading to major funding gaps that have affected<br />
protected areas and their management. There currently is<br />
almost no money to employ park rangers or to implement<br />
other measures to protect the forests inside Madagascar’s<br />
national parks, and forest degradation is going on without<br />
noticeable resistance from the relevant authorities. Despite<br />
the political crisis that affects most of the social and environmental<br />
activities of numerous NGOs, AEECL is still carrying<br />
out its research activities and support to the villagers surrounding<br />
the Sahamalaza - Iles Radama National Park. Since<br />
the establishment of the protected area in 2007, AEECL has<br />
been conducting, besides its research programme, different<br />
projects that aim to reduce the excessive environmental exploitation<br />
inside and around the park.As the major activity of<br />
the local population surrounding the Sahamalaza Peninsula<br />
National Park is rice-growing,every year AEECL organizes a<br />
rice-growing training course and rice-growing competition,<br />
using modern techniques in order to increase yield per ha<br />
and to decrease the use of slash and burn agriculture.To stop<br />
the ongoing overexploitation of the environment, environmental<br />
education is another important part of AEECL´s<br />
work.As many villages in Sahamalaza are unable to pay teachers,<br />
AEECL subsidizes teachers’ salaries to ensure the primary<br />
education of the local children. Additionally, leaflets<br />
about the Sahamalaza biodiversity and its importance are<br />
distributed. They inform and educate villagers about the importance<br />
of lemurs and other species for their forest ecosystem.<br />
To minimise bush fires and to protect the forest against<br />
uncontrolled fires, AEECL organizes firebreak programs<br />
around the Ankarafa Forest, close to the research camp,<br />
where during three days, hundreds of local people remove<br />
the grasses on a 7m wide strip around the forest fragments.<br />
Furthermore, several reforestation campaigns have been<br />
conducted, where villagers, including many teachers and<br />
their pupils,have planted trees around their villages with the<br />
help of AEECL.<br />
Because of all the factors described here, the protection of<br />
Sahamalaza’s unique flora and fauna continues to be a major<br />
challenge that has to be faced by the local human population<br />
with the help of Madagascar National Parks and foreign partners.<br />
Two essential parts of AEECL’s efforts to help meeting<br />
this challenge are to stimulate further scientific study of<br />
endangered lemurs and other wildlife at its research station<br />
in the Ankarafa Forest, especially by Malagasy students, and<br />
to enable the local human population around the Sahamalaza<br />
- Iles Radama National Park to sustainably use their natural<br />
resources.<br />
Acknowledgements<br />
We would like to thank Madagascar National Parks (MNP),<br />
especially the director of Sahamalaza - Iles Radama National<br />
Park, M. ISAIA Raymond, for their continuing collaboration.
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 9<br />
Thank you also to the DGEF and CAFF/CORE for granting<br />
us research permits for our work in Sahamalaza,and to Prof.<br />
RABARIVOLA Clément for his ongoing help. Tantely Ralantoharijaona<br />
and Bronwen Daniel,along with all Ankarafa field<br />
guides, contributed substantially to fighting forest fires and<br />
other environmental threats in Ankarafa in 2009. MS was<br />
funded by Bristol Conservation and Science Foundation,<br />
AEECL, Conservation International Primate Action Fund,<br />
Margot Marsh Biodiversity Foundation, Mohamed bin Zayed<br />
Species Conservation Fund,International Primatological Society<br />
and Christian-Vogel-Fonds.<br />
References<br />
Birkinshaw, C.R. 2004. Priority areas for plant conservation.<br />
Ravintsara 2(1): 14-<strong>15</strong>.<br />
Ruperti, F. 2007. Population density and habitat preferences<br />
of the Sahamalaza sportive lemur (Lepilemur sahamalazensis)<br />
at the Ankarafa research site, NW Madagascar.<br />
Unpublished MSc thesis, Oxford Brookes University, UK.<br />
82 p.<br />
Ruperti,F.;Smith,J.;Ratovonasy,L.;Thorn,J.2008.Sahamalaza<br />
Conservation Action Plan (SCAP).Unpublished report to<br />
the Association Européenne pour l’Etude et la Conservation<br />
des Lémuriens (AEECL). 17 p.<br />
Schwitzer, C.; Randriatahina, G.H. 2009. AEECL: Update on<br />
activities. Lemur News 14: 11-12.<br />
Schwitzer, N.; Randriatahina, G.H.; Kaumanns, W.; Hoffmeister,<br />
D.; Schwitzer, C. 2007. Habitat utilization of blue-eyed<br />
black lemurs,Eulemur macaco flavifrons (Gray,1867),in primary<br />
and altered forest fragments.Primate Conservation<br />
22: 79-87.<br />
Schwitzer, C.; Schwitzer, N.; Randriatahina, G.H.; Rabarivola,<br />
C.; Kaumanns, W. 2006. "Programme Sahamalaza": New<br />
perspectives for the in situ and ex situ study and conservation<br />
of the blue-eyed black lemur (Eulemur macaco flavifrons)<br />
in a fragmented habitat.Pp.135-149.In:C.Schwitzer;<br />
S.Brandt;O.Ramilijaona;M.Rakotomalala Razanahoera;D.<br />
Ackermand; T. Razakamanana; J. U. Ganzhorn (eds.). Proceedings<br />
of the German-Malagasy Research Cooperation<br />
in Life and Earth Sciences. Berlin: Concept Verlag.<br />
News and Announcements<br />
Madagascar conservationist wins international<br />
environmental prize<br />
Mr Rabary Desiré has been awarded the 2010 Seacology<br />
Prize (www.seacology.org/prize/index.htm) for his his tireless<br />
efforts to further forest conservation in northeastern Madagascar.Mr<br />
Desiré will receive the US$10,000 Prize on October<br />
7, 2010 at a ceremony in Berkeley, California.<br />
Rabary Desiré is recognized by many as a major conservation<br />
leader in northeastern Madagascar, and is a highlysought-after<br />
research and eco-tourism guide. With the money<br />
he makes from guiding,he buys forested land in order to<br />
protect it.Years of work have finally culminated in the establishment<br />
of his own small private nature reserve called<br />
Antanetiambo (antanetiambo.marojejy.com/Intro_e.htm),<br />
which means "on the high hill". It is perhaps the only reserve<br />
in northern Madagascar that has been entirely created from<br />
start to finish by a single local resident.<br />
According to Mr Desiré, "I am very happy to receive this<br />
award and I feel very lucky for myself and Madagascar. After<br />
many years of hard work and political instability,finally we are<br />
having some local conservation success. I plan to use these<br />
funds for such projects as reforestation, developing tourist<br />
Fig. 1: Rabary Desiré next to the sign for the Antanetiambo<br />
Nature Reserve he created.<br />
infrastructure and purchasing the land around Antanetiambo<br />
Nature Reserve to increase the size of the reserve and the<br />
amount of protected land in this region. This award will help<br />
preserve the precious biodiversity and high endemism of<br />
Madagascar,as well as fight the ongoing battle against massive<br />
deforestation and possible extinction of many beloved species...<br />
Thanks Seacology for giving me this prize. The whole<br />
region will never forget it."<br />
Read the full press release:<br />
www.seacology.org/news/display.cfm?id=4238<br />
Célébration du quinzième anniversaire<br />
du GERP (1994-2009)<br />
Jonah Ratsimbazafy*, Rose Marie Randrianarison,<br />
Muriel Nirina Maeder<br />
GERP, 34, Cité des Professeurs, Antananarivo 101,<br />
Madagascar<br />
*Corresponding author: gerp@wanadoo.mg<br />
Quinze ans se sont écoulés depuis la création, en 1994, de la<br />
Société de Primatologie malgache ou Groupe d’Etude et de<br />
Recherche sur les Primates de Madagascar (GERP). Elle fut<br />
fondée par dix Primatologues dont le Professeur Berthe<br />
Rakotosamimanana qui occupait à la fois le poste de Secrétaire<br />
Général du GERP et le Co-éditeur de la revue Lemur<br />
News jusqu’à sa disparition en 2005. De son vivant, elle<br />
désirait ardemment passer le flambeau au Docteur Jonah<br />
Ratsimbazafy pour le poste de Secrétaire Général du GERP<br />
qui, en 2006, a été mandaté à l’unanimité par les membres<br />
nationaux et internationaux du GERP au titre de Leader du<br />
GERP.<br />
L’Association compte aujourd’hui 169 membres et 20 d’entre<br />
eux sont de nationalité étrangère. La multidisciplinarité<br />
des membres du groupe (Primatologues, Anthropologues,<br />
Paléontologues, Ornithologues, Herpétologues, Spécialistes<br />
de Micromammifères et Mammifères, Parasitologistes, Botanistes,<br />
Géographes, Vétérinaires, Agro-forestiers, Biochimis-
Page 10 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
tes, Dessinateur, Financiers) apporte une importante potentialité<br />
dans l’accomplissement de la mission du GERP: transférer<br />
les compétences nécessaires à la préservation de la<br />
biodiversité pour les générations futures. Par ailleurs, les actions<br />
du GERP comprennent également la formation des<br />
pépinières de Primatologues, la mise en œuvre du plan de<br />
conservation des lémuriens, la contribution à l’amélioration<br />
des activités génératrices de revenu des communautés de<br />
base liées à la conservation,sans oublier l’éducation environnementale<br />
de la population cible.<br />
En 2007,l’attribution par le GERP du nom de Microcebus macarthurii<br />
à une nouvelle espèce découverte dans la forêt de<br />
Makira représentait un témoignage de reconnaissance au<br />
dévouement de la Fondation MacArthur. De plus, le GERP a<br />
depuis 2008 officiellement été mandaté par le MEFT/DGEF/<br />
DSAP comme Gestionnaire de la forêt de Maromizaha,pour<br />
que cette dernière devienne une Nouvelle Aire Protégée<br />
(NAP).Plus récemment encore,en février 2010,le prix "lifetime"<br />
décerné par l’IPS a été attribué à un membre scientifique<br />
du GERP en la personne du Docteur Alison Jolly.<br />
A l’occasion de son quinzième anniversaire, le GERP aura<br />
l’honneur d’organiser une conférence scientifique sur les<br />
lémuriens, à Antananarivo en novembre 2010.<br />
Conservation International’s Primate<br />
Action Fund:Projects funded March 2009<br />
to March 2010<br />
Anthony Rylands<br />
Conservation International, 2011 Crystal Drive, Suite 500,<br />
Arlington, VA 22202, USA, a.rylands@conservation.org<br />
Conservation International’s Primate Action Fund awards<br />
small grants (up to $5,000) to support projects and initiatives<br />
promoting the conservation of primates worldwide,focusing<br />
on Critically Endangered and Endangered species in their<br />
natural habitats (and most especially those included in the biennial<br />
listing of the World’s 25 Most Endangered Primates).<br />
Projects should contribute to at least one of the following<br />
themes: (1) enhancement of scientific understanding/knowledge<br />
of the target species/ecosystem; (2) improved protection<br />
of a key species, habitat, or a reserved area; (3) demonstration<br />
of economic benefits achieved through conservation<br />
of a species and its habitat,as compared to its loss;(4) increased<br />
public awareness or educational impact resulting<br />
from the project in question; (5) improved local capacity to<br />
carry out future conservation efforts through training or<br />
practical experience obtained through project participation;<br />
and (6) modification of inappropriate policies or legislation<br />
that previously led to species or habitat decline. Awards are<br />
given most frequently for population and distribution surveys,and<br />
ecological and behavioral studies pertinent to conservation<br />
initiatives for threatened species. Grants are also<br />
given that support genetic and taxonomic studies, publications,<br />
workshops for action plans and suchlike, and primate<br />
field courses. Some awards are given to help primate habitat-country<br />
primatologists attend the biennial congresses of<br />
the International Primatological Society. The fund does not<br />
support participation in academic courses.<br />
The Primate Action Fund comes from an annual award to<br />
Conservation International, Arlington, Virginia, USA, made<br />
by the Margot Marsh Biodiversity Foundation. It is managed<br />
jointly by Ms Ella Outlaw and Dr Anthony B.Rylands,both of<br />
CI’s Office of the President. Guidelines for application can be<br />
obtained by writing to Anthony Rylands (see Funding and<br />
Training section in this issue).<br />
Five grants were awarded to benefit lemur conservation in<br />
the March 2009 – March 2010 funding cycle. They were as<br />
follows: (1) Halting politically-induced deforestation in the<br />
short term to preserve the unique primate community of<br />
Tsinjoarivo, eastern central Madagascar–Mitchell T. Irwin,<br />
Fanomezantsoa, Jean-Luc Raharison and Marina Blanco; (2)<br />
Rapid survey and assessment of the northern sportive lemur,<br />
Lepilemur septentrionalis, in the Sahafary Region, Madagascar–Edward<br />
Louis Jr, Jean<br />
Ranaivoarisoa, John Zaonarivelo and Steig Johnson; (3) Support<br />
for the publication of the IUCN/SSC Primate Specialist<br />
Group newsletter and journal Lemur News, volume 14–Jörg<br />
U. Ganzhorn and Christoph Schwitzer; (4) Student training<br />
course “Field Methods in the Study of Primate Behavior and<br />
Ecology”,Kirindy forest,2010–Melanie Dammhahn,Peter M.<br />
Kappeler, Claudia Fichtel, Cornelia Kraus and Rodin Rasoloarison;<br />
and (5) Comparison of habitat requirements of the<br />
Data Deficient northern giant mouse lemur (Mirza zaza) in<br />
two differently degraded habitats, in Sahamalaza,northwestern<br />
Madagascar–Johanna Rode and Christoph Schwitzer.<br />
International Technical Meeting on Prolemur<br />
simus, 26-28 January 2010, Antananarivo,<br />
Madagascar<br />
The greater bamboo lemur Prolemur simus has long been considered<br />
to be one of the rarest primate species in the world.<br />
Up to 2007 only 60 individuals were known from the wild,<br />
and another 22 were in captivity (Wright et al.,2008;Primate<br />
Conservation 23: 5-17). Once widespread across Madagascar,<br />
more recent confirmed sightings were exclusively from<br />
south-eastern Madagascar, which led to the assumption that<br />
the species was extinct on the rest of the island. In 2008,<br />
Dolch et al. (Lemur News 13: 14-17) rediscovered P. simus in<br />
the Torotorofotsy wetlands, north of the Mangoro River.<br />
Since then, several extensive surveys have been conducted<br />
north and south of the Mangoro, and evidence of greater<br />
bamboo lemurs was found at several sites in the Ankeniheny-Zahamena<br />
Corridor, in the central region of the eastern<br />
rainforest (King and Chamberlan, 2010; Oryx 44: 167).<br />
In the context of developing a conservation action plan for<br />
the greater bamboo lemur, the Madagascar Fauna group organised,<br />
from 26-28 January 2010 at the motel d’Antananarivo,<br />
Anosy, an international technical meeting with the<br />
theme "Conservation of the critically endangered greater<br />
bamboo lemur Prolemur simus:What we know now,what we<br />
need to know and potential conservation strategies".Several<br />
members of the PSG contributed to this.<br />
The objectives of the meeting were 1) to share information<br />
about the current situation of the various groups/populations<br />
of Prolemur simus in the wild and in captivity; 2) to discuss<br />
the threats, the solutions and the conservation strategies<br />
for three groups - north of the Mangoro River (Torotorofotsy<br />
and the Ankeniheny-Zahamena corridor CAZ),<br />
south of the Mangoro River (south-east and the Fandriana-<br />
Vondrozo corridor COFFAV), and in captivity (Madagascar<br />
and Europe); and 3) to make a plan, short to long term, to<br />
move towards a conservation action plan for the species.<br />
With 28 participants,the meeting was well attended.Presentations<br />
were given by researchers studying P. simus in the wild<br />
and in captivity, representatives from the Ambatovy, Madagascar<br />
National Parks, the University of Antananarivo and
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 11<br />
conservation NGOs.While other potential P.simus sites still<br />
need to be explored, results from the most recent surveys<br />
suggest the total estimated size of the known population is<br />
between 221-346 individuals. Another 20 individuals are<br />
housed in one Malagasy (Parc Ivoloina Zoo) and several European<br />
zoos and and managed under the umbrella of an EEP.<br />
The following recommendations for the conservation of P.<br />
simus came out of the meeting:<br />
We need to achieve official/formal protection for all currently<br />
known P. simus habitat (using whatever status is appropriate<br />
to the site);<br />
Animals of the northern and southern populations (wild<br />
or captive) should not be mixed until the taxonomic situation<br />
is clarified;<br />
Faecal samples should be collected from all sites using a<br />
standard protocol (meeting participants agree to collaborate<br />
to achieve this);<br />
When animals are caught/immobilised the opportunity<br />
should be used to maximise the collection of samples;<br />
Bamboo plot data should be collected from all sites using<br />
a standard protocol (meeting participants agree to collaborate<br />
to achieve this);<br />
A health screening protocol should be applied whenever<br />
the opportunity arises;<br />
Sites in the Ankeniheny-Zahamena Corridor (CAZ) recently<br />
shown to harbour P. simus should be evaluated by<br />
2011 at the latest to assess population size;<br />
Maromiza and Lakato need to be evaluated for the presence<br />
of P. simus, and protected to ensure connectivity;<br />
We agree that assuring connectivity between Torotorofotsy<br />
and CAZ is a high priority,and that the area needs an<br />
integrated conservation plan involving all stakeholders –<br />
CI to drive the process under supervision of the Alaotra-<br />
Mangoro Forestry Commission;<br />
It is important to make P. simus a priority (conservation<br />
target) for the CAZ;<br />
Improved communication using a mailing list will be established,<br />
the "Prolemur Conservation Working Group";<br />
There are other sites that need to be surveyed for P.simus<br />
(a list of sites has already been identified);<br />
Maximising connectivity between P. simus sites is important;<br />
Local communities should be directly involved in P. simus<br />
conservation wherever possible;<br />
In case of a crisis scenario involving potential translocation,<br />
a technical strategy is needed consistent with IUCN<br />
guidelines;<br />
The EEP-Ivoloina exchange of P. simus is important to<br />
strengthen the global captive population;<br />
For the time being,it is not recommended that additional<br />
wild P. simus be added to the global captive programme,<br />
except in emergency;<br />
In the case of emergency, we recommend that animals go<br />
to PBZT if upgraded facilities have been installed; if not<br />
then they should go to Ivoloina;<br />
Based on the development of the global captive programme,<br />
integrated (metapopulation) management of P. simus<br />
should be considered;<br />
Another technical meeting should be held in January<br />
2011.<br />
The workshop was financially and technically supported by<br />
the Madagascar Fauna Group with additional contributions<br />
from Conservation International Madagascar.<br />
Lemur presentations at the 23rd Congress<br />
of the International Primatological<br />
Society, Kyoto, Japan<br />
Jonah Ratsimbazafy<br />
GERP, 34, Cité des Professeurs, Antananarivo 101, Madagascar,<br />
gerp@wanadoo.mg<br />
The 23rd Congress of the International Primatological Society<br />
(IPS) was held in Kyoto (Yoshida Main Campus),Japan on<br />
12th-18th September, 2010. This congress brought together<br />
more than 1,000 delegates from 56 countries. Twenty-eight<br />
talks and three posters were presented on lemur studies<br />
during that congress.<br />
I am also pleased to share with you the good news that the<br />
winner of the 2010 IPS Lifetime Award is Professor Alison<br />
Jolly who is an active member of GERP (Groupe d’Etude et<br />
de Recherche sur les Primates de Madagascar). The lemur<br />
lady,Prof.Jolly,has devoted her life to the conservation of the<br />
world’s primates. Education is one of the main activities that<br />
she never stops to discuss,as she found that the only chance<br />
to save the endangered lemurs of Madagascar is to provide<br />
the Malagasy children with tools enabling them to learn and<br />
love the creatures that exist in their backyards.<br />
I hope that even more lemur researchers will present the<br />
results of their work at the 24th IPS Congress in Mexico.<br />
Short Communications<br />
Preliminary conservation status assessment<br />
for the Data Deficient northern<br />
giant mouse lemur Mirza zaza<br />
Eva Johanna Rode 1,2, K. Anne-Isola Nekaris 2, Christoph<br />
Schwitzer 1*<br />
1Bristol Conservation and Science Foundation, c/o Bristol<br />
Zoo Gardens, Clifton, Bristol BS8 3HA, UK<br />
2Nocturnal Primate Research Group, School of Social Sciences<br />
and Law, Oxford Brookes University, OX3 0BP, UK<br />
*Corresponding author: cschwitzer@bcsf.org.uk<br />
Madagascar is one of the world’s most important biodiversity<br />
hotspots, underpinned by its large proportion of endemic<br />
species and high rates of deforestation.During the last<br />
decade, species diversity of Madagascar’s endemic lemurs<br />
has increased dramatically due to new discoveries and taxonomic<br />
revisions. This has resulted in the unusual situation of<br />
45 % of all Malagasy primate species being Red-Listed as Data<br />
Deficient (DD) by the IUCN. This is by far the highest such<br />
figure for any primate habitat country (by comparison, 13 %<br />
of all primates and <strong>15</strong> % of all mammals are Red-Listed as<br />
DD).The lack of species-specific knowledge makes it impossible<br />
to design effective conservation measures targeting<br />
these taxa. To help assign a conservation status to the DD<br />
northern giant mouse lemur Mirza zaza, described in 2005<br />
due to distinctive features in morphology, behaviour and<br />
genetics (Kappeler et al., 2005; Primate Report, 71, 3-26), we<br />
examined space requirements and group size of this small<br />
nocturnal lemur species during a three-month study (May-<br />
July 2010) and extrapolated our results to the taxon’s area of
Page 12 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
occupancy in order to estimate the size of its remaining<br />
population.<br />
Mirza zaza lives in dry forests of north-western Madagascar,<br />
one of the fastest declining habitats of the island, with a decrease<br />
in forest cover of 40 % from 1975 to 2000.The area of<br />
occurrence of the species is limited by the Maeverano River<br />
in the south and the Mahavavy River in the north.Combining<br />
forest cover data collected by the Madagascar Vegetation<br />
Mapping Project (www.vegmad.org) with data on group home<br />
range size and group size calculated from our study and additional<br />
literature, we calculated minimum and maximum estimates<br />
of total remaining population size. Since data for the<br />
Madagascar Vegetation Mapping Project were collected several<br />
years ago,we lowered the estimate of total available habitat<br />
according to the estimated annual rate of decline.Habitat<br />
decline may have accelerated since the onset of the political<br />
crisis in Madagascar in early 2009, which is not yet reflected<br />
in our estimates.Since a previous survey failed to detect Mirza<br />
zaza in several regions within the species’ area of occurrence,<br />
we applied different estimates of the percentage of<br />
suitable habitat actually inhabited by the species.Our calculations<br />
yielded the following estimates:<br />
Maximum estimate: The total area covered in dry forest<br />
within the area of occupancy of M. zaza is approximately<br />
1,650 km 2 . Assuming an occupancy of 80 %, group home<br />
ranges of 2 ha and group size of 4 individuals there would<br />
be max. 177,500 animals left in total.<br />
Minimum estimate: In order to reflect the long-term survival<br />
of the species in a very fragmented area, only fragments<br />
< 1km 2 and smaller fragments closer than 500 m to<br />
other, larger fragments (total area: 955 km 2 ) were considered.We<br />
chose 1 km 2 to allow a minimum viable population<br />
of 250 animals.If only 30 % of the habitat is inhabited,<br />
animals use group home ranges of 4 ha and live in groups<br />
of on average 2.3 animals, this leads to an estimate of<br />
16,500 individuals left.<br />
Mirza zaza should be assessed as Vulnerable (VU B2ab) since<br />
its area of occupancy in both estimates is lower than 2,000<br />
km 2. With several sites within the species’ distribution area<br />
found to be unoccupied, the remaining habitat being extremely<br />
fragmented with the smallest fragments unsuitable<br />
to support a viable population, and habitat vanishing quickly,<br />
M. zaza may become Endangered (EN B2ab) in the near<br />
future if its area of occupancy shrinks below 500 km 2.<br />
Our preliminary conservation status assessment used the<br />
best available data for Mirza zaza. More accurate estimates<br />
will be possible if more data become available, especially on<br />
percentage of occupancy. This method might be applied to<br />
other DD lemur species in order to gain initial assessments<br />
of their conservation status.<br />
An observation of the hairy-eared dwarf<br />
lemur, Allocebus trichotis, in the Lakato<br />
region, eastern Madagascar<br />
Erwan Lagadec 1, Steven M. Goodman 2*<br />
1Centre de Recherche et de Veille sur les maladies émergentes<br />
dans l’Océan Indien (CRVOI),GIP Cyclotron Réunion<br />
Océan Indien, 2 rue Maxime Rivière, 97492 Sainte Clotilde,<br />
Ile de la Réunion, France, and Centre National de la Recherche<br />
Scientifique, UMR5557 Ecologie Microbienne, Bât A.<br />
Forel, 43 bd du 11 novembre 1918, 69622 Villeurbanne<br />
CEDEX, France<br />
2Field Museum of Natural History, 1400 South Lake Shore<br />
Drive, Chicago, Illinois 60605, USA, and Association Vahatra,<br />
BP 3972, Antananarivo 101, Madagascar<br />
*Corresponding author: sgoodman@vahatra.mg<br />
Although a few decades ago the hairy-eared dwarf lemur<br />
(Allocebus trichotis) was considered "unquestionably the rarest<br />
of surviving lemurs" (Tattersall,1982,p.131),more recent<br />
field work has found this species to be widely distributed<br />
across portions of the eastern humid forests of Madagascar<br />
(e.g., Meier and Albignac, 1991; Rakotoarison, 1998; Schütz<br />
and Goodman, 1998; Goodman and Raselimanana, 2002).<br />
Since more than a decade, there have been numerous records<br />
of this species from the central portion of the eastern<br />
humid forests,and information is now available on aspects of<br />
its ecology and natural history (e.g.Rakotoarison et al.,1997;<br />
Garbutt,2000;Biebouw,2009;Ralison,2010).Here we add an<br />
additional record from the region of Lakato, an area from<br />
where this species had not been previously recorded.<br />
From 22-28 October 2010 we were part of a research group<br />
that conducted a biological inventory of a forest block in the<br />
Lakato area and in the southern portion of the Zahamena-<br />
Ankeniheny forest corridor. The specific study site was centered<br />
at the following locality,which served as the base camp<br />
for all inventory activities: Province de Toamasina, Alaotra-<br />
Mangoro Region, 14.5 km SW of Andasibe (Périnet) village,<br />
Ampasipotsy-Anivonimaro/Ambalafary Forest, 19°02’38"S,<br />
48°20’55"E, 995 m elevation.<br />
During a nocturnal survey on 28 October 2010, the first author<br />
observed and photographed an individual of A. trichotis.<br />
The animal appeared not to be accompanied by any conspecifics.<br />
The distinctive ear-tufts, characteristic of Allocebus,<br />
were clearly visible (Fig.1).The lemur was observed at 21h58<br />
and for about five minutes.The site was in partially disturbed<br />
lower montane forest, about 200 m away from the research<br />
camp and within a few meters of the dirt road connecting the<br />
RN 2 (connecting Antananarivo-Toamasina) and the village<br />
of Lakato. The animal was not particularly active and rested<br />
in the upper portion of a 4 m tall tree. As it was photographed,<br />
including the use of flash, the individual remained<br />
largely stationary, until it finally turned and moved off into<br />
another tree and into dense vegetation.<br />
During the course of nocturnal observations of forest animals<br />
within the study site,this was the only observation of A.<br />
trichotis. Each night numerous individuals of Microcebus cf.<br />
Fig. 1: Photo of Allocebus trichotis taken during the night of 28<br />
October 2010 in a forest block approximately halfway between<br />
the turn-off of RN 2 and the village of Lakato. The<br />
ear-tufts of this animal, diagnostic of this species, are readily<br />
visible in the photo.
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 13<br />
lehilahytsara were observed in close proximity. Although A.<br />
trichotis is now known to have a broad distribution across a<br />
good portion of the eastern humid forests, from lowland to<br />
montane forests (up to about 1,000 m),it occurs in low densities<br />
(Mittermeier et al., 2006).This factor might account for<br />
its absence in other forested sites surveyed within the<br />
Zahamena-Ankeniheny forest corridor (e.g., Schmid et al.,<br />
1999; Randrianabinina and Rasoloharijaona, 2006). However,<br />
continued surveying efforts at these sites will probably result<br />
in the finding that it occurs across the forest corridor.<br />
Acknowledgements<br />
The survey of the Lakato region was financed by a grant from<br />
the <strong>Vol</strong>kswagen Foundation.We are grateful to the Département<br />
de Biologie Animale, Université d’Antananarivo and<br />
the Direction du Système des Aires Protégées, Direction<br />
Générale de l’Environnement et des Forêts for permits to<br />
conduct this research.<br />
References<br />
Biebouw,K.2009.Home range size and use in Allocebus trichotis<br />
in Analamazaotra Special Reserve, central eastern Madagascar.<br />
Int. J. Primatol. 30: 367-386.<br />
Garbutt,N.2000.Brief observations of hairy-eared dwarf lemur<br />
(Allocebus trichotis) in Analamazaotra Special Reserve,<br />
eastern Madagascar. Lemur News 6: 37.<br />
Goodman, S.M.; Raselimanana, A.P. 2002. The occurrence of<br />
Allocebus trichotis in the Parc National de Marojejy. Lemur<br />
News 7: 21-22.<br />
Meier, B.; Albignac, R. 1991. Rediscovery of Allocebus trichotis<br />
Günther 1875 (Primates) in northeast Madagascar. Folia<br />
Primatol. 56: 57-63.<br />
Mittermeier,R.A.;Konstant,W.R.;Hawkins,A.F.A.;Louis,E.E.;<br />
Langrand, O.; Ratsimbazafy, J.; Rasoloarison, R.M.; Ganzhorn,<br />
J.U.; Rajaobelina, S.; Tattersall, I.; Meyers, D.M. 2006.<br />
Lemurs of Madagascar. Second edition. Conservation International,<br />
Washington, D.C.<br />
Rakotoarison,N.1998.Recent discoveries of the hairy-eared<br />
dwarf lemur (Allocebus trichotis). Lemur News 3: 21.<br />
Rakotoarison, N.; Zimmermann, H.; Zimmermann, E. 1997.<br />
First discovery of the hairy-eared dwarf lemur (Allocebus<br />
trichotis) in a highland rain forest of Eastern Madagascar.<br />
Folia Primatol. 68: 86-94.<br />
Ralison,J.M.2010.The lemurs of the Ambatovy-Analamay region.<br />
Malagasy Nature 3: 178-191.<br />
Randrianambinina, B.; Rasoloharijaona, S. 2006. Inventaires<br />
des lémuriens nocturnes dans la forêt pluviale de Maromizaha<br />
(Est de Madagascar). Lemur News 11: 9-11.<br />
Schmid,J.;Fietz,J.;Rakotobe,Z.L.R.1999.Lémuriens du corridor<br />
Mantadia-Zahamena, Madagascar. In: J. Schmid, L.E.<br />
Alonso (eds.). Une évaluation biologique rapide du corridor<br />
Mantadia-Zahamena, Madagascar. Bulletin of Biological<br />
Assessment 32: 61-72.<br />
Schütz, H; Goodman, S.M. 1998. Photographic evidence of -<br />
Allocebus trichotis in the Reserve Speciale d’Anjanaharibe-<br />
Sud. Lemur News 3: 21-22.<br />
Tattersall,I.1982.The primates of Madagascar.Columbia University<br />
Press, New York.<br />
When big lemurs swallow up small ones:<br />
Coquerel’s dwarf lemur as a predator of<br />
grey mouse lemurs and endemic rodents<br />
Susanne Schliehe-Diecks 1, Matthias Markolf 2, Elise<br />
Huchard 2*<br />
1Courant Research Center "Evolution of Social Behavior",<br />
Georg-August-University of Göttingen,Kellnerweg 6,37077<br />
Göttingen, Germany<br />
2Abteilung Verhaltensökologie and Soziobiologie,<strong>Deutsches</strong><br />
<strong>Primatenzentrum</strong>,Kellnerweg 4,37077 Göttingen,Germany<br />
*Corresponding author: ehuchard@gmail.com<br />
Predation has probably played a major role in the evolutionary<br />
history of lemurs, and specifically affects small nocturnal<br />
lemurs, which are heavily predated upon by a wide range of<br />
vertebrates, including carnivores (e.g., viverrid or domestic<br />
carnivores), birds (e.g. raptors, owls) or reptiles (e.g. Boidae)<br />
(Goodman, 2003). In contrast, lemur predation by other lemur<br />
species appears exceptional and highly opportunistic,<br />
with one observed case of predation of an infant Lemur catta<br />
by Eulemur fulvus (Pitts, 1995). However, such events might<br />
occur more regularly in other lemur species. Two indirect<br />
lines of evidence suggest that Coquerel’s dwarf lemur (Mirza<br />
coquereli) predates on closely related smaller mouse lemurs<br />
(Microcebus sp.) (Kappeler and Rasoloarison,2003).The first<br />
report is based on events where the partially eaten carcass of<br />
a gray mouse lemur (M. murinus) was found together with a<br />
live M. coquereli in a trap (Goodman, 2003). The second observation<br />
consists of an experimental confrontation of M.<br />
murinus with M. coquereli, both being kept in separate cages<br />
that were temporarily placed next to each other. In most<br />
experiments, mouse lemurs started alarm-calling at the<br />
Coquerel’s dwarf lemur and moved around in their cage in an<br />
agitated fashion (Fichtel,2009).Here,we present the first direct<br />
evidence of predation by wild M. coquereli upon gray<br />
mouse lemurs and endemic rodents (western tuft-tailed rats,<br />
Elliurus myoxinus).<br />
Study animals, study site and methods<br />
Coquerel’s dwarf lemurs (300 g;mean home range size:4 ha)<br />
occur in the western lowland forests and gray mouse lemurs<br />
(60 g;mean home range size:1.5 ha) can be found in most remaining<br />
forests in southern and western Madagascar (Kappeler<br />
and Rasoloarison,2003;Rasoloarison et al.,2000).Both<br />
species share several features.Both are nocturnal and omnivorous<br />
solitary foragers. They mainly feed on primary resources<br />
(fruits,gum,flowers,young leaves),insect secretions,<br />
small invertebrates and occasionally vertebrates (chameleons<br />
and lizards). Their diet displays seasonal fluctuation, as<br />
well as interspecific variation (Goodman, 1993, 2003), and<br />
the Coquerel’s dwarf lemur is reported to be slightly more<br />
carnivorous than the gray mouse lemur (Petter et al., 1977).<br />
In captivity, both species have been observed eating young<br />
rodents (Petter et al., 1977) although this has never been reported<br />
in the wild. Both species occur sympatrically in central<br />
western Madagascar with western tuft-tailed rats,a nocturnal,<br />
frugi- and granivorous and partially arboreal rodent<br />
(average body mass: 66 g) (Carleton, 2003).<br />
All following observations were made in the Forêt de<br />
Kirindy,a 12,500 ha forestry concession of the C.N.F.E.R.E.F.<br />
(formerly C.F.P.F.) Morondava. This dry deciduous forest is<br />
situated 60 km northeast of Morondava (44°39´E, 20°03´S).<br />
The predation of the western tufted-tail rat was witnessed<br />
during a focal observation of a Coquerel’s dwarf lemur which<br />
was equipped with a radio collar (Biotrack TW3). The observed<br />
mouse lemurs were similarly equipped with radio<br />
collars (Holohil Systems Ltd., BD-2C, 1.8 g), permitting behavioural<br />
observations of focal animals.<br />
Results and discussion<br />
The first observation reports the predation of a western<br />
tuft-tailed rat by an adult male M.coquereli in November 2006<br />
(Fig.1).The Coquerel’s dwarf lemur was found sitting on the<br />
ground at 20h17, feeding on a tufted-tail rat, and changed its<br />
position only to climb-up the vegetation from 1-3 m height<br />
and to recover the carcass when it fell to the ground. It devoured<br />
the whole carcass, including (cracked) bones. After<br />
finishing eating,the M.coquereli groomed its face and hands.
Page 14 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
Fig. 1: An adult Coquerel’s<br />
dwarf lemur (Mirza coquereli)<br />
in the Foret de Kirindy, Madagascar.<br />
The second observation<br />
reports an unsuccessful<br />
attack on an adult female<br />
gray mouse lemur (55 g,<br />
approx. age: 1 year and 9<br />
months) by a Coquerel’s<br />
dwarf lemur in October<br />
2009.At 21h24,the mouse<br />
lemur had been foraging<br />
high up in the vegetation<br />
(between 6 and <strong>15</strong> m) for<br />
at least 10 minutes, licking<br />
sugary insect secretions<br />
off leaves,when a M.coquereli,<br />
adult size, was spotted<br />
at the same height, about<br />
10m from the focal subject,<br />
slowly and silently<br />
moving towards the<br />
mouse lemur. Marking<br />
brief and frequent pauses<br />
in an apparently easy pro-<br />
gression into the canopy, its whole attitude strongly recalled<br />
the hunting cat, with a low head and a flexible body, apparently<br />
entirely focused on its prey.In less than 30 seconds,the<br />
Mirza was within 5 m of the mouse lemur,who kept feeding in<br />
the same location. While the Mirza approached within 2 m,<br />
the mouse lemur suddenly disappeared in an eclipse, quickly<br />
fleeing among the top and tiniest branches,and jumping from<br />
one slim branch to the next.The Coquerel’s dwarf lemur did<br />
not try to chase it.After 2 minutes out of sight,the mouse lemur<br />
was retrieved quietly feeding on tree exudates, 25 m<br />
away from its previous localization in its fleeing direction,and<br />
<strong>15</strong> m away from a frequently used sleeping site. Less than 10<br />
minutes later, the female was joined by a related female and<br />
both entered the tree hole together (21h37).<br />
Finally, an adult Coquerel’s dwarf lemur was observed feeding<br />
on a young male gray mouse lemur (body mass: 37 g;<br />
approx.age:2-3 months) in June 2010.The predation was recorded<br />
at 22h10,about two hours after behavioural data had<br />
been collected from the predated mouse lemur,which at the<br />
time showed no signs of injuries and displayed normal behaviour.The<br />
body of the gray mouse lemur was almost complete<br />
when the observer spotted the M. coquereli feeding on<br />
it,suggesting that the mouse lemur was killed shortly before.<br />
The Coquerel’s dwarf lemur was sitting with its prey in a tree<br />
of about 10m height, which stood 25 m away from the position<br />
where the grey mouse lemur was last spotted alive. It<br />
took about one hour to finish the entire carcass,interrupted<br />
by occasional vigilance scans of the surroundings.<br />
The frequency of such events is probably relatively low, and<br />
all reported observations happened during, or at the end of,<br />
the dry season in Kirindy. It is thus possible that predation<br />
pressure by M. coquereli increases at times of food scarcity,<br />
when alternative resources like fruits and invertebrates are<br />
rare or absent. However, it is also important to note that<br />
most observations took place during the dry season, when<br />
vegetation density is low in this dry, deciduous forest. This<br />
means that the timing of events reported here might simply<br />
reflect study methods. Nevertheless, this suit of anecdotal<br />
observations represents the first direct and unambiguous<br />
evidence for predation by the Coquerel’s dwarf lemur upon<br />
small nocturnal lemurs, as well as other mammals. Predation<br />
among other primate species is relatively rare. So far, only<br />
chimpanzees, orangutans, baboons, blue monkeys and capuchins<br />
have been observed preying upon other primates<br />
(Fichtel, in press). Our report provides evidence for the first<br />
case where a lemur species might commonly predate upon<br />
other lemurs.<br />
References<br />
Carleton,M.D.2003.Eliurus,Tufted-tailed rats.Pp.1373-1380.<br />
In:S.M.Goodman;J.Benstead (eds.). The Natural History<br />
of Madagascar.The University of Chicago Press,Chicago.<br />
Fichtel, C. 2009. Costs of alarm calling: lemur alarm calls<br />
attract fossas. Lemur News 14: 53-54.<br />
Fichtel,C.in press.Predation.In:J.Mitani:J.Call;P.M,Kappeler;<br />
R.Palombit;J.B.Silk;(eds.) The Evolution of Primate Societies.<br />
The University of Chicago Press, Chicago.<br />
Goodman,S.M.;O’Connor,S.;Langrand,O.1993.A review of<br />
predation on lemurs: implications for the evolution of<br />
social behavior in small, nocturnal primates. Pp. 51-66. In:<br />
P.M.Kappeler;J.U.Ganzhorn (eds.).Lemur social systems and<br />
their ecological basis. Plenum Press, New York.<br />
Goodman,S.M.2003.Predation on Lemurs.Pp 1221-1228.In:<br />
S. M. Goodman: J. Benstead (eds.). The Natural History of<br />
Madagascar. The University of Chicago Press, Chicago.<br />
Kappeler,P.M.;Rasoloarison,R.M.2003.Microcebus,mouse lemurs,<br />
tsidy. Pp 1310-13<strong>15</strong>. In: S. M. Goodman: J. Benstead<br />
(eds.).The Natural History of Madagascar.The University<br />
of Chicago Press, Chicago.<br />
Petter, J-J.; Albignac, R.; Rumpler, Y. 1977. Mammifères lémuriens<br />
(primates prosimiens). In: Faune de Madagascar,<br />
Paris.<br />
Pitts, A. 1995. Predation by Eulemur fulvus on an infant Lemur<br />
Catta at Berenty, Madagascar. Folia Primatol. 65: 169-71.<br />
Rasoloarison, R.M.; Goodman, S.; Ganzhorn, J.U. 2000. Taxonomic<br />
revision of mouse lemur (Microcebus) in the western<br />
portions of Madagascar.Int.J.Primatol.21:963-1019.<br />
Collective mobbing of a boa by a group of<br />
red-fronted lemurs (Eulemur fulvus rufus)<br />
Lennart Pyritz 1,2*, Tianasoa Andrianjanahary 1<br />
1Behavioral Ecology & Sociobiology Unit, German Primate<br />
Center, Kellnerweg 4, 37077 Göttingen, Germany<br />
2CRC "Evolution of Social Behavior", University of Göttingen,<br />
Germany<br />
*Corresponding author: LennartPyritz@gmx.net<br />
Key words: red-fronted lemurs, boa, predation, mobbing,<br />
anti-predator behaviour<br />
Introduction<br />
Collective anti-predator behaviour is one of the principal advantages<br />
of group-living (for mammals, e.g., Janzen, 1970; van<br />
Schaik, 1983). It can be broadly divided into two strategies<br />
and tactics employed before and after predator encounters<br />
(Caro,2005;Rahlfs and Fichtel, 2010;Fichtel, in press). While<br />
the former include predator-sensitive foraging and increased<br />
vigilance, mobbing occurs in several mammal species after<br />
detecting a predator (e.g., Tamura, 1989). Why animals<br />
engage in mobbing and who benefits from it in which way<br />
remains an unresolved question in animal behaviour (for<br />
reviews see Curio et al., 1978; Shields, 1984). Until today,<br />
published field observations of group-living lemurs mobbing<br />
a predator are rare (summarised in Scheumann et al., 2007).<br />
Regarding snakes, only three interactions have been described<br />
so far (Colquhoun,1993;Rakotondravony,1998;Burney,<br />
2002). Here, we report a prolonged mobbing display<br />
against a Madagascar ground boa (Acrantophis madagascariensis)<br />
by a group of red-fronted lemurs (Eulemur fulvus rufus)<br />
in Kirindy Forest. Observations like this may help to elucidate<br />
fundamental mechanisms of collective anti-predator behaviour<br />
by contributing to a pool of data on mobbing by particular<br />
pairs of prey and predators.
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page <strong>15</strong><br />
Observations<br />
The event was observed during regular behavioural observations<br />
of red-fronted lemurs in Kirindy Forest,60 km north of<br />
Morondava. It was the only snake-lemur interaction observed<br />
during the entire study period from November 2007<br />
to April 2010, in which four lemur groups were followed<br />
daily by one or two observers,respectively (> 4,000 h of observation<br />
data).Red-fronted lemurs live in multi-male,multifemale<br />
groups of 5-12 individuals (Pereira and Kappeler,<br />
1997;Wimmer and Kappeler,2002).The study group (B) that<br />
encountered the boa included 9 individually marked animals<br />
at the time (2 adult females, 5 adult males, 1 juvenile male, 1<br />
male infant).<br />
On March 1,2010,at 7.19 h,sudden alarm calls of several redfronted<br />
lemurs were heard in the study area known as CS7.<br />
Six individuals (2 adult females,2 adult males,1 juvenile male,<br />
1 male infant) could be identified after approaching the group<br />
to within 10 m.Three of them (2 adult females,1 adult male)<br />
emitted "Woofs" and "Huvvs",vocalisations typically uttered<br />
during predator encounters (Fichtel and Kappeler, 2002).<br />
The 5 individuals surrounded an approximately 2 m long<br />
Madagascar ground boa (Fig. 1) that was lying motionless on<br />
the ground. The lemurs sat at a height of 1-2 m, each about<br />
3 m away from the snake, wagging their tails vigorously, ex-<br />
Fig. 1: Madagascar ground boa in Kirindy. (Photo: Lennart<br />
Pyritz)<br />
cept the infant that kept a distance of 5 m during the entire<br />
event and did not display any vocalisations or tail-wagging.<br />
During the next 4 min,one of the adult males approached the<br />
front end of the boa twice, getting as close as 1-2 m. After 5<br />
min, he left the scene. During this time, one of the adult females<br />
also approached the snake up to within 2 m.When the<br />
male left,the second female started to quickly circuit the boa<br />
for 4 min,maintaining a distance of 2-3 m.After 14 min of several<br />
approaches and continuous alarm calls by 3-5 individuals,<br />
the boa moved for the first time,heading slowly away.The remaining<br />
adult male approached the moving snake also within<br />
2 m; also at its front end. About 1 min later, the boa had<br />
moved <strong>15</strong> m away, and the lemurs left in the opposite direction,still<br />
uttering grunts continuously.Once the boa was out<br />
of sight, the mobbing stopped and the lemurs` behaviour returned<br />
to baseline levels.<br />
Discussion<br />
The mobbing reaction of the group was strong and prolonged<br />
and included most of the group members.This is similar<br />
to the behaviour of a black lemur (Eulemur macaco<br />
macaco) group encountering a Madagascar boa at Ambato<br />
Massif, where the group mobbed the snake for <strong>15</strong>-20 min,<br />
and some individuals approached it as close as 1 m before<br />
finally leaving the location (Colquhoun, 1993). It is also noteworthy<br />
that females and males mobbed and approached the<br />
snake in equal measure as observed in a number of other<br />
species (e.g., Tamura, 1989; Ferrari and Ferrari, 1990; Tello et<br />
al., 2002). The infant maintained a larger distance to the boa<br />
and did not engage in the mobbing displays, however. Similar<br />
infant behaviour has also been reported for other primates<br />
(e.g., Ferrari and Ferrari, 1990) and might be due to a higher<br />
susceptibility to an attack due to smaller body size or a lack<br />
of innate experience regarding predator encounters and<br />
mobbing strategies (Curio et al., 1978; Fichtel, in press).<br />
The strong mobbing reaction of the lemurs might be explained<br />
by the hunting strategy of the snake.Boas are ambush<br />
hunters that usually abandon an attack as soon as they have<br />
been detected (Montgomery and Rand, 1978; Slip and Shine,<br />
1988). Therefore, it seems beneficial for prey animals to signal<br />
the ambush hunter quickly and distinctly that it has been<br />
detected. As boas do not pursue their prey after an unsuccessful<br />
attack, it is also unsurprising that the lemurs’ behaviour<br />
returned to baseline levels of anxiety shortly after departing<br />
the site of the predator encounter. In contrast,<br />
groups of red-fronted lemurs showed increased vigilance behaviour<br />
for at least 30-60 min after encountering a fossa<br />
(Cryptoprocta ferox;pers.comm.Jean-Pierre Tolojanahary and<br />
pers.observation by LP),which is probably due to the higher<br />
agility and climbing abilities of the largest mammalian carnivore.Furthermore,fossas<br />
have been observed to hunt cooperatively<br />
and pursue prey up to 45 min (Lührs and Dammhahn,<br />
2009).<br />
There are no quantitative data on predation rates of lemurs<br />
by snakes in Kirindy, only opportunistic observations (e.g.,<br />
Schülke, 2001; Eberle and Kappeler, 2008) that are biased by<br />
several factors, however. The low observation rate of boalemur<br />
interactions could be due to the reptiles` nocturnal<br />
lifestyle (Raxworthy,2003),so that most of the attacks would<br />
occur at night when no or only few observers are working in<br />
the forest. Furthermore, boas at Kirindy are only active during<br />
the rainy season from January to April, when observations<br />
are often limited by dense foliage and frequent rainfalls.<br />
Five of the six individuals taking part in the mobbing were related<br />
(1 adult female and her 4 offspring from the last 4<br />
years), while it is currently unknown whether the second<br />
adult male sired one of the two youngest group members.<br />
However, due to the small number of detailed observations<br />
of predator encounters it remains impossible to identify the<br />
ultimate causes of collective mobbing (kin defence/parental<br />
care, self-/group defence or cultural transmission of enemy<br />
recognition; Curio et al., 1978) in this species for the time<br />
being.<br />
Acknowledgements<br />
LP was supported financially by the Deutsche Forschungsgemeinschaft<br />
(DFG; KA 1082/16-1, FuE). We thank field assistant<br />
Jean-Pierre Tolojanahary for sharing his long-term observation<br />
experiences with us. We are also grateful to Peter<br />
Kappeler, Claudia Fichtel and Moritz Rahlfs for constructive<br />
and helpful comments on earlier drafts of the manuscript.<br />
References<br />
Burney, D.A. 2002. Sifaka predation by a large boa. Folia Primatologica<br />
73: 144-145.<br />
Caro, T. 2005. Antipredator defenses in birds and mammals.<br />
The University of Chicago Press, Chicago, London.<br />
Colquhoun, I.C. 1993. The socioecology of Eulemur macaco:<br />
A preliminary report. Pp. 11-23. In: P.M. Kappeler; J.U.<br />
Ganzhorn (eds.). Lemur social systems and their ecological<br />
basis. Plenum Press, New York and London.
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Curio, E.; Ernst, U.; Vieth, W. 1978. Cultural transmission of<br />
enemy recognition:one function of mobbing.Science 202:<br />
899-901.<br />
Eberle, M.; Kappeler, P.M. 2008. Mutualism, reciprocity, or kin<br />
selection? Cooperative rescue of a conspecific from a boa<br />
in a nocturnal solitary forager the gray mouse lemur.<br />
American Journal of Primatology 70: 410-414.<br />
Ferrari, S.F.; Ferrari, M.A.L. 1990. Predator avoidance behaviour<br />
in the buffy-headed marmoset, Callithrix flaviceps.<br />
Primates 31: 323-338.<br />
Fichtel, C. In press. Predation on primates. In: J.C. Mitani; J.<br />
Call;P.Kappeler;R.Palombit;J.Silk (eds.).The evolution of<br />
primate societies. University of Chicago Press, Chicago.<br />
Fichtel, C.; Kappeler, P.M. 2002. Anti-predator behaviour of<br />
group-living Malagasy primates: mixed evidence for a<br />
referential alarm call system. Behavioral Ecology and<br />
Sociobiology 51: 262-275.<br />
Janzen,D.1970.Altruism by coatis in the face of predation by<br />
boa constrictor. Journal of Mammalogy 51: 387-389.<br />
Lührs,M.L.;Dammhahn,M.2009.An unusual case of cooperative<br />
hunting in a solitary carnivore. Journal of Ethology<br />
DOI: 10.1007/s10164-009-0190-8.<br />
Montgomery, G.G.; Rand, A.S. 1978. Movements, body temperature<br />
and hunting strategy of a Boa constrictor. Copeia<br />
3: 532-533.<br />
Pereira, M.E.; Kappeler, P.M. 1997. Divergent system of agonistic<br />
behaviour in lemurid primates.Behavior 34:225-74.<br />
Rahlfs,M.;Fichtel,C.2010.Anti-predator behaviour in a nocturnal<br />
primate, the grey mouse lemur (Microcebus murinus).<br />
Ethology 116: 429-439.<br />
Rakotondravony, D.; Goodman, S.M.; Soarimalala, V. 1998.<br />
Predation on Hapalemur griseus griseus by Boa manditra<br />
(Boidae) in the Littoral Forest of Eastern Madagascar.<br />
Folia Primatologica 69: 405-408.<br />
Raxworthy, C.J. 2003. Boidae, Boas. Pp. 993–997. In: S.M.<br />
Goodman; J.P. Benstead (eds.). The Natural History of<br />
Madagascar. University of Chicago Press, Chicago.<br />
Scheumann, M.; Rabesandratana, A.; Zimmermann, E. 2007.<br />
Predation, communication, and cognition in lemurs. Pp.<br />
100-126. In: S. Gursky; K.A.I. Nekaris (eds.). Primate antipredator<br />
strategies. Springer, New York.<br />
Schülke, O. 2001. Social anti-predator behaviour in a nocturnal<br />
lemur. Folia Primatologica 72: 332-334.<br />
Shields,W.1984.Barn swallow mobbing:self-defence,collateral<br />
kin defence, group defence, or parental care? Animal<br />
Behaviour 32: 132-148.<br />
Slip, D.; Shine, R.1988. Feeding habits of the diamond python,<br />
Morelia s. spilota: ambush predation by a Boid snake. Journal<br />
of Herpetology 22: 323-330.<br />
Tamura, N. 1989. Snake-directed mobbing by the Formosan<br />
squirrel Callosciurus erythraeus thaiwanensis. Behavioral<br />
Ecology and Sociobiology 24: 175-180.<br />
Tello, N.S.; Huck, M.; Heymann, E.W. 2002. Boa constrictor attack<br />
and successful group defence in moustached tamarins,<br />
Saguinus mystax. Folia Primatologica 73: 146-148.<br />
Van Schaik, C. 1983. Why are diurnal primates living in<br />
groups? Behaviour 88: 120-143.<br />
Wimmer,B.;Kappeler,P.M.2002.The effects of sexual selection<br />
and life history on the genetic structure of redfronted<br />
lemur, Eulemur fulvus rufus, groups. Animal Behaviour 64:<br />
557-68.<br />
Response of two nocturnal lemurs (Microcebus<br />
murinus and Lepilemur leucopus)<br />
to a potential boiidae (Sanzinia madagascariensis)<br />
predator<br />
Krista Fish<br />
Department of Anthropology, The Colorado College, 14 E<br />
Cache La Poudre, Colorado Springs, CO 80903, USA,<br />
krista.fish@coloradocollege.edu<br />
Primates display an array of responses to predators, and<br />
differences in these responses were once thought to exist<br />
based on activity pattern. Solitary foraging, cryptic color-<br />
ation, and cryptic movements to remain hidden from predators<br />
were considered anti-predator adaptations among<br />
nocturnal primates while diurnal primates used large group<br />
size to enhance their ability to detect and defend against<br />
predators, give alarm calls to warn conspecifics of the presence<br />
of a predator,and flee from predators (Hill and Dunbar,<br />
1998;Isbell,1994;Stanford,2002).As research into nocturnal<br />
primate behavior expanded,results revealed that,like diurnal<br />
primates,nocturnal primates display a range of anti-predator<br />
behaviors upon encountering a predator (Fichtel, 2007).<br />
Nocturnal primates vary in the type of response (mobbing,<br />
alarm calling), height in the canopy, proximity to other<br />
individuals, and vigilance levels in the presence of different<br />
predators (Fichtel, 2007; Gursky, 2002, 2003, 2005, 2006;<br />
Schuemann et al.,2007;Schulke,2001).In particular,mobbing<br />
behaviors are well- documented in tarsiers (Tarsius sp.),<br />
mouse lemurs (Microcebus spp.), and fork-marked lemurs<br />
(Phaner furcifer) (Gursky, 2007; Eberle and Kappeler, 2008;<br />
Schulke, 2001). Mobbing includes close approach, touching,<br />
sniffing, and pouncing on the predator (Gursky, 2007). Interspecies<br />
mobbing of a predator occurs as well. Fork-marked<br />
lemurs and coquerel’s dwarf lemurs (Mirza coquereli) together<br />
mobbed a snake (Boa manditra) (Schulke, 2001).<br />
Several hypotheses have been put forth to explain the evolution<br />
of mobbing behavior (Eberle and Kappeler,2008):1) byproduct<br />
mutualism in which individuals defend others in the<br />
process of defending themselves, 2) reciprocity where animals<br />
obtain higher fitness by cooperating with others,and 3)<br />
kin selection whereby animals cooperate when they share<br />
common genes. A fourth hypothesis known as perception<br />
advertisement was developed as an explanation for the<br />
evolution of alarm calling and other mobbing behaviors in<br />
birds (Curio et al., 1978; Zuberbuhler et al., 1999) but has<br />
been extended to account for the presence of mobbing behaviors<br />
in primates (Gursky,2005).According to this hypothesis,<br />
alarm calling and mobbing are signals to the predator<br />
that the element of surprise has been lost. Snakes, leopards,<br />
and other animals that hunt by crypticity and rely on the element<br />
of surprise to capture prey are common recipients of<br />
alarm calling and mobbing.Research on both diurnal and nocturnal<br />
primates suggests that alarm calling and mobbing by<br />
these primates results in predators leaving an area (Zuberbuhler<br />
et al.,1999;Gursky,2006).Here I report the divergent<br />
responses of two species of nocturnal primates (Microcebus<br />
murinus and Lepilemur leucopus) to the same potential predator-<br />
a nocturnal boiidae snake (Sanzinia madagascariensis)<br />
and discuss implications for the above hypotheses based on<br />
these observations.<br />
Methods<br />
The observations reported here were made in the Ankoba<br />
gallery forest of Berenty Private Reserve in southern Madagascar.<br />
The encounters between the snake and primates<br />
were observed in the course of a six month study investigating<br />
the ecology of Microcebus murinus. During this study,<br />
trails within the reserve were walked to locate unhabituated<br />
mouse lemurs.When encountered,the time at which the encounter<br />
began, height, location, and activity of the mouse lemur<br />
were recorded continuously until the mouse lemur was<br />
out of sight for more than five minutes. The time, height, and<br />
activity of potential predators were also noted when they<br />
were encountered, but predators were not followed unless<br />
they were within proximity to a primate.<br />
Results<br />
While conducting walks to locate mouse lemurs in May 2009,<br />
a sportive lemur alarm call at 20:52 alerted me to the pres-
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 17<br />
ence of a boiidae snake later identified as Sanzinia madagascariensis.<br />
I saw two white-footed sportive lemurs (Lepilemur<br />
leucopus),one located at 4m height and the other at 5m<br />
height in a tree over the trail.The sportive lemurs were barking<br />
and looking at the snake. The snake was moving at<br />
approximately 4 m high in a tree and was attempting to cross<br />
a gap and move into a tree nearer to the tree in which the<br />
sportive lemurs were located.The attempt was unsuccessful<br />
as the snake slipped and almost fell out of the thin, terminal<br />
branches. The snake then moved down to 3.5 m height and<br />
began crossing the canopy gap along thicker branches. The<br />
sportive lemurs continued to alarm bark at the snake until<br />
21:02 when the snake turned away from the gap in the<br />
canopy and began moving lower down in the tree and away<br />
from the sportive lemurs. The sportive lemur alarm calling<br />
ended by 21:04 when the snake had traveled down the tree<br />
trunk to 3m in height. The sportive lemur lowest in the tree<br />
continued to watch the snake while the other sportive lemur<br />
moved to 8m height in its tree and began feeding.<br />
At 21:<strong>15</strong>, I was preparing to leave the area when I noticed a<br />
mouse lemur at 0.5 m in the same tree as the snake. The<br />
snake was at 3 m height in the tree and moving down the<br />
main trunk of the tree. The mouse lemur looked at me as it<br />
walked up the main trunk of the tree in the direction of the<br />
snake. The snake faced the mouse lemur, but the mouse<br />
lemur did not appear to notice the snake as it alternated<br />
looking in my direction with looking around its immediate<br />
area while foraging for insects. The mouse lemur continued<br />
to move up the tree until it was within 0.25 m of the snake<br />
where it paused and looked intently at the spot where the<br />
snake was located and then jumped backwards away from<br />
the snake. The mouse lemur then began moving around the<br />
tree at the same height (3 m) as the snake,jumping to terminal<br />
branches,and running along main branches while pausing<br />
to look at the snake. The mouse lemur’s movements took it<br />
from 1m to only a few cms in distance from the snake. The<br />
mouse lemur continued this pattern of running and pausing<br />
to examine the snake for 7 mins. The mouse lemur then began<br />
foraging for insects in a neighboring tree at 3-4 m height<br />
and within 1-2 m of the snake. While foraging, the mouse<br />
lemur would pause to look in the direction of the snake<br />
which remained motionless.The mouse lemur foraged in this<br />
manner for 4 mins until a sportive lemur alarm called. Prior<br />
to the alarm call, the mouse lemur was foraging approximately<br />
1m from the snake and the snake began moving down<br />
the tree trunk.At the sound of the sportive lemur alarm call,<br />
the mouse lemur jumped to 2 m distance from the snake and<br />
paused in its foraging to watch the snake. The snake remained<br />
motionless.A minute later the sportive lemurs alarm<br />
called again. The snake began moving down the trunk of the<br />
tree again and the mouse lemur moved to forage insects in<br />
trees that were approximately 3-5 m from the snake and at<br />
5m height.After 5 mins of foraging at this increased distance<br />
from the snake, the mouse lemur moved back into the tree<br />
where it was initially observed and foraged within 1m of the<br />
snake for 2 mins. The mouse lemur then moved close to the<br />
snake coming within less than 1m of the boa and running and<br />
jumping around the snake while pausing to watch it while<br />
standing bipedally. After 3 mins of remaining motionless<br />
while the mouse lemur ran, jumped and watched the snake,<br />
the snake began moving back up into the dense crown of the<br />
tree. The mouse lemur continued running and jumping<br />
around the snake and watching it at a distance of 0,5-1 m<br />
away as the snake moved into the crown of the tree. 8 mins<br />
later,the mouse lemur began foraging insects at 3-4 m height<br />
in the canopy and within 0,5-1 m from the snake with occasional<br />
glances in the direction of the snake.The mouse lemur<br />
then moved further away from the snake and foraged insects<br />
at 5m in height and approximately 5 m distance from the<br />
snake.At 21:46 the mouse lemur was out of sight and did not<br />
return by 21:51 when I left and continued the mouse lemur<br />
walk.A return visit to the location at 10:05 for species identification<br />
of the species revealed the presence of no mouse lemurs<br />
or Lepilemur in the vicinity.<br />
Discussion<br />
Eberle and Kappeler (2008) describe the successful mobbing<br />
of a Sanzinia madagascariensis by two female and one male<br />
mouse lemur (Microcebus murinus). The mobbing caused the<br />
snake to release a captured male mouse lemur. Subsequent<br />
genetic analysis revealed that the mobbing females were related<br />
to the attempted victim (Eberle and Kappeler, 2008).<br />
Additional encounters between mouse lemurs and snakes<br />
without captured prey did not elicit mobbing behaviors from<br />
the mouse lemurs (Eberle and Kappeler, 2008). Instead, the<br />
mouse lemurs sat approximately 2 m from the snake and<br />
watched it. (Eberle and Kappeler, 2008). The mobbing of the<br />
snake by relatives of a captured mouse lemur and the lack of<br />
mobbing behaviors by solitary mouse lemurs in Eberle and<br />
Kappeler’s study support the kin selection hypothesis for the<br />
evolution of mobbing behaviors. However, the solitary<br />
mouse lemur observed in this study displayed mobbing-like<br />
behaviors in the absence of kin or other individuals. Additionally,<br />
under experimental conditions, solitary mouse lemurs<br />
monitored model snakes and even locomoted towards<br />
the model predators (Rahlfs and Fichtel,2010).The presence<br />
of mobbing-like behaviors in solitary animals lends support<br />
to the by-product mutualism hypothesis. Mouse lemurs may<br />
display mobbing-like behaviors as an individual strategy and<br />
then when larger numbers of mouse lemurs contact a predator,<br />
the appearance of a group-defense strategy may result<br />
from multiple individuals pursuing the same strategy.<br />
The observations here also lend support to the perception<br />
advertisement hypothesis.The mouse lemur may have benefited<br />
by displaying mobbing behaviors toward the snake to let<br />
the snake know that it was being monitored and would not<br />
surprise the lone mouse lemur as it foraged.However,a prediction<br />
of the perception-advertisement hypothesis is that<br />
cryptic predators should flee when faced with alarm calls and<br />
mobbing as they cannot surprise prey in their vicinity. The<br />
alarm calling by the sportive lemurs and the mobbing-like<br />
behaviors of the mouse lemur observed in this study did not<br />
cause the snake to flee. Instead, the snake remained in the<br />
area while both the sportive lemurs and mouse lemur left<br />
the area.Size of the mobbing and alarm-calling group may be<br />
important. Tarsiers were more likely to retreat first if their<br />
mobbing group consisted of four or fewer individuals (Gursky,2007).The<br />
small group sizes here may have not caused the<br />
predator to flee,but may have served as an adequate warning<br />
that it had lost the element of surprise and should not<br />
expend energy in an attack.<br />
Of particular interest in these observations was the opportunity<br />
to view the responses of two different nocturnal primate<br />
species to the same predator. Such observations are<br />
infrequent in the literature. Schulke (2001) observed Phaner<br />
and dwarf lemur (Mirza coquereli) together mobbed Boa<br />
manditra. In the case of the sportive and mouse lemurs,<br />
neither showed the same behavioral response to the Sanzinia<br />
madagascariensis. The pair of Lepilemurs maintained a<br />
larger distance between themselves and the snake than did<br />
the mouse lemur. The mouse lemur did not vocalize in the<br />
vicinity of the snake, however, the sportive lemurs vocalized<br />
during their encounter with the snake.Mouse lemurs seldom<br />
respond to predator models with alarm calls in experimental
Page 18 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
conditions (Rahlfs and Fichtel,2010).Nocturnal primates and<br />
other mammals may not rely on alarm calls as an anti-predator<br />
strategy (Rahlfs and Fichtel, 2010) because early detection<br />
of predators at night is more difficult and solitary foraging<br />
may limit the usefulness of this strategy. The sportive<br />
lemurs however did alarm call in the presence of the snake.<br />
Alarm calls in nocturnal primates may function as a deterrent<br />
to the predator or to recruit conspecifics in defense against<br />
the predator (Rahlfs and Fichtel,2010).The exact function of<br />
the sportive lemur alarm calls in this study cannot be determined,<br />
but they do reveal a need for future studies to examine<br />
potential variation in alarm call behaviors between nocturnal<br />
species.Given variation in body size,diet,and sociality<br />
in nocturnal primates, we might expect variation in antipredator<br />
behaviors such as alarm calling among nocturnal<br />
species.<br />
The possibility exists that the mouse lemur may have recognized<br />
the alarm calls of the sportive lemur because it increased<br />
its distance from the snake, ceased foraging, and<br />
monitored the snake following an alarm call. Recognition of<br />
the alarm calls of other species has been documented for<br />
birds, small mammals, and primates including diurnal lemurs<br />
(Fichtel,2004;Rainey et al.,2004;Shriner,1998).However,the<br />
suggestion that mouse lemurs recognize sportive lemur<br />
alarm calls needs to be further investigated with field experiments.<br />
The ability of nocturnal primate species to recognize<br />
the alarm calls of other sympatric species would be beneficial<br />
to animals that frequently forage away from conspecifics.<br />
The few encounters documented between nocturnal primates<br />
and their predators describe a range of anti-predator<br />
responses that vary depending on the type of predator,proximity<br />
of conspecifics,and available vegetative cover.Such flexibility<br />
is interesting because nocturnal primates- especially<br />
mouse lemurs- are often used as living models for the ancestral<br />
primate. The anti-predator behaviors of nocturnal primates<br />
such as mouse lemurs may reflect the primitive<br />
primate or even primitive mammal condition (Stanford,<br />
2002). The range of anti-predator behavior described for<br />
nocturnal primates in this and other studies implies that the<br />
flexibility in anti-predator behaviors observed in primates<br />
have a more ancient origin than originally suspected or that<br />
mouse lemurs and sportive lemurs have developed divergent<br />
anti-predator behaviors that may not have been present in<br />
early mammals and primates. Wider use of experiments to<br />
explore the differing conditions which elicit variable responses<br />
to predators within and between nocturnal species<br />
will be necessary to develop a more complete understanding<br />
of ancestral versus derived anti-predator behaviors. Additionally,<br />
comparative research exploring variation in antipredator<br />
behaviors in other nocturnal mammal species will<br />
be needed.<br />
Acknowledgements<br />
I would like to thank Dr.Alison Jolly for her facilitation of this<br />
research project and the de Heaulme family for their permission<br />
to work at Berenty and their support of the project.I am<br />
grateful also to the staff of MICET (Madagascar Institut pour<br />
la Conservations des Ecosystèmes Tropicaux) for their assistance<br />
in travel and obtaining research permits and visas.<br />
Funding for this project was provided by Sigma Xi, The University<br />
of Colorado Museum, the University of Colorado<br />
Graduate Student Grants, and the American Society of<br />
Primatologists.<br />
References<br />
Curio,E.;Ernst,U.;Vieth,W.1978.The adaptive significance of<br />
avian mobbing.II.Cultural transmission of enemy recogni-<br />
tion in blackbirds: effectiveness and some constraints. Z.<br />
Tierpsychol. 48: 184-202.<br />
Eberle, M.; Kappeler, P.M. 2008. Mutualism, reciprocity, or kin<br />
selection? Cooperative rescue of a conspecific from a boa<br />
in a nocturnal solitary forager the gray mouse lemur.Am.J.<br />
Primatol. 70: 410-414.<br />
Fichtel, C. 2004. Reciprocal recognition of sifaka (Propithecus<br />
verreauxi verreauxi) and redfronted lemur (Eulemur fulvus<br />
rufus) alarm calls. Anim. Cogn. 7: 45-52.<br />
Fichtel, C. 2007. Avoiding predators at night: antipredator<br />
strategies in red-tailed sportive lemurs (Lepilemur ruficaudatus).<br />
Am. J. Primatol. 69: 611-624.<br />
Gursky, S. 2002. The behavioral ecology of the spectral tarsier,<br />
Tarsius specturn. Evol. Anth. 11: 226-234.<br />
Gursky, S. 2003. Predation experiments on infant spectral<br />
tarsiers (Tarsius spectrum) Folia. Primatol. 74: 272-284.<br />
Gursky, S. 2005. Predator mobbing in Tarsius spectrum. Internat.<br />
J. Primatol. 26(1): 207-221.<br />
Gursky, S. 2006. Function of snake mobbing in spectral tarsiers.<br />
Am. J. Phys. Anth. 129:601-608.<br />
Gursky, S. 2007. The Spectral Tarsier. Upper Saddle River, NJ:<br />
Pearson/Prentice Hall.<br />
Hill, R.A.; Dunbar, R.I.M. 1998. An evaluation of the roles of<br />
predation rate and predation risk as selective pressures<br />
on primate grouping behavior.Behaviour 135(4):411-430.<br />
Isbell, L.A. 1994. Predation in primates: ecological patterns<br />
and evolutionary consequences. Evol. Anth. 3(2): 61-71.<br />
Rahlfs, M.; Fichtel, C. 2010. Anti-predator behavior in a nocturnal<br />
primate, the grey mouse lemur (Microcebus murinus).<br />
Ethology 116: 429-439.<br />
Rainey HJ,Zuberbuhler K,Slater PJB.2004.Hornbills can distinguish<br />
between primate alarm calls. Proc R Soc Lond B<br />
271: 755-759.<br />
Scheumann, M.; Rabesandratana, A.; Zimmermann, E. 2007.<br />
Predation, communication, and cognition in lemurs. Pp.<br />
100-126. In: S. Gursky; K.A.I Nekaris (eds.). Primate Anti-<br />
Predator Strategies. Springer.<br />
Schulke, O. 2001. Social anti-predator behaviour in a nocturnal<br />
lemur. Folia Primatol. 72: 332-334.<br />
Shriner,W.M.1998.Yellow-bellied marmot and golden-mantled<br />
ground squirrel responses to heterospecific alarm<br />
calls. Anim. Behav. 55: 529-536.<br />
Stanford, C.B. 2002. Avoiding predators: expectations and<br />
evidence in primate antipredator behavior. Int J Primatol<br />
23(4): 741-757.<br />
Zuberbuhler, K.; Jenny, D.; Bshary, R. 1999. The predator deterrence<br />
function of primate alarm calls. Ethology105:<br />
477-490.<br />
Effective predation defence in Cheirogaleus<br />
medius<br />
Kathrin H. Dausmann<br />
Animal Ecology & Conservation, Biocentre Grindel, University<br />
Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg,<br />
Germany, kathrin.dausmann@uni-hamburg.de<br />
Besides one’s own death, the predation of offspring is the<br />
most severe loss of fitness possible.In species that invest significantly<br />
in their offspring it is therefore natural that this "expensive"<br />
offspring should be guarded to avoid predation.<br />
Nonetheless, and particularly in small animals, it is uncertain<br />
whether an adult can effectively defend its offspring when it<br />
is attacked by a larger predator.<br />
The fat-tailed dwarf lemur (Cheirogaleus medius) is a small<br />
(130 g), strictly nocturnal primate that occurs in the dry,<br />
deciduous woodlands of western Madagascar, and lives in<br />
social monogamous small family groups consisting of a reproductive<br />
male-female pair and their offspring from one or<br />
more breeding seasons. Males and females maintain lifelong<br />
pair bonds and usually separate only when one partner dies<br />
(Fietz, 1999; Müller, 1999; Fietz and Dausmann, 2003). The
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 19<br />
diet of C.medius consists mainly of fruits and varying proportions<br />
of arthropods depending on the season (Fietz and<br />
Ganzhorn, 1999). C. medius is unique among primates because<br />
it spends seven months hibernating during the cooler<br />
dry-season of the southern winter (April to October),when<br />
food and water availability are low (Petter,1978;Hladik et al.,<br />
1980; Dausmann et al., 2004). When resting during the day<br />
and when hibernating,the dwarf lemurs occupy tree hollows<br />
either alone or with members of the family group (Dausmann<br />
et al., 2005). In the wild, female C. medius usually give<br />
birth to twins and in most cases reproduction only takes<br />
place every second year (Fietz and Dausmann, 2006). C.<br />
medius only reproduce after delayed emigration from their<br />
family and successful occupation of their own territory (thus<br />
in their third year at the earliest;Fietz et al.,2000).In addition,<br />
their life span is restricted by their size and is usually<br />
between 4 and 11 years for territory holders, and so opportunities<br />
to reproduce are limited (for most animals between<br />
one and three). Therefore, every young is a costly and valuable<br />
investment. Avoiding predation of their young should<br />
therefore be a critical part of parental care.<br />
Predators of C. medius include raptors (Madagascar harrier<br />
hawk Polyboroides radiatus, Madagascar buzzard Buteo brachypterus,<br />
Madagascar long-eared owl Asio madagascariensis),<br />
mammals (Fossa Cryptoprocta ferox, Narrow-striped mongooses<br />
Mungodictis decemlineata), and snakes (Madagascar<br />
Ground Boa Acrantophis madagascariensis, Madagascar Tree<br />
Boa Sanzinia madagascariensis, Malagasy Cat-eyed Snake Madagascarophis<br />
colubrinus) (Dausmann,submitted). The choice<br />
of an appropriately sized tree hollow in which to give birth<br />
can reduce attacks from many of these predators with the<br />
exception of snakes which are able to enter any hollow that<br />
can be used by C. medius.<br />
C. medius leave their tree hollows at sunset to forage alone<br />
but both sexes defend their shared territory. After the birth<br />
of their offspring, both parents take turns in guarding the<br />
young in the tree hollow throughout the night, while the<br />
other one forages.As the young get older, the proportion of<br />
time that parents spend guarding them gradually declines<br />
(Fietz and Dausmann, 2003). At the age of about two weeks,<br />
both parents leave the hollow and return only occasionally.<br />
During this time, the young are particularly vulnerable to<br />
predation,since they are unguarded.Later,the young accompany<br />
the parents during their nightly excursions.<br />
In this note, I want to describe evidence that adult C. medius<br />
can repel larger predators and therefore guarding or at least<br />
remaining within hearing range of the tree hollow is an effective<br />
measure against predation of their offspring. I report an<br />
observation in which a snake (Madagascarophis colubrinus)<br />
tried to attack two C. medius young within a tree hollow but<br />
was successfully repelled by the mother.<br />
Our observation occurred in the Kirindy C.F.P.F.forest,a dry<br />
deciduous forest near the west coast of Madagascar (60 km<br />
north east of Morondava) during a focal animal survey of a<br />
female C. medius on January 31st. For a more detailed description<br />
of the area see Ganzhorn and Sorg (1996).<br />
The female was a mother of two young aged two weeks.The<br />
male of the pair was also being observed.The female left the<br />
tree hollow after sunset at 18:57 hours and the male<br />
followed at 19:01 hours.The two young were left alone in the<br />
hollow within a dead tree (Malagasy name: Mapingo). The<br />
entrance of the hollow was 3 m above ground. The female<br />
started her regular patrol of the territory border, but suddenly<br />
abandoned the patrol at 19:29 hours when about 80 m<br />
from the hollow. She returned quickly to the hollow in<br />
almost a straight line.On approach to the hollow,it was clear<br />
that the two young who had been left alone within the<br />
hollow were making loud and constant distress calls. On a<br />
branch of the same tree at a height of about 1.5 m above<br />
ground was a large M. colubrinus (> 1 m length) eying the<br />
hollow. Even though this crepuscular or nocturnal snake is<br />
mainly terrestrial, scansorial behaviour is possible. The female<br />
approached the snake to within a few centimetres and<br />
actively attacked it, and the snake responded by striking<br />
towards the female.Both the adult female and the juvenile C.<br />
medius were loudly vocalizing constantly.After three min,the<br />
female seized the tail of the snake with both hands and bit it<br />
about 10 cm from the end. The snake tried to drop to the<br />
ground, but remained dangling in the female’s teeth. After<br />
10 s of wriggling and repeated attempts to strike the female<br />
the snake fell to the ground and moved away quickly. The<br />
female descended to about 1 m, observed the ground for a<br />
few minutes and then spent <strong>15</strong> min agitatedly observing the<br />
surroundings at a height of about 3 m and inspecting the tree<br />
hollow containing the young,who were still loudly vocalizing.<br />
For the next 3.5 hours the female was moving rapidly around<br />
within the territory, which is very unusual for a C. medius,<br />
frequently returning and checking the tree hollow with the<br />
young.She finally carried leaves into a new tree hollow about<br />
50 m away and separately carried both young to the new tree<br />
hollow. She did not return to sleep in the original tree hole<br />
for the next two months, even though it had been used<br />
frequently prior to this encounter.It seems puzzling that the<br />
male of the pair did not come to help during the attack.Since<br />
the male was followed simultaneously we know that at the<br />
time of the attack he was less than 20 m from the tree hollow,<br />
and clearly within hearing range of the distress calls. He<br />
returned about 30 min after the attack where he met the<br />
female and groomed her while she and the young continued<br />
making distress calls. Since reproduction in this species is a<br />
fairly rare event even including extra pair copulations, the<br />
possibility of siring offspring is restricted, and the male<br />
should have been highly motivated to defend his young in<br />
order to increase his fitness. Considering the high (obligate)<br />
paternal investment in guarding the young and the life-long<br />
pair bond, C.medius have a surprisingly high rate (ca 40 %) of<br />
extra pair young (Fietz et al.,2000;Schwensow et al.,2007).It<br />
is thought that the male cannot discriminate between intra<br />
pair and extra pair young and therefore cares for any offspring<br />
of his pair-partner,so as not to jeopardize the survival<br />
of his own young. Alternatively, paternal care of extra pair<br />
young could be an indicator for male quality or simply a tactic<br />
to maintain his bond with the female and so securing future<br />
mating possibilities in such a long-lasting relationship (Fietz<br />
and Dausmann, 2003). Genetic analyses showed that the<br />
male ("social father") of our observation was indeed only the<br />
genetic father of one offspring, but not the other (Schwensow<br />
et al., 2007). However, even if he was able to distinguish<br />
kin from non-kin, he should still have defended the tree hollow<br />
in order to protect his one own offspring.Interestingly,in<br />
the weeks before these observations, the male and the<br />
female had always spent the daily resting period together in<br />
the same tree hollow. However, the day after the predation<br />
attempt they slept apart, the male in the old, and the female<br />
together with the offspring in the new tree hollow. We<br />
cannot judge whether the male was unable to find the female<br />
in the new tree hollow, whether he chose to rest in the old<br />
hollow,or was prevented from entering the new tree hollow.<br />
Conclusion<br />
Clearly,the surveillance of offspring either directly within the<br />
tree hollow (additionally offering thermoregulatory advan-
Page 20 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
tages simultaneously; Fietz and Dausmann, 2003), or by staying<br />
within hearing range, does offer protection from attack<br />
by at least some predators even in this small species of primate.<br />
Acknowledgements<br />
This study was carried out under the "Accord de Collaboration"<br />
between Madagascar National Parks (MNP, formerly<br />
ANGAP), the University of Antananarivo and the University<br />
of Hamburg. We thank MNP, Chantal Andrianarivo, Jocelyn<br />
Rakotomala, Domoina Rakotomalala, the late Olga Ramilijaona<br />
and Daniel Rakotondravony for their collaboration and<br />
support. We acknowledge the authorization and support of<br />
this study by the Ministère de l’Environnement, des Eaux et<br />
Forêts et du Tourisme, MNP and the University of Antananarivo.C.Thurner<br />
was an invaluable observation companion.<br />
The study was financed by DFG (Ga 342/14) and DAAD.<br />
References<br />
Dausmann, K.H. submitted. Spoilt for choice – Choice of<br />
hibernacula and its influence on predation and energy expenditure<br />
during hibernation in Cheirogaleus medius. In: J.<br />
Masters; F. Génin (eds.). Leaping Ahead.<br />
Dausmann, K.H.; Glos, J.; Ganzhorn, J.U.; Heldmaier, G. 2004.<br />
Hibernation in a tropical primate. Nature 429: 825-826.<br />
Dausmann, K.H.; Glos, J.; Ganzhorn, J.U.; Heldmaier, G. 2005.<br />
Hibernation in the tropics:lessons from a primate.J Comp<br />
Physiol B 175: 147-<strong>15</strong>5.<br />
Fietz, J. 1999. Monogamy as a rule rather than exception in<br />
nocturnal lemurs: The case of the fat-tailed dwarf lemur,<br />
Cheirogaleus medius. Ethology 105: 259-272.<br />
Fietz,J.;Dausmann,K.H.2003.Costs and potential benefits of<br />
parental care in the nocturnal fat-tailed dwarf lemur (Cheirogaleus<br />
medius). Folia Primatol 74: 246-258.<br />
Fietz, J.;Dausmann,K.H.2006.Big is beautiful: fat storage and<br />
hibernation as a strategy to cope with marked seasonality<br />
in the fat-tailed dwarf lemus (Cheirogaleus medius). Pp.<br />
97-111. In: L. Gould; Sauther, M. L. (eds.). Lemurs: Ecology<br />
and Adaptation. Springer, Berlin Heidelberg New York.<br />
Fietz, J; Ganzhorn, J. U. 1999. Feeding ecology of the hibernating<br />
primate Cheirogaleus medius: how does it get so fat?<br />
Oecologia 121: <strong>15</strong>7-164.<br />
Fietz,J.;Zischler,H.;Schwiegk,C.;Tomiuk,J.;Dausmann,K.H.;<br />
Ganzhorn,J.U.2000.High rates of extra-pair young in the<br />
pair-living fat-tailed dwarf lemur, Cheirogaleus medius.Behav<br />
Ecol Sociobiol 49: 8-17.<br />
Ganzhorn,J.U.;Sorg,J.P.1996.Ecology and economy of a tropical<br />
dry forest in Madagascar. Primate Report 46-1, Göttingen.<br />
Hladik, C.M.; Charles-Dominique, P.; Petter, J. J. 1980. Feeding<br />
strategies of five nocturnal prosimians in the dry forest of<br />
the west coast of Madagascar.Pp.41-73.In:P.Charles-Dominique;H.M.Cooper;A.Hladik;C.M.Hladik;E.Pages;G.F.<br />
Pariente;A.Petter-Rousseaux;J.J.Petter;A.Schilling (eds.).<br />
Nocturnal Malagasy Primates:ecology,physiology and behaviour.<br />
Academic Press, New York.<br />
Müller, A.E. 1999. Social organization of the fat-tailed dwarf<br />
lemur (Cheirogaleus medius) in North-western Madagascar.<br />
Pp. 139-<strong>15</strong>7. In: B. Rakotosamimanana;H.Rasaminanana;<br />
J.U. Ganzhorn; S.M. Goodman (eds.). New Directions<br />
in Lemur Studies. Kluwer Academic/Plenum Publishers,<br />
New York.<br />
Petter, J.J. 1978. Ecological and physiological adaptations of<br />
five sympatric nocturnal lemurs to seasonal variations in<br />
food production. Pp. 211-223. In: D.J. Chivers; J. Herbert<br />
(eds.). Recent Advances in Primatology. Academic Press,<br />
New York.<br />
Schwensow, N.; Fietz, J.; Dausmann, K.H.; Sommer, S. 2007.<br />
Neutral versus adaptive variation in parasite resistance:<br />
importance of MHC-supertypes in a free-ranging primate.<br />
Heredity 99: 265-277.<br />
Lepilemur feeding observations from<br />
Northern Madagascar<br />
Andrew J. Lowin<br />
Society for Environmental Exploration / Frontier, 50-52<br />
Rivington Street, London EC2A 3QP, United Kingdom,<br />
research@frontier.ac.uk<br />
Lepilemur ankaranensis is the most northerly distributed<br />
member of the genus Lepilemur, with a range that extends<br />
south from Montagne d’Ambre National Park (Mittermeier<br />
et al.,2008).The behaviour and ecology of Lepilemur is poorly<br />
understood (Ratsirarson et al.,1987);this report summarises<br />
some preliminary observations of L. ankaranensis.<br />
Observations took place in a forest fragment (09°23.6E,<br />
46°07.3’S) 70km south of Antsiranana (Diego Suarez), near<br />
the town of Anivorano, west of the Route Nationale 6. The<br />
site is situated approximately 4 km south of the Mt.d’Ambre<br />
Park limit (Fig. 1). The area is heavily degraded, with only<br />
pockets of secondary dry deciduous forest remaining.<br />
Fig. 1: Study site in northern Madagascar.<br />
Casual feeding observations of L. ankaranensis took place<br />
from August 2009 to March 2010 (excluding the month of<br />
December). Observations were made in the first hour after<br />
sun set, with animals located using a flashlight. They were<br />
then followed,and any feeding bouts were recorded,with the<br />
plant species and food item noted. During this time, 32 % of<br />
all observations were of L. ankaranensis feeding on fruits,<br />
whereas all other observations were of leaf feeding. Five<br />
plant families were utilized for their fruits during the study:<br />
Moraceae, Verbenaceae, Rubiaceae, Pittosporaceae, and one<br />
that was not identified.<br />
As Lepilemur are thought to be predominantly folivorous<br />
(Ganzhorn et al., 2004; Thalmann and Ganzhorn, 2003), this<br />
proportion of fruit consumption seems to be high as compared<br />
to other Lepilemur species. For example, Thalmann<br />
(2001) found that during their study of L.edwardsi,only 0.3 %<br />
of 229 feeding bouts were feeding on fruits.<br />
Also during this study, L. ankaranensis was observed feeding<br />
on fruits with both Eulemur coronatus and Eulemur sanfordi in<br />
the same tree,also feeding on the same fruit,with no signs of<br />
aggression shown between any of the animals. A second<br />
study took place in June 2010, six dusk-till-dawn follows<br />
were carried out on consecutive nights, for a total of 64.3<br />
hours. Again, animals were located at dusk with a flashlight<br />
and followed until they returned to their sleeping sites in the
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 21<br />
morning.During this time,175 feeding observations were recorded,and<br />
no fruit was consumed.During this second study,<br />
a focal animal was observed to be chased out of a feeding<br />
tree by a female E.coronatus.The female E.coronatus then began<br />
eating the unripe fruits of the tree.It thus seems that during<br />
times of fruit abundance L. ankaranensis utilize fruits as a<br />
food resource along with several other lemur species occurring<br />
in the area. However, when food resources were not<br />
abundant in the dry season, only leaves were eaten and<br />
interspecific competition appears to be higher.<br />
On several occasions during this second study, leaf stems<br />
were snapped from trees and white tree exudates were consumed.<br />
Latex exudates are thought to be a toxic defence<br />
mechanism and therefore usually avoided by primates (Glander,1994),but<br />
latex feeding by Colobus spp.has also been observed<br />
(Mckey, 1978). Other lemur species, such as Phaner<br />
furcifer (Petter et al., 1975;Petter,1978;Thalmann,2006) and<br />
Mirza coquereli (Hladik,1979),are also known to feed on tree<br />
exudates. A review of the literature on exudate feeding in<br />
primates by Coimbra-Filho and Mittermeier (1977) suggested<br />
that tree exudates, in addition to simple sugars, protein,<br />
and minerals,may also provide a source of calcium.However,<br />
the latter authors also suggested that for most primates<br />
exudate feeding was rare and of little nutritional importance.<br />
This short report highlighted some behaviors of Lepilemur<br />
ankaranensis,a relatively poorly studied member of the Lepilemur<br />
genus. Further field work is required to examine in detail<br />
the previously discussed observations and to improve<br />
our knowledge of this species.<br />
References<br />
Coimbra-Filho, A.F.; Mittermeier, R.A. 1977. Tree-gouging,<br />
exudate-eating and the "short-tusked" condition in Callithrix<br />
and Cebuella. Pp. 105-1<strong>15</strong>. In: D.G. Kleiman (ed.). The<br />
Biology and Conservation of the Callitrichidae. Smithsonian<br />
Institution Press, Washington, D. C.<br />
Ganzhorn,J.U.;Pietsch,T.;Fietz,J.;Gross,S.;Schmid,J;Steiner,<br />
N. 2004. Selection of food and ranging behavior in a sexually<br />
monomorphic folivorous lemur: Lepilemur ruficaudatus.<br />
Journal of Zoology 263: 393-399.<br />
Glander, K.E. 1994. Nonhuman primate self-medication with<br />
wild plant foods.Pp.239-256.In:N.L.Etkin (ed.).Eating on<br />
the wild side: The Pharmacologic, Ecologic, and Social Implications<br />
of Using Nncultigens. University of Arizona<br />
Press, Tuscon.<br />
Hladik, C.M. 1979. Diet and ecology of prosimians. Pp. 307-<br />
357. In: A. Doyle; R.D. Martin (eds.). The Study of Prosimian<br />
Behavior. Academic Press, New York and London.<br />
Mckey,D.1978.Plant Chemical Defences and the Ranging Behaviour<br />
of Colobus Monkeys in African Rainforests. Ph.D.<br />
thesis, University of Michigan, Ann Arbor.<br />
Mittermeier,R.A.;Ganzhorn,J.U.;Konstant,W.R.;Glander,K.;<br />
Tattersall, I.;Groves,C.P.;Rylands,A.B.;Hapke,A.;Ratsimbazafy,<br />
J.; Mayor, M.I.; Louis, E.E.; Rumpler, Y.; Schwitzer, C.;<br />
Rasoloarison, R.M. 2008. Lemur diversity in Madagascar.<br />
International Journal of Primatology 29: 1607-1656.<br />
Petter, J.J. 1978. Ecological and physiological adaptations of<br />
five sympatric nocturnal lemurs to seasonal variations in<br />
food production. Pp. 211-223. In: D.J. Chivers; J. Herbert<br />
(eds.). Recent Advances in Primatology, <strong>Vol</strong>. 1: Behavior.<br />
Academic Press, New York and London.<br />
Petter,J.J.;Schilling,A.;Pariente,G.1975.Observations on the<br />
behavior and ecology of Phaner furcifer. Pp. 209-218. In: I.<br />
Tattersall; R.W. Sussman (eds.). Lemur Biology. Plenum<br />
Press, New York.<br />
Ratsirarson, J.; Anderson, J.; Warter, S.; Rumpler, Y. 1987. Notes<br />
on the Distribution of Lepilemur septentrionalis and Lepilemur<br />
mustelinus in Northern Madagascar. Primates 28:<br />
119-122.<br />
Thalmann, U. 2001. Food resource characteristics in two<br />
nocturnal lemurs with different social behavior: Avahi<br />
occidentalis and Lepilemur edwardsi.International Journal of<br />
Primatology 22: 287-324.<br />
Thalmann, U.; Ganzhorn, J.U. 2003. The Sportive Lemurs, genus<br />
Lepilemur.In:S.M.Goodman;J.Benstead (eds.).Natural<br />
History of Madagascar. The University of Chicago Press,<br />
Chicago.<br />
Thalmann, U. 2006. Lemurs - Ambassadors for Madagascar.<br />
Madagascar Conservation and Development 1: 4-8.<br />
Hypotheses on ecological interactions<br />
between the aye-aye (Daubentonia madagascariensis)<br />
and microhylid frogs of the<br />
genus Platypelis in Tsaratanana bamboo<br />
forest<br />
Andolalao Rakotoarison 1*, Solohery A. Rasamison 1,<br />
Emile Rajeriarison 2,David R.Vieites 3,Miguel Vences 4<br />
1Département de Biologie Animale, Université d’Antananarivo,<br />
BP 906, Antananarivo 101, Madagascar<br />
2Research assistant, Ranomafana National Park, BP 2, Fivondronana,<br />
Ifanadiana, Ranomafana 312, Madagascar<br />
3Museo Nacional de Ciencias Naturales-CSIC, C/José Gutiérrez<br />
Abascal 2, 28006 Madrid, Spain<br />
4Zoological Institute, Technische Universität Braunschweig,<br />
Spielmannstr. 8, 38106 Braunschweig, Germany<br />
*Corresponding author: andomailaka@gmail.com<br />
The aye-aye (Daubentonia madagascariensis) is the most distinctive<br />
of all lemurs.It is the only known living species of the<br />
Daubentoniidae (Simon and Meyer, 2001). The hands of the<br />
aye-aye are highly specialised,with long and slender third fingers<br />
that are used for precise grooming, mainly at face level,<br />
to get food into the mouth with rapid movements,and to tap<br />
on the bark of tree trunks to detect insect larvae or other<br />
arthropods (Goix,1993).When an aye-aye locates a cavity,it<br />
will anchor the upper incisors into the wood and then gnaw<br />
away at the wood with the lower incisors to make a pit<br />
(Erickson, 1995a, 1994). This unique manner of foraging for<br />
arthropods leaves traces of biting on the wood cover which<br />
are often used to ascertain the presence of the species even<br />
without an actual sighting (Duckworth,1993 and own observations<br />
of one of us, ER). During a recent herpetological inventory<br />
on the Tsaratanana massif in northern Madagascar,<br />
we noticed bamboo holes that were possibly caused or enlarged<br />
by foraging aye-aye,and we observed frogs living inside<br />
these cavities. Here we report these observations and posit<br />
a number of hypotheses on the possible ecological interactions<br />
among these species, with the goal of stimulating further<br />
studies.<br />
During a herpetological inventory in Tsaratanana (the highest<br />
mountain massif of Madagascar,which rises up to 2876 m<br />
above sea level) one of us (AR) carried out an ecological<br />
study on frogs of the genus Platypelis (Mycrohylidae: Cophylinae),<br />
from the 9th to the 22nd of June 2010. Specifically, we<br />
worked in a mountain forest bordering the temporary pond<br />
locally called Matsabory Maiky (S 14°09’04.09"- E 48°57’<br />
26.06" – 2,066 m elevation) - corresponding to campsite 2 on<br />
the trail from Mangindrano to the Maromokotro peak. The<br />
observed Platypelis occupy a specific microhabitat: the species<br />
live and breed inside the bamboo internodes which contain<br />
water and are accessible through small external holes.<br />
These frogs have endotrophic development: their non-feeding<br />
tadpoles develop inside the water retained in the tree<br />
holes and bamboo internodes. Based on a comparison with<br />
type material and DNA barcoding, we ascertained that the<br />
encountered Platypelis belong to two species described from
Page 22 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
the Tsaratanana massif: P. tsaratananaensis (most common)<br />
and the much larger P. alticola (more rare). Detailed data on<br />
the ecology and reproductive biology of these frogs will be<br />
published elsewhere. Approximately 754 bamboo trunks, at<br />
five different study sites, were inspected around the campsite.<br />
These sites each had four plots of 10 x 10 m areas. Out<br />
of the 754 trunks,we discovered in 162 of them,a total number<br />
of 204 internode segments; small rounded holes that<br />
were most probably made by insects like Dinoderus minutus<br />
(Delobel and Tran, 1993). According to these authors,<br />
Dinoderus minutus deposit eggs in bamboo internodes in<br />
which their larvae develop (Fig. 1d).<br />
At one of the sites (ca. 600 m east of the pond), we discovered<br />
some bamboo stems with remarkably different kinds of<br />
holes which allowed access to the hollow cavity of the<br />
internodes. Parts of the bamboo had been damaged in an irregular<br />
way. This appeared similar to what has already been<br />
described as typical damage caused by the gnawing activity of<br />
the aye-aye, whereby a freshly ripped-back piece is still attached<br />
and solid (Duckworth, 1993) (Fig. 1a-b).<br />
On 20 of the 281 bamboo trunks at this study site,we found<br />
similar damages,with a total of 71 holes which were more or<br />
less oval and measured 5.2-29.7 mm vertically and 1.5-<br />
8.2 mm horizontally. The diameter measurements of the<br />
non-damaged bamboo trunks were 5.3-54.2 mm, and those<br />
with holes were 5.3-48.9 mm.On several bamboo trunks we<br />
observed such holes in various internodes (1-6 m above the<br />
Fig.1:Traces of animals on the trunks of bamboo at the study<br />
site: (a) bamboo internode segment with an upper and a<br />
lower node attributed to the aye-aye; (b) segments of two<br />
bamboo trunks with a hole attributed to the aye-aye on the<br />
right and bite traces on the left; (c) traces attributed to<br />
aye-aye upper and lower incisors on a "virgin" bamboo trunk<br />
segment; (d) typical regular-shaped hole in a bamboo segment<br />
attributed to insects.<br />
Fig. 2: Animals observed within bamboo segments: (a) frogs:<br />
various specimens of Platypelis tsaratananaensis in one segment;<br />
(b) spider; (c) myriapod; (d) insect (cockroach).<br />
ground),and some internodes had an upper and a lower hole.<br />
Most importantly for the hypotheses drawn below, on some<br />
of the trunks without holes, we observed clear traces of<br />
gnawing that probably represent the upper and lower incisors<br />
of the aye-aye (Fig. 1b). According to our observations,<br />
80 % of all the holes found in the study site were caused by<br />
the activity of insects, and 20 % by the aye-aye.<br />
Bamboo internodes accessible by both kinds of holes were<br />
populated by Platypelis frogs as well as a variety of insects,spiders<br />
and centipedes (Fig. 2). At the study site where the bite<br />
traces ascribed to the aye-aye were discovered, the altogether<br />
282 holes (putatively made by insects) contained: 61<br />
Platypelis distributed in 24 different holes, 12 insects in 8 different<br />
holes, and 2 myriapods in 2 holes. In the 71 holes<br />
ascribed to the activity of the aye-aye, we observed 30<br />
Platypelis in 11 holes, 4 insects in 3 holes, and 0 centipedes.<br />
Based on these observations, we posit the following (partly<br />
alternative) hypotheses which require verification and further<br />
study:<br />
(1) We are confident that the observed marks at one of our<br />
study sites, similar to those noted by Duckworth (1993),are<br />
indeed caused by the activity of the aye-aye. Fresh bamboo<br />
stems are externally smooth and very strong, and it seems<br />
unlikely that any other mammal or even a bird could cause<br />
such damage. However, the possibility that these holes may<br />
be made by rats (such as Rattus rattus (which we collected at<br />
Matsobory Maiky), or Brachytarsomys) needs to be excluded<br />
by direct observations.<br />
(2) We assume that the aye-aye will typically search for bamboo<br />
internodes which already have small holes made by insects.This<br />
is because in such internodes there is a high likelihood<br />
of finding prey.In addition to insect larvae and other arthropods,<br />
tree-hole breeding frogs like Platypelis may also be<br />
consumed. In areas with high bamboo density, these frogs<br />
may constitute an import part of the aye-aye diet. If proven,<br />
this fact - that aye-ayes may eat frogs in addition to invertebrates<br />
- would be an interesting discovery in terms of<br />
Primatology.<br />
(3) Alternatively, the aye-aye may also gnaw holes into previously<br />
untouched bamboo segments. The bite traces we en-
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 23<br />
countered in such "virgin" internodes support this hypothesis.Reasons<br />
for this might either be a search for drinking water,<br />
or the search and detection of insect larvae which develop<br />
inside these internodes and which have not yet made a<br />
hole to emerge.<br />
(4) As a fourth and highly speculative hypothesis, the aye-aye<br />
may gnaw holes into "virgin" bamboo segments (or increase<br />
the size of pre-existing holes) as part of a long-term feeding<br />
strategy in which such holes are produced to make the bamboo<br />
segment suitable for colonization by arthropods and<br />
frogs. This would enable the aye-aye to "harvest" its food<br />
during a subsequent visit to the site several days later. Obviously,<br />
such a foresighted feeding strategy in a basal primate<br />
would be of extreme interest,but we are aware that alternative<br />
and more probable explanations exist.<br />
Detailed testing of these hypotheses will require long-term<br />
observations in an area of dense growth of large bamboo,<br />
probably including the deployment of a large number of camera<br />
traps and possibly hair traps to obtain evidence of aye-aye<br />
activity. Carrying out such studies at the site in Matsabory<br />
Maiky is difficult. It should be noted that the Tsaratanana<br />
massif is difficult to access. However, alternative sites, e.g. at<br />
Marojejy (Duckworth, 1993) might contain a large population<br />
of Platypelis (albeit other species) as well, and could be<br />
surveyed more systematically.<br />
Acknowledgements<br />
A warm thanks to the Directorate of Waters and Forests<br />
and the Head of the Offices of Madagascar National Parks at<br />
Mangindrano and Ambanja for the research permits on<br />
Tsaratanana. We are also indebted to the many people who<br />
have logistically assisted in our expedition, especially to the<br />
local guides (Faly and Levaovao) from Mangindrano.<br />
References<br />
Delobel, A.; Tran, M. 1993. Les coléoptères des denrées alimentaires<br />
entreposées dans les regions chaudes. Faune<br />
tropical XXXII: 98-99.<br />
Duckworth,J.W.1992.Feeding damage left in bamboos,probably<br />
by aye-ayes (Daubentonia madagascariensis).International<br />
Journal of Primatology 14: 927-931.<br />
Erickson, C.J. 1994. Tap-scanning and extractive foraging in<br />
aye-ayes, Daubentonia madagascariensis. Folia Primatologica<br />
62: 125-135.<br />
Erickson, C.J. 1995a. Feeding sites for extractive foraging by<br />
the aye-aye,Daubentonia madagascariensis.American Journal<br />
of Primatology 35: 235-240.<br />
Goix,E.1993.L’utilisation de la main chez le aye-aye en captivité<br />
(Daubentonia madagascariensis) (Prosimiens, Daubentoniidés).<br />
Mammalia 57: 171-188.<br />
Milliken,G.W.;Ward,J.P.;Erickson,C.J.1991.Independent digit<br />
control in foraging by the aye-aye (Daubentonia madagascariensis).<br />
Folia Primatologica 56: 219-224.<br />
Simon,E.L.;Meyer,D.,2001.Folklore and beliefs about the aye<br />
aye (Daubentonia madagascarienis) Lemur News 6:11-16.<br />
Discovery of crowned sifaka (Propithecus<br />
coronatus) in Dabolava, Miandrivazo, Menabe<br />
Region<br />
Josia Razafindramanana 1*, Rija Rasamimanana 2<br />
1Groupe d’Etude et de Recherche sur les Primates de Madagascar<br />
(GERP), Lot 34 Cité des Professeurs Fort Duchesne,<br />
Ankatso, Antananarivo 101, Madagascar.<br />
2Pan African Mining Madagascar (PAMM), Lot 137 II AN<br />
Analamahitsy, Antananarivo 101, Madagascar.<br />
*Corresponding author: r_josia@hotmail.com<br />
Key words: Propithecus coronatus, Dabolava, distribution,<br />
lemurs, Indridae<br />
The crowned sifaka Propithecus coronatus was until recently<br />
regarded as one of four subspecies of P. verreauxi, family<br />
Indridae, which occur throughout western and southern<br />
Madagascar (Muller et al., 2000; Mittermeier et al., 1994; Tattersall,<br />
1986; Wilmé and Callmander, 2006). Recent taxonomic<br />
revisions (Mittermeier et al., 2008) have promoted all<br />
four subspecies to full species status (Mittermeier et al.,<br />
2006). However, there is considerable debate about the validity<br />
of P. coronatus, and especially its relationship with P.<br />
deckeni (Mittermeier et al.,2008),due to the physical similarities<br />
and close geographical distributions of these taxa,including<br />
apparent sympatry at some sites (e.g. Tattersall, 1986;<br />
Curtis et al.,1998;Muller et al.,2000;Groves,2001;Thalmann<br />
et al., 2002).<br />
P. coronatus was previously assigned to the IUCN conservation<br />
rating "Critically Endangered",but has since been moved<br />
into the "Endangered" category; nevertheless, the distribution<br />
range and the ecology of this species are not yet well<br />
understood (IUCN, 2008). Crowned sifakas are diurnal, and<br />
their habitat is characterised by dry deciduous forests and<br />
mangroves (Petter and Andriatsarafara, 1987). They live in<br />
groups of two to eight individuals, with home ranges from<br />
1.2–1.5 ha.They feed mainly on buds,green fruits and mature<br />
leaves (Muller,1997).It is known that they reproduce seasonally,<br />
with females giving birth every 2-3 years (Curtis et al.,<br />
1998; Mittermeier et al., 2006). Compared to other lemurs,<br />
their reproduction rate is very slow, making recovery of<br />
small populations even more problematic.<br />
The newly discovered crowned sifaka population is situated<br />
at Amboloando (UTM WGS 84,N 7822351 E 580189) in the<br />
Commune of Dabolava in central Madagascar,and is the most<br />
southerly record of the species.Amboloando lies about 4 km<br />
from Dabolava village, and 40 km to the southeast of Miandrivazo.Amboloando<br />
comprises 7 ha of dry semi-deciduous,<br />
secondary forest that exhibits the characteristics of riverine<br />
forests, consisting of deciduous as well as evergreen trees<br />
such as Acacia sp.,Nastus sp.and Macaranga sp.The altitude is<br />
about 600 m above sea level,and the area is characterized by<br />
a clearly defined wet and dry season.The sifaka population is<br />
composed of a single group,which constituted six adults and<br />
one juvenile when first discovered in June 2009 (Razafindramanana,<br />
2009). One of the adult males disappeared later in<br />
the year, presumed dead, leaving six individuals remaining.<br />
The animals appear to be classic P.coronatus (Fig.1),but some<br />
individuals show pelage colour variation, with dark fur on<br />
their back and arms (Fig.2).Behavioural studies of the group<br />
are underway,and a preliminary community-based conservation<br />
program has been established at the site, involving several<br />
organisations including GERP, The Aspinall Foundation,<br />
SAHA and Pan-African Mining Madagascar.Forests in Amboloando<br />
and the surrounding area are heavily degraded. Different<br />
factors threaten the survival of this species in Madagascar:in<br />
contrast to the other sites such as Anjamena (Muller,2000),hunting<br />
does not occur in Amboloando,partly due<br />
to the sifaka being regarded as holy by the local people.<br />
Therefore,other threats such as habitat destruction through<br />
slash-and-burn agriculture to make way for pasture for livestock,charcoal<br />
production,and mining exploitation affect the<br />
sifaka group.<br />
Surveys in the vicinity of Dabolava suggest that this is the<br />
only group of P.coronatus remaining in that area,despite local<br />
people claiming that other groups were present between 5<br />
and 10 years ago. Therefore, it appears that habitat destruc-
Page 24 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
Fig.1:Crowned sifaka in the Amboloando<br />
Forest (top).<br />
Fig. 2: Crowned sifaka with dark colour<br />
on its arms (right).<br />
Fig. 3: Map showing the area of discovery and survey in Dabolava.<br />
tion for local livelihoods has resulted in the almost complete<br />
extirpation of crowned sifaka in the area, probably due to a<br />
combination of habitat loss and food scarcity.<br />
Some studies describe the range of P.coronatus as broadly restricted<br />
to the region between the rivers Betsiboka and<br />
Mahavavy (Muller et al., 2000; Wilmé and Callmander, 2006),<br />
with a population density of 173 individuals/km² at Anjamena<br />
(Muller, 2000). In an analysis of the distribution of lemurs in<br />
central western Madagascar, Thalman and Rakotoarison<br />
(1994) suggested that the faunal region bounded by the<br />
Betsiboka,the central highlands,the river Tsiribihina and the<br />
Mozambique Channel can be divided into four sub regions.<br />
These sub regions are separated by the three rivers: Mahavavy,<br />
Manambaho and Manambolo, but are interconnected<br />
with the Bongolava Massif.The discovery of a crowned sifaka<br />
population in Dabolava, which is located in the south of the<br />
central highlands sub region, confirms the hypothesis that<br />
the historical range of this species might spread along the<br />
central highlands of Madagascar. The record of a group of<br />
crowned sifaka in Andranotonga, slightly north of the Mahajilo<br />
River, was cited by Tattersall (1986). The present report<br />
appears to be the first location of P.coronatus to the south of<br />
this river.<br />
The Mahajilo is a tributary of the Tsiribihina<br />
River, which is considered to<br />
represent the north-western limit of P.<br />
verreauxi (Mittermeier et al., 2006). P.<br />
coronatus is therefore unlikely to be<br />
found much further south or southwest<br />
than Dabolava. Only 80 km<br />
south-west of Dabolava, a population<br />
of P. verreauxi is known from Ambatolahy<br />
(SAHA,2009),which lies within<br />
the Ambararata/Londa protected area<br />
complex (Fig. 3). Another tributary of<br />
the Mahajilo, the Mania River, lies between<br />
Ambatolahy and Dabolava and<br />
may therefore represent the distributional<br />
limit of P. coronatus and P. verreauxi<br />
in the south of Madagascar. A<br />
conservation programme and restoration<br />
of the remaining habitat with<br />
the local people are needed to save<br />
this population of P. coronatus.<br />
Acknowledgments<br />
I thank GERP – Groupe d’Etude et de<br />
Recherche sur les Primates de Madagascar,<br />
The Aspinall Foundation, Cotswold<br />
Wildlife Park, SECAS, Belfast<br />
Zoo, Besancon Museum and Parc<br />
Zoologique de Paris for funding this<br />
research. Many thanks to Dr Jonah<br />
Ratsimbazafy and Tony King for discussions<br />
about the project implementation.I<br />
am grateful for permission and<br />
assistance in the field from the Direction<br />
Régionale de l’Environnement et<br />
des Forêts and the Commune of<br />
Dabolava.I also thank Pan African Mining<br />
Madagascar for providing accommodation<br />
in their lovely camp site.<br />
References<br />
Curtis, D.J.; Velo, E.-O.; Raheliarisoa; Zaramody, A.; Muller, P.<br />
1998.Surveys on Propithecus verreauxi deckeni,a melanistic<br />
variant, and P.v. coronatus in Northwest Madagascar. Oryx<br />
32: <strong>15</strong>7-163. Groves, C.P. 2001. Primate taxonomy. Smithsonian<br />
Institution Press, Washington, D.C.<br />
Hawkins,A.F.A.;Durbin,J.C.;Reid,D.B.1998.The primates of<br />
the Baly Bay area,north-western Madagascar.Folia Primatologica<br />
69: 337-345.<br />
IUCN. 2008. 2008 IUCN Red list of threatned species.<br />
www.iucnredlist.org.<br />
Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.;<br />
Mast,R.1994.Lemurs of Madagascar.1st ed.Conservation<br />
International, Washington, D.C.<br />
Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis E.E.;<br />
Langrand, O.; Ratsimbazafy, J.; Rasoloarison, R.; Ganzhorn,<br />
J.U.; Rajaobelina, S.; Tattersall, I.; Meyers, D.M. 2006.<br />
Lemurs of Madagascar. 2nd ed. Conservation International,<br />
Washington, D.C.<br />
Mittermeier,R.A.;Ganzhorn,J.U.;Konstant,W.R.;Glander,K.;<br />
Tattersall, I.;Groves,C.P.;Rylands,A.B.;Hapke,A.;Ratsimbazafy,<br />
J.; Mayor, M.I.; Louis Jr., E.E.; Rumpler, Y.; Schwitzer,<br />
C.; Rasoloarison, R.M. 2008. Lemur diversity in Madagascar.<br />
International Journal of Primatology 29 (6): 1607-<br />
1656.<br />
Muller,P.1997.The behaviour and ecology of the crowned sifaka<br />
(Propithecus verreauxi coronatus) in north west Madagascar.<br />
Unpublished Ph.D. thesis, University of Zurich.
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 25<br />
Muller,P.A.;Velo,E.-O.;Raheliarisoa;Zaramody A.;Curtis.D.J.<br />
2000. Surveys of five sympatric lemurs at Anjamena,<br />
northwest Madagascar. African Journal of Ecology 38:<br />
248-257.<br />
Petter, J.-J.; Andriatsarafara. F. 1987. Conservation Status and<br />
distribution of lemurs in the west and northwest of Madagascar.<br />
Primate Conservation 8: 169-171.<br />
Razafindramanana, J. 2009. Propithecus coronatus on the verge<br />
of extinction:Help to save them:GERP,Antananarivo,Madagascar,<br />
6.<br />
SAHA. 2009. Crèation de Nouvelle Aire Protegee pour le<br />
complexe Ambararata/Londa.<br />
Tattersall,A.I.1986.Notes on the distribution and taxonomic<br />
status of some species of Propithecus in Madagascar. Folia<br />
Primatologica 46: 51-63.<br />
Thalmann, U.; Rakotoarison, N. 1994. Distribution of lemurs<br />
in central western Madagascar, with a regional distribution<br />
hypothesis. Folia Primatologica 63: <strong>15</strong>6-161.<br />
Thalmann, U.; Kümmerli, R.; Zaramody, A. 2002. Why Propithecus<br />
verreauxi deckeni and P. v. coronatus are valid taxa –<br />
quantitative and qualitative arguments. Lemur News 7:<br />
11-16.<br />
Wilme, L.; Callmander M.W. 2006. Les populations reliques<br />
de primates: les Propithèques. Lemur News 11: 24-31.<br />
Inferences about the distant past in Madagascar<br />
Elwyn L. Simons<br />
Duke Lemur Center, Division of Fossil Primates, Dept. Evol.<br />
Anthropology, Duke University, 3705 Erwin Road, Durham,<br />
NC 27705, USA, esimons@duke.edu<br />
From Etienne de Flacourt (1658), the following is an English<br />
translation, from the original French, of an entry in his book<br />
"Histoire de la Grande Isle Madagascar:"<br />
"Tretretretre or Tratratratra. It is a large animal like a calf of<br />
two years old, with a round head and the face of a man: the<br />
fore feet are like those of a monkey (or ape),and the hind feet<br />
also.It has curly (or frizzy) hair,a short tail and ears like those<br />
of a man. It resembles the "Tanacht" described by Ambroise<br />
Paré. It can be seen near the pond of the Lipomami [tribe]<br />
and in that region is where it can be found. It is a highly<br />
solitary animal,the people of that country have a great fear of<br />
it and flee from it as it also does from them." From this context,it<br />
is not clear whether Flacourt actually had seen this animal.<br />
It is well known that in Madagascar village people tend to<br />
name lemurs after the sounds they make, following a sort of<br />
onomatopoeic pattern for animal names such as occurs in<br />
the case of the cuckoo bird in English. For instance, the<br />
ground predator alarm call of species of genus Propithecus is a<br />
loud "si-i-fak!" cry and so the name of this animal in the Malagasy<br />
language is "Sifaka".The mouse lemur makes a chittering<br />
alarm call and has the name "T’sit-sihy".The Avahi,a nocturnal<br />
lemur, has one call that sounds like the word "avahi!" In the<br />
case of Flacourt’s animal the name Tretretretre or Tratratratra<br />
is definitely onomatopoeic and sounds like an alarm<br />
bark–it is not unlike the bark alarm call of the southeastern<br />
Madagascan lemur Propithecus edwardsi,or that of another lemur<br />
related to it, Indri indri, or even the alarm bark of the<br />
chimpanzee.<br />
Hence, I have often considered this term, or name, to be a<br />
"fossil" sound and it seems likely that it would have been a<br />
replication, by members of the Lipomami tribe of the alarm<br />
call of this giant lemur when they told Flacourt about the animal.<br />
The location of the Lipomami region in southeastern<br />
Madagascar is known today (Tattersall,1982).Thus Flacourt’s<br />
name for the animal may be the only known "fossil" sound.<br />
There are frequent references by various scientists all agreeing<br />
that Flacourt must have been describing,in the above passage,<br />
one of the giant extinct lemurs; but which one? All lemurs<br />
have hands and feet like those of monkeys, but wavy<br />
hair is more restricted–mainly to members of the Indriid<br />
group or taxonomic family [this family includes only species<br />
of the extant genera Propithecus,Indri,and Avahi] and incidentally<br />
they all have rounded ears–like those of a man. One<br />
large giant extinct lemur of the south and southwest of the<br />
island is known as Palaeopropithecus ingens and taxonomists<br />
generally agree that genus Palaeopropithecus is related to the<br />
family Indriidae (Orlando et al., 2008), where curly or frizzy<br />
hair occurs. A number of scientists have speculated that<br />
Flacourt’s Tretretretre lemur belonged to the genus Megaladapis<br />
(Tattersall, 1982; Mittermeier et al., 1994), which also<br />
occurs as fossils from the southern part of the island, but<br />
species of this genus have a large snout and could never be<br />
described as having a round head. Moreover, the distal ends<br />
of the nasal bones in species of this genus are elongated and<br />
expanded and, in life, there must have been an expanded or<br />
bulbous nose or even a trunk. Hence, one could never say<br />
that the creature had "a face of a man." Differing from Megaladapis,Palaeopropithecus<br />
ingens does have a rounded humanlike<br />
head with forward directed eyes and a small face.In addition<br />
to all these other features,the living species Indri indri or<br />
babacoot,is the only lemur that has a short tail.In addition to<br />
this,the most complete skeleton of Palaeopropithecus,recovered<br />
by a Duke expedition in 1983, and the only associated<br />
skeleton of this animal ever found includes a sacrum that diminishes<br />
posteriorly and could only hold a very small and<br />
short tail.In opposition to all these conclusions,it can be said<br />
that Palaeopropithecus ingens could not possibly be construed<br />
to have been the size of a calf of two years in age–nor,in fact,<br />
would any of the extinct giant lemurs have been that large.<br />
Nevertheless, it is well known that exaggeration surrounds<br />
stories about such little known animals, and also Malagasy<br />
cattle tend to be small. For these reasons it would appear<br />
that the Tretretretre was a Palaeopropithecus species–a conclusion<br />
also implied by Godfrey and Jungers (2002). A year<br />
later these authors reconfirm the same position (Godfrey<br />
and Jungers, 2003).<br />
Between 1994 and the year 2003 teams from the Duke<br />
Lemur Center excavated fossils at two caves called Akomaka<br />
and Ankilitelo in southwestern Madagascar on the Mikoboka<br />
Plateau north of Tulear. This region is Madagascar’s most<br />
extensive and stratigraphically thickest calcareous plateau.<br />
Discoveries made at both of these caves show that the<br />
southwestern part of the island was inhabited comparatively<br />
recently by several giant lemurs including Palaeopropithecus<br />
ingens and Megaladapis edwardsi. More importantly, these<br />
species lived relatively recently (Godfrey and Jungers, 2002),<br />
as evidenced by radiocarbon dating based on specimens<br />
from Ankilitelo (Megaladapis at 630 ± 50 years B.P. and<br />
Palaeopropithecus, 510 ± 80 years B.P. The latter of these<br />
dates (calculated in 1996) ranged from 1406 to <strong>15</strong>66 years<br />
AD and falls into historic times. These are relatively recent<br />
ages,not so far from the date of Flacourt’s observation of the<br />
Tretretretre which could have been at any time after he was<br />
named Governor of Ft.Dauphin Madagascar in 1648;approximately<br />
350 years ago. A more recent date determined in<br />
2008 on a Cryptoprocta bone from Ankilitelo gives a similar<br />
age to that of the Palaeopropithecus, estimated as between<br />
1408 and 1488 (Simons,1997;Muldoon et al.,2009;Muldoon<br />
and Simons, 2007). This suggests that the small mammals<br />
accumulated more or less contemporaneously with the giant<br />
lemurs. [The cave name, Ankilitelo, means "at the three kili<br />
(tamarind) trees" but no such trees grow there now. Mala-
Page 26 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
Fig. 1: Harpagophytum grandidieri burr or "hitchhiker" from<br />
southern Madagascar.These barbs are approximately 9.5 cm<br />
across.<br />
gasy caves can be named for a nearby settlement or village.At<br />
present there is no nearby hamlet with such a name but in<br />
the past there could have been. Kili trees often grow in<br />
villages there.]<br />
At this point another speculation can be introduced.Pastoral<br />
grass burning has reduced present day forests on the Mahafaly<br />
Plateau, north of Tulear, to the ridges of hills and many<br />
present day plant species occurring in them must be the<br />
same as those of 500-1000 years ago. The remaining forests<br />
near Ankilitelo represent both spiny thicket and succulent<br />
woodland regions and examples of the dry deciduous forest<br />
are nearby (Simons, 1997; Muldoon et al., 2009). In general,<br />
grass burning in that region, near the Ankilitelo cave, is arrested<br />
from spreading by outcrops of limestone on the<br />
slopes of hills. In addition, it would appear that surviving forests<br />
on hilltops of the Ankilitelo region remain as they were a<br />
few hundred years ago.The surface finds of small mammals in<br />
Ankilitelo (34 species) do reflect those of a few hundred<br />
years ago–when the giant lemurs existed and when the cave<br />
was serving as a natural trap–these environmental conditions<br />
were similar to those of the present.The fauna suggests<br />
that, perhaps, the region then was slightly more humid, and<br />
definitely the forests of the region have undergone fragmentation<br />
(Simons, 1997; Muldoon and Simons, 2007). Each such<br />
natural trap cave serves as a window in time because, as the<br />
solution cavities open mainly from below,the fauna that fall in<br />
will only begin to collect when there is a surface opening.<br />
More dates are being determined for fossils from Ankilitelo<br />
but the window concerned here did not open long enough<br />
ago that the 34 small mammal species are different from<br />
those now extant in the region or relatively nearby.The commonest<br />
giant lemur at Ankilitelo is the awkwardly constructed<br />
Palaeopropithecus that presumably could not support<br />
itself on all fours on the ground. Once having fallen to<br />
earth it would have been restricted to swimming, sloth-like<br />
motions and this perhaps explains why so many P. ingens fell<br />
into the pit. Also, Godfrey and Jungers (2003) report a Malagasy<br />
tradition (p. 258) that an "ogre with the body of an animal<br />
but the face of a human" could be made helpless on<br />
smooth rock surfaces. Such clumsiness of the sloth-like<br />
Palaeopropithecus would account both for its abundance in<br />
Ankilitelo and its presumed inabilities on the ground. Whatever<br />
the pelage of this animal was like,or indeed that of any of<br />
the giant lemurs, their fur could not have resisted picking up<br />
burrs and other hitchhikers from the southwestern forests<br />
as has been noted with modern lemurs (personal observa-<br />
tions of Michelle Sauther).A present day student of behavior,<br />
Sauther,has observed individual Lemur catta that had become<br />
entangled with the large seeds of what is often called Uncarina<br />
grandidieri–but more properly,because of an earlier date<br />
of description, this species should be assigned to genus Harpagophytum–meaning<br />
"snatcher plant". This plant of southwestern<br />
Madagascar has amazing "hitchhikers" about 2.5 to 3<br />
inches across each of which has 30/35 protruding spines<br />
approximately 1.5 inches long (see Fig.l).Each of these spines<br />
is, in turn, tipped by 4 recurving fishhook-like projections.<br />
This huge seed pod is something that it seems would only<br />
have evolved to be transported by a much larger animal than<br />
any now extant on the Island–presumably a giant lemur like<br />
species of such southern genera as Palaeopropithecus, Megaladapis,<br />
or Archaeolemur or even transportation by the elephant<br />
bird (see below).<br />
Modern botanists report that the pasty pollen of this plant is<br />
spread by pollen-eating beetles, who, after feeding on pollen<br />
from the anthers, get it all over themselves and when covered<br />
by pollen fly from flower to flower where pollen is<br />
transmitted from them to the stigma.This sort of pollination<br />
may be the principal fertilization process, but lemur transport<br />
of these seed burrs does occur today (personal observations<br />
of Michelle Sauther), and must have also done so in<br />
the past. Working at the Beza Mahfaly Special Reserve, in<br />
southwestern Madagascar,Sauther has seen Lemur catta individuals<br />
with Harphagophytum burrs stuck on the face, feet,<br />
and tail. However, she has not seen them attached to Propithecus<br />
verreauxi; a second larger, diurnal lemur species which<br />
occurs at Beza but is more arboreal than L. catta. These dry<br />
seed pods would naturally attach to the skin, not necessarily<br />
fur,of any passing animals and be carried while attached until<br />
its spines were broken enough for the seed to drop off.It was<br />
recently suggested that dispersal of these seed pods might<br />
have been carried out primarily by the extinct elephant birds<br />
of Madagascar (Midgley and Illing, 2009). The authors presenting<br />
this view argue that the mature fruit more often accumulate<br />
on the ground as "trample burrs" and so are more<br />
likely to stick to the feet of these extinct giant ratites than to<br />
fur of arboreal animals. I suspect, however, that the giant lemurs<br />
did not always stay high up in trees but were often on<br />
or near to the ground. The mature terminal hooks of the<br />
Harpagophytum (Uncarina) burr have evolved so as to attach<br />
to any extremity,not necessarily fur.Also it is of interest that<br />
these plants are often called the "Mouse-trap tree" or "Grapple<br />
tree". These species belong in the sesame family (Pedaliaceae)<br />
and typically constitute shrubs or small trees. It is<br />
told that Malagasy people sometimes collect and put together<br />
bunches of these seeds and place cheese or other attractants<br />
at the center of the bunch. They then use this device<br />
to trap rats and mice: Hence, the origin of the common<br />
name.<br />
References<br />
Etienne de Flacourt.1658.Histoire de la grande Isle Madagascar,<br />
2 vols. Chez G. de Lvyne, Paris.<br />
Godfrey, L. R.; Jungers, W. L. 2002. Quaternary fossil lemurs.<br />
Pp. 1-530. In W.C. Hartwig (ed.). The Primate Fossil<br />
Record:Cambridge Studies in Biological and Evolutionary<br />
Anthropology No.33.Cambridge Univ.Press,Cambridge,<br />
UK.<br />
Godfrey,L.R.;Jungers,W.L.2003.The Extinct Sloth Lemurs of<br />
Madagascar. Evol. Anth. 12: 252-263.<br />
Midgley, J. J; Illing, N. 2009. Were Malagasy Uncarina fruits dispersed<br />
by the extinct elephant bird? So. Af. J. Sci. 105<br />
(11/12): 467-499.<br />
Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.;<br />
Mast, R.B. 1994. Lemurs of Madagascar. Conservation<br />
International, Washington, D.C., USA.
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 27<br />
Muldoon, K.M.; de Blieux, D.D.; Simons, E.L.; Chatrath, P.S.<br />
2009.The subfossil occurrence and paleoecological significance<br />
of small mammals at Ankilitelo cave,southwestern<br />
Madagascar. Journ. Mammology 90 (5): 1111-1131.<br />
Muldoon,K.M.;Simons,E.L.2007.Ecogeographic Size Variation<br />
in Small-Bodied Subfossil Primates From Ankilitelo,<br />
Southwestern Madagascar.Am.J.Phys.Anth.134:<strong>15</strong>2-161.<br />
Orlando, L.; Calvignac, S.; Schnebelen, C.; Douady, C.J.; Godfrey,<br />
L.R.; Hänni, C. 2008. DNA from extinct giant lemurs<br />
links archaeolemurids to extant indriids.BMC Evolutionary<br />
Biology 8: 121.<br />
Simons,E.L.1997.Lemurs:Old and New.In S.M.Goodman;B.<br />
D.Patterson (eds.). Natural Change and Human Impact in<br />
Madagascar. Smithsonian Inst. Press, Washington, D.C.<br />
Tattersall, I. M. 1982. The Primates of Madagascar. Columbia<br />
Univ. Press.<br />
Husbandry guidelines for mouse lemurs<br />
at Paris Zoo<br />
Delphine Roullet<br />
Parc Zoologique de Paris, MNHN, 53 avenue de Saint Maurice,<br />
75012 Paris, France, roullet@mnhn.fr<br />
There are two species of mouse lemur in captivity in Europe:<br />
the grey mouse lemur, Microcebus murinus, and the Goodman’s<br />
mouse lemur, Microcebus lehilahytsara (Pes, 2009). The<br />
European captive population of grey mouse lemurs was<br />
established at the end of the 1960s and is now composed of<br />
165 individuals (778.77.10; Pes, 2009),distributed in 29 institutions.<br />
The population of this species is of unknown origin.<br />
According to recent morphological measurements (Pes,<br />
2009) and preliminary results of mtDNA studies (Roos,<br />
2008, in Pes, 2009), this population can be divided into two<br />
pure lineages:one composed of pure breed animals from the<br />
region of Vohimena (SW Madagascar),and the other of pure<br />
breed animals from the region of Mandena (SE Madagascar).<br />
A third lineage is composed of hybrids between the two pure<br />
lineages (Pes, 2009).<br />
The European captive population of Goodman’s mouse lemurs<br />
was established in 2005 and is currently composed of<br />
62 individuals (33.29; Pes, 2009) distributed in two institutions.<br />
The first animals were imported from the area of<br />
Andasibé, Madagascar. They were recognized as a new species<br />
when they arrived in Europe (Rübel, pers. comm.).<br />
The Parc Zoologique de Paris has a success story with the<br />
grey mouse lemurs. The most important group arrived in<br />
December 1990 and was composed of 79 individuals. The<br />
origin of the animals that arrived in the 1980s is unknown.<br />
According to the analysis of the European captive population<br />
(conducted by Tomas Pes for the ESB), the animals coming<br />
from Paris appear to be hybrids between the two pure lineages<br />
described above (Pes, pers. comm.).<br />
The first births occurred in 1991 only a few months after the<br />
arrival of the first animals.During the period of 1991 to 2004,<br />
when the last grey mouse lemurs eventually left the zoo after<br />
the closing of the nocturnal area, there had been a total of<br />
224 successful births (young surviving longer than two<br />
months), with an 86.5 % birth success rate. 1994 was the<br />
most prolific year with 56 successful births. The colony of<br />
grey mouse lemurs in Paris occasionally reached more than<br />
<strong>15</strong>0 individuals.<br />
Before 2001, we didn’t know much about the animals, especially<br />
the composition of the groups.Eleven females were the<br />
founders of the colony in Paris. Since the identity of the fathers<br />
was not recorded, the filiations were only built from<br />
the females. Potentially,28 males could have been the founders<br />
of the colony.<br />
Before 2002, most of the females lived alone in small cages<br />
and were introduced to males (of various group sizes) only<br />
during the few days of oestrus. The females were kept isolated<br />
again afterwards.The young were separated from their<br />
mothers just after their weaning to join a young animals<br />
group.<br />
In 2002 we decided to implement some changes in the management<br />
of the colony in order to improve the wellbeing of<br />
the animals: to increase the space available to them, to rearrange<br />
the enclosures according to the wild habitat of the animals,<br />
to carry out enclosure enrichment, and to re-constitute<br />
the groups to make them more similar to the ones observed<br />
in the wild. The following husbandry guidelines were<br />
established according to the new management of the colony<br />
set up in 2002.<br />
Only single sex groups could be seen by the public.The breeding<br />
groups were kept in a separated building. Moreover,<br />
from 2002 onwards,we limited the number of births (around<br />
10 per year) in order to be able to keep all the animals in<br />
good conditions (and no longer in small cages as had been<br />
done in the past).<br />
Facility standards<br />
1. Enclosure<br />
Size: The enclosure should have a minimum total floor size<br />
of 4 m² with a minimum height of 2 m for both male and female<br />
groups.For a mother with her young,the enclosure can<br />
be smaller during the first month.After this period,the young<br />
start to explore their environment and need more space.<br />
Temperature: 20°C (18-22° C). Not below 18° C. Below<br />
this temperature, the animals enter torpor. Torpor can also<br />
be provoked by intense stress such as prolonged capture of<br />
an animal.<br />
Inside Humidity: 50-70 %.<br />
Lighting and photoperiod:Similar to that found in Madagascar<br />
or Europe but the photoperiod must vary during the<br />
year for breeding.<br />
Furniture: Dense environment with thin branches and<br />
leaves. The animals need to have many places to hide from<br />
people and also from each other when they live in groups,especially<br />
when the animals are unrelated.<br />
Nest box (see Fig. 1):<br />
Size: 12x12x12 cm<br />
Entry diameter: 5 cm<br />
It’s very important to provide one nest box per animal,in<br />
different places, even if they sleep together. This allows<br />
them to be alone if they want to be.<br />
Fig.1:Grey mouse lemurs (Microcebus murinus) in nest box at<br />
Paris Zoo. (Photo: F.-G. Grandin, MNHN)
Page 28 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
2. Feeding and watering<br />
Two feeding spots are needed if there are more than two<br />
adults in the group.Additional feeding spots are required according<br />
to the size of the group.If there are not enough nest<br />
boxes or feeding spots,the animals can be very aggressive towards<br />
one another, sometimes causing serious wounds,<br />
mostly to the tail. One water bottle per enclosure is sufficient.<br />
Diet for 1 animal/day:<br />
Monday: 1 tea-spoon of gruel* +3folivorous pellets<br />
Tuesday: 1/8 of apple + 2 slices of carrot + 3 folivorous pellets<br />
Wednesday:1/8 of apple + 1/8 of pear + 3 folivorous pellets<br />
Thursday: 1 tea-spoon of gruel* +3folivorous pellets<br />
Friday:1/8 of apple + 2 slices of carrot + 3 folivorous pellets<br />
Saturday: 1 tea-spoon of gruel* +3folivorous pellets<br />
Sunday: 1 slice of banana + 1/8 of mango + 5 mealworms + 3<br />
folivorous pellets<br />
*Gruel composition: folivorous pellet powder + milk<br />
powder + baby cereals + yolks + cottage-cheese + juice of<br />
squeezed oranges + vitamins (every Monday).<br />
For overweight animals (weight >100 g):from Tuesday to Saturday<br />
we provide only 2 slices of carrot and 3 folivorous pellets.<br />
No change for Sunday and Monday.<br />
For underweight animals (weight
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 29<br />
females may be more interested in food during the breeding<br />
season than the males which,in turn,may be more interested<br />
in the females (Roullet, 1998).<br />
The enrichment made with a branch covered in fruit juice<br />
was the most used by the animals:This is a branch (diameter<br />
of around 5 cm, length of 30-40 cm) in which two trenches<br />
are dug along its length. The branch is fixed onto the mesh<br />
roof. We apply the fruit juice in the trenches with a brush<br />
(Roullet,1998).The animals spend their time licking the juice<br />
out of the branch as they would do with sap in wild (Martin,<br />
1973).<br />
Mixed-species exhibits<br />
Paris Zoo experienced 2 successful combinations:with ayesayes<br />
(Daubentonia madagascariensis) and greater tenrecs<br />
(Tenrec ecaudatus).We tried to put the mouse lemurs in with<br />
slow lorises (Nycticebus coucang), but without success:<br />
wounds were observed on the mouse lemurs’ tails, so they<br />
were removed from the exhibit.<br />
References<br />
Martin, R.D. 1973. A review of the behaviour and ecology of<br />
the lesser mouse lemur (Microcebus murinus). Pp. 1-68. In:<br />
R.P. Michael; J.H. Crook (eds.). Comparative ecology and<br />
behaviour of primates. Academic Press, London, UK.<br />
Pes, T. 2009. The European Studbook of Grey Mouse Lemur<br />
(Microcebus murinus). Zoo and Botanical Garden Plzen.<br />
Roullet, D. 1998. Effet d’un enrichissement physique sur les<br />
comportements agonistiques et exploratoires de plusieurs<br />
groupes de microcèbes murins (Microcebus murinus)<br />
en captivité. DESS d’Ethologie Appliquée et de Chronobiologie<br />
du Comportement. Université Paris XI.<br />
Articles<br />
Diurnal lemur density in the national<br />
park parcel Ivontaka Nord, UNESCO<br />
Biosphere Reserve of Mananara-Nord<br />
Marta Polasky Lyons<br />
School for International Training, Fort Dauphin, Madagascar;<br />
and Carleton College, Northfield, Minnesota, USA,<br />
marta.lyons@gmail.com<br />
Abstract<br />
Here I present a recent diurnal lemur density study performed<br />
in the Biosphere Reserve of Mananara Nord, conducted<br />
between the dates of November 9 and 22, 2008 in<br />
Ivontaka Nord; part of Mananara-Nord National Park. Densities<br />
were calculated using transect walks, and other information<br />
was gathered through interviews with local people<br />
and national park staff. The density of Eulemur fulvus albifrons<br />
appears to be over twice that recorded in other areas<br />
(Table 1), while the density of Varecia variegata variegata<br />
appears to be low,perhaps due to the latter’s preference for<br />
undisturbed habitat and past problems with overhunting. In<br />
addition, Eulemur rubriventer was found to inhabit the park,<br />
though its range was previously thought not to extend east<br />
into the biosphere. Having been, in the past, under strong<br />
pressures from local inhabitants,Ivontaka Nord represents a<br />
disturbed low altitude rainforest;however this report shows<br />
that the lemur populations within the parcel may be recovering.<br />
Introduction<br />
Background<br />
The Biosphere Reserve of Mananara-Nord was created in<br />
1989. It is one of 533 UNESCO (the United Nations Educational,<br />
Scientific, and Cultural Organization) Biospheres<br />
around the world.The Biosphere Reserve of Mananara Nord<br />
covers 144,000 ha,with 23,000 ha being devoted to a terrestrial<br />
park,and another 1,000 ha to a marine park.The terrestrial<br />
national park is split into three separate parcels. Going<br />
from north to south these parcels are: Ivontaka Nord,<br />
Ivontaka Sud, and Verezanantsoro. The parcel of Ivontaka<br />
Nord covers an area of only 827 ha.This is small compared to<br />
the other two parcels; Ivontaka Sud and Verezanantsoro,<br />
which are 1,300 ha and 20,685 ha respectively (Fig. 1). These<br />
parcels are connected by forest sections that are not protected<br />
by the national park system (ANGAP, 2005).<br />
Fig.1:National Park of Mananara-Nord.Three parcels in dark<br />
shade,going from north to south:Ivontaka Nord (where this<br />
survey was carried out), Ivontaka Sud, Verezanantsoro.<br />
Source: MNP.<br />
Threats<br />
It is estimated that approximately 1.9 to 2.2 % of the primary<br />
forest within the biosphere is cleared every year, usually for<br />
rice cultivation through tavy (slash and burn agriculture)<br />
(ANGAP, 2005). Besides just destroying necessary habitat,<br />
this deforestation further splits already extremely fragmented<br />
sections of primary forest.Another concern is illegal<br />
(and legal) selective extraction. The population within the<br />
biosphere relies on wood for constructing their houses and<br />
fuel for cooking.In the villages most cooking is done over an<br />
open flame using collected wood.The wood does not simply<br />
go to the villages on the periphery of a forest,but is also collected<br />
for sale in urban centers and areas farther away from<br />
the forest.
Page 30 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
Another large concern in the biosphere has to do with<br />
poaching. Until recently, hunting was thought to not be of<br />
great concern in Madagascar, being far below the menace<br />
caused by deforestation (Goodman et al., 2003); however,<br />
that is quickly proving incorrect. Unlike in other areas of the<br />
country where eating lemurs is considered fady (taboo), the<br />
Betsimisaraka ethnic group that makes up the majority of inhabitants<br />
within the biosphere reserve is only known to have<br />
fady related to the eating of Indri indri,and even this taboo has<br />
been shown not to be universal (Mittermeier et al., 2006). In<br />
the area people have classically used laly (traditional lemur<br />
traps) and firearms in order to hunt lemurs for consumption<br />
(ANGAP, 2005). Lemurs within the reserve have been a traditional<br />
source of protein for villagers whose diets are based<br />
on the staple rice.<br />
Site<br />
The field camp was situated within the town of Ambodivoandrozana<br />
approximately 17 km south of Mananara Nord. The<br />
village is inaccessible by vehicle but rests less than 2 km from<br />
the national park parcel of Ivontaka Nord.Between the town<br />
and the national park lies another forest managed by the<br />
COBA (communaute de base) for conservation and local subsistence<br />
needs. This forest is split into three zones; Beantohiravina<br />
(21.7 ha), Betsingiala (11.52 ha), and Ambahinkarabo<br />
(41.06 ha). These zones are separated by areas of<br />
agricultural land and secondary forest. A Gestion Contractualisée<br />
des forêts (GCF) in 2005 transferred certain management<br />
and use rights to the local community. It is mostly<br />
low altitude primary growth rainforest, intermixed with<br />
zones of savoka (secondary growth). The parcel of Ivontaka<br />
Nord is made up of both primary forest and disturbed habitat.From<br />
past species inventories,it was believed that two diurnal<br />
lemur species (Eulemur albifrons and Varecia v.variegata)<br />
and multiple nocturnal species, including Microcebus rufus,<br />
Lepilemur mustelinus, Avahi laniger, and Daubentonia madagascariensis<br />
among others inhabited this national park.<br />
Methods<br />
A survey of diurnal lemur species was conducted in the low<br />
altitude (between 250 and 300 m above sea level) rainforest<br />
in the national park of Ivotaka Nord and adjoining community-managed<br />
forest (GCF).The survey was conducted from<br />
November 9-22,2008.Within the national park parcel,a preestablished<br />
2 km transect set up by ANGAP was followed,in<br />
addition to other transects along preexisting paths throughout<br />
the two forests. Additional permanent transects were<br />
not established because of time constraints and the desire to<br />
limit the impact on the habitat. Instead, distance was measured<br />
by walking at a constant pace along preexisting paths<br />
(approximately 900 m per hour). This pace was calculated<br />
using the 2 km transect, and variation in speed was used to<br />
create a range for densities. The total of 17 transects were<br />
walked, ranging from 600-2,500 m in length, with a total distance<br />
walked of approximately 30 km.<br />
For each transect the date, start and end time, weather, and<br />
location were recorded.When a lemur was spotted,the species<br />
was noted along with the group size,time of day,distance<br />
on transect, distance from path, habitat type, and the GPS<br />
coordinates. Density was then calculated by the number of<br />
individual lemurs/area. Area was based on the total length of<br />
transects walked multiplied by double the average perpendicular<br />
distance of lemur from the path (Whitesides et al.,<br />
1988; Norscia et al., 2006). Common methods for this type<br />
of primate study include using a 50 % criterion for falloff<br />
distance based on histograms (Whitesides et al., 1988; John-<br />
son and Overdorff,1999;Erhart and Overdorff,2008) or the<br />
program DISTANCE (Quemere et al., 2010). However, given<br />
time constraints it was not possible to collect an adequate<br />
amount of data to perform these tests.Based on earlier studies<br />
on similar species in similar habitats we used a falloff distance<br />
of 20 m (Irwin, 2001). In addition to transect walks,<br />
interviews were conducted with employees of the ANGAP<br />
office of Mananara Nord and local people of the town of Ambodivoandrozana.<br />
Within the village, the interviews were<br />
done with prominent members of the community including<br />
members of Slow Food (an agriculture movement within the<br />
biosphere) and the COBA.<br />
Results<br />
The observed population density for Eulemur albifrons was<br />
46.13 ± 2.32 individuals/km 2 within the park (Tab.1).The average<br />
group size observed was 7 individuals. Multiple groups<br />
were observed with females carrying babies on their back,<br />
and overall this species was found in a variety of different<br />
habitats, both dense and sparse primary forest, and even on<br />
the edge of secondary growth. Through a combination of<br />
data collected in the forest and interviews with local people,<br />
we found that though Eulemur albifrons frequents multiple<br />
habitats,even leaving the forest to eat crops,they mainly rest<br />
within the National Park.<br />
Two other diurnal lemur species were observed within the<br />
parcel, Varecia variegata variegata and Eulemur rubriventer. V. v.<br />
variegata was observed to have a density of 1.06 ± 0.02 individuals/km<br />
2 (Tab. 1). This species was only observed on one<br />
occasion and only one individual was seen. According to the<br />
local guide and vice president of the COBA,eight individuals<br />
of this species exist within the parcel, a group of five and a<br />
group of three (F. Frejes, personal communication). This implies<br />
a density of 0.97 individuals/km 2.From traces observed<br />
on the ground (eaten fruit) and calls heard,it seems clear that<br />
this lemur spends the majority of its time within the limits of<br />
the parcel as it prefers deep valleys with tall trees; a habitat<br />
not found in the more disrupted community managed forest.<br />
Tab.1:Density of diurnal lemurs within the parcel of Ivontaka<br />
Nord. Densities were calculated of the three species observed<br />
within the parcel, Eulemur albifrons, Eulemur rubriventer,and<br />
Varecia v.variegata.A 50 % falloff distance was used<br />
for perpendicular distances over 20 m (Whitesides et al.,<br />
1988; Irwin et al., 2000) Error bars represent the error in<br />
pace of transect walked and error in observation of animals<br />
from path.<br />
Species Number of<br />
individuals<br />
encountered<br />
Eulemur<br />
albifrons<br />
Eulemur<br />
rubriventer<br />
Varecia v.<br />
variegata<br />
Number of<br />
groups<br />
encountered<br />
Calculated density<br />
(individuals/km2)<br />
31 5 46.13 ± 2.32<br />
3 1 6.40 ± 0.38<br />
1 1 1.06 ± 0.02<br />
Eulemur rubriventer, being a species believed to live west of<br />
the biosphere in higher altitude rainforest, was thus not on<br />
the initial list of lemurs to be found in the area.From the one<br />
sighting of three individuals in a dense part of the primary<br />
forest, the calculated density is 6.40 ± 0.38 individuals/km 2<br />
for the parcel (Tab. 1). No sightings were made of this individual<br />
outside of the parcel,and as its eating habits closely mirror<br />
that of E.albifrons,it was not possible to tell the difference<br />
between traces found on the ground.
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 31<br />
Discussion<br />
The parcel of Ivontaka Nord is considered to be an example<br />
of disturbed primary forest.Anthropogenic effects are much<br />
stronger in Ivontaka Nord than in other parcels,because it is<br />
the smallest of the three, and the closest to the town of<br />
Mananara. Before the creation of the national park, the area<br />
of Ivontaka Nord was frequently used as a place to grow<br />
crops through tavy,harvest wood for construction and cooking,and<br />
hunt lemurs.For these reasons,the lemur population<br />
of the parcel was at one time diminished to the point of localized<br />
extinction of the critically endangered Varecia v.variegata,<br />
in addition to the localized extinctions of Indri indri and Hapalemur<br />
griseus (J. Betsiahilika, personal communication). Yet,<br />
because it is connected through a corridor of community<br />
managed forest to the less disturbed parcel of Ivontaka Sud<br />
(Fig.1),in 2002,V.v.variegata migrated into the GCF corridor<br />
between Ivontaka Sud and Ivontaka Nord, and in 2005 these<br />
lemurs could again be found within the parcel. However, the<br />
species of Indri indri and Hapalemur griseus are not believed to<br />
currently inhabit the parcel, though they can be found in the<br />
other two parcels within the biosphere (J. Betsiahilika, personal<br />
communication).<br />
Within the parcel, there was convincing evidence to suggest<br />
that the lemurs might be recovering even more than previously<br />
known. During the two weeks of transects, Eulemur<br />
rubriventer was observed within the parcel, even though its<br />
recorded range ends west of the biosphere reserve. In addition,traces<br />
of the distinct eating habits (shredding the stems<br />
of tall plants whilst stripping off foliage) of Hapalemur griseus<br />
were found within the savoka. Additionally, there were reports<br />
of Indri indri spending time in the corridor between<br />
Ivontaka Sud and Ivontaka Nord (J. Betsiahilika, personal<br />
communication).<br />
Through field observations, E. albifrons was found to have a<br />
population twice that found in other areas such as Masoala<br />
(Mittermeier et al.,2006).They frequent both the parcel and<br />
community forest. During the season in which this study<br />
took place,when most of the fruit within the parcel is not yet<br />
ripe, many Eulemur albifrons were reported by local villagers<br />
to be exiting the parcel in search of other cultivated fruits<br />
like lychee and banana. V. v. variegata on the other hand<br />
appeared to not leave the protected parcel.It is possible that<br />
they adapt their feeding habits based on the season and thus<br />
do not eat cultivated fruits (Ratsimbazafy, 2002).<br />
The observed density of V. v. variegata was not congruent<br />
with the report of the local guide (F.Frejes,personal communication).<br />
This species, like Eulemur rubriventer, was observed<br />
on only one occasion. Because of time constraints, surveys<br />
had to be conducted over a two week period and with only<br />
the use of one field team. For more complete and definitive<br />
findings, a survey needs to be done over a longer period of<br />
time, possibly across different seasons. Also, it would be<br />
beneficial to conduct the same type of study after dark.<br />
Finally, it would be helpful to determine lemur densities in<br />
the other two parcels, and the community managed forests<br />
in-between.<br />
Conclusion<br />
It is currently of the utmost importance to make sure that<br />
the integrity of forested sections outside the realm of the<br />
parcel remains protected.These areas represent a buffer between<br />
the fragile low altitude rainforest and an ever expanding<br />
human population. The corridor formed by these community-managed<br />
forests, between the three national park<br />
parcels, is a priority area. It has already been shown to provide<br />
a bridge to facilitate migration between the three parcels,which<br />
is critical in a country where forest fragmentation<br />
is proving detrimental to the gene flow of lemurs (Louis et al.,<br />
2006).<br />
Through the work of the biosphere reserve,significant steps<br />
have already been taken to encourage conservation. Both<br />
government officials and villagers appear to be working together<br />
to promote a healthy ecosystem. Already with the<br />
promotion of crops such as vanilla,cloves,and coffee,as an alternative<br />
to other more environmentally negative livelihoods,<br />
villagers say they have seen an improvement both in<br />
their lives and the forest health (Desana and Berger,personal<br />
communication). However, people have needs; the agricultural<br />
inhabitants of these rural areas need both to grow their<br />
food and to have wood and other materials for performing<br />
everyday tasks. It is possible to improve both the lives of the<br />
people in the area and decrease their negative impact on<br />
their environment through simple strategies such as rice intensification<br />
and promoting more efficient cooking methods.<br />
Maybe then populations of Indri indri, Hapalemur griseus, and<br />
Propithecus diadema,all species that at one time inhabited the<br />
area, will return to the parcel of Ivontaka Nord.<br />
Acknowledgements<br />
This study was carried out under a Memorandum of Understanding<br />
between SIT and ANGAP Mananara for an internship<br />
for the author in November 2008.SIT thanks the Ministry<br />
of Higher Education and Scientific Research and the University<br />
of Antananarivo for the ongoing collaboration under<br />
which SIT Study abroad operates. This study could not have<br />
been possible without the help of Barry Ferguson and Jim<br />
Hansen of SIT in addition to the staff in the ANGAP office of<br />
Mananara, including Willy Mora, Jocelyn Bezara, Justin Besiahilika,<br />
and Jean Cristophe Josoa.<br />
References<br />
ANGAP. 2005. Plan de Gestion de la Conservation-Parc<br />
National Mananara Nord.<br />
COBA FMAA. 2005. Fifanekena Famindra-Pitantanana ny<br />
Atiala sy ny Harena Voajanahary azo Havaozina ao Ambodivoandrozana.<br />
Erhart, E.M.; Overdorff, D.J. 2008. Population demography<br />
and social structure changes in Eulemur fulvus rufus from<br />
1988 to 2003.American Journal of Physical Anthropology<br />
136: 183-193.<br />
Goodman, S.; Raselimanana, A. 2003. Hunting of wild animals<br />
by Sakalava of the Menabe region: a field report from<br />
Kirindy-Mite. Lemur News 8: 4-6.<br />
Irwin, M.T.; Samonds, K.E.; Raharison, J. 2001. A biological inventory<br />
of the lemur community of Réserve Spéciale de<br />
Kalambatritra, south-central Madagascar. Lemur News 6:<br />
24-28.<br />
Johnson, S.E.; Overdorff, D.J. 1999. Census of brown lemurs<br />
(Eulemur fulvus sspp.) in southeastern Madagascar:<br />
Methods-testing and conservation implications. American<br />
Journal of Primatology 47: 51-60.<br />
Louis, E.E. jr.; Coles, M.S.; Andriantompohavana, R.; Sommer,<br />
J.A.; Engberg, S.E.; Zaonarivelo, J.R.; Mayor, M.I.; Brenneman,<br />
R.A. 2006. Revision of the mouse lemurs (Microcebus)<br />
of Eastern Madagascar. International Journal of<br />
Primatology 27: 347-389.<br />
Mittermeier, R.; Tattersall, I.; Konstant, W.R.; Nash, S.D. 2006.<br />
Lemurs of Madagascar. Conservation International, Washington<br />
D.C.<br />
Norscia,I.;Rahanitriniaina,O.G.;Jolly,A.;Donati G.2006.Preliminary<br />
survey of lemur density in the semimontane rainforest<br />
of Anka,Fort-Dauphin region.Lemur News 11:14-<br />
16.<br />
Quemere, E.; Champeau, J.; Besolo, A.; Rasolondraibe, E.;<br />
Rabarivola,C.;Crouau-Roy,B.;Chikhi,L.2010.Spatial Variation<br />
in Density and Total Size Estimates in Fragmented<br />
Primate Populations: The Golden-Crowned Sifaka (Propithecus<br />
tattersalli). American Journal of Primatology 72:<br />
72-80.
Page 32 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
Ratsimbazafy, J.H.; Ramarosandratana, H.V.; Zaonarivelo, R.J.<br />
2002. How do black-and-white ruffed lemurs still survive<br />
in highly disturbed habitat? Lemur News 7: 7-10.<br />
Sur le Commerce International des Especes de Faune et de<br />
Flore Sauvage. 2005. Loi N 2005-018.<br />
Whitesides, G.H.; Oates, J.F.; Green, S.M.; Kluberdanz, R.P.<br />
1988. Estimating primate densities from transects in a<br />
West African rain forest; a comparison of techniques.<br />
Journal of Animal Ecology 57: 345-367.<br />
Distribution of Prolemur simus north of<br />
the Mangoro-Nosivolo River – how far<br />
north do we really have to look?<br />
Rainer Dolch 1*, Erik R. Patel 2, Jonah H. Ratsimbazafy<br />
3,4, Christopher D. Golden 5, Tianasoa Ratolojanahary<br />
1, Jean Rafalimandimby 1, Jonathan L. Fiely 1<br />
1Association MITSINJO, Lot 104 A, Andasibe 514, Madagascar<br />
2Cornell University, 211 Uris Hall, Ithaca, NY 14850, USA<br />
3Durrell Wildlife Conservation Trust,BP 8511,Antananarivo<br />
101, Madagascar<br />
4GERP, 34 Cité des Professeurs, Fort Duchesne, Antananarivo<br />
101, Madagascar<br />
5University of California, Department of Environmental<br />
Science,Policy and Management,Mulford Hall #3114,Berkeley,<br />
CA 94720, USA<br />
*Corresponding author: rdolch@gmx.de<br />
Introduction<br />
While the Mangoro-Nosivolo river system is a recognized<br />
biogeographical divide for several lemur species (Goodman<br />
and Ganzhorn, 2004), this pattern does not hold for the<br />
Greater Bamboo Lemur (Prolemur simus). Findings from<br />
numerous subfossil sites indicate that the historical distribution<br />
of P. simus, now one of the rarest Malagasy primates<br />
(Wright et al.,2008,2009),once encompassed most of Madagascar<br />
(Godfrey and Vuillaume-Randriamanantena, 1986).<br />
During the last <strong>15</strong>0 years, documented sightings of the species<br />
became more and more scarce and by the middle of the<br />
last century it was already feared extinct (e.g. Napier and<br />
Napier, 1967).<br />
More recent discoveries came solely from southeastern<br />
Madagascar (Petter et al., 1977; Meier and Rumpler, 1987),<br />
which led to the unspoken assumption that P.simus had been<br />
extirpated from the rest of the island. Despite the fact that<br />
the last collected specimen of P. simus had come from<br />
Mananara in 1876 (Godfrey and Vuillaume-Randriamanantena,<br />
1986), not a single individual had been found north of<br />
the Mangoro river for more than 130 years, before Dolch et<br />
al.(2004,2008) rediscovered the species in Torotorofotsy,in<br />
the commune of Andasibe.<br />
In order to investigate further into the distribution and abundance<br />
of P. simus north of the Mangoro, several extensive<br />
surveys have recently been, and are currently being, conducted<br />
(King and Chamberlan, 2010). As preliminary results<br />
of these surveys are trickling in, accounts of P. simus from<br />
inhabitants of these regions also multiply rapidly and await<br />
verification.<br />
We do not aspire to anticipate survey results,but we believe<br />
that summarizing our current knowledge of P.simus north of<br />
the Mangoro is crucial for the planning of future surveys and<br />
conservation strategies.<br />
Methods<br />
We gathered and compiled information on P. simus north of<br />
the Mangoro deduced from our own research,from reports<br />
that villagers brought to our attention, and from anecdotal<br />
evidence in existing literature.The ten localities from where<br />
information was available included (from south to north) the<br />
Marolambo area, the Brickaville area, the western parts of<br />
the Ankeniheny-Zahamena Corridor (CAZ), Zahamena, the<br />
Soanierana-Ivongo area, Ambatovaky, Marotandrano, Mananara,<br />
Makira, and Marojejy (Fig. 1).<br />
Fig. 1: Localities from where information on P. simus was collected.<br />
Results<br />
Results of our compilation are summarized in Tab.1.Of all localities<br />
north of the Mangoro examined,two (Brickaville and<br />
western CAZ) have steadfast records of P.simus based on independently<br />
confirmed sightings. One locality (Soanierana-<br />
Ivongo) has a record based on a single observation, whereas<br />
for four others (Marolambo, Zahamena, Ambatovaky, Makira)<br />
evidence is only based on reports of villagers. The final<br />
three (Marotandrano, Mananara, Marojejy) do not have any<br />
records. Details are given below.<br />
Tab. 1: Potential localities for P. simus north of the Nosivolo-<br />
Mangoro.<br />
Region evidence based on<br />
Marolambo area reports by villagers<br />
Brickaville area sightings, confirmed<br />
Western CAZ sightings, confirmed<br />
Zahamena reports by villagers<br />
Soanierana-Ivongo area sightings, unconfirmed<br />
Ambatovaky reports by villagers<br />
Marotandrano (no evidence)<br />
Mananara (mo evidence)<br />
Makira reports by villagers<br />
Marojejy (no evidence)<br />
Marolambo area. A hotspot of endemic fish species richness,<br />
the Nosivolo and lower Mangoro rivers have recently<br />
received increased attention by researchers and conservationists<br />
alike.While working in this area,we received several<br />
accounts of villagers on P. simus from 2006-2009. Reports<br />
claiming the occurrence of P. simus come from 4 communes<br />
along the Nosivolo-Mangoro river, and focus (from west to<br />
east) on Ambohimilanja, Betampona, Marolambo and Ambinanidilana.<br />
In order to verify these accounts, a preliminary<br />
survey is currently being conducted within The Aspinall<br />
Foundation’s "Saving Prolemur simus" project (Ratsimbazafy,<br />
2010).
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 33<br />
Brickaville area. A similar survey has already been conducted<br />
for the Brickaville area. A total of 6 sites in isolated<br />
fragments in the communes (from west to east) of Fanasana,<br />
Anivorano and Fetraomby have been found containing P.<br />
simus. Details are given by Ravaloharimanitra et al. (2010).<br />
Western CAZ.In the course of the same survey,12 sites in<br />
the western parts of the Ankeniheny-Zahamena corridor<br />
have been found containing P. simus (Ravaloharimanitra et al.,<br />
2010). These findings follow earlier reports from villagers<br />
that had claimed its presence (e.g.Schmid and Alonso,2005).<br />
The communes where P. simus has been confirmed include<br />
(from south to north) Andasibe, Morarano-Gare, Fierenana,<br />
and Didy.<br />
Zahamena. Ganzhorn (2004) states that a report of the<br />
presence of P. simus in the PN Zahamena was brought to his<br />
attention in 1995, but that it "was questioned and eventually<br />
withdrawn".Information on a possible occurrence of P.simus<br />
in Zahamena, obviously derived from that report, is mentioned<br />
in Godfrey et al. (1997).<br />
Soanierana-Ivongo area. In their little noticed bulletin,<br />
the Association de Défense de la forêt d’Ambodiriana report<br />
that trainee Coralie Ebert, while studying the woolly<br />
lemurs (Avahi laniger) of this forest, claims to have observed<br />
an individual of Prolemur simus (ADEFA, 2009). An earlier<br />
lemur survey of the area (Beaucent and Fayolle, 2008) has<br />
not yielded evidence of P.simus.The forêt d’Ambodiriana lies<br />
just 30 km to the north of Soanierana-Ivongo,a region where,<br />
according to Mittermeier et al. (2006), halogodro and bokombolobe<br />
are still used as local names for P. simus.<br />
Ambatovaky.Ambatovaky is an area that has received only<br />
little attention due to its difficult accessibility. In the early<br />
1990s, a lemur survey was conducted by Evans et al. (1993-<br />
1994). They did not find tangible evidence for P. simus, but<br />
state that "local people indicated that there existed until recently<br />
a lemur which fed on giant bamboo along the Sandrangato<br />
and/or Marimbona rivers,known as alakoto or halokoto".<br />
Marotandrano.Lying to the northwest of Ambatovaky,Marotandrano<br />
has even received less attention than the former.<br />
A lemur survey by Ralison (2006) did not indicate presence<br />
of P. simus.<br />
Mananara. The last specimen of P. simus to be collected<br />
from north of the Nosivolo-Mangoro river came from an<br />
area close to Mananara (Godfrey and Vuillaume-Randriamanantena,1986).Although<br />
the exact collection locality can not<br />
be traced (due to unsuited transcription of its name by the<br />
collector J.P.Audebert),the assumption that P.simus may still<br />
occur in the forests around Mananara was still put forward<br />
by Nicoll and Langrand (1989). No evidence for P. simus in<br />
Mananara has been produced since.<br />
Makira. Being Madagascar’s largest continuous rainforest<br />
(317,000 ha), lemur surveys in Makira are not easy to conduct.<br />
Two years of intensive surveys by Rasolofoson et al.<br />
(2007) and Ratelolahy and Raivoarisoa (2007) have not uncovered<br />
any evidence of P.simus.Similarly,during seven years<br />
of relying on trusting relationships with hunters, Golden<br />
(2009) has not come across P. simus among the 23 mammal<br />
species hunted for consumption throughout southern,western,<br />
northern, and eastern Makira. However, villagers living<br />
adjacent to the newly discovered Antohaka Lava forest at the<br />
edge of northeastern Makira (20 km south of Andrakata on<br />
Marojejy’s southeastern border) have reported recent sightings<br />
of a large bamboo lemur with ear tufts known locally as<br />
bokombolobe.Unfortunately,several months of systematic lemur<br />
surveying of the Antohaka Lava forest between August<br />
and December 2009 did not confirm these reports, despite<br />
an exceptional primate diversity documented inthat area<br />
(Patel, 2009).<br />
Marojejy. A lemur survey by Sterling and McFadden (2000)<br />
found no evidence of P. simus. Alleged observations of bamboo<br />
lemurs other than Hapalemur griseus by tourists may be<br />
attributed to the possible presence of H. occidentalis, rather<br />
than P. simus (R. Mittermeier, pers. comm.). Moreover, during<br />
nine years of research on Propithecus candidus in Marojejy,no<br />
local reports or sightings of P.simus have been received (Patel,<br />
2009).<br />
Discussion<br />
Despite the scarcity of information, growing evidence supports<br />
that P. simus may still be widespread in Madagascar<br />
north of the Mangoro river. Since the species occupies large<br />
home ranges (Dolch et al., unpubl. data), appears to travel at<br />
night due to possible cathemerality (Santini-Palka, 1994), is<br />
cryptic, and often silent when unhabituated, it is conceivable<br />
that it has been overlooked in the past.However,because the<br />
P.simus vocal repertoire is distinct and extensive (Bergey and<br />
Patel,2008) and its feeding traces on giant bamboo (Cathariostachys<br />
madagascariensis) are unmistakable,attention to such<br />
indirect evidence of P. simus presence should be focused<br />
upon in all surveys.<br />
The report from Makira, if confirmed, is especially interesting,<br />
since the northernmost former record for the species<br />
(other than from subfossils) comes from Antongil Bay<br />
(Schwarz, 1931).<br />
Our experience shows that accounts of villagers are mostly<br />
reliable, and that people usually have a good sense of what<br />
animal species do or do not occur in their vicinity.Therefore,<br />
integrating local people is crucial for further studies into<br />
Prolemur distribution. Logically, collaboration with local people<br />
is one conservation recommendation given by the Prolemur<br />
Conservation Working Group (Madagascar Fauna<br />
Group, 2010).<br />
Based on information compiled,we tentatively conclude that<br />
P. simus is still more widespread than previously thought.<br />
Without sufficient data, given persisting threats to the habitats<br />
in which it occurs and our incomplete understanding of<br />
habitat requirements for the species,we do not dare say that<br />
a larger distribution area contributes in any way to relieving<br />
the species from extinction pressure. Unfortunately, P. simus<br />
still has to be considered one of the most threatened primates<br />
in the world.<br />
Acknowledgements<br />
We thank Tokiniaina Hobinjatovo for helping with literature<br />
research and Coralie Ebert for additional information. We<br />
thank all individuals that have shared their observations and<br />
made available the information presented here. We would<br />
also like to thank the Margot Marsh Biodiversity Fund, and<br />
the National Geographic Society Conservation Trust Award<br />
#C135-08.<br />
References<br />
ADEFA. 2009. Stages. Bulletin d’Information de l’Association<br />
de Défense de la forêt d’Ambodiriana 25: 1-2.<br />
Beaucent,S.;Fayolle,M.2008.Etude de la communauté de lémuriens<br />
de la forêt d’Ambodiriana,NE Madagascar.Lemur<br />
News 13: 28-32.<br />
Bergey,C.;Patel. E.R.2008.A preliminary vocal repertoire of<br />
the Greater Bamboo Lemur (Prolemur simus): classification<br />
and contexts. Nexus 1: 69-84.<br />
Dolch, R.; Fiely, J.L.; Ndriamiary, J.N.; Rafalimandimby, J.; Randriamampionona,R.;Engberg,S.E.;Louis,E.E.Jr.2008.Confirmation<br />
of the greater bamboo lemur, Prolemur simus,<br />
north of the Torotorofotsy wetlands,eastern Madagascar.<br />
Lemur News 13: 14-17.<br />
Dolch, R.; Hilgartner, R.D.; Ndriamiary, J.N.; Randriamahazo,<br />
H.2004.The grandmother of all bamboo lemurs:evidence
Page 34 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
for the occurrence of Hapalemur simus in fragmented<br />
rainforest surrounding the Torotorofotsy marshes, central<br />
eastern Madagascar. Lemur News 9: 24-26.<br />
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Primatological Society (IPS), and Conservation<br />
International (CI), Arlington, VA.<br />
Enquête préliminaire de la distribution<br />
des lémuriens de bambou dans et autour<br />
du Corridor forestier Fandriana-Vondrozo,<br />
Madagascar<br />
Andry Rajaonson 1,2, Maherisoa Ratolojanahary 2,<br />
Jonah Ratsimbazafy 1, Anna Feistner 3, Tony King 2*<br />
1Groupe d’Etude sur les Primates de Madagascar (GERP),<br />
Lot 34 Cité des Professeurs, Fort Duchesne, Ankatso, Antananarivo<br />
101, Madagascar<br />
2The Aspinall Foundation, BP 7170 Andravoahangy, Antananarivo<br />
101, Madagascar<br />
3Centre ValBio, BP 33 Ranomafana, Ifanadiana 312, Madagascar<br />
(present address: anna@feistner.com)<br />
*Corresponding author: tonyk@aspinallfoundation.org<br />
Mots-clés: Prolemur simus, Hapalemur aureus, Varecia variegata<br />
editorum, bambou<br />
Introduction<br />
Le Grand Hapalémur (Prolemur simus) mangeur de bambou,<br />
est classé comme étant "gravement menacé CR" sur la liste<br />
rouge de l’Union Internationale pour la Conservation de la<br />
Nature (UICN,2009).Il est aussi l’une des quatre espèces de<br />
Madagascar faisant partie des 25 primates considérés<br />
comme les plus menacés au monde (Mittermeier et al.,2007,<br />
2009). Des individus capturés dans les années 1800 venaient<br />
d’une région plus étendue, ce qui laisse supposer que son<br />
habitat a diminué.Les sites de subfossiles avec des squelettes<br />
identiques à celui de P. simus sont nombreux, impliquant une<br />
distribution encore plus vaste à une époque très ancienne<br />
(Godfrey et Vuillaume-Randriamanantena, 1986; Godfrey et<br />
al., 2004).<br />
Actuellement, la distribution géographique de l’espèce semble<br />
très étroite.On pense que P.simus est seulement présent<br />
dans quelques fragments de forêt tropicale humide près de la<br />
côte Est de Madagascar (Mittermeier et al.,2006 ;Dolch et al.,<br />
2008;Wright et al.,2008).Wright et al.(2008) résume la crise
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 35<br />
actuelle de l’espèce: sur les 70 localités étudiées, la présence<br />
de P.simus est confirmée seulement sur 11 d’entre elles,à une<br />
altitude comprise entre 121 et 1600 m.<br />
La plupart des sites connus abritant P.simus se trouvent dans<br />
ou autour du Corridor Fandriana-Vondrozo, dans le Sud-Est<br />
du pays (Andriaholinirina et al.,2003;Meier et Rumpler,1987;<br />
Sterling et Ramaroson, 1996; Wright et al., 1987, 2008; Mittermeier<br />
et al., 2009). Deux autres espèces de lémuriens de<br />
bambou vivant dans le corridor, Hapalemur aureus et H.<br />
griseus, sont aussi menacées (Mittermeier et al., 2006; IUCN,<br />
2009).Notons que P. simus est le plus grand des lémuriens de<br />
bambou,avec un pelage gris brun (comme H. griseus).Il se distingue<br />
facilement des deux autres espèces par des touffes de<br />
poils blancs sur les oreilles. Sa face est aussi plus allongée, et<br />
on le trouve souvent au sol, alors que les autres espèces y<br />
sont rarement (Wright et al., 1987). Selon l’étude de Tan<br />
(1999) à Ranomafana, le régime alimentaire de P. simus est<br />
constitué à 95 % d’une seule espèce de bambou Cathariostachys<br />
sp. (ou volohosy dans le dialecte local malgache), 3 %<br />
d’autres espèces de bambous et de graminées,0,5 % de fruits<br />
et 1,5 % d’autres éléments (principalement de la terre et des<br />
champignons).<br />
La présente étude a été organisée dans le cadre du Projet<br />
Varibolomavo proposé par The Aspinall Foundation (TAF).<br />
Ce projet veut mettre en place des actions rapides, efficaces<br />
et collaboratives pour sauver Prolemur simus. Plus précisément,<br />
le deuxième objectif du projet est d’organiser une<br />
étude de la distribution et de l’abondance de P. simus (TAF,<br />
2008, 2009; King and Chamberlan, 2010). Par conséquent, le<br />
but de cette étude était de contribuer à réaliser ces objectifs,<br />
dans et autour du corridor Fandriana-Vondrozo, par a) la<br />
récolte des connaissances indigènes locales sur les distributions<br />
des lémuriens; et b) la recherche des signes de présence<br />
des lémuriens de bambou. Nous présentons ici un<br />
résumé des résultats de l’étude, exposés de façon plus<br />
détaillée par Ratolojanahary et al. (2009).<br />
Méthodes<br />
Entre les 27 novembre 2008 et 25 mai 2009, nous avons enquêté<br />
dans 14 zones situées dans et autour du Corridor<br />
Fandriana-Vondrozo (Tab. 1, Fig. 1). Pour chaque commune,<br />
des entretiens avec les autorités locales ont eu lieu.Des collaborations<br />
avec ces personnes ont permis d’organiser les<br />
réunions villageoises pour mener les enquêtes participatives,<br />
à l’aide de photos des espèces de lémuriens supposées<br />
coexister dans ce couloir forestier (Prolemur simus, Hapalemur<br />
aureus, H. griseus, Eulemur rufus, E. rubriventer, Propithecus<br />
edwardsi, Varecia variegata editorum, Microcebus rufus,<br />
Cheirogaleus major, Avahi laniger, Lepilemur microdon, Daubentonia<br />
madagascariensis). Les appellations locales des différentes<br />
espèces connues par les communautés villageoises<br />
ont été relevées lors de chaque enquête.De plus,nous avons<br />
utilisé la méthode de cartographie participative (Jones et al.,<br />
2005) durant la réunion dans les communes de Mahazoarivo,<br />
Iandraina, Sahamadio et Evato. Suite aux résultats des enquêtes<br />
villageoises, nous avons visité des forêts et sites<br />
intéressants dans la région, toujours accompagnés par un<br />
guide local et des agents de recherche du Centre ValBio de<br />
Ranomafana. Nous nous arrêtions tous les 25 mètres pour<br />
relever la localisation des bambous et des espèces de lémuriens,<br />
ainsi que les signes de présence de ces dernières.<br />
La présence des espèces de lémuriens était révélée soit par<br />
l’observation directe (animal vu), soit par l’observation indirecte<br />
(signes de nourrissage,excréments ou vocalisation).La<br />
recherche des signes de nourrissage des lémuriens de bambou<br />
était faite dans les zones de bambous, et les signes<br />
Tab. 1: Les zones visitées pendant l’enquête.<br />
Zone Sites visités Dates<br />
Zones situées dans le corridor forestier<br />
Ambendrana 1 27-28 nov 2008<br />
Amindrabe 1 29 nov - 1 déc 2008<br />
Ambodiara 1 11-13 déc 2008<br />
Antarehimamy 1 14-16 déc 2008<br />
Antaranjaha 4<br />
29 jan - 6 fév 2009 (dont Tsianivoho<br />
et Ambolomadinika)<br />
Manambolo 1 9-11 mai 2009<br />
Zones situées autour du corridor forestier<br />
Mananjary 4 14-16 jan 2009<br />
Sahalanona 9 17-23 jan 2009<br />
Manakara 1 25 jan 2009<br />
Mahazoarivo 2 18-19 mai 2009 (dont Ifasy)<br />
Iandraina 1 21 mai 2009<br />
Sahamadio 1 22-23 mai 2009<br />
Evato 0 24-25 mai 2009<br />
Mahafasa 0 25 mai 2009<br />
14 zones 27 sites 27 nov 2008 - 25 mai 2009<br />
Fig. 1: Les sites visités lors de l’enquête dans et autour du<br />
Corridor forestier Fandriana-Vondrozo.<br />
étaient examinés précautionneusement afin d’identifier l’espèce<br />
qui en était responsable.Prolemur simus préfère surtout<br />
les bambous de grand diamètre, et les parties de bambou<br />
privilégiées varient avec les saisons.Entre juillet et novembre,<br />
P. simus consomme principalement la moelle tendre de<br />
bambou géant (Tan,1999),après avoir ouvert la tige en deux<br />
et l’avoir déchirée en petits morceaux (Wright et al., 1987).<br />
Ainsi, l’échantillon à rechercher devrait être des tiges déchirées<br />
sans ou avec peu de moelle.Par contre,entre novembre<br />
et avril, il se concentre sur les jeunes pousses (Tan, 1999),<br />
donc les échantillons devraient être des bouts de jeunes<br />
pousses de bambou géant. Avec de l’expérience, il est égale-
Page 36 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
ment possible de distinguer les signes laissés sur les feuilles<br />
de bambou. P. simus se nourrit des jeunes feuilles matures et<br />
ne consomme pas la partie tranchante de la feuille. Par<br />
contre, Hapalemur aureus et H. griseus consomment uniquement<br />
la base de la feuille, H. aureus déchirant la gaine de<br />
chaume à l’aide de ses dents avant de manger les branches.<br />
Résultats et Interprétations<br />
Espèces de lémuriens recensées<br />
Les populations locales ont souvent des noms vernaculaires<br />
distincts pour les différentes espèces de lémuriens.En outre,<br />
nous avons remarqué qu’ils ne pouvaient pas toujours identifier<br />
les espèces sur les photos, alors qu’ils pouvaient les reconnaître<br />
dans la nature.Les informations récoltées lors des<br />
enquêtes villageoises doivent donc être utilisées avec précaution<br />
et sont toujours à vérifier sur le terrain. Durant les<br />
vérifications, nous n’avons trouvé qu’un seul site présentant<br />
des signes de nourrissage de Prolemur simus (Tab. 2). Par<br />
contre, nous avons effectué 39 observations (directes et<br />
indirectes) de Hapalemur griseus, dans 10 des 12 zones visitées<br />
(Tab. 2), et huit observations (indirectes) de H. aureus<br />
(Tab.3).Toutes ces observations ont été faites dans les zones<br />
situées au sein du corridor forestier, mais aucune dans ses<br />
alentours (Tab. 2). Cette espèce a laissé des signes de nour-<br />
Tab. 2: Espèces de lémuriens rencontrées dans chaque zone.<br />
Zone Hapalemur Hapalemur Prolemur Autres espèces<br />
griseus aureus simus<br />
Zones situées dans le corridor forestier<br />
Ambendrana vu & signes signe<br />
E rubriventer (vu)<br />
Microcebus sp. (nid)<br />
Amindrabe vu & signes signe<br />
signe P. edwardsi (vu et entendu)<br />
(environ 1 an) D. madagascariensis (signes)<br />
Ambodiara signes signes V. variegata (entendu)<br />
Antarehimamy signes signes V. variegata (entendu)<br />
Antaranjaha signes<br />
signes &<br />
entendus<br />
V. variegata (entendu)<br />
E. rufus (vu)<br />
Manambolo signes E. rufus (vu) E. rubriventer (vu)<br />
Zones situées autour du corridor forestier<br />
Sahalanona vus & signes M. rufus et A. laniger à vendre<br />
Mananjary vus & signes<br />
Microcebus sp. (nid)<br />
Cheirogaleus sp. (signes)<br />
Manakara E. rufus (vu) A. laniger (vu)<br />
Mahazoarivo vus<br />
Iandraina signes<br />
Sahamadio signes<br />
Tab. 3: Observations de Hapalemur aureus faites pendant l’étude.<br />
Zone Remarque Latitude Longitude Altitude (m)<br />
Ambendrana signe de nourrissage S 21º 22’ 22.7" E 047º 20’ 46.5" 1182<br />
Amindrabe signe de nourrissage S 21º 24’ 13.1" E 047º 22’ 47.7" 1070<br />
Ambodiara signe de nourrissage S 21º 53’ 27.0" E 047º 21’ 18.9" 825<br />
Antarehimamy signe de nourrissage S 21º 54’ 47.5" E 047º 20’ 38.4" 1074<br />
Antaranjaha signe de nourrissage S 21º 58’ 20.3" E 047º 20’ 16.7" 828<br />
Antaranjaha entendu des cris S 21º 58’ 25.3" E 047º 20’ 17.7" 783<br />
Antaranjaha signe de nourrissage S 21º 58’ 39.1" E 047º 19’ 43.8" 786<br />
Manambolo signes de nourrissage S 22º 04’ 06.2" E 046º 59’ 27.5" 1238<br />
Tab. 4: Observations de Varecia variegata faites pendant l’étude.<br />
Zone Remarque Latitude Longitude Altitude (m)<br />
Ambodiara entendu des cris S 21º 53’ 17.4" E 047º 21’ 42.3" 500<br />
Ambodiara entendu des cris S 21º 53’ 17.7" E 047º 21’ 34.5" 825<br />
Antarehimamy entendu des cris S 21º 55’ 00.4" E 047º 22’ 10.3" 489<br />
Antaranjaha entendu des cris S 21º 58’ 23.6" E 047º 20’ <strong>15</strong>.4" 828<br />
Antaranjaha entendu des cris S 21º 58’ 31.5" E 047º 20’ 13.6" 743<br />
rissage et émis des cris. La présence de huit autres espèces<br />
de lémuriens a également été constatée (Tab. 2). L’une d’entre<br />
elles, Varecia variegata editorum, est une sous-espèce<br />
gravement menacée selon l’UICN (2009), et les détails de<br />
toutes les observations de cette espèce sont présentés dans<br />
le Tab. 4.<br />
Résultats par zone située dans le corridor forestier de Fandriana-<br />
Vondrozo<br />
Ambendrana: Le village d’Ambendrana (S21°22’44.9" E 047°<br />
18’31.0",altitude 1121 m) est placé sous l’autorité de la commune<br />
rurale d’Androy et situé à une vingtaine de kilomètres<br />
au sud du Parc National de Ranomafana. Ce village est entouré<br />
de rizières localisées tout autour du corridor.La forêt<br />
d’Ambendrana a une superficie de 1.496 hectares et est<br />
gérée par la communauté de base depuis 2003. Au cours de<br />
l’enquête, les villageois n’ont reconnu que 3 espèces de<br />
lémuriens des 12 présentées sur les photos, notamment<br />
l’espèce Hapalemur griseus. D’après nos observations, la<br />
forêt d’Ambendrana est perturbée. Cependant, nous avons<br />
pu localiser quelques groupes de lémuriens, dont un groupe<br />
de H. griseus, et des signes de nourrissage.<br />
Amindrabe: La forêt d’Amindrabe a une superficie de 5.800<br />
hectares et est également gérée par la communauté de base<br />
depuis 2003.Cette forêt est située à 5,7 km du village d’Ambendrana.<br />
Le Fokontany Amindrabe (S21°<br />
23’14.8" E047°21’ 46.4", altitude 1096 m)<br />
fait également partie de la commune ru-<br />
rale d’Androy et comprend plusieurs villages.<br />
Durant l’enquête, les villageois ont<br />
reconnu 5 espèces de lémuriens, dont H.<br />
griseus. Pendant l’expédition dans le site<br />
d’Amindrabe, deux anciens signes de<br />
nourrissage (vieux d’environ un an d’après<br />
nos constatations) de P.simus ont été trouvés<br />
sur le tronc d’une espèce de bambou<br />
localement appelé <strong>Vol</strong>otsangana (S21º24’<br />
22.5", E047º23’07.2", altitude 1055 m).<br />
Nous avons également trouvé deux<br />
groupes de H. griseus, des Propithecus edwardsi<br />
et des signes d’alimentation de Daubentonia<br />
madagascariensis, attestant de la<br />
grande diversité de ce site en espèces de<br />
lémuriens.<br />
Ambodiara: Le Fokontany d’Ambodiara<br />
(S21°54’41.3", E047°23’29.2, altitude<br />
346 m) existe depuis 1910 et est composé<br />
de 8 villages. Le village d’Ambodiara est<br />
situé à 5,9 km, c’est-à-dire à environ 3<br />
heures de marche à l’ouest d’Ikongo. Au<br />
cours de l’enquête, les villageois ont reconnu<br />
9 espèces de lémuriens,dont Hapalemur<br />
aureus, H. griseus et également Prolemur<br />
simus. D’après nos observations, la<br />
forêt d’Ambodiara est perturbée. Nous<br />
n’avons pas trouvé P.simus sur ce site,mais<br />
nous avons constaté la présence de Cathariostachys<br />
sp. Par contre, Varecia variegata<br />
editorum abonde dans cette localité, et<br />
nous avons trouvé des signes de nourrissage<br />
de H. aureus et H. griseus.<br />
Antarehimamy: Situé dans le district<br />
d’Ikongo, le village d’Antarehimamy (S21°<br />
55’59.2",E047°22’17.6",altitude 410 m) se<br />
situe à 3,16 km au Nord-Est d’Ambodiara<br />
et à 9,01 km à l’ouest de la commune<br />
rurale d’Ikongo.Lors de l’enquête,les villa-
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 37<br />
geois ont reconnu 7 espèces de lémuriens,mais aucun lémurien<br />
de bambou.Néanmoins,nous avons trouvé des signes de<br />
nourrissage de Hapalemur aureus et H. griseus. Nous avons<br />
également remarqué l’abondance de Varecia variegata editorum<br />
sur le site.<br />
Antaranjaha/Ambolomadinika/Tsianivoho:Située dans le district<br />
d’Ikongo, la commune d’Ambolomadinika gère 12<br />
Fokontany, dont Antaranjaha et Tsianivoho. Le Fokontany<br />
Antaranjaha (S21°59’42.3",E047°25’40.2") est situé à 3,7 km<br />
au Sud-Ouest d’Ambolomadinika. Dans ce site, les villageois<br />
n’ont pas reconnu de lémuriens de bambou. Cependant, des<br />
cris de Hapalemur aureus ont été entendus dans la forêt à<br />
<strong>15</strong>0 m environ de notre campement,c’est-à-dire à Ankazondrano.A<br />
Marofototra,situé à 30 minutes du Fokontany d’Antaranjaha,<br />
toutes les jeunes pousses de bambou Cathariostachys<br />
sp.étaient coupées.Ce sont des signes de nourrissage de<br />
H. griseus. De plus, un villageois a confirmé avoir trouvé un<br />
groupe de H. griseus comprenant 12 individus à cet endroit.<br />
Nous avons également entendu des cris de Varecia variegata<br />
editorum, vu un groupe de Eulemur rufus, et trouvé un piège à<br />
lémuriens dans la forêt. Dans la commune rurale d’Ambolomadinika,<br />
on remarque beaucoup de zones agricoles déboisées.<br />
Malheureusement, les lémuriens de bambou sont menacés<br />
à cause de la coupe massive de bambous dans ces<br />
zones et la chasse pratiquée par les habitants de la Commune.<br />
Manambolo:Située dans la région de Fianarantsoa,la forêt de<br />
Manambolo (S22°04’06.2",E046°59’27.5",1238 m) se trouve<br />
dans le Fokontany de Morafeno, commune rurale de Sendrisoa.<br />
La gestion de la forêt est assurée par le FI.TE.MA<br />
(FIkambanan’ny TEraky MAnambolo) et concerne cinq villages:<br />
Mandamako, Mahavita, Ambinda, Ankazobe, Ampidira.<br />
La survie de la population locale dépend largement de l’agriculture,<br />
l’élevage et la production du rhum traditionnel.<br />
Cependant, la culture sur brûlis est encore pratiquée sur la<br />
lisière forestière. Lors de l’enquête, les villageois ont reconnu<br />
4 espèces de lémuriens, dont Hapalemur griseus et H.<br />
aureus. La vérification en forêt nous a révélé des signes de<br />
nourrissage de H.aureus, et nous avons vu directement Eulemur<br />
rufus et E. rubriventer.<br />
Résultats par zone située autour du corridor forestier de Fandriana-Vondrozo<br />
Mananjary: Notre campement à Tsararivotra (S21°10’41.8",<br />
E048°13’19.6", altitude 39 m) était situé à 23 km au nordouest<br />
de la ville de Mananjary. Le site de Tsararivotra est<br />
inclus dans le Fokontany de <strong>Vol</strong>omborona Asakatara et fait<br />
partie de la commune de Morafeno Mananjary.Nous n’avons<br />
trouvé que Hapalemur griseus, Cheirogaleus major et Microcebus<br />
rufus dans cette zone.<br />
Sahalanona: La commune de Sahalanona fait partie du<br />
District d’Ikongo et inclut 9 Fokontany (Sahalanona, Mahaly,<br />
etc.). La population est composée d’agriculteurs, d’éleveurs<br />
et de pêcheurs. Le village de Sahalanona (S22°03’19.2"<br />
E047°37’ 12.2", altitude 129 m) existe depuis environ 300<br />
ans. Malgré l’abondante présence de bambous, dont le<br />
bambou géant Cathariostachys sp., nous n’avons trouvé que<br />
Hapalemur griseus dans cette zone. Cette espèce est menacée<br />
par la chasse que pratiquent les villageois. D’autres<br />
espèces de lémuriens sont également en danger car elles<br />
sont aussi chassées et vendues par les villageois, notamment<br />
Avahi laniger (chassé pour l’alimentation et l’usage domestique)<br />
et Microcebus rufus (dont le prix est de 5.000 Ariary<br />
par individu).<br />
Manakara: Le village d’Ambila se trouve à 17 km au nord de<br />
Manakara. Le Fokontany Ambila fait partie de la commune<br />
d’Ambila (S22°00’11.6", E047°58’19.9") de la région de Manakara.Notre<br />
observation a été effectuée directement dans<br />
la forêt de Tsiazombazaha située à 10 km du village d’Ambila.<br />
L’enquête n’a pas eu lieu dans ce site car il n’y avait plus de<br />
village (principalement notre cible) autour de la forêt. Cette<br />
forêt est gérée par la communauté de base du Fokontany<br />
d’Ambila.A cet endroit,nous n’avons pas trouvé de bambou,<br />
et avons trouvé seulement deux espèces de lémuriens, Avahi<br />
laniger et Eulemur rufus, après vérification dans la forêt.<br />
Mahazoarivo/Ifasy: Située dans la région de Farafangana, la<br />
commune de Mahazoarivo (S22°39’49.0", E047°18’42.4",<br />
222 m) fait partie du corridor forestier, et la population pratique<br />
l’agriculture et l’élevage. L’exploitation des ressources<br />
minières, surtout des pierres précieuses, représente une<br />
source de revenus importante pour la population. Lors de<br />
l’enquête, les villageois n’ont reconnu que deux espèces de<br />
lémuriens,Hapalemur griseus et Microcebus rufus.Nous avons<br />
visité deux forêts dans cette commune,à Mitimboto (Fokontany<br />
de Mahazoarivo) et Ifasy ou Mahafasy (Fokontany Mahatsara)<br />
où deux groupes de H. griseus ont été vus sur chaque<br />
site. A Ifasy (S22°39’13.0", E47°14’56.1"), des individus de H.<br />
griseus de très grande taille ont été localisés, similaires à<br />
Prolemur simus, mais l’absence des touffes de poils blancs sur<br />
les oreilles nous a permis de faire la distinction. Nous avons<br />
également remarqué que le nom local de H. griseus était<br />
différent à Mitimbato et Ifasy,respectivement Varibolo madinika<br />
et Varibolo vaventy.<br />
Iandraina: Le Fokontany d’Iandraina fait partie de la commune<br />
Rurale de Vohimasy. Il se situe à <strong>15</strong> km au nord-ouest<br />
de Farafangana. La forêt de Befoza et celle d’Ambolosy<br />
(S22°46’07.0", E047°41’07.0", 53 m) se trouvent dans ce<br />
fokontany. Les populations sont constituées principalement<br />
d’agriculteurs et d’éleveurs. La pratique des cultures vivrières<br />
constitue l’activité principale. Contrairement aux<br />
autres sites que nous avons visités dans le sud de la zone<br />
d’étude, nous avons trouvé une population de Cathariostachys<br />
sp. à Ambolosy.<br />
Sahamadio: Située dans la région de Farafangana, cette zone<br />
est plus ou moins enclavée (absence d’infrastructure routière)<br />
et même la circulation et le transport de produits<br />
locaux s’effectuent toujours par pirogue.La commune rurale<br />
de Sahamadio (S22°31’13.4", E047°35’02.8", altitude 27 m)<br />
dépend beaucoup de l’agriculture. L’existence de signes de<br />
nourrissage dans la forêt de Sahamadio nous a permis d’établir<br />
que Hapalemur griseus,localement appelé "varibolo" y est<br />
présent.L’enquête effectuée au niveau de la commune rurale<br />
d’Ambalatany a également confirmé la présence de lémuriens<br />
de bambou de grande taille dans la forêt d’Ambalakazaha.<br />
Evato:Dans la commune d’Evato (S 22°36’42",E 047°41’20"),<br />
dans la région de Farafangana, le développement des différentes<br />
infrastructures est remarquable,citons comme exemple<br />
les hôpitaux, écoles, marchés et routes en bon état. Un<br />
bloc de forêt primaire se trouve à Iaboloha dans cette<br />
commune. Notre enquête nous a donné des informations<br />
sur la présence de plusieurs espèces de lémuriens dans cette<br />
forêt. Ce site mérite donc d`être visité pour une prochaine<br />
vérification.<br />
Mahafasa: Dans cette zone située également dans la région<br />
de Farafangana, ce qui reste de forêt primaire est en général<br />
la forêt de bambou,un endroit où se trouvent des tombeaux.<br />
Etant donné la situation actuelle de sécurité, nous n’avons<br />
pas obtenu la permission de visiter cette forêt sacrée de<br />
bambou. Cette dernière recouvre une grande surface, environ<br />
3 km de longueur et jusqu’à 100 m de largeur,et pourrait<br />
être importante en tant qu’habitat de lémuriens.
Page 38 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
Discussion<br />
La série d’expéditions menée le long du Corridor forestier<br />
Fandriana - Vondrozo nous a permis d’évaluer provisoirement<br />
la répartition des lémuriens de bambou. Concernant<br />
Prolemur simus,un seul signe de nourrissage a été identifié,et<br />
ce signe remontait à un an, confirmant les résultats des<br />
études précédentes qui indiquent que l’espèce a une distribution<br />
fragmentée dans la région (Wright et al., 2008). Pour<br />
Hapalemur aureus (espèce menacée EN), la découverte de<br />
l’évidence de sa présence sur six zones,toutes dans le corridor<br />
forestier, est encourageante car cela implique une large<br />
distribution dans celui-ci, bien que l’espèce ne semble pas<br />
exister en-dehors.Hapalemur griseus (espèce vulnérable VU)<br />
a été trouvé dans presque toutes les zones visitées, dans le<br />
corridor forestier mais également dans des zones éloignées<br />
de ce dernier.<br />
La menace principale pour les espèces de bambou dans la<br />
région du corridor Fandriana-Vondrozo est la destruction<br />
des habitats naturels et leur conversion en champs de culture.Cette<br />
technique est appelée "agriculture sur brûlis".Par<br />
conséquent, cette pression entraîne la raréfaction et même<br />
la disparition des espèces autochtones de bambous. Malgré<br />
la présence de bambous à l’intérieur du corridor, la persistance<br />
de la pratique du tavy,les coupes de bambous en permanence<br />
et surtout la chasse aux lémuriens mettent en péril<br />
la survie des espèces de lémuriens. En outre, la taille de la<br />
forêt du Corridor Fandriana-Vondrozo est petite par rapport<br />
aux autres corridors forestiers du pays. Sa largeur est<br />
très réduite surtout dans sa partie sud, et sur la photo<br />
aérienne, la voûte forestière apparaît très ouverte. Tous ces<br />
facteurs menacent la viabilité des populations de lémuriens<br />
vivant dans le corridor, et tout particulièrement les espèces<br />
présentant une distribution fragmentée telles que P. simus.<br />
Actuellement, beaucoup de lémuriens sont chassés et vendus<br />
par les villageois (exemple:Sahalanona,Antaranjaha).Les<br />
forêts de bambous sont fragmentées et isolées les unes des<br />
autres, ce qui laisse à penser que ces lémuriens de bambou<br />
sont réellement en danger. En outre, les utilisations des<br />
bambous dans la région sont nombreuses. La population locale<br />
utilise les différentes espèces de bambou suivant leur<br />
taille pour la construction des maisons, particulièrement<br />
pour les toitures, les murs, et des clôtures. Les bambous<br />
servent également à fabriquer du matériel pour les usages<br />
quotidiens, parmi lesquels les paniers à fruits, volailles, écrevisses<br />
et anguilles. Enfin, ils permettent de transporter des<br />
bagages. Aussi, les espèces de bambous de plus grand diamètre<br />
sont utilisées comme récipient pour transporter de<br />
l’eau. La conséquence négative de l’utilisation des bambous<br />
est minime par rapport à la destruction des habitats. Exemple:le<br />
Corridor d’Ampitsinjovabe (site d’Antarehimamy) est<br />
une bonne localité pour trouver H. griseus, H. aureus et V.<br />
variegata editorum, mais ces trois espèces sont menacées à<br />
cause de la chasse et des coupes sélectives de bois pratiquées<br />
par les habitants résidant autour du corridor.<br />
Pour la conservation de Prolemur simus,il faudrait accroître la<br />
taille des aires protégées en y incluant les forêts de bambous,<br />
et restaurer les fragments d’habitats isolés au sein d’un<br />
paysage agricole déboisé, afin d’équilibrer la valence écologique,<br />
c’est-à-dire la zone supportable pour l’espèce (en<br />
pratiquant une reforestation de bambou). Cependant, d’une<br />
façon générale, il y a un besoin immédiat de sensibilisation,<br />
pour conscientiser la population aux problèmes de coupe de<br />
bambous,de tavy et de chasse des lémuriens,afin d’assurer la<br />
survie d’espèces de lémuriens dans et autour du Corridor<br />
Fandriana-Vondrozo. Finalement, les sites d’Ambodiara, Mahazoarivo<br />
(Alafady, Ranomena), Ambalakazaha et Mahafasa<br />
sont recommandés pour une nouvelle vérification de la présence<br />
ou non des lémuriens de bambou. En effet, la population<br />
locale semble être convaincue d’avoir trouvé P. simus à<br />
ces endroits.<br />
Remerciements<br />
Nos vifs remerciements vont: au Ministère de l’Environnement,<br />
des forêts et du Tourisme, à la Direction Générale de<br />
l’Environnement et des forêts, et à la Direction du Système<br />
des Aires Protégées, Madagascar, pour leur accord et la<br />
délivrance de l’autorisation de recherche (permis n°279/08/<br />
MEFT/SG/DGEF/DSAP/SSE); à The Aspinall Foundation, GB,<br />
pour le financement de l’enquête dans le cadre du Projet<br />
"Sauver Prolemur simus";au Groupe d’Etude et de Recherche<br />
sur les Primates de Madagascar (G.E.R.P) et son personnel<br />
administratif; au Centre International de Formation pour la<br />
Valorisation de la Biodiversité (Centre ValBio) et son personnel<br />
administratif; à l’ICTE et Conservation International,<br />
Antananarivo,pour leurs conseils et entire collaboration;aux<br />
communes, Fokontany, et COBAS des zone visitées pour<br />
leurs amabilité et collaboration; et enfin, aux assistants de<br />
recherche du Centre ValBio à Ranomafana, Justin Rakotonjatovo,<br />
Dominique Razafindraibe, Jean-Guy Razafindraibe,<br />
Aime-Victor Tombotiana et Telo Albert, et au chauffeur de<br />
The Aspinall Foundation,Mohamad Mbaraka,pour leur assistance<br />
sur le terrain.<br />
Références<br />
Andriaholinirina, V.N.; Fausser J.L.; Rabarivola, J.C. 2003. Etude<br />
comparative de Hapalemur simus (Gray,1870) de deux<br />
sites de la province autonome de Fianarantsoa, Madagascar:forêt<br />
dégradée d’Ambolomavo et forêt secondaire de<br />
Parc National de Ranomafana. Lemur News 8: 9-13.<br />
Dolch, R.; Fiely, J.L.; Ndriamiary, J.N.; Rafalimandimby, J.; Randriamampionona,R.;Engberg,S.E.;Louis,E.E.Jr.2008.Confirmation<br />
of the greater bamboo lemur, Prolemur simus,<br />
north of the Torotorofotsy wetlands,eastern Madagascar.<br />
Lemur News 13: 14-17.<br />
Godfrey, L.; Vuillaume-Randriamanantena, M. 1986. Hapalemur<br />
simus: endangered lemur once widespread. Primate<br />
Conservation 7: 92-96.<br />
Godfrey, L.R.; Simons, E.L.; Jungers, W.L.; DeBlieux, D.D.;<br />
Chatrath,P.S.2004.New discovery of subfossil Hapalemur<br />
simus, the greater bamboo lemur, in western Madagascar.<br />
Lemur News 9: 9-11.<br />
Jones,J.P.G.;Andriahajaina,F.B.;Hockley,N.J.;Balmford,A.;Ravoahangimala,<br />
O.R. 2005. A multidisciplinary approach to<br />
assessing the sustainability of freshwater crayfish harvesting<br />
in Madagascar.Conservation Biology 19:1863-1871.<br />
King, T.; Chamberlan, C. 2010. Conserving the critically endangered<br />
greater bamboo lemur Prolemur simus.Oryx 44:<br />
167.<br />
Meier, B.; Rumpler, Y. 1987. Preliminary survey of Hapalemur<br />
simus and of a new species of Hapalemur in eastern Betsileo,<br />
Madagascar. Primate Conservation 8: 40-43.<br />
Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis E.E.;<br />
Langrand, O.; Ratsimbazafy, J.; Rasoloarison, R.; Ganzhorn,<br />
J.U.; Rajaobelina, S.; Tattersall, I.; Meyers, D.M. 2006. Lemurs<br />
of Madagascar. 2nd ed. Conservation International,<br />
Washington, D.C.<br />
Mittermeier, R.A.; Ratsimbazafy, J.; Rylands, A.B.; Williamson,<br />
L.;Oates,J.F.;Mbora,D.;Ganzhorn,J.U.;Rodriguez-Luna,E.;<br />
Palacios, E.; Heymann, E.W.; Cecilia, M.; Kierfull, M.; Yongcheng,L.;Supriatna,J.;Roos,C.;Walker,S.;Aguiar,J.M.2007.<br />
Primates in Peril: The World’s 25 Most Endangered Primates<br />
2006-2008. Primate Conservation 22: 1-40.<br />
Mittermeier, R.A.; Wallis, J.; Rylands, A.B.; Ganzhorn, J.U.;<br />
Oates, J.F.; Williamson, E.A.; Palacios, E.; Heymann, E.W.;<br />
Kierulff,M.C.M.;Yongcheng,L.;Supriatna,J.;Roos,C.;Walker,<br />
S.; Cortés-Ortiz, L.; Schwitzer, C. 2009. Primates in<br />
Peril: The World’s 25 Most Endangered Primates 2008-<br />
2010. Primate Conservation 24: 1-57.<br />
Ratolojanahary,M.;Rajaonson,A.;Ratsimbazafy,J.;Feistner,A.;<br />
King, T. 2009. Identification des sites prioritaires pour la
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 39<br />
conservation de Prolemur simus par la récolte des connaissances<br />
indigènes locales sur les distributions respectives<br />
du bambou et des hapalémurs dans et autour du corridor<br />
Fandriana-Vondrozo: Rapport Final. GERP / The Aspinall<br />
Foundation / Centre ValBio, Madagascar. 25 pp.<br />
Sterling E.J.; Ramaroson, M.G. 1996. Rapid assessment of the<br />
primate fauna of the eastern slopes of the Réserve Naturelle<br />
Intégrale d’Andringitra, Madagascar. In: S.M. Goodman<br />
(ed.), A Floral and Faunal Inventory of the Eastern<br />
Slopes of the RPserve Naturelle Intégrale d’Andringitra,<br />
Madagascar, with Reference to Elevational Variation. Fieldiana<br />
Zoology 85: 293-305.<br />
TAF 2008. Projet Varibolomavo: Sauver Prolemur simus -Objectifs<br />
et actions proposées. The Aspinall Foundation,<br />
Port Lympne Wild Animal Park, Kent, GB. 4 pp.<br />
TAF 2009. Projet Varibolomavo: Sauver Prolemur simus -Premiers<br />
résultats et actions immédiates. The Aspinall Foundation,<br />
Antananarivo, Madagascar. 6 pp.<br />
Tan, C.L, 1999.Group Composition, Home Range Size, and<br />
Diet of Three Sympatric Bamboo lemur species (genus<br />
Hapalemur) in Ranomafana National Park, Madagascar.<br />
International Journal of Primatology 20(4): 547-566.<br />
UICN 2009. IUCN Red List of Threatened Species. Version<br />
2009.2. www.iucnredlist.org.<br />
Wright P.C.; Daniels P.S.; Meyers, D.M.; Overdorff, D.J.; Rabesoa,<br />
J.A. 1987. Census and study of Hapalemur and Propithecus<br />
in Southeastern Madagascar.Primate Conservation<br />
8: 84-88<br />
Wright,P.C.;Johnson,S.E.;Irwin,M.T.;Jacobs,R.;Schlichting,P.;<br />
Lehman, S.; Louis, E.E. Jr.; Arrigo-Nelson, S.J.; Raharison,<br />
J.-L.; Rafalirarison, R.R.; Razafindratsita, V.; Ratsimbazafy, J.;<br />
Ratelolahy, F.J; Dolch, R.; Tan, C. 2008. The Crisis of the<br />
Critically Endangered GreaterBamboo Lemur (Prolemur<br />
simus). Primate Conservation 23: 5-17.<br />
Effect of red ruffed lemur gut passage on<br />
the germination of native rainforest plant<br />
species<br />
Onja H. Razafindratsima 1,2*, Emilienne Razafimahatratra<br />
1<br />
1Department of Animal Biology, University of Antananarivo,<br />
Madagascar<br />
2Department of Ecology and Evolutionary Biology, Rice University<br />
– MS 170, 6100 Main St., Houston, TX 77005, USA<br />
(current affiliation)<br />
*Corresponding author: ohr1@rice.edu, onjhar@hotmail.com<br />
Key words: seed dispersal, germination success, Varecia<br />
rubra, primate, corridor restoration, Masoala<br />
Abstract<br />
Like much of Madagascar’s remaining rainforest,the forest of<br />
Masoala National Park is facing severe threats from deforestation<br />
and fragmentation. The remaining fragmented areas<br />
are connected by degraded corridors which are important<br />
for biological exchange. Frugivorous animals such as lemurs<br />
may have an important role in the restoration of such degraded<br />
areas through seed dispersal. Unfortunately,no studies<br />
have been carried out before concerning the role lemurs<br />
play in the restoration of the largest corridor in Masoala,<br />
Ambatoledama. This study explores the effect of seed passage<br />
inside the gut of the frugivorous red-ruffed lemur<br />
(Varecia rubra) on the germination of some native tropical<br />
plants with the aim to understand the capacity of V. rubra to<br />
help in the restoration of the Ambatoledama corridor. We<br />
planted seeds of nine plant species that we collected from V.<br />
rubra’s fresh feces in a nursery to compare with seeds that<br />
we extracted manually from corresponding fruits. The germination<br />
of seeds was monitored each month after planting<br />
them. Results showed that defecated seeds had overall a significantly<br />
higher germination rate than non-passed seeds.<br />
Thus, lemur ingestion of seeds has the capacity to improve<br />
seed germination of several species and some plants require<br />
the physiological treatment inside the gut to germinate. Results<br />
suggested that restoration projects in the area including<br />
the Ambatoledama corridor should take into account the<br />
important role Varecia rubra plays in the regeneration of the<br />
forest and corridor.Management actions that increase movement<br />
and protection of animals moving into and out of the<br />
corridor will be important for the long term success of the<br />
project.<br />
Introduction<br />
The rainforest of the Masoala Peninsula suffers greatly from<br />
loss and fragmentation caused by the human population<br />
living around the area.The forest is subdivided into different<br />
fragments, connected by corridors of degraded habitat<br />
which are Ambatoledama,Analambolo and Ilampy (Holloway,<br />
1997). Corridors are vital for enabling gene flow and dispersal<br />
of wildlife among habitat fragments (Mech and Hallett,<br />
2001). The largest of these is the Ambatoledama corridor,<br />
which connects two large parcels of the forest (Fig. 1). The<br />
restoration of this corridor is critical for safeguarding wildlife<br />
populations in the fragments and for preserving gene flow<br />
between fragments (Mech and Hallett, 2001; Haddad et al.,<br />
2003). To restore this degraded corridor, it is necessary to<br />
plant native trees or to encourage zoochory (biological dispersal<br />
of seeds through animal defecation) (Duncan and<br />
Chapman, 2002; Neilan et al., 2006). Since 1997, Madagascar<br />
National Parks (MNP) and the Wildlife Conservation Society<br />
(WCS) have established a restoration project in the Ambatoledama<br />
corridor by planting native fruiting trees (Holloway,1997)<br />
with the aim of attracting frugivorous vertebrates<br />
which will in turn carry seeds into the degraded parts of the<br />
forest and into forest clearings. Unfortunately, no studies<br />
have previously been carried out to shed light on the importance<br />
of frugivorous animals,especially lemurs,in the reforestation<br />
of the Ambatoledama corridor.Unlike the majority of<br />
tropical forests,the diversity of the frugivorous bird community<br />
in Madagascar is impoverished, and therefore primates<br />
are the principal dispersers of its tropical trees (Goodman,<br />
1997; Dew and Wright, 1998; Ganzhorn et al., 1999; Bleher<br />
and Böhning-Gaese,2001).Ten lemur species are indentified<br />
as living in the Masoala Forest (Mittermeier et al., 2006); one<br />
of which (Varecia rubra) is endemic to this region and has Endangered<br />
status (IUCN, 2008), and can be found in both the<br />
corridor habitat and adjacent forest fragments (Razakamaharavo<br />
et al., 2010). Previous studies demonstrated that<br />
Varecia variegata is an effective disperser in the southeastern<br />
rainforests (Dew and Wright,1998).However,we know very<br />
little about the potential role of V.rubra for regeneration and<br />
restoration of the corridor habitat in Ambatoledama.<br />
In this study, we explored the germination success of seeds<br />
defecated by Varecia rubra in order to understand their capacity<br />
for seed dispersal and potential impact on the restoration<br />
of the degraded rainforest corridor at Ambatoledama.<br />
Our objective was to shed light on the role of this species in<br />
forest regeneration. Understanding their influence on tree<br />
germination is particularly important given the threatened<br />
status of this species. This paper tested the hypothesis that<br />
gut passage of seeds by Varecia rubra facilitates seed germination.<br />
Our prediction was that lemur-gut-passed seeds have a<br />
higher germination rate than non-passed seeds because of<br />
the physiological treatment affecting the seed coat inside the<br />
gut.
Page 40 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
Materials and methods<br />
Field site<br />
This study was carried out at the Ambatoledama corridor<br />
(S<strong>15</strong>°27’ E050°01’) on the north-eastern part of the Masoala<br />
Peninsula. Ambatoledama connects Masoala National Park<br />
with Makira National Forest to the West. Its forest has undergone<br />
significant deforestation but restoration projects<br />
have augmented Ambatoledama such that it now forms a 1<br />
km wide corridor of secondary forest (Hekkala et al., 2007;<br />
Razakamaharavo et al.,2010).It consists of a dense evergreen<br />
rainforest with an altitude ranging from 300 to 700 m. The<br />
forest is mostly characterized by the presence of tree species<br />
of the Pandanacea, Ebenaceae, Clusiaceae, Euphorbiaceae,<br />
Sapotaceae and Rubiaceae families (Martinez, unpublished).<br />
Study species<br />
Varecia rubra belongs to the family Lemuridae (Gray, 1821)<br />
and is one of two species recognized within the genus (Mittermeier<br />
et al., 2006). V. rubra is only found on the Masoala<br />
peninsula and it is classified by the World Conservation Union<br />
(IUCN) as Endangered (IUCN, 2008). V. rubra is a largesized<br />
diurnal species with a body length ranging from 43 to<br />
57 cm (Vasey, 2003) and has a typically frugivorous diet<br />
(Rigamonti, 1993; Vasey, 1997). They currently inhabit both<br />
the corridor habitat and the adjacent protected areas (Razakamaharavo<br />
et al., 2010) and are thus potentially important<br />
for regeneration of the corridor habitat.<br />
Field experiment<br />
Focal animals were followed for 3-5 days per week from<br />
dawn to dusk (from 0600 hours to 1800 hours) to collect<br />
fresh fecal samples (Dew and Wright,1998;Kaplin and Moermond,1998;Stevenson,2000;Poulsen<br />
et al.,2001;Link and Di<br />
Fiore, 2006). Each fecal sample was washed and filtered<br />
through a 1-mm sieve (Stevenson,2000).Seeds were extracted<br />
and then identified with the help of local research guides<br />
and an expert local botanist familiar with the Masoala flora.<br />
We planted gut-passed seeds and control seeds that were<br />
extracted manually from fruits in an outdoor nursery adjacent<br />
to the corridor at Ambatoledama. The nursery consisted<br />
of two "flower beds" of 11.2 m 2: one for defecated<br />
Fig. 1: Location of the Ambatoledama corridor.<br />
seeds and the other one for non-passed seeds. Following<br />
methods used by the conservation agents of MNP in Ambatoledama,<br />
a sunshade of 80 cm height, composed of Longoza<br />
leaves (Afromomum angustifolium) was placed above each<br />
flower bed to imitate the closed canopy of the forest. Also,<br />
the soil of the nursery was mixed with fertile soil from cultivated<br />
field. Seeds were placed in the soil mixture and<br />
covered by 1 mm-thick river sand to keep a constant temperature.<br />
An equal number of seeds were planted within each species<br />
per treatment. However, the numbers varied between species<br />
depending on how many seeds were collected from<br />
lemur feces. The germination of seeds was assessed each<br />
month after planting.<br />
Data analysis<br />
We performed a paired t-test to test for differences between<br />
the germination rate of lemur-gut-passed and nonpassed<br />
seeds, an ANOVA analysis to test if the two factors<br />
(seed species and treatment) had effects on the germination<br />
rate of the seeds and to determine whether there was interaction<br />
between these factors. We analyzed the germination<br />
of each species in order to assess the influence by lemur gut<br />
passage, with Pearson test using contingency tables, which<br />
was adjusted with Bonferroni correction for multiple comparisons<br />
(Sokal and Rohlf, 1995).<br />
Results<br />
In total, 268 fresh fecal samples from three individuals of<br />
red-ruffed lemur were collected during 58 days of observation.<br />
The fecal samples contained fleshy fruit parts, stalks,<br />
leaves, soil and fecal liquid. 95.52 % of these contained seeds,<br />
to some of which fleshy fruit parts were still attached. 906<br />
seeds of more than 1mm size were extracted. A majority of<br />
them were intact with minor scarification.They represented<br />
34 different plant species that belong to <strong>15</strong> Families.Based on<br />
our collected sample, the most common seed species found<br />
in lemur defecations were the nine species we chose to<br />
study here (Tab. 1). In the nursery, we planted a total of 390<br />
defecated seeds and compared them with 398 non-passed<br />
seeds.<br />
Lemur-gut-passed seeds had significantly higher germination<br />
rates overall than non-passed seeds (t=3.284,df=8,p=0.011).<br />
Passed seeds had a germination rate<br />
of 64.61 %, whereas non-passed<br />
seeds had a rate of 39.69 %.For each<br />
species, seeds that had been defecated<br />
had a higher germination rate<br />
than non-passed seeds, except for<br />
Tsilaitra (Tab. 1). This pattern was<br />
driven primarily by four species, including<br />
Antaivaratra,Matahobaratra,<br />
Tsilaitra, and Vongobe species.<br />
In a two factor analysis of variance<br />
for seed germination, there was a<br />
significant interaction between the<br />
species of seeds and their treatment<br />
(passed or non-passed) (F= 4.2004,<br />
p
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 41<br />
Tab. 1: List of species studied with their germination rate. Sample sizes are represented in<br />
brackets. The star on p-values corresponds to their significance (Pearson test) after a<br />
Bonferroni correction for multiple tests.<br />
# Malagasy Scientific Family Germination rate Pearson test<br />
name name<br />
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*<br />
2 Hazondronono Sideroxylon Sapotaceae 80.00 (n = 10) 60.00 (n = 10) 0.952 0.3291<br />
3 Karaka Pandanus Pandanaceae 40.00 (n = 20) <strong>15</strong>.00 (n = 20) 3.135 0.0766<br />
4 Matahobaratra Garcinia sp. Clusiaceae 58.06 (n = 31) 00.00 (n = 31) 25.364
Page 42 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
Dokolahy, J. 2005. Etude de la régénération naturelle des<br />
savoka du pont forestier d’Ambatolaidama en vue d’une<br />
restauration forestiere.Unpubl.DEA thesis,ESSA University<br />
of Antananarivo, Madagascar.<br />
Duncan,R.S.;Chapman,C.A.2002.Limitations of animal seed<br />
dispersal for enhancing forest succession on degraded<br />
lands. Pp. 437-450. In: D.J. Levey; W.R. Silva; M. Galetti<br />
(eds.). Seed Dispersal and Frugivory: Ecology, Evolution<br />
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1999. Lemurs and the regeneration of dry deciduous forest<br />
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Bay of Antongil in northeastern Madagascar. Primate<br />
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Holloway, L. 1997. Catalysing natural regeneration of rainforest:<br />
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to Wildlife Conservation Society, Madagascar.<br />
Howe, H.F. 1986. Seed dispersal by fruit-eating birds and<br />
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Howe, H.F.; Smallwood, J. 1982. Ecology of seed dispersal.<br />
Annual Review of Ecology and Systematics 13: 201-228.<br />
Into,I.2009.Investigation into the illegal felling,transport and<br />
export of precious wood in the SAVA region of Madagascar.<br />
Global Witness and the Environmental Investigation<br />
Agency, Inc. Report.<br />
IUCN. 2008. Red List of Threatened Species.<br />
www.iucnredlist.org.<br />
Kaplin, B.A.;Lambert,J.E. 2002.Effectiveness of Seed Dispersal<br />
by Cercopithecus Monkeys: Implications for Seed Input<br />
into Degraded Areas.Pp.351-364.In:D.J.Levey;W.R.Silva;<br />
M. Galetti (eds.). Seed Dispersal and Frugivory: Ecology,<br />
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UK.<br />
Kaplin,B.A.;Moermond,T.C.1998.Variation in seed handling<br />
by two species of forest monkeys in Rwanda. American<br />
Journal of Primatology 45: 83-101.<br />
Link, A.; Di Fiore, A. 2006. Seed dispersal by spider monkeys<br />
and its importance in the maintenance of neotropical<br />
rain-forest diversity. Journal of Tropical Ecology 22: 235-<br />
246.<br />
McKey,D.1975.The ecology of coevolved seed dispersal systems.Pp.<strong>15</strong>9-191.In:L.E.Gilbert,P.H.Raven(eds.).Coevolution<br />
of Animals and Plants. University of Texas Press,<br />
Austin, TX, USA.<br />
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corridors: a genetic approach. Conservation Biology <strong>15</strong>:<br />
467-474.<br />
Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis, E.E.;<br />
Langrand, O.; Ratsimbazafy, J.; Rasoloarison, R.; Ganzhorn,<br />
J.U.;Rajaobelina,S.;Tattersall, I. 2006.Conservation International<br />
Tropical Field Guide Series: Lemurs of Madagascar.<br />
Conservation International, Washington, DC, USA.<br />
Neilan, W.; Catterall, C.P.; Kanowski, J.; McKenna, S. 2006. Do<br />
frugivorous birds assist rainforest succession in weed<br />
dominated old field regrowth of subtropical Australia?<br />
Biological Conservation 129: 393-407.<br />
Poulsen, J.R.; Clark, C.J.; Smith, T.B. 2001. Seed dispersal by a<br />
diurnal primate community in the Dja Reserve, Cameroon.<br />
Journal of Tropical Ecology 17: 787-808.<br />
Razakamaharavo, V.R.; McGuire, S.M.; Vasey, N.; Louis, E.E.;<br />
Brenneman, R.A. 2010. Genetic architecture of two red<br />
ruffed lemur (Varecia rubra) populations of Masoala National<br />
Park. Primates 51: 53-61.<br />
Rigamonti, M.M. 1993. Home range and diet in red ruffed<br />
lemurs (Varecia variegata rubra) on the Masoala Peninsula,<br />
Madagascar. Pp. 25-40. In: P.M. Kappeler; J.U. Ganzhorn<br />
(eds.). Lemur social systems and their ecological basis.<br />
Plenum Press, New York, USA.<br />
Schupp, E.W. 1993. Quantity, quality and the effectiveness of<br />
seed dispersal by animals. Plant Ecology 107: <strong>15</strong>-29.<br />
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massacre. Madagascar Conservation & Development 4<br />
(2): 98-102.<br />
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of statistics in biological research. WH Freeman, New<br />
York, USA.<br />
Stevenson, P.R. 2000. Seed dispersal by woolly monkeys (Lagothrix<br />
lagothricha) at Tinigua National Park, Colombia:<br />
dispersal distance, germination rates, and dispersal quantity.<br />
American Journal of Primatology 50(4): 275-289.<br />
Vasey, N. 1997. Community ecology and behavior of Varecia<br />
variegata rubra and Lemur fulvus albifrons on the Masoala<br />
Peninsula, Madagascar. Unpubl. Ph.D. thesis, Washington<br />
University, St. Louis, USA.<br />
Vasey, N. 2003. Varecia, ruffed lemurs. Pp. 1332-1336. In: S.M.<br />
Goodman; J.P. Benstead (eds.). The Natural History of<br />
Madagascar. Chicago University Press, Chicago, USA.<br />
Wehncke, E.V.; Dalling, J.W. 2005. Post-dispersal seed removal<br />
and germination of selected tree species dispersed by<br />
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37: 73-80.<br />
Feeding ecology of the crowned sifaka<br />
(Propithecus coronatus) in a coastal dry<br />
forest in northwest Madagascar (SFUM,<br />
Antrema)<br />
Claire Pichon 1*, Rivo Ramanamisata 2, Laurent Tarnaud<br />
1, Françoise Bayart 1, Annette Hladik 1, Claude<br />
Marcel Hladik 1, Bruno Simmen 1<br />
1UMR 7206, Eco-anthropologie et Ethnobiologie, Centre<br />
National de la Recherche Scientifique,and Museum National<br />
d’Histoire Naturelle, 4 avenue du Petit Château, 91800 Brunoy,<br />
France<br />
2Département de Biologie Animale,Faculté des Sciences,B.P.<br />
906, Université d’Antananarivo, Antananarivo (101), Madagascar<br />
*Corresponding author: cpichon@mnhn.fr<br />
Key words: diet, primate, activity budget, forest composition<br />
The crowned sifaka (Propithecus coronatus; Milne-Edwards,<br />
1871) inhabits dry forests, riparian forests and mangroves of<br />
northwest Madagascar. Originally believed to occur in a restricted<br />
area between the Mahavavy River in the southwest<br />
(where it overlaps with P.deckenii) and the Betsiboka River in<br />
the northeast (which separates it from P. coquereli), sightings<br />
west of the Mahavavy River and along the Bongolava Massif<br />
suggest that the distribution of this medium-sized species is<br />
wider (Tattersall, 1986; Thalmann et al., 2002). The distribution<br />
and the taxonomic status of crowned sifakas have long<br />
been debated, but the combination of morphological and<br />
biogeographic evidence supports considering it as a valid<br />
species (Thalmann et al., 2002; Mittermeier et al., 2006;<br />
Groves and Helgen, 2007; Mittermeier et al., 2008). Considered<br />
as Endangered (A2 c,d) by the IUCN (2010), populations<br />
of crowned sifakas were estimated not to exceed 1,000<br />
individuals in the wild.However,recently discovered populations<br />
in restricted fragmented forests extend the species’
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 43<br />
distribution range farther towards the Southwest. A "Biocultural<br />
Project" was therefore initiated at Antrema (a site<br />
located in the Mahajanga region) in 2000 to promote sustainable<br />
management (Gauthier et al., 2001).The project aims at<br />
preserving a coastal environment in which crowned sifakas<br />
occur in high densities while allowing villagers, mainly fishermen,<br />
to use natural products of the environment with parsimony<br />
and to benefit from technical and economical help.The<br />
project also aims at promoting local socio-cultural rules and<br />
a way of life that tends to respect the forest environment,including<br />
useful plants and several sympatric lemur species<br />
(Propithecus coronatus, Eulemur fulvus, Eulemur mongoz, Lepilemur<br />
sp., Microcebus murinus). The site contains three of the<br />
Northwest’s typical ecosystems (dry semi-deciduous forest,<br />
mangrove swamp, savanna), which suffer moderate anthropogenic<br />
pressure (Gauthier et al., 2001). Owing to local<br />
beliefs, especially the sacred (“masina”) nature of sifakas, the<br />
Sakalava community plays a central role in this conservation<br />
process (Harpet et al.,2000,2008).In this context,a few studies<br />
started investigating the behavior in relation to habitat<br />
and food supply of the lemur species of Antrema (Gauthier et<br />
al., 1999; Razafimahefa, 2001; Ramanikirahina, 2004). However,a<br />
detailed analysis of the feeding ecology and population<br />
densities of P. coronatus is still lacking. We present here preliminary<br />
data on the plant species composition of the whitesand<br />
coastal forest inhabited by a dense population of sifakas<br />
(among other prosimian species) and on the feeding ecology<br />
of sifaka groups censused since 2008.<br />
Methods<br />
Study site<br />
The Antrema station is a coastal area of 12,300 ha located on<br />
the left riverside of the Betsiboka estuary, northwestern<br />
Madagascar (<strong>15</strong>°42’-<strong>15</strong>°50’S, 46°-46°<strong>15</strong>’E; Gauthier et al.,<br />
2001). The region undergoes a distinct dry season of 7<br />
months from April to October.The mean annual rainfall (n =<br />
9 years) in the Mahajanga region is 1,410 mm (with a peak in<br />
January-February), with irregular rainfall during the dry season.With<br />
an annual mean of 27° C,temperature is highest in<br />
October and lowest between June and August (Airport of<br />
Mahajanga, 2000-2009).<br />
Although the Antrema area has been traditionally protected<br />
by the local Sakalava beliefs, forest areas where studies are<br />
conducted are fragmented. After two first surveys in November<br />
2007 and April-May 2008, we decided to establish<br />
the study site at Badrala (<strong>15</strong>°45.665’S, 46°12.300’E). With<br />
about 24 ha just behind the littoral dune,this non-sacred forest<br />
site offers suitable conditions to study the socioecology<br />
of sifakas and the dynamics of a dry forest in Madagascar.The<br />
forest there is partly split by a sandy open dune that sifaka<br />
groups can cross easily. Tree logging occurs at low intensity<br />
(with few selected species for defined use, e.g. for boats or<br />
coffins) and small trees are sporadically cut for fences and<br />
house building. We studied floristic composition by inventorying<br />
trees along four North/South-oriented line transects,<br />
10 m-wide each, that were roughly perpendicular to the sea<br />
front.Within this 0.73 ha,we tagged each tree > 10 cm diameter<br />
at breast height (DBH) with plastic labels, recorded<br />
their DBH, the number of stems and their vernacular name.<br />
Likewise, we counted woody lianas and herbaceous vines<br />
> 1 m high within eight 10 x 10 m (800 m 2) plots regularly<br />
spaced along the transects. Plant species were sampled and<br />
dried for later botanical identification.<br />
Sifaka population density<br />
In order to locate and identify groups of P. coronatus in<br />
Badrala, we initially mapped groups encountered during<br />
repeated transect walks. We drew individuals’ facial masks<br />
for each of the followed groups, noted their sex from visual<br />
inspection of the genitalia and other external characteristics<br />
(cysts, scars, damaged ears, fur colour), and took pictures.<br />
Knowledge gained progressively of groups and individuals<br />
allowed us to provide a preliminary estimate of population<br />
density for the Badrala site.<br />
Behavioral data collection<br />
We collected behavioral data during 4 periods (06 to 21 July<br />
2008; 11 November to 12 December 2008; 05 April to 06<br />
June 2009;17 October to 22 November 2009). Most groups<br />
were already accustomed to the sporadic presence of local<br />
people. Once we could observe animals at close distances,<br />
we followed each group successively over 2 to 5-day periods,<br />
from 06:30 to 18:30 hours.<br />
We used the instantaneous scan-sampling method (Altmann,<br />
1974) to study group activity budget.Every 5 minutes,we recorded<br />
the individuals’ activity using one of the following categories:resting<br />
(immobile,with eyes open or closed),moving<br />
(more than 0.5 m),foraging (searching for a food item),feeding<br />
(processing or chewing a food item), social activity (displaying<br />
agonistic and affiliation behaviors with other individuals)<br />
and other miscellaneous behaviors. We noted the plant<br />
part and species eaten by individuals.<br />
Besides recording activity budgets,we determined diet from<br />
mouthful counts converted into weight of ingested matter<br />
(Hladik, 1977) for 2 periods: April-June 2009 and October-<br />
November 2009.We estimated food intake in focal individuals<br />
that were followed continuously for 30 minutes each.<br />
Observations were alternated across males and females (excluding<br />
juveniles) within groups.<br />
Results<br />
Forest composition<br />
Plant families occurring at Badrala are presented separately<br />
for trees and lianas/vines in Fig.1.To date,91 tree and liana or<br />
vine species have been identified at least at the family level,<br />
and taxonomic identification of <strong>15</strong> more putative species is<br />
still in progress.The 5 richest families in terms of the number<br />
of species are Fabaceae, Sapindaceae, Ebenaceae, Euphorbiaceae<br />
and Apocynaceae.The most dominant tree species are<br />
Strychnos decussata, Vitex beraviensis, Mimusops occidentalis,<br />
Baudouinia fluggeiformis and Macphersonia gracilis that represent<br />
almost one third of total basal area and tagged trees.<br />
Combretum coccineum, Hypoestes sp., Landolphia perrieri and<br />
Reissantia sp.accounted for more than one third of the lianas<br />
and vines.<br />
Density of trees inventoried on the 4 transects (n=486) corresponds<br />
to 666 inds. ha 1 with a total basal area of 14.5 m²<br />
ha -1.We found a high density of woody lianas and herbaceous<br />
vines in the 800m² plots (n=373).<br />
Sifaka population density and group composition<br />
Groups at Badrala have 1-3 breeding adult males, 1-4 breeding<br />
adult females, and 1-4 immature offspring. We encountered<br />
between <strong>15</strong> and 20 groups at this site. Based on current<br />
recognition of individuals within these groups, a minimum<br />
estimate of 300 inds. km 2 was calculated. Mean size of<br />
focal groups was 4.3 ± 1.8 individuals (n=16).<br />
Diet and activity pattern<br />
Sifakas consumed at least 60 plant species from 32 families.<br />
Tab. 1 lists major food species eaten. During the dry season,<br />
14 plant species represented 75 % of the diet whereas only 7<br />
species were the main food resource in the wet season.
Page 44 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
Tab. 1: Food species accounting for 50 and 75 % of the diet of Propithecus coronatus during the dry season and the wet season.<br />
Eaten plant species are listed in decreasing order and their abundance in transects and plots (see text) is indicated.<br />
Family Species Vernacular name Items Abundance (%)<br />
Dry season<br />
Lamiaceae Vitex beraviensis Mojiro yl 10,7<br />
Fabaceae Baudouinia fluggeiformis Manjakabentany yl ml 4.7<br />
Sapotaceae Mimusops occidentalis Natofotsy yl stems 3.9<br />
Anacardiaceae Operculicarya gummifera Atokonjo ml buds 3.5 <br />
Sapindaceae Majidea zanguebarica Tsipopoka yl ml fl 2.3<br />
Oleaceae Noronhia boinensis Tsilaitra beravina yl 1.9<br />
Moraceae Trilepisium occidentalis Kililo ml 1.2<br />
Sphaerosepalaceae Rhopalocarpus lucidus Hazondringitra yl fr 1.0<br />
Melastomataceae Warneckea sp. Voatrotrokoala yl 0.6 75 %<br />
Burseraceae Commiphora sp. Arofy fr buds 0.4<br />
Fabaceae Bussea perrieri Morango ml 0.2<br />
Olacaceae Olax dissitiflora Ambiotsy ml 0.2<br />
Moraceae Ficus pyrifolia Nonika fr<br />
Unidentified - RR80 ml<br />
Wet season<br />
Anacardiaceae Abrahamia deflexa Motsovavy yl fl 3,1<br />
Anacardiaceae Abrahamia sp. Manavodrevo buds yl fl 1.2<br />
Fabaceae Chadsia flammea Fanamohazo buds yl fl 0.8 50 %<br />
Sapotaceae Capurodendron gracilifolium Natoboay buds yl fr 0.2<br />
Apocynaceae Landolphia perrieri Vahipira yl 6.8 75 %<br />
Combretaceae Terminalia sp. Taly buds yl 3.1<br />
Anacardiaceae Operculicarya gummifera Atokonjo yl 3.5<br />
yl: young leaves; ml: mature leaves; fl: flowers; fr: fuits<br />
Fig.1:Abundance of plant families plotted in<br />
decreasing number of individuals among a)<br />
trees with DBH>10cm (based on transects;<br />
0.73 ha) and b) woody lianas and herbaceous<br />
vines >1 m height (based on plots;<br />
0.08 ha).Striped bars refer to the plant families<br />
with the highest number of species.
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 45<br />
58%<br />
(a) Dry season (b) Wet season<br />
9%<br />
2%<br />
1%<br />
30%<br />
25%<br />
11%<br />
6%<br />
1%<br />
Sifakas were highly folivorous during both seasons, supplementing<br />
their diet with flowers, fruits, vegetative buds, and<br />
sometimes young stems (Fig. 2). They consumed more mature<br />
leaves and fruits in the dry season and more flowers in<br />
the wet season.<br />
The activity budget of the sifakas is presented in Fig. 3. Although<br />
‘resting’ predominated throughout the study,<br />
individuals rested more in the dry season than in the wet season.Inversely,they<br />
travelled less and engaged in feeding activities<br />
more often during the wet season.<br />
Discussion<br />
The sifaka density was found to be high in the dry forest of<br />
Antrema, with a minimum estimate far above the 173 inds.<br />
km 2 found in the riparian forest of Anjamena (Muller et al.,<br />
2000) or for other sifaka species in dry or wet forests<br />
(O’Connor, 1988; Ganzhorn, 1992). This high density might<br />
be related to some peculiar characteristics of the forest in<br />
terms of food quantity and/or food quality available to this<br />
prosimian species. However,tree basal area was not particularly<br />
high compared with other dry forests in Madagascar<br />
and Mayotte (Hladik, 1980; Simmen et al., 2005). It is not yet<br />
clear also whether high density is related to a putative low<br />
predation pressure.To our knowledge,no sightings or traces<br />
of viverrid carnivores have been reported; large raptors and<br />
boas would be the only predators that could affect the<br />
demography of the Antrema sifaka population (Garbutt,<br />
2007; Sinclair and Legrand, 2008).<br />
As regards their feeding behavior, crowned sifakas fed primarily<br />
on leaves from a few tree, liana and vine species, and<br />
supplemented their diet with a wide range of secondary<br />
items as commonly occurs in other Propithecus species<br />
(Meyers and Wright, 1993; Simmen et al., 2003; Lehman and<br />
Mayor, 2004; Irwin, 2008). Although this species remained<br />
57% Young leaves<br />
Mature leaves<br />
Flowers<br />
Fruits<br />
Other<br />
Fig.2:Food categories in the diet of Propithecus coronatus during the dry season<br />
(a) and the wet season (b).<br />
29,4%<br />
(a) Dry season<br />
0,8% 5,9%<br />
50,0%<br />
(b) Wet season<br />
3,9%<br />
35,8%<br />
5,3%<br />
46,6%<br />
Resting<br />
Locomotion<br />
Foraging<br />
Feeding<br />
6,8%<br />
7,1%<br />
3,6% 4,8%<br />
Social<br />
Other<br />
Fig. 3: Activity budget of Propithecus coronatus during the dry season (a) and the<br />
wet season (b).<br />
folivorous during our study,its diet changed<br />
with seasons. Young leaves were the preferred<br />
food type in the early wet season,<br />
while mature leaves were the dominant one<br />
in the beginning of the dry season. In addition,<br />
P. coronatus ate a higher proportion of<br />
liana and vine parts during the wet season.<br />
Crowned sifakas also followed the typical<br />
activity pattern of other sifaka species<br />
(Norscia et al.,2006;Patel,2006;Charrier et<br />
al., 2007), spending most of their time resting<br />
and devoting a substantial amount of<br />
time to feeding activities and locomotion.<br />
Activity budget nevertheless changed with<br />
seasons. It is generally suggested that the<br />
cool dry season represents a period of food<br />
scarcity for animals,which they compensate<br />
for by reducing their energy expenditure,<br />
travelling less and resting more. In a recent<br />
joint research project, the content of litter<br />
traps regularly distributed along the transects<br />
was collected and weighted every two<br />
weeks throughout one year. It was found<br />
that plant species could be grouped according<br />
to their temporal pattern of leaf loss<br />
(Ranaivoson et al., 2010; see also Razakanirina,<br />
2010). Several trees, lianas and vines<br />
lost their leaves more or less regularly<br />
throughout the dry season while others<br />
were characterized by delayed leaf loss or<br />
on the contrary by precocious leaf fall.One<br />
consequence is that leaves are available<br />
throughout the year, although as different sets of species<br />
varying in quantity,diversity,and presumably,nutritional quality.This<br />
at least could explain why sifakas are able to increase<br />
the diversity of consumed plants (and adopt a more opportunistic<br />
strategy) during the dry season, a period normally<br />
described by the scarcity of food resources.<br />
Future work on seasonal variations in the diet’s nutritional<br />
and chemical content will allow us to examine the role of<br />
qualitative aspects in food choices (Moss,1991;Dearing et al.,<br />
2000) and further examine potential differences between<br />
genders with regard to the importance of energy conservation<br />
for female sifakas (Wright, 1999; Charrier et al., 2007).<br />
Conclusion<br />
Better knowledge of the ecology and the villagers’ social perception<br />
of this flagship species may contribute to conservation<br />
of other diurnal lemurs, by incorporating the villagers’<br />
symbolic perception of their natural environment. Investigating<br />
the interactions between this species and plants of the<br />
coastal dry forest ecosystem will undoubtedly result in<br />
better conservation decisions for Antrema. From an evolutionary<br />
ecology standpoint, the studies we have planned for<br />
the next years at Antrema will also contribute to better<br />
identify the selective pressures that have been driving the<br />
evolution of prosimian typical life-history traits such as<br />
reproductive synchronization or dominance-based feeding<br />
priority of females over males in gregarious species (Wright,<br />
1999; Dewar and Richard, 2007).<br />
Acknowledgements<br />
We thank the Malagasy Institutions that authorized to collect<br />
and export the plant samples,the Ministère de l’Environnement,<br />
des Eaux et forêts et du Tourisme. We also thank<br />
Antrema’s project staff for assistance with the field work as
Page 46 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
well as Master degree students S.Razakanirina,T.Ranaivoson,<br />
V. Randriantoposon and L. Razafindramahatra for helping<br />
with data collection. Special thanks to the specialists of the<br />
Dept of Phanerogamy,J.N.Labat,P.Phillipson and Pete Lowry,<br />
for helping with plant identification. Finally, we thank C.A.<br />
Gauthier, E. Roger, D. Rakotondravony and H. Razafindraibe<br />
for logistic support and collaborative work, and E.G. Leigh<br />
for helpful comments on an earlier draft. This study was<br />
funded by the UMR 7206 - CNRS,and was conducted under<br />
the "Convention cadre de cooperation" between the Université<br />
d’Antananarivo and the Museum National d’Histoire<br />
Naturelle, Paris.<br />
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Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 47<br />
Effet de la dégradation de l’habitat sur la<br />
consommation alimentaire d’Eulemur<br />
rubriventer dans deux sites: Talatakely et<br />
Vatoharanana,du Parc National de Ranomafana<br />
Laingoniaina H.Rakotonirina 1,2*,Germain J.Spiral 1,2,<br />
Jonah H. Ratsimbazafy 1,2, Soanorolalao Ravelonjanahary<br />
1,Raharizelina Ralaiarison 1,2,Stacey Tecot 3,Alex<br />
Hall 4, Tricia Calhoon 4, Gisèle R. Randria 1,2<br />
1Département de Paléontologie et d’Anthropologie Biologique,<br />
Faculté des Sciences, B.P. 906, Université d’Antananarivo,<br />
Madagascar<br />
2Groupe d’Etudes et de Recherche sur les Primates de Madagascar<br />
(GERP)<br />
3Department of Anthropology, University of Stony Brook,<br />
USA<br />
4<strong>Vol</strong>untary field assistant<br />
*Contact de l’auteur principal: laingoniaina2000@yahoo.fr<br />
Mots clés:Eulemur rubriventer,dégradation,habitat,consommation<br />
alimentaire, Ranomafana, Madagascar<br />
Introduction<br />
La grande île est potentiellement riche en matière de biodiversité<br />
et est par conséquent renommée pour sa remarquable<br />
richesse écologique, biologique et génétique (Ganzhorn<br />
et al.,2001).Cette richesse qui est gravement menacée<br />
par la diminution et la destruction immuables des habitats<br />
naturels de nombreuses espèces fait de Madagascar un des<br />
huit "hotspots" les plus considérés de notre planète (Ganzhorn<br />
et al., 2001).<br />
A l’échelle mondiale, la menace la plus grave pour la population<br />
des primates est la destruction et la dégradation de leur<br />
habitat, notamment les forêts tropicales qui hébergent aujourd’hui<br />
environ 90 % des primates non humains du monde<br />
(Mittermeier et al., 2006, 2010). Les lémuriens malgaches ne<br />
font pas exception à cette constatation. La dégradation des<br />
forêts affecte la biologie générale des lémuriens car non<br />
seulement elles leur fournissent des abris et de la nourriture,<br />
mais aussi elles servent de supports à la locomotion et aux<br />
différentes activités de ces animaux (Razafimahazo, 2001).<br />
Selon Randriatahina en 2001, la fragmentation de l’habitat<br />
affecte en premier lieu la distribution et la dispersion de la<br />
nourriture.Certains facteurs influencent le rythme d’activité<br />
et le budget-temps des primates:il s’agit surtout des facteurs<br />
écologiques majeurs tels que la structure de l’habitat,le type<br />
d’alimentation (Zaonarivelo, 1999). Par ailleurs, Dunbar<br />
(1988) affirme que les primates pourraient augmenter leur<br />
déplacement journalier pour trouver de la nourriture ou<br />
inversement en vue d’économiser leur énergie.<br />
Notre présent travail est axé sur la corrélation entre l’habitat<br />
et l’alimentation des lémuriens.Les lémuriformes montrent<br />
un degré de variabilité en ce qui concerne la spécialisation<br />
aux régimes alimentaires.La plupart d’entre-eux (les<br />
Lémuridés,les Mégaladapidés,les Indridés) se spécialisent au<br />
régime végétarien. Cependant, la proportion de feuilles, de<br />
fleurs, et de fruits consommés varie suivant les espèces et<br />
sous-espèces,d’une région à une autre,et de saison en saison<br />
(Richard, 1978). Selon Zaonarivelo (1999), des facteurs écologiques<br />
influencent les comportements des lémuriens et la<br />
perturbation de leur habitat affecte leur organisation sociale<br />
et l’exploitation des ressources alimentaires.<br />
En tenant compte de toutes ces observations, nous avons<br />
effectué une étude concernant l’effet de la dégradation de<br />
l’habitat sur la consommation alimentaire d’Eulemur rubriventer<br />
dans deux sites: Talatakely et Vatoharanana du Parc National<br />
de Ranomafana, dans la province de Fianarantsoa.<br />
Il y a lieu de souligner qu’Eulemur rubriventer, une espèce<br />
hautement frugivore (Overdorff, 1993), dispose d’une haute<br />
importance écologique car elle participe activement à la dispersion<br />
des graines dans la région du Sud-Est de Madagascar,<br />
en particulier dans le Parc National de Ranomafana. A cet<br />
égard, bien que l’animal soit encore classé dans la catégorie<br />
vulnérable selon la liste rouge de l’UICN (Mittermeier et al.,<br />
1994,2006,2010),il a besoin d’ une action de conservation.<br />
C’est la raison pour laquelle le parc national de Ranomafana a<br />
été choisi comme notre station de recherche car par rapport<br />
aux autres régions de l’île, les lémurs à ventre roux y<br />
sont les plus répandus (Mittermeier et al.,2006);et leur habitat<br />
présente un degré variable de dégradation.<br />
Compte tenu de cette variation du degré de dégradation et<br />
de perturbation du milieu de vie d’Eulemur rubriventer dans le<br />
Parc National de Ranomafana, nous pouvons avancer une<br />
hypothèse selon laquelle la consommation alimentaire n’est<br />
pas statistiquement différente entre celle de Vatoharanana<br />
et celle de Talatakely.<br />
Ce projet a été réalisé dans le cadre de la collaboration interdépartementale<br />
entre l’Université d’Antananarivo, l’ICTE/<br />
MICET,le MNP et l’Université de Texas.Ainsi,le présent travail<br />
qui vise en la conservation des lémurs à ventre roux a<br />
comme objectifs d’inventorier les différentes espèces de<br />
plantes consommées par Eulemur rubriventer,de comparer le<br />
régime alimentaire adopté dans chaque site d’étude, de<br />
déterminer les caractéristiques des plantes consommées<br />
dans les deux sites d’études à dégradations différentes.<br />
Site d’études<br />
Le parc National de Ranomafana se trouve dans le Sud Est de<br />
Madagascar.Sa superficie est de 41.600 ha.Ce parc se localise<br />
au Nord-Est de Fianarantsoa, à 70 km à l’Ouest de l’Océan<br />
Indien et à 400 km d’Antananarivo.Il est situé entre 47°18’ à<br />
47°37’ Est de longitude et 21°2’ à 21°25’ Sud de latitude. La<br />
température moyenne annuelle est de l’ordre de 21°C selon<br />
Turk en 1995. Quant à la pluviosité, Overdorff a affirmé en<br />
1996 que la pluie y est saisonnière avec une précipitation<br />
moyenne de 2000 mm. Deux sites ont été choisis pour<br />
effectuer notre travail de recherche. Il s’agit de:<br />
Talatakely:milieu perturbé et plus dégradé.Il est situé à environ<br />
10 mn de marche de la poste de garde et de contrôle<br />
du Parc d’Ambodiamontana. Ce site de 1020 m d’altitude<br />
(Brady et al., 1996), est caractérisé par une visite fréquente<br />
de touristes. Notons également que, à cause des abattages<br />
intensifs des arbres par les bûcherons (Kremen, 1992), la<br />
forêt de Talatakely se trouve fortement dégradée.<br />
Vatoharanana: un milieu moins perturbé et moins dégradé.<br />
Cet endroit est à 1090 m d’altitude et se trouve à 2<br />
heures de marche de Talatakely. Ce champ de forêt était exploité<br />
par les bûcherons il y a 25 ans (Brady et al.,1996).Mais,<br />
la dégradation était moins intense que celle de Talatakely<br />
(Kremen, 1992). La visite des touristes dans cette station<br />
d’études est également moins fréquente. Vatoharanana est<br />
donc un site moins perturbé par rapport à Talatakely.<br />
Les deux stations de recherche sont représentées dans la<br />
figure 1.<br />
Espèces étudiées<br />
Afin de répondre à toutes nos questions, une espèce hautement<br />
frugivore, qui participe activement à la dispersion des<br />
graines (Overdorff,1993),a fait l’objet de notre étude.Il s’agit<br />
du Lémur à ventre roux ou Eulemur rubriventer. Généralement,cette<br />
espèce de taille moyenne vit en petit groupe de 2
Page 48 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
Fig. 1: Localisation des deux sites d’études dans le Parc<br />
National de Ranomafana.<br />
à 5 individus. Elle consomme également des feuilles, des<br />
fleurs, de la terre, des mille-pattes. Selon Overdorff (1996),<br />
l’animal mange beaucoup plus de fruits et moins de feuilles.<br />
Eulemur rubriventer présente un dimorphisme sexuel au<br />
niveau de la morphologie. En effet, le ventre des femelles est<br />
clair;tandis que la poitrine et la partie inférieure du corps du<br />
mâle sont visiblement colorées en marron roux. Le mâle<br />
diffère également de la femelle par la présence de tâche<br />
blanche sur le coin interne des yeux (Dague et Petter,1988).<br />
En ce qui concerne notre étude, nous avons suivi cinq<br />
groupes de lémurs à ventre roux dont trois à Talatakely et<br />
deux à Vatoharanana.<br />
Suivi écologique<br />
Le suivi écologique proprement dit était précédé de la familiarisation<br />
de tous les groupes d’études. A ce propos, durant<br />
cinq mois d’études sur terrain,du mois de décembre 2003 au<br />
mois d’avril 2004, la fréquence d’observation était de cinq<br />
jours par semaine. L’observation s’étale de sept à douze<br />
heures dans la matinée et de treize à quinze heures dans<br />
l’après midi. Au total, l’équipe a suivi cinq groupes pour les<br />
sites de Talatakely et de Vatoharanana, pendant 588 heures<br />
45 minutes et 54 secondes.<br />
La méthode d’enregistrement continu de données a été<br />
adoptée.Elle nous donne des informations plus fiables et plus<br />
pratiques par rapport à une méthode d’enregistrement instantané<br />
(Martin et Bateson, 1986). En outre, nous avons<br />
collecté les données sur l’alimentation en suivant la méthode<br />
de "focal animal sampling" (Altmann, 1974). Durant ces observations,nous<br />
avons enregistré les informations suivantes:<br />
Site d’études: Talatakely ou Vatoharanana;<br />
Groupe d’étude;<br />
Focal animal;<br />
Date de l’observation;<br />
Heure d’observation (Début et fin);<br />
Temps dépensé (Début et fin) à la consommation alimentaire<br />
et aux autres activités;<br />
Nom de chaque espèce consommée (plantes ou autres);<br />
Parties consommées des plantes: fruit, feuille, fleur;<br />
Etat des parties comsommées (fruit immature ou mûr,<br />
jeune feuille, feuille mature, fleur ouverte ou fermée);<br />
Piste la plus proche.<br />
Analyses statistiques<br />
Test de similarité entre deux échantillons<br />
Ce test sert à vérifier la similarité entre le régime alimentaire<br />
adopté par les lémurs à ventre roux de Talatakely et celui de<br />
Vatoharanana.Il se base sur la valeur du coefficient de Jaccard<br />
(Brower et al.,1990).Ce coefficient est donné par la formule<br />
suivante:<br />
CC: Coefficient de Jaccard<br />
S1: Effectif d’espèces végétales, animales et autres dans le<br />
régime de l’espèce de Talatakely (ET);<br />
S2: Effectif d’espèces végétales, animales et autres dans le<br />
régime d’Eulemur rubriventer de Vatoharanana (EV);<br />
C:Effectif d’espèces végétales,animales et autres communes<br />
(ET et EV)<br />
Par souci de conformité, nous avons adopté les échelles<br />
suivantes:<br />
0-40 %: faible similarité entre les deux régimes;<br />
40-60 %: similarité moyenne entre les deux régimes;<br />
60-80 %: grande similarité entre les deux régimes;<br />
80-100 %: forte similarité entre les deux régimes.<br />
Test de Chi-deux:<br />
Cette méthode sert à comparer la durée moyenne journalière<br />
(en minute) consacrée à la consommation alimentaire<br />
des lémurs à ventre roux de Talatakely et de Vatoharanana.A<br />
cet effet, les variables utilisées sont les durées moyennes<br />
journalières dépensées à la consommation des fruits, des<br />
feuilles, des fleurs et autres (sol, eau, champignon, insectes,…);<br />
et ce dans des intervalles de temps bien déterminés;<br />
c’-est-à-dire, entre 7 et 8h, 8 et 9h, 9 et 10h, 10 et 11h, 11 et<br />
12h, 13 et 14h et finalement entre 14 et <strong>15</strong>h. Plus précisément,<br />
elle nous permet de vérifier si la différence entre la<br />
consommation de ces aliments est statistiquement significative<br />
ou non dans les deux milieux.<br />
Résultats<br />
Temps dédié à l’activité alimentaire<br />
Fig. 2 montre l’allure générale de la durée moyenne journalière<br />
en ce qui concerne la consommation alimentaire<br />
générale des lémurs à ventre roux du milieu plus dégradé de<br />
Talatakely et celle du site moins dégradé de Vatoharanana.<br />
Selon cette figure,la prise de nourriture débute entre 7 et 8h.<br />
Concernant le site de Talatakely,elle dessine un pic entre 9 et<br />
10h. Cette activité diminue jusqu’à 12h, puis remonte pour<br />
atteindre le maximum vers 14 à <strong>15</strong>h. Quant à l’Eulemur rubriventer<br />
de Vatoharanana,le pic de l’alimentation se situe entre<br />
8 et 9h. La courbe diminue jusqu’à 12h environ. Ensuite, une<br />
légère remontée de l’activité est constatée jusqu’à <strong>15</strong>h envi-
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 49<br />
ron. L’hypothèse nulle est acceptée car l’analyse statistique<br />
( 2= 9,95 ;avec ddl=6,et p>0,05) indique une différence non<br />
significative concernant la prise de la nourriture entre<br />
chaque intervalle de temps.<br />
20<br />
18<br />
16<br />
14<br />
12<br />
10<br />
8<br />
6<br />
4<br />
2<br />
0<br />
7-8h 8-9h 9-10h 10-11h 11-12h 13-14h 14-<strong>15</strong>h<br />
Intervalle de temps<br />
Talatakely Vatoharanana<br />
Fig. 2: Temps consacré à la consommation alimentaire pour<br />
Eulemur rubriventer dans les deux sites d’étude: Talatakely et<br />
Vatoharanana.<br />
Durée en minute<br />
Similarité entre les régimes alimentaires d’Eulemur rubriventer du<br />
site dégradé de Talatakely et de celui de la station moins dégradée<br />
de Vatoharanana<br />
L’inventaire des espèces consommées par les lémurs à<br />
ventre roux dans les deux sites nous a permis de calculer le<br />
coefficient de Jaccard afin de tester s’il existe ou non une<br />
similarité de régimes dans les deux milieux. Nous avons<br />
recensé 52 espèces de plantes qui sont utilisées comme<br />
source de leur nourriture pour le site de Talatakely. En<br />
revanche, 60 espèces sont inventoriées dans le site de<br />
Vatoharanana.Il est à noter que 35 d’entre elles sont à la fois<br />
consommées à Talatakely et à Vatoharanana. La liste des<br />
espèces végétales utilisées par ces animaux comme source<br />
de leur nourriture (avec la durée de consommation correspondante)<br />
est résumée dans le tableau récapitulatif suivant.<br />
Tab. 1: Liste des espèces végétales consommées (61) par<br />
Eulemur rubriventer, avec la durée de consommation correspondante,<br />
dans les deux sites d’étude.<br />
Nom<br />
malagasy<br />
Ramy<br />
Kalafambakaka<br />
Durée (en mn)<br />
Genre Famille Fr Fe Fl Tala Vato<br />
Canarium<br />
madagascariensis<br />
Burseraceae + +<br />
Oncostemum sp. Myrsinaceae + + + 23,57 242,55<br />
Vakoana Pandanus sp. Pandanaceae + + + 21,40<br />
Nonoka Ficus sp. Moraceae + 24,02 269,75<br />
Vahimberana<br />
Strongylodon c<br />
raveniae<br />
Fabaceae + 2,03 189,72<br />
Voara Ficus sp. Moraceae + 70,08 39,45<br />
Voara rano Ficus botryoides Moraceae + + 23,13 36,<strong>15</strong><br />
Sandramy Protorhus sp.<br />
Anacardiaceae<br />
+ 17,08<br />
Voandavenona<br />
+ 12,70 87,63<br />
Famakilela Ficus sp. Moraceae + + 3,65 7,55<br />
Tavolo malady Cryptocarya<br />
acuminata<br />
Lauraceae + 98,32<br />
Vahitamboro Danais sp. Rubiaceae + + 19,10 12,97<br />
Mahanoro<br />
Streblus<br />
dimepate<br />
Moraceae + 62,58<br />
Tsirika Pandanus sp. Pandanaceae + 38,00 1,67<br />
Rotra Syzygium sp. Myrtaceae + 27,28<br />
Apana Ficus sp. Moraceae + 8,070 135,<strong>15</strong><br />
Sira Neodypsis sp. Arecaceae + 36,30 21,47<br />
Rotra mena Syzygium sp. Myrtaceae + 9,020 17,32<br />
Fandramanana Aphloia<br />
theaeformis<br />
Flacourtiaceae<br />
+ 127,82<br />
Velatra spécial Ruellia sp. Acanthaceae + 68,68 27,05<br />
Durée (en mn)<br />
Nom<br />
malagasy<br />
Genre Famille Fr Fe Fl Tala Vato<br />
Faritraty Memecylon sp.<br />
Melastomataceae<br />
+ 16,83 101,57<br />
Maranitratoraka<br />
Vernonia sp. Asteraceae + 68,40<br />
Vahivoraka<br />
Mendoncia<br />
avani<br />
Mendonciaceae<br />
+ + 169,48<br />
Fatsikahitra Alberta humblotii Rubiaceae + + 9,03 425,63<br />
Albizia Albizia chinensis Leguminosae + 29,67<br />
Rotra fotsy Syzygium sp. Myrtaceae + 30,97<br />
Tongoalahy Bakerella sp.<br />
Loranthaceae<br />
+ + 3,25 25,68<br />
Kalafana<br />
spécial<br />
Oncostemum<br />
botryoides<br />
Myrsinaceae + + + 7,67 130,45<br />
Hafipotsy Grewia sp. Tiliaceae + 35,92<br />
Fanalamangidy +<br />
Vavaporetaka Melanophylla<br />
crenata<br />
Melanophyllaceae<br />
+ 26,05 16,58<br />
Kaboka Voacanga sp. Apocynaceae + 86,73 66,30<br />
Sandramy fot-<br />
Protorhus sp.<br />
syAnacardiaceae<br />
+ 20,32<br />
Apaliala Treculia africana Moraceae + 14,85<br />
Ramandriona Dilobeia thouarsii Proteaceae + 7,<strong>15</strong> <strong>15</strong>,28<br />
Andriambo.<br />
lamena<br />
Menispermaceae<br />
+ 138,73<br />
Bararata Gaertnera sp. Rubiaceae + 26,75 11,62<br />
Nato jabo<br />
Mammea<br />
vatoensis<br />
Clusiaceae + 69,60<br />
Tavilona Vernonia sp. Asteraceae + 4,07<br />
Malanimanta<br />
Apodytes<br />
thouvenotii<br />
Icacinaceae + 3,52<br />
Voantsosoka + 45,08<br />
Lambinanala Nuxia sp. Loganiaceae + 17,88<br />
Rahiaka<br />
Chrysophyllum<br />
boivinianum<br />
Sapotaceae + 6,30 663,50<br />
Amontana Ficus lutea Moraceae + 22,42<br />
Goavy<br />
Psidium<br />
cattleianum<br />
Myrtaceae + 589,02<br />
Fanorafa<br />
Euphorbiaceae<br />
+ 173,10<br />
Kimba spécial Symphonia sp. Clusiaceae + 3,23<br />
Veso<br />
Terminalia tetranoCombretaraceae + 2,02<br />
Vahirano Cissus sp. Vitaceae + <strong>15</strong>,73 104,17<br />
Fohaninasity Psychotria sp. Rubiaceae + 260,93<br />
Rohindambo Smilax anceps Smilacaceae + 5,67<br />
Sary<br />
Potameia<br />
chartacea<br />
Lauraceae + 36,60<br />
Kalamasina<br />
Embelia madagascariensis<br />
Myrsinaceae + 10,43<br />
Ambora<br />
Tambourissa<br />
thouvenotii<br />
Monimiaceae + 10,85<br />
Harongana<br />
Harungana mada-<br />
Clusiaceae<br />
gascariensis<br />
+ 21,45<br />
Nato spécial Sideroxylon sp. Sapotaceae + 1,82<br />
Holatra Champignon +<br />
Amboralahy<br />
Decarydendron he-<br />
Monimiaceae<br />
lenae<br />
+ 10,33<br />
Sandramy<br />
Mena<br />
Protorhus sp.<br />
Anacardiaceae<br />
+ 1,50<br />
Inconnue 1,37 8,82<br />
Champignon 6,60<br />
Tala = site de Talatakely; Vato = site de Vatoharanana; Fr = fruits; Fe = feuilles ;<br />
Fl = fleurs.; *: consommé<br />
Le calcul du coefficient de Jaccard offre une valeur de 0,45;<br />
soit 45 %. Il en découle que le régime alimentaire d’Eulemur<br />
rubriventer des deux sites présente une similarité moyenne.<br />
Aussi, la différence entre le régime d’Eulemur rubriventer des<br />
deux milieux peut être révélée par la constatation de la<br />
durée consacrée à la consommation de chaque catégorie<br />
alimentaire telle que les fruits, les fleurs, les feuilles et bien<br />
d’autres (Cf. Tab. 2 et Fig. 2).<br />
En se basant sur ce tableau récapitulatif (Tab.2) et sur la Fig.2,<br />
nous constatons que dans le milieu dégradé de Talatakely, la<br />
consommation des fruits s’avère très importante par rapport<br />
à celle du milieu moins dégradé de Vatoharanana. De<br />
plus en comparant avec l’espèce du site de Talatakely,celle de
Page 50 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
la station de Vatoharanana comble beaucoup plus sa nourriture<br />
avec des fleurs, des feuilles, et d’autres types d’aliments.<br />
Tab. 2: Comparaison de la consommation journalière de<br />
chaque catégorie alimentaire d’Eulemur rubriventer dans les<br />
deux sites durant la période d’observation (en minute).<br />
Sites Fruit Fleur Feuille Autres<br />
Talatakely 80,80 0,52 4,54 0,12<br />
Vatoharanana 72,58 8,32 11,16 1,63<br />
Durée de la consommation<br />
(en minute)<br />
90<br />
80<br />
70<br />
60<br />
50<br />
40<br />
30<br />
20<br />
10<br />
0<br />
Fruit Fleur Feuille Autres<br />
Catégorie alimentaire<br />
Talatakely Vatoharanana<br />
Fig.3:Allure de la consommation journalière de chaque catégorie<br />
alimentaire d’Eulemur rubriventer dans les deux sites<br />
durant la période d’observation.<br />
Discussion<br />
Eulemur rubriventer est un lémurien hautement frugivore.<br />
Mais la proportion des fruits qu’il consomme varie suivant le<br />
degré de perturbation,de dégradation,de l’altitude du milieu,<br />
ainsi que de la saison. Notons également que la consommation<br />
des différentes catégories alimentaires dépend de la<br />
disponibilité des ressources alimentaires (Rasolofonirina,<br />
2001). Lors de notre étude, nous constatons que dans le<br />
milieu dégradé de Talatakely, la consommation des fruits<br />
s’avère très importante que dans le milieu moins dégradé de<br />
Vatoharanana.De plus,en comparant avec l’espèce du site de<br />
Talatakely, celle de la station de Vatoharanana comble beaucoup<br />
plus sa nourriture avec des fleurs, des feuilles, et d’autres<br />
types d’aliments. Ceci est dû certainement à la saison<br />
cyclonique durant laquelle le vent est violent.Comme le site<br />
de Vatoharanana est beaucoup plus élevé par rapport à Talatakely,il<br />
s’avère logique qu’il est beaucoup plus affecté par ce<br />
vent violent.En effet,les fruits deviennent rares car beaucoup<br />
d’entre eux tombent par terre. Selon Zaonarivelo (1999),<br />
pendant la période de crise, Varecia variegata variegata augmente<br />
le taux de folivorie même si les feuilles sont des aliments<br />
de compensation.Cette stratégie adoptée par l’animal<br />
lors de la période de crise est également observée chez<br />
d’autres espèces de lémuriens de la forêt dense humide<br />
(Ganzhorn, 1988) entre-autres l’Eulemur rubriventer,leEulemur<br />
mongoz qui se nourrissent de fleurs en plus des fruits et<br />
des feuilles durant la période de floraison (Sussman, 1975).<br />
Par ailleurs,Garbutt (1999) argumente que les fruits constituent<br />
la majeure partie de l’alimentation d’Eulemur rubriventer.<br />
Mais quand ils ne sont pas disponibles,les feuilles et les fleurs<br />
sont aussi consommées (Rasolofonirina, 2001). Notons à la<br />
même occasion que les primates adoptent différentes stratégies<br />
pour affronter le manque de nourriture de base (le<br />
fruit pour notre cas) soit en augmentant le temps de la<br />
recherche de nourriture en se déplaçant beaucoup, soit en<br />
acceptant de consommer des aliments de basse qualité<br />
(Zaonarivelo, 1999).<br />
Par rapport au milieu dégradé,l’abondance des fruits dans le<br />
milieu moins dégradé est évidente.Mais en tenant compte de<br />
l’étude phénologique mensuelle des plantes recensées, ce<br />
n’est pas toujours le cas; car plusieurs facteurs pourraient<br />
agir sur ce milieu.Ainsi le climat,en particulier la pluviosité et<br />
le cyclone, influence la qualité et la quantité de la nourriture<br />
disponible (Dajoz, 1985). En effet, durant la période cyclonique,<br />
l’altitude de la station joue un rôle important sur les<br />
arbres à semences. Autrement dit, plus l’altitude d’un milieu<br />
est élevée,plus le vent agit directement sur les arbres et plus<br />
les fruits tombent. Par conséquent, les arbres portent moins<br />
de fruits. L’animal est obligé de se rabattre sur d’autres<br />
catégories alimentaires comme les feuilles, les fleurs, les<br />
champignons, les insectes pour pouvoir combler l’insuffisance<br />
de nourriture de base.Ce cas se rencontre dans le site<br />
de Vatoharanana qui est considéré comme milieu moins<br />
dégradé et qui est situé à une altitude plus élevée (1090 m)<br />
par rapport au site dégradé de Talatakely qui se trouve à une<br />
altitude de 1020 m (Brady et al, 1996). Concernant le site de<br />
Talatakely, la dispersion des fruits dans l’espace est insuffisante<br />
à cause de la dégradation de ce milieu.Ainsi,au lieu de<br />
combler sa nourriture par d’autres types d’aliments, il est<br />
contraint à se déplacer loin afin de consommer de la nourriture<br />
de haute qualité qui lui apporte beaucoup plus d’énergie<br />
comme le glucide, le lipide et les protéines (Zaonarivelo,<br />
1999).<br />
Finalement, en se basant sur les données phénologiques,<br />
nous avons constaté que la consommation de chaque catégorie<br />
alimentaire varie suivant leur disponibilité mensuelle<br />
dans chaque mileu. Au mois de décembre, par exemple,<br />
l’Eulemur rubriventer de Vatoharanana consomme beaucoup<br />
plus de fleurs que de fruits par rapport à celui de Talatakely<br />
car pendant ce mois, les fleurs y sont disponibles. Remarquons<br />
également que l’espèce de la station de Talatakely ne<br />
consomme que des fruits durant la fructification de l’espèce<br />
introduite de goyave (Psidium cattleianum). Ce type de fruit<br />
est très apprécié par l’animal.<br />
Le régime alimentaire d’Eulemur rubriventer est moyennement<br />
similaire dans les deux milieux étudiés.<br />
Cependant, quelques espèces de plantes consommées par<br />
l’animal sont propres à chaque site. L’espèce introduite de<br />
goyave Psidium cattleianum se rencontre uniquement à Talatakely.<br />
Notons que l’introduction de cette espèce marque la<br />
dégradation de cette station. Par contre, l’animal de Vatoharanana<br />
a l’opportunité de consommer, par exemple, l’espèce<br />
Mammae vatoensis qui est endémique à ce site. Cette<br />
similarité moyenne entre les deux régimes implique que la<br />
dégradation du site de Talatakely n’est pas encore poussée à<br />
l’extrême (Randriamahaleo,2005).En effet,il est classé parmi<br />
les sites moyennement dégradés. Selon Tam-Alkis (1997), le<br />
site de Talatakely est séparé de Vatoharanana par une barrière<br />
biogéographique (rivière Fompohonona). Il paraît que<br />
la dispersion des graines de part et d’autre de cette rivière<br />
est empêchée. Voilà pourquoi certaines espèces de plantes,<br />
utilisées comme source alimentaire, caractérisent uniquement<br />
l’un de ces sites. Par conséquent son régime varie<br />
suivant le site.<br />
Conclusion<br />
Cette étude nous a permis de fournir plus d’informations sur<br />
la relation entre la dégradation de l’habitat d’Eulemur rubriventer<br />
et sa consommation alimentaire.L’investigation révèle<br />
une similarité moyenne entre le régime alimentaire adopté<br />
par l’animal de Talatakely et celui de Vatoharanana. L’espèce<br />
semble être principalement frugivore quelque soit le site.<br />
Cependant, la proportion de consommation varie suivant le<br />
milieu. A Talatakely, par exemple, ce lémur à ventre roux se
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 51<br />
nourrit beaucoup plus de fruits par rapport à celui de Vatoharanana<br />
où il comble sa nourriture avec des fleurs, des<br />
feuilles, et bien d’autres catégories alimentaires. Ceci est dû<br />
sans doute au passage des deux cyclones (Elita et Gafilo)<br />
dans la région durant la période d’étude. En fait, Vatoharanana<br />
se trouve à une altitude très élevée par rapport à Talatakely.En<br />
effet,les vents violents agissent directement sur les<br />
arbres fruitiers, conduisant ainsi l’insuffisance des fruits.<br />
Aussi, la consommation de Psidium cattleianum (Myrtaceae),<br />
qui est une espèce introduite propre à Talatakely semble être<br />
très importante dans ce milieu. Par contre, Chrysophillum<br />
boivinianum s’avère être la plus appréciée par l’animal de<br />
Vatoharanana. Toutes ces constatations nous conduisent à<br />
dire que la consommation alimentaire d’Eulemur rubriventer<br />
paraît être conditionnée par la disponibilité de la nourriture<br />
dans chacun des sites visités et par l’état de l’habitat.<br />
Remerciements<br />
Nous tenons à remercier le Centre Valbio, le MICET représentés<br />
respectivement par le Professeur Patricia Wright et le<br />
Docteur Benjamin Andriamihaja pour leur soutien et leur<br />
collaboration durant la longue haleine de travail dans le Parc<br />
National de Ranomafana. Nos vifs remerciements s’adressent<br />
également à tous les guides de recherche pour leur<br />
assistance (Telo Albert, Victor, Koto, Nirina) et à tout ce qui<br />
contribue, de près ou de loin à la réalisation de ce projet.<br />
Références bibliographiques<br />
Altmann, J. 1974. Observational study of behavior: Sampling<br />
methods. Behavior 49: 227-267.<br />
Brady, L.H.; Jenkens, R.; Kauffmann, J.; Rabearivony, J.; Raveloson,<br />
G.; Rowcliffe, M. 1996. Madagascar Expedition 93.<br />
Final Report. Univ. East Anglia. Norwich, U.K.<br />
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de DEA d’Anthropologie Biologique. DPAB, Faculté des<br />
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de Paléontologie et d’Anthropologie Biologique, Faculté<br />
des Sciences, Université d’Antananarivo.<br />
Observations of terrestrial latrine behaviour<br />
by the southern gentle lemur Hapalemur<br />
meridionalis in the Mandena littoral<br />
forest, southeast Madagascar<br />
Timothy M. Eppley* and Giuseppe Donati<br />
Nocturnal Primate Research Group,Department of Anthropology<br />
and Geography, Oxford Brookes University, Gipsy<br />
Lane, OX3 0BP, Oxford, UK<br />
*Corresponding author: eppleyti@gmail.com<br />
Key words: southern gentle lemur, Hapalemur meridionalis,<br />
defecation, latrines, Mandena<br />
Latrine behaviour is defined as the non-random selection of<br />
a specific defecation site,and although it is rarely described in<br />
primates it is well known among other mammalian species as<br />
a form of olfactory communication (Irwin et al.,2004).Olfactory<br />
compounds, which convey chemical signals, are transmitted<br />
via scent-producing skin gland secretions, saliva, and/<br />
or waste products (Epple, 1986). Olfaction signals may be<br />
advantageous as they are not limited spatially and temporally,<br />
allowing individuals of a predominately visual communication
Page 52 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
to receive signals when distant from the source (Schilling,<br />
1979;Irwin et al.,2004).Olfactory signals may transmit information<br />
pertaining to environmental familiarisation (Schilling,<br />
1979), reproductive behaviour and condition (Epple, 1986),<br />
territorial demarcation (Mertl-Milhollen,1979),and inter- or<br />
intra-group spacing (Schilling, 1979; Epple, 1986).<br />
Previous literature has discussed latrines by varying in location<br />
(arboreal, terrestrial, or subterranean), and being further<br />
analysed by volume of faeces and spatial distribution<br />
(Schilling, 1980; Boonstra et al., 1996; Irwin et al., 2004; Pouvelle<br />
et al., 2009). This behaviour appears to be well studied<br />
within other mammals but for primates, especially strepsirhines,<br />
latrine use is only sparsely mentioned within much<br />
broader scope research (Irwin et al.,2004).In this report we<br />
present our observations of this peculiar behaviour exhibited<br />
by the southern gentle lemur, Hapalemur meridionalis,a<br />
threatened primate that occurs in southeast Madagascar.We<br />
will use our observations to review the hypotheses offered<br />
thus far to explain latrine use,i.e.advertisement of sexual cycle,predation<br />
avoidance,intra- group and inter-group spacing<br />
in the context of the fragmented littoral forest.<br />
Methods<br />
This research was conducted from May to July 2008, on the<br />
southern gentle lemur within the Mandena littoral forest<br />
(24°95’S 46°99’E), a coastal forest in southeast Madagascar.<br />
The littoral forest is among the most endangered ecosystems<br />
in Madagascar (Bollen and Donati, 2006) and Mandena<br />
is a protected conservation zone encompassing 230 hectares<br />
of fragmented and partially degraded littoral forest<br />
interspersed with marsh and swamp. Three groups of H.<br />
meridionalis (mean = 5.7 ind/group) were habituated and<br />
followed daily from dawn to dusk with 62 hours of observation<br />
recorded (Eppley and Donati, in press). These lemurs<br />
are of particular interest, as they do not subsist exclusively<br />
on bamboo like the majority of their congeners. Rather, the<br />
southern gentle lemur exhibits a dietary predilection for<br />
terrestrial (turf) grasses while displaying a unique grazing<br />
behaviour (Eppley and Donati, in press). Although the main<br />
research being conducted focused on the feeding ecology of<br />
these animals, opportunistic observations of latrine behaviour<br />
were collected ad libitum. In addition to habitat characteristics<br />
and GPS waypoints taken at the feeding and resting<br />
sites of three groups, latrine locations were also recorded.<br />
Spatial analyses were carried out with ArcMap version 9.3,<br />
with the outermost feeding and resting site waypoints for<br />
each group being used to create the minimum polygon for<br />
their respective ranging areas.<br />
Results<br />
On three separate occasions an entire group of foraging H.<br />
meridionalis were witnessed descending to the ground and<br />
defecating in succession either near or under a high-rooted<br />
tree of different species.Latrine bouts were recorded ad libitum<br />
on a single occasion for Group B (four individuals) and<br />
twice for Group C (seven individuals). Observations took<br />
place within the forest fragment, and were never witnessed<br />
in open canopy areas. The three occurrences of this terrestrial<br />
latrine behaviour were observed shortly after individuals<br />
had awoken from a midday resting bout (between 11:00<br />
and 14:00) at distances approx. 20 m from the resting site.<br />
The three locations where the latrine behaviour was exhibited<br />
are shown in Figure 1.Little overlap between the ranging<br />
areas of the three groups was observed with 0.92 ha (7.74 %)<br />
calculated to exist between groups A and B,0.18 ha (1.68 %)<br />
between groups A and C, and 0.06 ha (0.73 %) between<br />
groups B and C. After an individual had defecated, they<br />
ascended three to four meters and continued feeding.Subsequently,<br />
the group followed an order of sequential defecation,<br />
where by one conspecific would defecate and the next<br />
would rapidly follow.The individuals remaining at an elevated<br />
height always appeared vigilant, scanning the surrounding<br />
area and sometimes continuing to forage, while the conspecific<br />
was on the ground. Upon inspection of two of the<br />
defecation sites, accumulations of hardened faecal matter<br />
were identified. Dissimilarly, the third site consisted of only<br />
fresh faeces with no sign of old faecal matter.<br />
Fig. 1: Ranging areas of the three observed H. meridionalis<br />
groups within the northeast corner of the Mandena littoral<br />
forest. The three latrine sites were observed within the territorial<br />
boundary buffer zones of Groups A & C and B & C.<br />
Discussion<br />
Among primates,latrine behaviour has been recorded (Table<br />
1) in Alouatta seniculus (Gilbert, 1997; Feeley, 2005; Neves et<br />
al., 2009; Pouvelle et al., 2009), Ateles geoffroyi (Notman et al.,<br />
2009), as well as the strepsirhines: Cheirogaleus major, C.<br />
medius, Lepilemur leucopus (Charles-Dominique and Hladik,<br />
1971; Russell, 1977), L. microdon, L. ruficaudatus, Hapalemur<br />
aureus, H. griseus, Prolemur simus, and Lemur catta (Irwin et al.,<br />
2004). With the exception of Neotropical primates, however,minimal<br />
research has been conducted to understand its<br />
function within primates and latrine utilisation is often a matter<br />
of debate. In Ranomafana National Park, for example, H.<br />
griseus have been observed to occasionally use the same<br />
resting/sleeping sites and since they often defecate very soon<br />
after they wake up (Tan,pers.comm.),an accumulation of faecal<br />
material under these trees may appear inadvertently<br />
latrine-like (Notman et al., 2009; Pouvelle et al., 2009). Thus,<br />
latrine use by H.griseus may be anecdotal within this population.<br />
Irwin et al. (2004) presented four non-mutually exclusive<br />
hypotheses for the possible adaptive function of primate<br />
latrines:1) advertisement of sexual cycle,2) predation avoidance,<br />
3) intra-group spacing, 4) and inter-group spacing.<br />
The conveyance of scent-marks is well known to advertise<br />
sexual activity and receptivity (Asa, 2008). If this holds true<br />
for H. meridionalis, latrines should be used more frequently<br />
during the breeding seasons. It has been reported that H.<br />
griseus mate between June and July, experiencing a gestation<br />
length of approximately 137 days (Tan, 2006). Though our<br />
research potentially overlapped with the mating season, we<br />
made no observations of mating during the study period.
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 53<br />
Thus, we had no possibilities to compare the mating season<br />
with pre- or post-mating periods, as the hypothesis testing<br />
would require. However, since two of the latrines at Mandena<br />
appeared to be utilized long-term, similar to the findings<br />
of Irwin et al.(2004),we conclude that the advertisement<br />
of sexual receptivity is unlikely the sole function of lemur<br />
latrines.<br />
Tab. 1: Observed primates that have exhibited terrestrial<br />
latrine behaviour. Adapted from Irwin et al. (2004).<br />
Species Localities References<br />
Hapalemur<br />
meridionalis<br />
Mandena Conservation<br />
Zone<br />
This study<br />
Hapalemur<br />
griseus<br />
Analamazaotra<br />
Special Reserve<br />
Irwin et al. (2004)<br />
Prolemur<br />
simus<br />
Ranomafana<br />
National Park<br />
P. Wright (in Irwin et al., 2004)<br />
Lemur catta Isalo National Park<br />
J. Jernvall and P. Wright<br />
(in Irwin et al., 2004)<br />
Lepilemur<br />
leucopus<br />
Beza Mahafaly<br />
Special Reserve<br />
L. Nash (in Irwin et al., 2004)<br />
Lepilemur Manombo Special J. Ratsimbazafy (in Irwin et al.,<br />
microdon Reserve<br />
2004)<br />
Lepilemur sp.<br />
(?microdon)<br />
Kalambatritra<br />
Special Reserve<br />
Irwin et al. (2004)<br />
Lepilemur<br />
ruficaudatus<br />
Kirindy Forest<br />
J.U. Ganzhorn (in Irwin et al.,<br />
2004)<br />
Ateles<br />
geoffroyi<br />
Runaway Creek Nature<br />
Preserve, Belize<br />
Notman et al. (2009)<br />
Alouatta<br />
seniculus<br />
Nouragues Reserve,<br />
French Guiana<br />
Pouvelle et al. (2009)<br />
Gentle lemurs have a particularly effective predator avoidance<br />
strategy including camouflage from cryptic pelage, rapid<br />
flight behaviour,and potential cathemeral activity pattern<br />
(Mutschler et al., 1999; Curtis et al., 2006; Tan, 2006). Several<br />
potential predators of Hapalemur exist in the littoral forest.<br />
There have been documented cases of fossa Cryptoprocta<br />
ferox preying on H. griseus (Goodman and Pidgeon, 1999;<br />
Sterling and McFadden,2000).The Madagascar tree boa Sanzinia<br />
madagascariensis (= Boa manditra) also prey on Hapalemur<br />
spp.(Goodman et al.,1993;Rakotandravany et al.,1998),<br />
and several aerial predators (Madagascar harrier hawk Polyboroides<br />
radiatus, Frances’s sparrowhawk Accipiter francesii,<br />
Henst’s goshawk Accipiter henstii,common barn owl Tyto alba,<br />
and the Madagascar long-eared owl Asio madagascariensis)<br />
represent a threat to medium-sized lemurs (Goodman et al.,<br />
1993; Wright, 1997; Karpanty and Goodman 1999). In fact,<br />
the concealment of Hapalemur faeces under large high-rooted<br />
trees may theoretically act as a safeguard against predation<br />
by impairing the ability of a predator to detect the prey<br />
population (Boonstra et al.,1996;Irwin et al.,2004).Although<br />
these observations are in accord with the anti-predator idea,<br />
single faecal deposits were also detected at indiscriminate<br />
locations. Thus, more data are necessary to test the hypothesis<br />
of latrine behaviour as an anti-predator strategy.<br />
Intra-group spacing has also been suggested to advertise<br />
proximal resource use and assist in inter-individual spacing<br />
(Kruuk,1992).In accord with Irwin et al.(2004),however,it is<br />
unlikely that Hapalemur latrine behaviour is used for intragroup<br />
spacing, as they live in cohesive family units.<br />
The territorial demarcation hypothesis suggests that scentmarks<br />
are placed around home range boundaries to act as a<br />
delineation of the territory, i.e. inter-group spacing (Mertl-<br />
Millhollen, 1979; Lewis, 2005). In fact, it is evident from our<br />
observations that H.meridionalis chose defecation sites in the<br />
narrow areas of overlap with neighbouring conspecifics<br />
groups (Fig. 1). If this adaptive function holds true, latrine<br />
behaviour might be even more common in areas of dense<br />
population (Irwin et al.,2004),such as the forest fragments of<br />
Mandena (Eppley and Donati, in press).<br />
Although the exhibition of preferred, non-random defecation<br />
sites is most likely multifactorial,latrines in Mandena appear<br />
to best fulfil the function of inter-group spacing. Therefore,<br />
latrines may be a low-energy behavioural response to<br />
the ecological challenge of defending resources with minimal<br />
rates of agonism (Irwin et al.,2004).In the future,more quantitative<br />
studies should focus on seasonal and spatial exhibition<br />
of latrine use to verify whether this behaviour is intrinsically<br />
linked to territorial delineation and resource defence in<br />
lemurs.<br />
Acknowledgements<br />
We would like to thank the Commission Tripartite of the<br />
Malagasy government, Ministère de l’Environnement, des<br />
Eaux et forêts of the Malagasy government,the University of<br />
Antananarivo, and CAFF/CORE for permission to conduct<br />
research, as well as the Malagasy Institute for the Conservation<br />
of Tropical Environments (MICET) for all of their logistical<br />
assistance. Financial support was provided partly by the<br />
Chester Zoo (NEZS) and QMM.We would also like to thank<br />
the QMM Environmental Team, most especially Manon Vincelette,<br />
Jean-Baptiste Ramanamanjato, Johny Rabenantoandry,<br />
Faly Randriatafika, and Christophe Rambolamanana for<br />
all of their advice and logistical help. We are grateful to Jörg<br />
Ganzhorn for all of his continuous support and scientific advice.Thank<br />
you to the entire staff of the Oxford Brookes Primate<br />
Conservation MSc program, especially Simon Bearder,<br />
Anna Nekaris, and Vincent Nijman. We greatly appreciate<br />
the GIS assistance of Maureen Mullen. My sincere gratitude<br />
goes to my field guide Robertin "Tintin" Ravelomanantsoa<br />
and research assistant Abi Coleman for their companionship<br />
and tireless help in the marecage.<br />
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Lemurs: Ecology and Adaptation. Springer, New York.<br />
Wright, P.C.1997.Impact of predation risk on the behaviour<br />
of Propithecus diadema edwardsi in the rain forest of Madagascar.<br />
Behaviour 135: 483-512.<br />
Wright, P.C. 1999. Lemur traits and Madagascar ecology:<br />
coping with an island environment. Yearbook of Physical<br />
Anthropology 42: 31-72.<br />
Conservation des lémuriens via la protection<br />
de leurs habitats et le développement<br />
communautaire dans les corridors<br />
de Betaolana et Tsaratanana-Betaolana,<br />
région de SAVA<br />
Lala Razafy Fara 1*, Iarilanto Andriamarosolo 2<br />
1WWF Madagascar & Western Indian Ocean PO, BP 738<br />
Antananarivo 101, Madagascar<br />
2WWF Andapa, BP 28, Andapa 205, Madagascar<br />
*Corresponding author: frazafy@wwf.mg<br />
Contextes<br />
Ecologique:<br />
Madagascar est connu pour sa haute valeur en biodiversité.<br />
Sa flore et sa faune ont une valeur d’endémicité très élevée<br />
(au dessus de 80 %) due entre autres à son insularité. Madagascar<br />
est donc plus riche en espèces endémiques comparées<br />
à d’autres continents du monde.Le microclimat de l’île a<br />
permis que d’une région à une autre, la flore et la faune constituent<br />
une richesse spectaculaire à part.<br />
Pour la partie Nord de Madagascar, la flore luxuriante, avec<br />
des espèces endémiques du genre Dalbergia, est encore à<br />
découvrir.Le WWF a travaillé dans la région de SAVA pour la<br />
mise en place de deux Aires Protégées (AP),Parc National de<br />
Marojejy et Réserve Spéciale d’Anjanaharibe Sud, actuellement<br />
sous gestion du Madagascar National Parks. Pour le<br />
WWF,la conservation du flux génétique implique le maintien<br />
et la restauration de la connectivité écologique.Ce maintien<br />
peut être le plus important paramètre le long des pentes<br />
altitudinales car cette connectivité est actuellement très<br />
rare au sein de l’écorégion de l’Est.De plus,il peut constituer<br />
un refuge pour la biodiversité vu les changements du climat<br />
(Erdmann et al., 2005).<br />
Eu égard aux efforts déjà investis dans la protection des deux<br />
AP (Parc de Marojejy et de la Réserve Spéciale d’Anjanaharibe<br />
Sud), le WWF a continué ses efforts de conservations<br />
dans les deux corridors forestiers (Betaolana et Tsaratanana-Betaolana).<br />
Dans ces localités, les espèces endémiques<br />
sont aussi très remarquables comme le palmier Marojejya<br />
insignis et la fougère Asplenium marojyense. La forêt dense et<br />
humide de cette partie de l’île abrite une multitude de faune<br />
à découvrir et à protéger.<br />
La forêt occupe encore 35,60 % de la région de SAVA (données<br />
images satellites de 2000),ce taux est élevé par rapport<br />
à d’autres régions de Madagascar.La richesse en biodiversité<br />
de cette zone est démontrée par de nombreuses études,sur<br />
la flore et sur la faune, conduites dans cette zone.<br />
Pour ces deux corridors Betaolana et Tsaratanana-Betaolana,<br />
les inventaires effectués par Rajaonson et Rakotonirina<br />
(2007) pour le WWF ont montré l’existence de 10 espèces<br />
de lémuriens pour le corridor Betaolana, et de sept espèces<br />
pour le corridor Tsaratanana Betaolana, 84 espèces d’oiseaux<br />
pour l’ensemble des deux corridors. En comparaison<br />
avec les inventaires des lémuriens effectués par Goodman et<br />
al. (2003), la région d’Andapa, incluant les AP de Marojejy et<br />
d’Anjanaharibe-Sud et le corridor forestier de Betaolana,<br />
possèdent une richesse spécifique en communautés de lémuriens<br />
et tient une place importante en biodiversité à Madagascar.La<br />
synthèse des études déjà effectuées dans la zone a<br />
recensé 12 espèces de lémuriens dont: Microcebus rufus,<br />
Cheirogaleus major, Allocebus trichotis, Phaner furcifer, Avahi<br />
laniger, Lepilemur mustelinus, Daubentonia madagascariensis,<br />
Hapalemur griseus griseus, Eulemur rubriventer, Eulemur fulvus
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 55<br />
albifrons, Propithecus candidus, Indri indri. Parmi ces espèces<br />
citées, celles non rencontrées dans les deux corridors sont:<br />
Indri indri et Phaner furcifer (pour les deux corridors Betaolana<br />
et Tsaratanana-Betaolana) et Allocebus trichotis pour le<br />
corridor Tsaratanana Betaolana).<br />
Parmi les espèces de lémuriens, le Simpona (Propithecus candidus)<br />
est une des 25 espèces de primates les plus menacées<br />
dans le monde (Mittermeier et al., 2005), d’où l’intérêt de se<br />
focaliser sur sa conservation. Certains auteurs (Petter et al.,<br />
1979; Tattersall, 1982) pensent que l’habitat du Simpona<br />
pourrait éventuellement s’étendre vers le Sud jusqu’aux<br />
forêts de la péninsule de Masoala. Il est par contre prouvé<br />
que le versant Ouest de Anjanaharibe-Sud, actuellement<br />
sans statut de protection, héberge une population importante<br />
de Simpona. La population de Simpona de la région<br />
Andapa et donc mondiale est estimée à une valeur comprise<br />
entre 100-1000 individus (Mittermeier et al., 1994). Des<br />
inventaires effectués par le WWF vers la fin 2006 ont relevé<br />
que le Simpona se rencontre aussi vers le Nord du Corridor<br />
de Betaolana en allant vers le Nord Ouest du côté du massif<br />
de Tsaratanana.<br />
Social et économique:<br />
Les deux corridors forestiers, Betaolana et Tsaratanana<br />
Betaolana, couvrent approximativement une superficie respective<br />
de 16 500 ha et de 130 900 ha et sont très riches en<br />
forêt.En se référant aux délimitations sur la carte 1,les deux<br />
corridors renferment encore jusqu’à 90 % de forêts. Situées<br />
sur des sols ferralitiques, les forêts des deux corridors sont<br />
localisées entre les altitudes 850 et 1600 m pour Betaolana<br />
et 800 à 2280 m d’altitude pour Tsaratanana-Betaolana. Les<br />
deux corridors jouissent encore du climat humide de la<br />
région de SAVA.<br />
Les deux corridors appartiennent aux districts d’Andapa<br />
(pour le Corridor de Betaolana) et de Bealanana (pour le<br />
corridor de Tsaratanana-Betaolana). Les populations des<br />
communes locales vivant à la périphérie de ces corridors<br />
sont estimées, en 2009, à 72 521 pour le corridor Betaolana<br />
et à 41 333 pour le corridor Tsaratanana-Betaolana (à partir<br />
des données d’INSTAT 1993).<br />
Dans la région de SAVA, les forêts dans les deux corridors<br />
Betaolana et Tsaratanana Betaolana font partie des domaines<br />
forestiers de l’Etat. Ils n’ont de ce fait pas de statuts<br />
particuliers (Garreau et Manantsara, 2003). Toutefois, le<br />
WWF, connaissant l’importance de la flore et la faune dans<br />
cette zone, a mis en œuvre de 2005 à 2008 deux projets<br />
spécifiques pour la conservation des lémuriens et de leurs<br />
habitats dans la région de SAVA. Dans le cadre des projets<br />
mis en œuvre par le WWF qui se sont succédés dans la région<br />
de SAVA, celui-ci a travaillé avec les populations locales<br />
pour mettre ces deux corridors sous statuts de Nouvelles<br />
Aires Protégées. Pour le Corridor Betaolana, huit communautés<br />
de bases ayant reçu des contrats de transferts de gestion<br />
des lots de forêts ont déjà déposé leur manifestation<br />
d’intérêt dans ce sens en 2008.Les différentes étapes et procédures<br />
à respecter suivent actuellement normalement leur<br />
cours.<br />
Cet article, eu égard à tous les acquis dans le cadre des<br />
projets du WWF, est une capitalisation des expériences de<br />
WWF Madagascar pour les projets de conservation des<br />
lémuriens et de leurs habitats dans le Nord de Madagascar.<br />
L’un des deux projets est connu sous son appellation courte,<br />
projet Simpona. Son but est en effet de mettre en exergue<br />
cette espèce phare bien qu’elle ne soit pas la seule à être<br />
protégée dans le cadre des projets du WWF.<br />
Approche de conservation<br />
Les sites des deux projets<br />
La région de SAVA se trouve dans l’ex Province d’Antsiranana<br />
au Nord de Madagascar. Les deux projets du WWF<br />
dans cette localité ont été conduits dans deux corridors<br />
forestiers entre les Aires Protégées Tsaratanana (Réserve<br />
Naturelle Intégrale), Marojejy (Parc National) et Anjanaharibe<br />
Sud (Réserve Spéciale).La carte suivante montre la situation<br />
générale de ces localités.<br />
Les objectifs de conservation<br />
A partir de la connaissance des richesses de cette localité,<br />
des pressions et menaces sur les espèces cibles,la finalité des<br />
deux projets était de freiner l’utilisation irrationnelle de la<br />
forêt tout en construisant un environnement où les populations<br />
locales pourront vivre en harmonie avec la nature.<br />
Découlant de cette finalité,les objectifs des deux projets ont<br />
été de: 1) Mieux connaître le niveau de menace sur les<br />
lémuriens et sur leurs habitats; 2) Elaborer des scénario de<br />
zonages et de gestion; 3) Conscientiser et éduquer les<br />
communautés locales concernant les menaces et la protection<br />
des lémuriens; 4) Initier de nouveaux transferts de<br />
gestion des forêts auprès des communautés locales de base<br />
(sur la partie Ouest du Corridor de Betaolana); 5) Protéger<br />
et/ou restaurer les habitats des lémuriens; 6) Renforcer la<br />
protection des lémuriens par l’extension et la création de<br />
nouvelles aires protégées; 7) Vulgariser et promouvoir les<br />
produits de substitution des principaux produits forestiers;<br />
8) Procéder au suivi d’évaluation des méthodes et procédures<br />
de gestion internes des associations des forêts; 9)<br />
Restaurer les terrains défrichés de la périphérie du corridor.<br />
En bref,les deux projets essaient de renforcer les conditions<br />
requises pour la conservation à long terme de la biodiversité<br />
via la conservation des fonctions écologiques des deux corridors<br />
(Betaolana et Tsaratanana-Betaolana) de manière participative.<br />
Diagnostic participatif: Mieux connaitre pour mieux<br />
protéger<br />
Cartographie<br />
A partir des connaissances issues de la littérature, une cartographie<br />
simplifiée basée sur des cartes topographiques<br />
(échelles 1/100 000) et des interprétations des images satellites<br />
2000 a été élaborée. L’objectif est de partir des limites<br />
des forêts à partir de ces images pour délimiter des zones où<br />
les inventaires forestiers vont être réalisés.De même,de par<br />
cette technique,des zonages forestiers et des cartes d’occupation<br />
des sols sont élaborés.Le zonage forestier se base sur<br />
les états de dégradations des forêts. Les unités semblables<br />
ont été groupées dans une même catégorie définie pour un<br />
objectif spécifique (conservation, enrichissement / restauration,<br />
droit d’usage).<br />
Inventaire des lémuriens par des primatologues<br />
Pour mieux intégrer les populations locales dans l’importance<br />
de la conservation des lémuriens,elles ont été invitées<br />
à participer aux inventaires des lémuriens.Leur connaissance<br />
de base, combinée aux connaissances scientifiques de deux<br />
primatologues recrutés ont été pour renforcer les acquis<br />
dans le cadre de la littérature.Cette approche a aussi permis<br />
d’élaborer des documents de base pour le suivi des lémuriens<br />
et de choisir des espèces pour la restauration<br />
forestière.
Page 56 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
Fig. 1: Carte de localisation générale des sites.<br />
Etudes des pressions et menaces sur les lémuriens et leurs<br />
habitats<br />
Les études sur les pressions et menaces ont été conduites<br />
selon des enquêtes informelles et formelles et selon des observations<br />
directes.Les enquêtes informelles servent à la fois<br />
à gagner la confiance des populations locales et à recueillir le<br />
maximum d’informations. Les enquêtes formelles sont utilisées<br />
par les consultants primatologues pour une meilleure<br />
représentativité des réponses. Les membres de l’équipe de<br />
WWF,à mesure de l’avancement de la conduite des activités<br />
dans le cadre des deux projets, ne cessent de faire des<br />
recoupements entre toutes les réponses obtenues.<br />
La participation des membres des communautés de base est<br />
indispensable dans les deux types d’enquêtes pour qu’ils se<br />
sentent responsables et pour confronter les réponses au fur<br />
et à mesure que des biais sont observés. Les méthodes<br />
classiques de recherches participatives utilisées sont du<br />
genre MARP: Méthode Accélérée de Recherche Participative<br />
et PALM: Participatory Learning Methods ou Méthode<br />
d’apprentissage Participatives. Les membres de l’équipe du<br />
WWF ont reçu auparavant des formations sur ces méthodes.<br />
Dans le cadre de leur travail, ces méthodes sont<br />
conduites lors des premières approches au sein d’une nouvelle<br />
communauté demandant à gérer une ressource naturelle<br />
dans leur terroir.<br />
Responsabilisation sous forme de transfert de<br />
gestion<br />
Inventaire forestier et zonage<br />
Dans chaque localité de travail avec une communauté, des<br />
inventaires forestiers ont été conduits.La finalité est de connaitre<br />
les potentialités des différents types de forêts en<br />
produits forestiers ligneux et non ligneux. Une uniformisation<br />
des méthodes de relevés a été réalisée. A partir des<br />
cartes de zonages élaborées, les inventaires ont été concentrés<br />
dans les zones de conservations et des droits<br />
d’usage.Des parcelles imbriquées sont définies.Elles ont des<br />
surfaces de 100 x 100 m pour des diamètres de 1,30 m (dhp)<br />
supérieur ou égal à 30 cm;deux parcelles de 25 x 25 m pour<br />
les arbres de dhp compris entre 10 cm à 30 cm; et deux<br />
parcelles de 5x5m.Lesdimensions de ces parcelles peuvent<br />
varier en fonction du relief (si le terrain est trop accidenté,la<br />
réduction des dimensions est préconisée) et leur nombre<br />
varié.Mais quelques soient les variantes,une moyenne de un<br />
hectare par type de forêt est requise pour les gros diamètres.<br />
Elaboration de plan d’aménagement<br />
En vue de l’obtention des contrats de transfert de gestion<br />
des forêts auprès du Ministère des forêts, chaque communauté<br />
devrait établir des plans d’aménagement des forêts.
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 57<br />
Les plans d’aménagement suivent les directives proposées<br />
par la Direction Générale des forêts. Le WWF,pour l’élaboration<br />
de ces plans,a surtout veillé à ce que les prélèvements<br />
respectent le principe de la durabilité à savoir: ne prélever<br />
que le volume correspondant aux accroissements moyens<br />
annuels. Pour un meilleur équilibre entre les besoins des<br />
populations locales, des reboisements en espèces introduites<br />
sont aussi proposés dans ces plans d’aménagement,<br />
mais dans des parcelles en dehors des forêts naturelles.<br />
Appropriation sous forme de contrôle<br />
L’approche d’appropriation sous forme de contrôle a aussi<br />
été développée pour atteindre les objectifs des deux projets.<br />
Elle consiste à encourager les membres des communautés à<br />
l’établissement de parcelles d’observation en forêt en se<br />
basant sur les pistes existantes et sur les signes visibles de<br />
pressions (coupe, pièges). Des représentants dynamiques au<br />
sein de chaque communauté sont ainsi élus et font des visites<br />
(contrôles) régulières, parfois inattendues, en forêts. Ces<br />
visites sont organisées pour observer la biodiversité<br />
(floraison des plantes,augmentation du nombre des groupes<br />
de lémuriens observés, passage d’un oiseau ou autre type<br />
d’animal faisant la particularité de leur forêt etc.) et pour<br />
constater si des pressions sur les lémuriens et la forêt sont<br />
encore présentes. Ces membres notent leurs observations<br />
dans des cahiers réservés pour ces observations et tout<br />
autre évènement qu’ils jugent importants (rencontre avec<br />
d’autres personnes, etc.).<br />
Mesures d’accompagnement<br />
Le WWF défini les mesures d’accompagnement de toute<br />
activité développée pour contribuer à l’amélioration des<br />
conditions de vie des populations locales. Ces activités doivent<br />
à la fois réduire les pressions sur la biodiversité en générale<br />
et sur les cibles de conservation en particulier (forêt<br />
naturelle et lémurien) et compenser les efforts de conservations<br />
entrepris. L’approche consiste d’abord à analyser les<br />
résultats de toutes les études préalables aux transferts de<br />
gestion des forêts. A partir de ces analyses, les potentialités<br />
de chaque terroir (à partir des cartes d’occupation des sols<br />
et des rendements obtenus pour chaque spéculation engagée)<br />
dans lesquels vivent les communautés sont dégagées.<br />
Les mesures à développer cherchent ainsi à augmenter les<br />
sources de revenus des paysans et à améliorer leur alimentation<br />
pour une meilleure santé.<br />
Résultats et discussions<br />
Inventaires forestiers et zonages<br />
Au total, 14 inventaires forestiers (correspondants au nombre<br />
de transfert de gestion des forêts) ont été effectués dans<br />
la région de SAVA. La potentialité des forêts est variable. Le<br />
nombre des arbres de dhp > 30 varie de 200 à 400/ha. Au<br />
total, 200 espèces de plantes (ligneuses et non ligneuses)<br />
sont inventoriées. D’un site à l’autre, le nombre des espèces<br />
ligneuses inventoriées varie de 45 à 87.<br />
Les espèces les plus fréquemment rencontrées sont: Tambourissa<br />
religiosa,Weinmannia rutembergii,Zantoxyllon mananarense,<br />
Chrysophyllum boivinianum, Canarium madagascariensis,<br />
Symphonia fasciculata, Diospyros aff.ambilensis, Macaranga<br />
decaryana, Erythroxylum sphaeranthum, Brachylaena merana,<br />
Syzygium emirnense, Uapacca densiflora, Ocotea cymosa…<br />
En fonction de l’éloignement des forêts par rapport aux villages,<br />
leur état de dégradation diffère. Cette observation a<br />
permis de faire trois grande classifications: forêt naturelle<br />
plus ou moins intacte,classée à protéger ou à conserver (for-<br />
mant donc le noyau dur);forêt partiellement dégradée,localisée<br />
encore en plein cœur de la forêt,classée comme forêt à<br />
restaurer;forêt naturelle à faible potentialité, due à un degré<br />
d’écrémage localisée à la périphérie des lisières, classée<br />
comme zone de cantonnement de droit d’usage.<br />
Transfert de gestion des forêts<br />
Le WWF a pu mettre en place dans le cadre de ces deux projets<br />
14 transferts de gestion. Pour chaque communauté, la<br />
surface totale des forêts dans ces transferts varie de 300 à<br />
5000 ha (dépendant du taux de couverture forestière dans le<br />
territoire de chaque village d’appartenance de la communauté).<br />
Les zonages des forêts sont décrits précédemment.<br />
En principe, les contrats sont établis pour trois ans. Ensuite<br />
une évaluation devra se faire par le Service Forestier en partenariat<br />
avec les communes d’appartenance des communautés<br />
gestionnaires des forêts.Etablis à partir de 2007,certains<br />
contrats nécessitent ainsi une évaluation à partir de cette<br />
année ou au plus tard en début de l’année prochaine.<br />
Au total,les 14 transferts de gestion des forêts ont permis de<br />
sécuriser sous la gestion des COBA,27 000 ha de forêts.Ces<br />
forêts incluent tous les types de forêts à différents usages définis<br />
auparavant (conservation, restauration, droit d’usage).<br />
Les valeurs de la restauration pour le WWF sont présentées<br />
dans son document de la vision de la biodiversité (Erdmann<br />
et al., 2005) qui a défini 40 aires comme Aires Prioritaires de<br />
conservation de par leur valeur en biodiversité. Les études<br />
sur leur état de dégradation ont relevé aussi que 23 de ces aires<br />
auront probablement besoin d’importante restauration<br />
(< 20 % de forêt),si elles doivent entièrement concourir à la<br />
conservation de la biodiversité. Dans tous les sites de transferts<br />
de gestion des forêts, le WWF met ainsi l’accent sur<br />
l’importance de la restauration. Les communautés avec l’encadrement<br />
du WWF, des observations sur terrain et des<br />
inventaires effectués définissent ainsi des zones de restauration<br />
dont la superficie varie d’une communauté à une<br />
autre.Les espèces utilisées pour ces restaurations des forêts<br />
dégradées sont essentiellement des essences autochtones.<br />
Leur choix est justifié par leur emplacement (héliophile pour<br />
les zones très ouverts et/ou périphérie de la forêt; nomade<br />
pour les zones sous couvert des espèces héliophiles) ou par<br />
leur utilisation (construction, nourriture des lémuriens).<br />
Suivis et contrôles<br />
Le WWF, pour faciliter l’uniformisation des suivis et contrôle<br />
en matière des lémuriens, a élaboré un livret sur les<br />
lémuriens avec la photo des espèces et une description<br />
sommaire des espèces.L’idée de départ était d’exploiter ces<br />
livrets peu avant la fin des projets pour les analyser. Il a<br />
pourtant été constaté que les représentants des communautés<br />
n’ont pas utilisé les livrets,mais les a bien classés dans<br />
leur valise. Leur explication est que le livret (en couleur) est<br />
trop beau pour être amené et abimé en forêt à cause de<br />
l’humidité. Néanmoins, les suivis et contrôles ont été effectivement<br />
effectués par les communautés. De plus, les données<br />
ont été stockées soit dans des cahiers à part, soit dans<br />
leur tête.<br />
Dans le cadre des suivis et contrôles instaurés de manière<br />
participative,des suivis et contrôles à part doivent aussi être<br />
faits par les organismes techniques d’appuis des communautés.<br />
En effet, à travers les expériences de l’élaboration du<br />
livret, il a été constaté que les instructions et formations<br />
n’ont pas été suivies correctement.Les agents des deux projets,<br />
ne se sont donc rendus compte qu’un peu tardivement<br />
de la nécessité de faire aussi des suivis et contrôles rapprochés.
Page 58 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
Table 1: Caractéristiques des localités d’observation des lémuriens.<br />
Localités Andrakengy Andasipiro Ambodivoara Ambodimandresy<br />
Période 20 Novembre - 6 Décembre - 19 Novembre - 5 Décembre -<br />
d’observation 3 Décembre 2006 17 Décembre 2006 3 Décembre 2006 17 Décembre 2006<br />
Coordonnées S 14° 18’ 53.4’’ S 14° 12’ 10.0’’ S 14° 32’ 05.0’’ S 14° 32’ 05.0’’<br />
géographiques E 049< 16’ 38.3’’ E 049< 22’ 34.8’’ E 049< 26’ 42.1’’ E 049< 30’ 21.1’’<br />
Les suivis et contrôles effectués par les paysans ont quand<br />
même permis de localiser des sites d’observations de l’espèce<br />
Propithecus candidus (Simpona).Patel, en échangeant les<br />
données avec l’équipe des projets du WWF a publié en 2009<br />
que l’espèce Propithecus candidus est aussi rencontrée dans le<br />
Corridor de Betaolana et de Tsaratanana-Betaolana. Quelques<br />
caractéristiques des localités où les inventaires ont été<br />
effectués et où cette espèce a été observée sont synthétisées<br />
dans le tableau 1.<br />
Les alternatives aux pressions et menaces<br />
Les pressions sont définies comme étant les activités causant<br />
des impacts négatifs aussi bien sur les forêts que sur les<br />
lémuriens. Ces activités peuvent être légales ou illégales. Par<br />
contre, les menaces sont des activités pouvant apparaître<br />
dans le futur et pouvant avoir des impacts négatifs sur les<br />
cibles (dans le cadre de ce projet les cibles sont les lémuriens<br />
et leur habitat).<br />
Les analyses,effectuées dans le cadre de ces deux projets ont<br />
montré que les pressions et les menaces sur les cibles sont<br />
principalement constituées de: la déforestation causée essentiellement<br />
par les cultures sur brûlis, la dégradation des<br />
forêts engendrée par les prélvements diverses (bois de construction<br />
ou autre matériels pour la construction tels que les<br />
lianes et les bambous, cueillette de miel), la chasse moderne<br />
et le piège traditionnel. La déforestation tue à la fois les<br />
lémuriens et détruit leur habitat. Il en est de même des<br />
pièges traditionnels mais à un degré moindre.<br />
Pour arrêter les pressions sur les forêts, les deux projets du<br />
WWF ont donc analysé les activités pouvant remplacer<br />
celles formant une pression et constituant une menace dans<br />
le futur.En bref les activités développées sont:l’amélioration<br />
de l’exploitation de l’espace par l’agroforesterie; la promotion<br />
des cultures maraichères,l’amélioration des cultures de<br />
riz sur les bas fonds étroits par les Systèmes de Riziculture<br />
Améliorés (SRA),la promotion des briques pour la construction<br />
des maisons et le reboisement des espèces à croissance<br />
rapide; l’apiculture, la pisciculture et l’amélioration de l’élevage<br />
des volailles.<br />
Elaboration d’une stratégie régionale pour la conservation des<br />
lémuriens<br />
Les principales cibles de conservations des deux projets sont<br />
la forêt humide et les lémuriens. A part les transferts de<br />
gestion,les deux projets ont aussi été conduits pour éduquer,<br />
informer et sensibiliser les populations locales (partant des<br />
élèves dans les écoles primaires aux écoles secondaires mais<br />
aussi l’ensemble des populations concernées dans les communes).<br />
Pour la sauvegarde des lémuriens, les engagements des parties<br />
prenantes sont recherchés à travers l’élaboration et la<br />
mise en œuvre d’une stratégie régionale pour la conservation<br />
des lémuriens. Dans cette stratégie, les activités de<br />
chaque partie sont définies de manière à minimiser les<br />
dépenses monétaires afin de les rendre réalisables.<br />
Pour une meilleure intégration ou engagement de chaque<br />
entité, une Association des Amis des lémuriens a été créée.<br />
Cette association est mise en réseau via un site web aux<br />
amoureux des lémuriens dans le monde. La vision a été<br />
développée pour le long terme. Toutefois,<br />
le WWF s’est rendu comte<br />
qu’avec l’isolement de chaque localité,<br />
les réponses instantanées et directes<br />
des membres aux intéressés via le site<br />
web sont très limités et précaires. En<br />
effet,pour se connecter,les élèves doivent<br />
aller à Andapa; et même si cer-<br />
tains élèves d’Andapa sont concernés, le reflexe avec cette<br />
haute technologie nécessite encore un encadrement rapproché<br />
pour être efficace.<br />
Pour mieux unir les efforts,une Union Régionale des Associations<br />
de gestion des forêts, appelée aussi "Gestion Unie du<br />
Corridor de Betaolana" a aussi été créé. Malgré les efforts<br />
investis pour créer cette Union, elle est restée au stade de<br />
constitution (dépôt de dossier) au moment où les phases des<br />
deux projets sont terminées. Toutefois, elle est engagée et<br />
sera reprise dans les autres projets du WWF dans cette<br />
localité. Il est donc souligné ici l’importance des encadrements<br />
par des organismes promoteurs dans le long terme ou<br />
du moins à moyen terme pour obtenir de meilleurs impacts<br />
dans les phases du projet.<br />
Les parties prenantes définies dans ce document<br />
sont composées par les partenaires techniques et les autres<br />
organismes et/ou associations travaillant dans les domaines<br />
de l’environnement, les communautés de base, les organes<br />
de décentralisation et de déconcentration de l’Etat (les<br />
régions, les communes, les districts et les Fokontany).<br />
Conclusions<br />
La capitalisation des deux projets du WWF dans le Nord a<br />
permis de comprendre les efforts encore à fournir dans le<br />
cadre de la conservation des lémuriens et de leur habitat.Les<br />
deux projets conduits dans la région de SAVA sont complémentaires,<br />
ceci a permis une uniformisation des approches<br />
techniques et scientifiques. Les deux projets ont enrichis les<br />
données sur la diversité de la zone en flore et en faune.Ils ont<br />
pu se réaliser de manière participative. En effet, ils ont été<br />
très bien accueillis par les populations locales du fait que ces<br />
projets étaient majoritairement axés sur les activités de<br />
développement pour atteindre leurs objectifs de conservation.<br />
En conclusion, les défis à lancer doivent se concentrer sur<br />
deux aspects.<br />
Aspect technique: des études plus poussées sur la flore et la<br />
faune sont à conduire. En effet, les découvertes à faire sont<br />
encore immenses vu l’étendue du massif forestier.<br />
Aspect développement: des activités sont engagées et n’en<br />
sont qu’au début de leur mise en œuvre. Il serait nécessaire<br />
dans le futur de faire des évaluations de ces projets en<br />
termes d’impact sur la conservation des habitats et des<br />
lémuriens ainsi que sur l’amélioration des conditions de vie<br />
de la population locale.<br />
Ces deux projets ne sont pas des projets de recherches.<br />
Toutefois, ils ont pu être enrichis par les interventions des<br />
scientifiques consultants d’une part et des équipes du WWF<br />
d’autre part.<br />
Remerciements<br />
Nos vifs remerciements s’adressent aux bailleurs qui ont<br />
financés les projets du WWF Madagascar dans la région de<br />
SAVA. Ils ont permis de conduire à la fois des études scientifiques<br />
et des activités de développement.Ces bailleurs sont:<br />
WWF Allemagne, WWF Suède et la Conservation Internationale.<br />
Bien qu’à faible volume en terme de fonds, nous<br />
tenons aussi à remercier le WWF Danemark pour des fonds<br />
spécifiquement alloués à l’élaboration des support de com-
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 59<br />
munication (poster,teeshirt,banderoles etc.) pour une meilleure<br />
compréhension des aspects de la conservation auprès<br />
des paysans en particulier et des parties prenantes en<br />
général.<br />
Bibliographie<br />
Erdmann, T.K.; Rambeloarisoa, G.; Ratsifandrihamanana, N.;<br />
Ranaivomanantsoa,A.F.;Abraham,A.;Powell,G.;Allnutt,T.<br />
2005. Vision sur la Biodiversité de l’Ecorégion de la forêt<br />
Humide de Madagascar. Rapport interne pour le WWF-<br />
International et le WWF Madagascar and West Indian<br />
Ocean Programme Office.<br />
Patel,E.R.2009.Silky Sifaka,Propithecus candidus,1871.In:R.A.<br />
Mittermeier, J. Wallis, A.B. Rylands, J.U. Ganzhorn, J.F.<br />
Oates,E.A.Williamson,E.Palacios,E.W.Heymann,M.C.M.<br />
Kierulff, Y.Long,J. Supriatna,C.Roos,S.Walker,L.Cortés-<br />
Ortiz, C. Schwitzer (eds.). Primates in Peril: The World’s<br />
25 Most Endangered Primates 2008-2010. IUCN/ SSC<br />
Primate Specialist Group (PSG), International Primatological<br />
Society (IPS), and Conservation International (CI),<br />
Arlington, VA. 84pp.<br />
Garreau,J.M.;Manantsara,A.2003.The protected-area complex<br />
of the Parc National de Marojejy and the Reserve<br />
Speciale d’Anjanaharibe-Sud. Pp. 1451-1458. In: S.M.<br />
Goodman, J.P. Benstead (eds.). The Natural History of<br />
Madagascar. University of Chicago Press, Chicago, USA.<br />
Goodman,S.M.;Raherilalao,M.J.; Rakotomalala,D.;Raselimanana,A.;Schütz,H.;Soarimalala,V.2003.Les<br />
Lémuriens.Pp.<br />
279-286.In:S.M.Goodman,L.Wilmé (eds.).Nouveaux résultats<br />
d’inventaires biologiques faisant référence à l’altitude<br />
dans la région des massifs montagneux de Marojejy<br />
et d’Anjanaharibe-Sud. Recherches pour le Développement,<br />
série sciences biologiques no. 19. Centre d’Information<br />
et de Documentation Scientifique et Technique,<br />
Antananarivo, Madagascar.<br />
Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyer, D.;<br />
Mast,R.1994.Lemurs of Madagascar.Conservation International,<br />
Washington, D.C., USA.<br />
Mittermeier, R.A.; Valladares-Padua, C.; Rylands, A.B.; Eudey,<br />
A.A.; Butynski, T.M.; Ganzhorn, J.U.; Kormos, R.; Aiguiar, J.<br />
M.; Walker, S. 2005. The World’s 25 Most Endangered<br />
Primates 2004-2005. Lemur News 10: 3-6.<br />
Petter,J.J.;Charles-Dominique,P.1979.Vocal communication<br />
in prosimians. Pp. 247-305. In: G.A. Doyle and R.D. Martin<br />
(eds.).The Study of Prosimian Behavior.Academic Press.<br />
Rajaoson,A.;Rakotonirina,L.H.2007.Rapport pour le WWF;<br />
Inventaire des lémuriens et des forêts dans le Corridor de<br />
Betaolana et le Corridor Tsaratanana-Marojejy. 89p.<br />
Tattersall, I. 1982. The Primates of Madagascar. Columbia<br />
University Press, New York, USA.<br />
Genetic diversity in ten Indri (Indri indri)<br />
populations compared to other lemur<br />
species<br />
John Zaonarivelo 1,Rick Brenneman 2*,Rambinintsoa<br />
Andriantompohavana 3, Edward E. Louis, Jr. 2,3<br />
1University of North Madagascar, Antsiranana, Madagascar<br />
2Center for Conservation and Research, Omaha’s Henry<br />
Doorly Zoo,3701 South 10th Street,Omaha,NE 68107,USA<br />
3Madagascar Biodiversity Partnership, Antananarivo, 101<br />
Madagascar.<br />
*Corresponding author: rabr@omahazoo.com<br />
Genetic diversity is considered by most to be the key to long<br />
term survival and the maintenance of the evolutionary trajectory<br />
of a species.Genetic variation at loci under selection<br />
gives the species as a whole the mechanisms with which to<br />
respond or adapt to environmental changes.Lemurs in general<br />
are poorly studied with respect to establishing baseline<br />
parameter estimates for genetic diversity.Only limited popu-<br />
lation genetics studies exist on the genera Propithecus, Avahi,<br />
Varecia, Eulemur, Microcebus, and Mirza (Tab. 1). To date, the<br />
genus Indri is depauperate of population genetic data that<br />
would help better understand the genetic diversity harbored<br />
in its populations.<br />
Tab.1:Lemur studies using multilocus microsatellite suites to<br />
estimate genetic diversity as observed (HO) and expected<br />
(HE) heterozygosity levels.<br />
Species PopuSam- Loci HO HE Reference<br />
lationsples Indri indri 2 20 20 0.654 0.766 Zaonarivelo<br />
et al., 2007b<br />
Propithecus<br />
deckeni<br />
2 20 14* 0.790 0.851 Lei et al., 2008a<br />
P. deckeni 1 10 18* 0.776 0.776 Lei et al., 2008b<br />
P. coquereli 1 25 20 0.635 0.771 Rakotoarisoa<br />
et al., 2006a<br />
P. candidus 2 18 17* 0.648 0.614 McGuire et al., 2009<br />
P. coronatus 1 10 18* 0.771 0.774 Lei et al., 2008b<br />
P. diadema 2 20 13* 0.818 0.814 Ramarokoto<br />
et al., 2008<br />
P. edwardsi 2 20 12* 0.681 0.618 Bailey et al., 2009<br />
P. verreauxi 3 30 13 0.670 0.712 Rakotoarisoa<br />
et al., 2006b<br />
P. tattersalli 2 20 16* 0.673 0.683 Razafindrakoto<br />
et al., 2008<br />
P. tattersalli **3 75 13<br />
Quéméré et al.,<br />
0.699 0.682<br />
2009<br />
P. tattersalli 9 224 13<br />
Quéméré et al.,<br />
0.690 0.660<br />
2010<br />
Avahi laniger 5 37 22 0.640 0.838 Andriantompohavana<br />
et al., 2004<br />
A. occidentalis<br />
1 7 22 0.514 0.586 Andriantompohavana<br />
et al., 2004<br />
Varecia<br />
rubra<br />
2 32 <strong>15</strong> 0.616 0.618 Razakamaharavo<br />
et al., 2010<br />
V. variegata<br />
variegata<br />
4 35 25 0.337 0.506 Louis et al., 2005<br />
Eulemur<br />
cinereiceps<br />
2 21 16* 0.598 0.641<br />
Tokiniaina et al.,<br />
2009<br />
E. collaris 4 40 10* 0.617 0.576 Ranaivoarisoa<br />
et al., 2010<br />
E. sanfordi 5 54 11* 0.562 0.567 Ramanamahefa<br />
et al., 2010a<br />
E. coronatus 6 80 11* 0.636 0.673 Ramanamahefa<br />
et al., 2010b<br />
E. rubriventer<br />
2 12 20 0.531 0.643 Andriantompohavana<br />
et al., 2007<br />
Lemur catta 1 24 7*<br />
Zaonarivelo et al.,<br />
0.837 0.838<br />
2006<br />
Microcebus<br />
ravelobensis<br />
8 205 7<br />
Olivieri et al.,<br />
0.6<strong>15</strong> 0.605<br />
2007<br />
M. ravelobensis<br />
12 187 8<br />
Radespiel et al.,<br />
0.708 0.734<br />
2008<br />
M. bongolensis<br />
3 45 8<br />
Olivieri et al.,<br />
0.557 0.565<br />
2008<br />
M. danfossi 7 78 8 0.628 0.662 Olivieri et al., 2008<br />
Mirza<br />
coquereli<br />
***1 69 7<br />
Markolf et al.,<br />
0.712 0.799<br />
2008<br />
* Heterozygosity averages calculated using loci with null allele<br />
frequency estimates less than 0.10.<br />
** Estimated over genetic clusters, not actual populations<br />
*** Samples taken from 1993-2006.<br />
The Indri (Indri indri, Gmelin, 1788), or Babakoto as it is<br />
known in most of eastern Madagascar,is the largest extant lemur<br />
(Powzyk and Thalmann, 2003). Indri are primarily midlevel<br />
forest folivores preferentially feeding on immature<br />
leaves and somewhat also on mature leaf matter, flowers,<br />
fruits,seeds and even bark when necessary (Britt et al.,2002).<br />
The Babakoto is currently threatened by the rapid reduction
Page 60 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
of forest cover and fragmentation of suitable habitat (Harper<br />
et al., 2007) which limits the species’ density and range<br />
(Glessner and Britt, 2005). Although protected, the Babakoto<br />
is also threatened by subsistence hunting pressure and<br />
bush meat trade (Golden, 2005). Designated as Endangered<br />
according to the IUCN Red List of Threatened Species<br />
(IUCN, 2008), I. indri is currently split into two subspecies,<br />
Indri indri indri (Gmelin, 1788) and Indri i. variegatus (Gray,<br />
1872). Here, we present population genetic parameter estimates<br />
from populations along the entire range of the species<br />
analyzed from nuclear microsatellite multilocus genotypes.<br />
Methods<br />
Samples were collected from 106 Indri from 10 sites across<br />
the geographical range of the species (Fig. 1). From north to<br />
south,the forests represented in the collection were Anjanaharibe-Sud<br />
Special Reserve, Marotandrano Special Reserve,<br />
Ambatovaky Special Reserve,Zahamena Special Reserve and<br />
National Park, Betampona Nature Reserve, Anjozorobe<br />
Regional Forest Reserve, Mantadia National Park, Analamazoatra<br />
Special Reserve (Andasibe),Maromizaha Classified<br />
Forest, and Anosibe an’ala Classified Forest. The elevations<br />
of the sampling sites ranged from lowland forests (Anosibe<br />
An’ala, 125 m asl) to highland forests (Anjozorobe, 1358 m<br />
asl).<br />
Fig. 1: Map of Madagascar indicating the study areas.<br />
Immobilization and collection<br />
All lemurs investigated in this study were free-ranging and<br />
were immobilized with a CO2 powered DAN-INJECT (Brrkop,<br />
Denmark) Model JM rifle propelling Pneu-Darts (Williamsport,<br />
PA) loaded with 10 mg/kg estimated body weight<br />
of Telazol ® (Fort Dodge).We recorded the location (within 6<br />
m accuracy) of all of the immobilized lemurs using a global<br />
positioning system (GPS) device. Each individual was transported<br />
back to the base camp where complete morphometric<br />
data were taken (Zaonarivelo et al., 2007a). Whole<br />
blood (1.0 cc per kilogram) from the femoral artery and<br />
2.0 mm skin biopsies from the ear pinnae were collected<br />
from each sedated lemur (Junge and Louis, 2002). A Home<br />
Again ® (Home Again Pet Recovery Service, East Syracuse,<br />
NY) microchip was placed subcutaneously between the scapulae<br />
of each lemur to positively identify individuals re-captured<br />
during any future immobilizations. Following data and<br />
sample collection, an injection of lactated Ringer’s solution<br />
was administered subcutaneously to support maintenance<br />
requirements and to dissipate the effect of the Telazol ®.Animals<br />
were monitored for three hours post recovery then<br />
released according to the capture GPS coordinates.<br />
Data generation<br />
Ear punches were dissected into quarters and DNA was<br />
extracted using standard PCI/Chloroform procedures (Sambrook<br />
et al., 1989). Approximately 50 ng of genomic DNA<br />
was used for each PCR reaction. Multilocus genotypes were<br />
generated from a suite of 20 Indri-specific microsatellite loci<br />
as described in Zaonarivelo et al. (2007b). The genotype file<br />
was checked for typographical errors,scoring errors,stutter<br />
bands and allele dropout with Micro-Checker (van Oosterhout<br />
et al., 2004) and Microsatellite Analyser (MSA; Dieringer<br />
and Schlötterer, 2002). We used CERVUS (version 2.0,<br />
Marshall et al.,1998;Slate et al.,2000) to identify loci with excessive<br />
null allele frequency estimates (nf > 0.10) and to estimate<br />
polymorphic information content of the loci.Moderate<br />
(0.05 < nf < 0.20) and high (0.20 < nf) null allele frequencies<br />
can have significant effects on population genetics parameter<br />
estimates (Chapuis and Estoup, 2007). The process of redesigning<br />
primer pairs is both costly and time consuming;<br />
therefore,we opted to delete problematic loci from the data<br />
set. We deleted eight loci with moderate null allele frequencies<br />
(nf > 0.1) to reduce the bias from misclassification of null<br />
heterozygotes as homozygotes (Callen et al., 1993; Hoffman<br />
and Amos, 2005) and to control the variance of parameter<br />
estimates (Chapuis and Estoup, 2007). The accepted loci<br />
were verified for independence of linkage disequilibrium<br />
(with Bonferroni-adjusted P-values) in FSTAT (Goudet,1995,<br />
2001).<br />
Hardy-Weinberg exact tests (Guo and Thompson, 1992)<br />
were performed by locus and population in Genepop (version<br />
4.0, Raymond and Rousset, 1995). Initially, we used the<br />
default settings for the MCMC estimation of HWE then<br />
increased the batch size from 100 to 250 to reduce the standard<br />
error of the P-value to below 0.01. Genetic diversity<br />
was measured as observed heterozygosity (HO) and expected<br />
heterozygosity (HE). In addition, the number of effective<br />
migrants was estimated globally and pair-wise using the private<br />
allele method. We used FSTAT to calculate the total<br />
number of alleles (k), mean number of alleles (MNA), and<br />
rarefacted allelic richness (AR; Leberg, 2002) by locus and<br />
population.Allelic richness estimates the allelic diversity in a<br />
data set based on the population with the fewest number of<br />
individuals contributing genotypes by locus. This is an unbiased<br />
comparison of allelic diversity since populations with<br />
more contributors provide a greater opportunity to capture<br />
more alleles from lower frequency occurrences. Wright’s Fstatistics<br />
were estimated in FSTAT for within population<br />
similarity (FIS) and between population differences (FST)<br />
according to Weir and Cockerham (1984).<br />
The effective population sizes were estimated with the linkage<br />
disequilibrium (LD) option in NeEstimator (Peel et al.,<br />
2004; Hill, 1981; Waples, 1991). We tested all populations<br />
having met the minimum statistical threshold required (n =<br />
20 genes or 10 individuals) for the presence of bottleneck<br />
events using Bottleneck (version 2.0, Cornuet and Luikart,
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 61<br />
1996; Luikart et al., 1998; Piry et al., 1999) under the Infinite<br />
Alleles Model (IAM; Kimura and Crow, 1964), the Stepwise<br />
Mutation Model (SMM;Ohta and Kimura,1973),and the Two<br />
Phase Model (TPM;di Rienzo et al.,1995).We varied the proportion<br />
of the single step contribution to the TPM to identify<br />
the P < 0.05 threshold of significance.The program identifies<br />
populations with an excess of heterozygosity relative to<br />
mutation-drift equilibrium which is indicative of a reduction<br />
in the effective population size (Maruyama and Fuerst,1985).<br />
An estimate of relationships among all individuals sampled at<br />
each forest was done in SPAGeDi (Hardy and Vekemans,<br />
2002),then compared to a simulation of known pedigreed individuals.<br />
The analysis was performed to calculate the relationship<br />
coefficients described in Queller and Goodnight<br />
(1989) in the absence of spatial data.<br />
Results<br />
Genetic diversity as mean number of alleles ranged from<br />
6.08-8.92 per population. Using the rarefacted allelic richness,<br />
the range lowered to 5.87-7.67. The expected heterozygosity<br />
ranged from 0.77-0.86 (P > 0.05;Fig.2) with an average<br />
of 0.81, while the observed heterozygosity ranged from<br />
0.65-0.84 (P < 0.05; Fig. 3) with an average of 0.74. The number<br />
of effective breeders in the sampled populations averaged<br />
between 12.6 and 39.6 per population (Tab. 2).<br />
Results from Bottleneck showed that none of the 10 populations<br />
deviated from a mutation-drift equilibrium under the<br />
SMM.Three populations,Anjanaharibe Sud,Ambatovaky and<br />
Anjozorobe did not show evidence of population bottleneck<br />
under the IAM either.The rest of the populations were significant<br />
for bottleneck events under the IAM and varying proportions<br />
of single step contributions under the TPM.<br />
The frequencies of the relationship coefficients estimated<br />
using SPAGeDi were overlaid upon a simulation generated<br />
from known pedigreed data so that each of the population’s<br />
relative distribution of relationships could be compared with<br />
parent offspring, full sibling, half sibling and unrelated relationship<br />
coefficient distributions (Fig. 4). The data indicated<br />
that the sampling was from individuals that were somewhat<br />
related more than the unrelated individuals in the reference<br />
simulation. Inbreeding can also be due to background relatedness<br />
where an increased allelic identity by descent is a result<br />
from bottleneck events in the population’s history.Relationship<br />
coefficient distributions sup-<br />
port the assumption that the individuals<br />
sampled were often from family<br />
groups. All of these sources may potentially<br />
be due to the effects of habitat<br />
fragmentation which is certainly<br />
the case in the Anosibe an’ala population<br />
where the habitat is so fragmented<br />
that although multiple family<br />
groups were encountered, they were<br />
found in isolated forest fragments.<br />
Discussion<br />
Of the 10 Indri populations sampled,<br />
six (Anjanaharibe Sud, Ambatovaky,<br />
Zahamena, Betampona, Mantadia, and<br />
Anosibe an’ala) deviated from HWE<br />
with an excess of homozygotes.Considering<br />
inbreeding as potential cause,<br />
five of the populations (Anjanaharibe<br />
Sud, Ambatovaky, Betampona, Mantadia,<br />
and Andasibe) had relatively<br />
high FIS and one (Anosibe an’ala) had<br />
Y<br />
.90<br />
.80<br />
.70<br />
.60<br />
55DD<br />
A B C D E F G H I J<br />
X<br />
Fig. 2: Ranges of expected heterozygosities with 95 % confidence<br />
intervals: A) Anjanaharibe Sud; B) Marotandrano; C)<br />
Ambatovaky; D) Zahamena; E) Betampona; F) Anjozorobe;<br />
G) Mantadia;H) Andasibe;I) Maromizaha; J) Anosibe an’ala.<br />
4DF4<br />
Y<br />
.90<br />
.85<br />
.80<br />
.75<br />
A B C D E F G H I J<br />
X<br />
Fig. 3: Ranges of observed heterozygosities with 95 % confidence<br />
intervals: A) Anjanaharibe Sud; B) Marotandrano; C)<br />
Ambatovaky; D) Zahamena; E) Betampona; F) Anjozorobe;<br />
G) Mantadia;H) Andasibe;I) Maromizaha; J) Anosibe an’ala.<br />
Tab. 2: Population genetic parameter estimates for 10 populations comprised of n<br />
samples each derived from 12 microsatellite loci for number of alleles (k), the mean<br />
number of alleles (MNA), allelic richness (AR), probability of satisfying Hardy-Weinberg<br />
Equilibrium (HWE), observed (HO) and expected (HE) heterozygosities, inbreeding<br />
estimate (FIS), the number of effective breeders (Neb) estimated with the<br />
linkage disequilibrium method and 95 % confidence interval,and results from the Bottleneck<br />
test under the infinite allele model (IAM) and the two phased model (TPM)<br />
with proportion of multistep mutations contributing to the P < 0.05 significance level.<br />
n k MNA AR HWE HO HE FIS Neb 95% CI IAM TPM<br />
ANJ 10 80 6.67 6.41 * 0.67 0.77 0.135 23.0 16.2-37.8 NS NS<br />
TAND 10 73 6.08 5.87 NS 0.75 0.77 0.024 21.9 <strong>15</strong>.1-36.9 ** 70<br />
VAK 11 76 6.33 5.92 NS 0.69 0.77 0.121 20.1 14.7-30.4 NS NS<br />
ZAH 14 107 8.92 7.65 NS 0.81 0.86 0.060 39.6 28.8-61.3 ** 10<br />
BET 10 79 6.58 6.37 ** 0.73 0.80 0.093 18.3 13.5-27.1 ** 20<br />
ANJZ 10 83 6.92 6.70 NS 0.84 0.79 -0.066 20.9 <strong>15</strong>.1-32.5 NS NS<br />
TAD 10 96 8.00 7.67 NS 0.75 0.85 0.120 20.1 <strong>15</strong>.3-28.3 ** 70<br />
DASI 11 81 6.75 6.40 NS 0.73 0.81 0.095 12.6 10.2-16.2 ** 5<br />
MIZA 10 89 7.42 7.17 NS 0.83 0.84 0.021 <strong>15</strong>.9 12.4-21.3 ** 5<br />
ANOSIBE 10 92 7.67 7.38 * 0.65 0.85 *0.236 28.8 19.9-49.3 ** NS<br />
* P < 0.05, ** P < 0.001; Anjanaharibe-Sud(ANJ), Marotandrano (TAND), Ambatovaky (VAK),<br />
Zahamena (ZAH), Betampona (BET), Anjozorobe (ANJZ), Mantadia (TAD), Andasibe (DASI).<br />
Maromizaha (MIZA), Anosibe an’ala (ANOSIBE).
Page 62 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
Frequency<br />
Frequency<br />
Frequency<br />
Frequency<br />
Frequency<br />
Frequency<br />
Frequency<br />
0,4<br />
0,3<br />
0,2<br />
0,1<br />
0<br />
0,4<br />
0,3<br />
0,2<br />
0,1<br />
0<br />
0,4<br />
0,3<br />
0,2<br />
0,1<br />
0<br />
0,4<br />
0,3<br />
0,2<br />
0,1<br />
0<br />
0,4<br />
0,3<br />
0,2<br />
0,1<br />
0<br />
0,4<br />
0,3<br />
0,2<br />
0,1<br />
0<br />
0,4<br />
0,3<br />
0,2<br />
0,1<br />
0<br />
Anjanaharibe-Sud<br />
-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9<br />
Relationship Coefficients<br />
Morontandrano<br />
-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9<br />
Relationship Coefficients<br />
Ambatovaky<br />
-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9<br />
Relationship Coefficients<br />
Zahamena<br />
-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9<br />
Relationship Coefficients<br />
Betampona<br />
-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9<br />
Relationship Coefficients<br />
Anjozorobe<br />
-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9<br />
Relationship Coefficients<br />
Mantadia<br />
-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9<br />
Relationship Coefficients<br />
Frequency<br />
Frequency<br />
Frequency<br />
0,4<br />
0,3<br />
0,2<br />
0,1<br />
0<br />
0,4<br />
0,3<br />
0,2<br />
0,1<br />
0<br />
0,4<br />
0,3<br />
0,2<br />
0,1<br />
0<br />
Andasibe<br />
-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9<br />
Relationship Coefficients<br />
Maromizaha<br />
-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9<br />
Relationship Coefficients<br />
Anosibe an'ala<br />
-0,5 -0,3 -0,1 0,1 0,3 0,5 0,7 0,9<br />
Relationship Coefficients<br />
Parent-Offspring Full Sibling Half Sibling<br />
Unrelated All Populations<br />
Fig. 4: Distributions of relationship coefficients estimated in<br />
each population overlaid on a simulation of 10,000 known<br />
genotypes and pedigreed relationships (Queller and Goodnight,<br />
1989).<br />
a significantly high FIS estimate. Considering relationship<br />
among the samples as one manner of capturing inbreeding,all<br />
populations show some degree of relationship above what<br />
would be expected if the sampled individuals were unrelated.<br />
This is not surprising since samples were collected as found<br />
and this could include individuals that are members of a family<br />
group which is supported by the relationship coefficient<br />
distributions in Fig. 2.<br />
All populations demonstrated some degree of recent reduction<br />
in the effective population sizes.The bottlenecks did not<br />
appear to have been a single global event as different populations<br />
showed differing degrees to which the bottlenecks<br />
were detected. Using Lawler’s (2008) 4*Neb*generations,<br />
the bottlenecks detected would have had an expected window<br />
of occurrence of up to 250 to 800 years ago, within the<br />
timeframe of human encroachment. Hence, we postulate<br />
that among other things, anthropogenic disturbances,<br />
whether habitat destruction or subsistence hunting, may<br />
have influenced the demographic reduction that we detected<br />
in the bottleneck test. These baseline values could be useful<br />
in long-term or future studies to determine genetic health<br />
trends over time under various forest or habitat conditions.<br />
Genetic diversity is considered to be the most important<br />
factor in determining the genetic health of a species or population.Among<br />
the lemurs,there is little information in the literature<br />
that addresses the genetic diversity estimations for<br />
multiple populations of a given species. Ranaivoarisoa et al.<br />
(2010) found the observed heterozygosity in three of four<br />
Eulemur collaris populations to be higher than the expected<br />
heterozygosity. In E. coronatus, Ramanamahefa et al. (2010a)
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 63<br />
found the expected heterozygosity to be higher in five out of<br />
six populations,one comparison was significantly higher (P <<br />
0.01).In E.sanfordi,Ramanamahefa et al.(2010b) found that in<br />
all populations sampled,the observed heterozygosities were<br />
higher than the expected heterozygosities. Lastly, Quéméré<br />
et al. (2009), found that their study estimated the expected<br />
heterozygosity level for several populations of P. tattersalli to<br />
be higher than the estimate found by Razafindrakoto et al.<br />
(2008) using a different marker suite in different populations.<br />
Other studies (included in Tab. 2) have estimated heterozygosity<br />
levels for one or two populations of various lemur<br />
species in recent years.While these estimates provide some<br />
general sense of genetic diversity, they are not standardized<br />
(e.g. not the same markers were used in each study) so the<br />
estimates are contingent on the polymorphic information<br />
content and amplification quality in each independent study.<br />
What we do see is a relative trend that the observed<br />
heterozygosities are in general lower, but usually not significantly,<br />
than the expected heterozygosities under the assumptions<br />
of HWE.<br />
In this study on I. indri, we found the differences among the<br />
HO estimates to differ but not significantly and the differences<br />
among the HE estimates to differ with low significance<br />
(P < 0.05). The average estimates for heterozygosity, thus<br />
genetic diversity, were in the upper range of those found in<br />
limited population studies on other lemur species. These<br />
estimate trends provide the basis for future and integrative<br />
studies where multiple species might be considered in sympatric<br />
zones to investigate the overall genetic health of the<br />
biodiversity and to better understand the effects that humans<br />
may be having on the evolutionary potential of lemurs.<br />
Acknowledgements<br />
We acknowledge the Ministry of Environment and Eaux &<br />
Forets, Madagascar National Parks, U.S. Fish & Wildlife Service,<br />
Professor Gisele and the Department of Paleontology<br />
and Anthropology, University of Antananarivo for their help.<br />
This project would not have been possible without the support<br />
of the staff, guides, and drivers of Henry Doorly Zoo<br />
and the Madagascar Biodiversity Partnership. We also wish<br />
to thank the Theodore F.and Claire M.Hubbard Family Foundation,<br />
Bill and Berniece Grewcock, the Ahmanson Foundation,<br />
the James Family Foundation and the Hawks Family<br />
Foundation for their support of this project.<br />
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Brenneman,R.A.;Louis,E.E.Jr.2007b.Isolation and characterization<br />
of twenty microsatellite marker loci from the<br />
Indri (Indri indri) genome. Mol. Ecol. Notes 7: 25-28.<br />
Zaonarivelo, J.R.; Andriantompohavana, R.; Shore G.D.; Engberg,<br />
S.E.; McGuire, S.M.; Louis, E.E. Jr.; Brenneman, R.A.<br />
2006.Characterization of 21 microsatellite marker loci in<br />
the ring-tailed lemur (Lemur catta). Conserv. Genet. 8(5):<br />
1209-1212.<br />
Verreaux’s sifaka fur condition in the<br />
spiny forest of southern Androy<br />
Ivan Norscia 1*, Jean Lambotsimihampy 2, Elisabetta<br />
Palagi 1,3<br />
1Natural History Museum, UniversitB di Pisa, Via Roma, 79,<br />
56011, Calci (PI), Italy<br />
2Berenty Village, Antandroy, Madagascar<br />
3Unit of Cognitive Primatology and Primate Center,Institute<br />
of Cognitive Sciences and Technologies, The National Research<br />
Council (CNR), Rome, Italy<br />
*Corresponding author: norscia@hotmail.com<br />
Résumé<br />
Les conditions du pelage des animaux peuvent représenter<br />
un moyen fiable et un indicateur non invasif pour comprendre<br />
l’état de santé d’une population et distinguer des segments<br />
différents de la même population.En 2008 nous avons<br />
effectué un recensement de sifaka (Propithecus verreauxi)<br />
dans les forêts riveraines de la réserve de Berenty (foret<br />
galerie et de transition de Malaza et forêt secondaire aux
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 65<br />
espèces allochtones de Ankoba) et dans six portions de<br />
forêt épineuse qui sont inclues dans le domaine privé de<br />
Berenty. Nous avons relevé l’état du pelage des animaux<br />
selon trois conditions: fourrure intègre (niveau 1), fourrure<br />
faiblement endommagée (ponctuée par des petites zones,<br />
sans pelage, couvrant moins de 30 % du corps; niveau 2),<br />
fourrure fortement endommagée (pelage manquant sur une<br />
surface supérieure à 30 %;niveau 3).Nous avons aperçu seulement<br />
quatre sifaka au niveau de pelage 3 et, par conséquence,<br />
nous avons pu évaluer statistiquement les différences<br />
seulement entre les niveaux de pelage 1 et 2, en comparant<br />
forêt épineuse et les forêts riveraines, soi au niveau<br />
des groups d’animaux (n= 41) soi au niveau des zones recensée<br />
(n= 9). Même si avons détecté un nombre significativement<br />
plus haut de sifaka avec la fourrure faiblement<br />
endommagée dans le domaine épineuse,la nature et surtout<br />
l’entité du dommage indiquent que les conditions du pelage<br />
n’arrivent pas vraiment à différentier des segments distincts<br />
dans la population de sifaka de Berenty.<br />
Introduction<br />
An index of coat condition can be a non-invasive tool for<br />
tracking health and stress at the population level (Jolly,<br />
2009a). In fact, pelage growth can be directly influenced by<br />
the proximate stimulus of light (acting through neuro-endocrine<br />
pathways), by the nutritional status, and indirectly by<br />
temperature and behavior (Ling, 1970). Two main functions<br />
of fur are a) insulation, which allows conservation of body<br />
heat, thus reducing energy expenditure and food requirements;<br />
and b) shielding, which protects day-active mammals<br />
from excessive heat load from solar radiation (Scott et al.,<br />
2001; Kenagy and Pearson, 2000).<br />
Here, we considered coat condition of Propithecus verreauxi<br />
(Verreaux’s sifaka) as a possible indicator of the "health status"<br />
of animals in different habitats and investigated whether<br />
it could provide information on possible population stress in<br />
the poorly investigated spiny forest of south Madagascar.<br />
The dry spiny forest of southern Madagascar is a thorny environment,both<br />
metaphorically and literally speaking.Listed as<br />
one of the 200 most important ecological regions in the<br />
world,it harbors the highest level of plant endemism in Madagascar<br />
(Elmqvist et al., 2007). In spite of its importance, the<br />
spiny forest is underrepresented in terms of protection and<br />
conservation programs (Fenn, 2003; Ganzhorn et al., 2003;<br />
Seddon et al.,2000).To fill,at least in part,this gap,we investigated<br />
sifaka fur condition in different spiny forest parcels<br />
inside the Berenty Estate (Androy region,south Madagascar)<br />
and compared it with sifaka inhabiting the riverine forest<br />
areas inside the Berenty Reserve, a habitat much richer in<br />
staple food for lemurs.<br />
Methods<br />
Study site, survey technique, and fur condition evaluation<br />
In March-April 2008, a comprehensive sifaka survey was conducted<br />
in the Berenty Estate, covering 134 ha of spiny forest<br />
and 60 ha of non-spiny forest areas. The Berenty Estate is located<br />
in the semi-arid Androy Region (rainfall averages less<br />
than 500 mm per year). The spiny forest is usually 3 to 6 m in<br />
height with dwarf and xerophyte plants,and emerging trees of<br />
the Family Didieraceae that may reach more than 10 m in<br />
height,such as the keystone species Allouadia procera (Elmqvist<br />
et al., 2007). We performed the survey in all accessible spiny<br />
forest parcels (sacred areas, used as a cemetery, cannot be<br />
accessed by anyone except for local family clans) and in three<br />
riverine areas of the Berenty Reserve (on the Mandrare river),<br />
comprising a northern section (occupied by the 40 ha secondary<br />
forest of Ankoba dominated by the exotic legume species<br />
Pithecellobium dulce;S 24.99°;E 46.29°) and a southern section<br />
(Malaza: S 25.01°; E 46,31°) (Fig. 1 shows study locations). Inside<br />
Malaza we considered the 7 ha gallery forest (dominated<br />
by tamarinds;Tamarindus indica) and the front-transitional forest<br />
(13 ha) (Jolly et al.,2006).In all the areas considered in this<br />
study,logging and hunting are prohibited,and the fossa (Cryptoprocta<br />
ferox) isabsent.<br />
Fig.1:Study site location:Berenty reserve (solid outline;white<br />
area:scrub;diagonal lines:Ankoba and Malaza riverine forests)<br />
and spiny forest fragments (black areas): 1 = Spiny Malaza; 2 =<br />
Spiny Reserve 1;3 = Spiny Reserve 2;4 = West Rapily;5 = Fragment<br />
X; 6 = Anjapolo, about 13 km north-west of Berenty.<br />
Dashed outlines include degraded spiny and/or scrub areas.<br />
The rest of the territory (white) is covered by pasture and<br />
sisal fields. (Map based on Google Earth satellite view).<br />
We performed the survey via walking, at a speed of about 1<br />
km/h,along preexisting trails and through forest paths chosen<br />
ad hoc to have visibility of at least 50 m to the right and left (to<br />
avoid pseudoreplication we followed Norscia and Palagi,<br />
2008).<br />
During the census we evaluated the fur condition of each individual<br />
lemur.We scored coat condition on a 3-point scale:coat<br />
undamaged, with fur fully covering the body (level 1); ruffled<br />
coat,with fur punctuated by small areas of reduced/missing fur<br />
(on less than 30 % of the body, especially on elbows and/or<br />
knees;level 2);patchy coat,usually with black skin areas clearly<br />
visible due to reduced/missing fur (on more than 30 % of the<br />
body, especially on elbows/knees, external sides of forearms<br />
and thighs, fingers and toes; level 3) (Fig. 3).<br />
Statistics<br />
We performed the analyses at group or at forest site level.<br />
Owing to the small sample size (n
Page 66 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
in the spiny and two in the non-spiny forest areas). Thus, we<br />
considered only level 1 (undamaged coat) and 2 (moderately<br />
missing fur) for the analyses and found that the proportion of<br />
individuals with level-1 fur was significantly higher in the nonspiny<br />
than in the spiny areas both in the analysis per forest<br />
site (Exact Mann-Whitney U test, nnon-spiny=3, nspiny=6, Z=<br />
-2.35, p=0.024) (Fig. 2) and in the analysis per group (Mann-<br />
Whitney U test, nnon-spiny=31, nspiny=11, Z=-3.26, p=0.001).<br />
FUR FUR CONDITION<br />
CONDITION<br />
0.90<br />
0.80<br />
0.70<br />
0.60<br />
0.50<br />
0.40<br />
0.30<br />
0.20<br />
SPINY FOREST RIVERINE<br />
AREAS FOREST AREAS<br />
Fig. 2: Difference in fur condition (proportion of individuals<br />
showing level-1 fur) between the spiny and riverine forest areas.<br />
The difference is significant (p
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 67<br />
Fenn, M.D. 2003. The spiny forest ecoregion. Pp. <strong>15</strong>25-<strong>15</strong>30.<br />
In: S.M. Goodman; J.P. Benstead (eds.). The Natural History<br />
of Madagascar. University of Chicago Press, Chicago.<br />
Ganzhorn, J.U.; Goodman, S.M.; Dehgan, A. 2003. Effects of<br />
forest fragmentation on small mammals and lemurs. Pp.<br />
1228-1234. In: S.M. Goodman; J.P. Benstead (eds.). The<br />
Natural History of Madagascar. University of Chicago<br />
Press, Chicago.<br />
Jolly, A. 2009a. Coat conditions of ringtailed lemurs, Lemur<br />
catta, at Berenty Reserve, Madagascar: I. Differences by<br />
age, sex, density and tourism, 1996-2006. Am. J. Primatol.<br />
71: 191-198.<br />
Jolly, A. 2009b. Coat conditions of ringtailed lemurs, Lemur<br />
catta, at Berenty Reserve, Madagascar: II. Coat and tail<br />
alopecia associated with Leucaena leucocephala. Am. J.<br />
Primatol. 71: 199-205.<br />
Jolly, A; Koyama, N; Rasamimanana, H; Crowley, H; Williams,<br />
G.2006.Berenty Reserve:a research site in southern Madagascar.Pp.32-42.In:A.Jolly,R.W.Sussman,N.Koyama,H.<br />
Rasamimanana (eds.). Ringtailed Lemur Biology: Lemur<br />
catta in Madagascar. Springer Verlag, New York.<br />
Kenagy, G.J.; Pearson, O.P. 2000. Life with fur and without:<br />
experimental field energetics and survival of naked meadow<br />
voles. Oecologia 122: 220-224.<br />
Ling, J.K. 1970. Pelage and molting in wild mammals with<br />
special reference to aquatic forms.Q.Rev.Biol.45:16-54.<br />
Mundry, R; Fischer, J. 1998. Use of statistical programs for<br />
nonparametric tests of small samples often leads to<br />
incorrect P values: examples from Animal Behaviour.<br />
Anim. Behav. 56: 256-259.<br />
Norscia, I.; Palagi, E. 2010. Fragment quality and distribution<br />
of the arboreal primate Propithecus verreauxi in the spiny<br />
forest of south Madagascar. J. Trop. Ecol. DOI:<br />
10.1017/S0266467410000519.<br />
Norscia,I.;Palagi E.2008.Berenty 2006:census of Propithecus<br />
verreauxi and possible evidence of population stress. Int. J.<br />
Primatol. 29: 1099-11<strong>15</strong>.<br />
Scott, D.W.; Miller, W.H.; Griffin, C.E. 2001. Structure and<br />
function of skin.Muller and Kirk’s Small Animal Dermatology,<br />
6th ed. W.B. Saunders, Philadelphia.<br />
Seddon, N.; Tobias, J; Yount, J; Ramanampamonjy, J.M.;<br />
Butchart,S;Randrianizahana H.2000.Conservation issues<br />
and priorities in the Mikea forest of south-western Madagascar.<br />
Oryx 34: 287-304.<br />
Siegel,S.;Castellan,N.J.Jr.1988.Nonparametric Statistics for<br />
the Behavioral Sciences, Second edition. MacGraw-Hill,<br />
New York.<br />
Rediscovery of Sibree’s dwarf lemur in<br />
the fragmented forests of Tsinjoarivo,<br />
central-eastern Madagascar<br />
Marina B. Blanco<br />
Department of Anthropology, University of Massachusetts,<br />
240 Hicks Way, Amherst, MA 01003, USA,<br />
mbblanco@anthro.umass.edu<br />
The recent genetic confirmation of a rare dwarf lemur<br />
species, C. sibreei, at Tsinjoarivo is bitter-sweet. The excitement<br />
of reporting the first known living population of this<br />
species is tainted by conservation concerns, as the forest<br />
fragment in which Sibree’s dwarf lemurs were captured is<br />
highly disturbed and targeted for illicit logging. This species,<br />
like many others inhabiting rapidly degrading forests, faces<br />
the serious threat of extinction.<br />
Taxonomic background of the genus, first field discovery,<br />
and subsequent recognition of C. sibreei<br />
During the 19th century,the small nocturnal lemurs of Madagascar<br />
were clumped in a chaotic array of species and genera.<br />
For most of the 20th century,however,dwarf lemurs (Cheirogaleus)<br />
were classified in only two species: the eastern C.<br />
major and the western C.medius (Schwarz,1931).Around the<br />
turn of the century, Groves (2000) conducted a taxonomic<br />
revision of the genus on the basis of morphological analysis<br />
of museum specimens and increased the species number to<br />
seven: C.medius,C.adipicaudatus,C.major,C.ravus,C.crossleyi,<br />
C. minusculus and C. sibreei. This last species, in fact, had been<br />
originally described by the Swiss naturalist Forsyth Major in<br />
1896 during one of his expeditions to Madagascar (Forsyth<br />
Major, 1896). He had named it Chirogale sibreei in honor of<br />
fellow naturalist James Sibree,who had spent more than fifty<br />
years in Madagascar and had written extensively about its<br />
people, fauna, flora and geology. Forsyth Major published<br />
measurements of an individual "obtained from the neighbourhood<br />
of Ankeramadinika," a locality vaguely described<br />
by its discoverer as "one day’s journey to the east of Antananarivo",but<br />
in fact a well-known village at the time,located in<br />
the central high plateau on the road that connected Antananarivo<br />
to Mahatsara on the east coast (Capitaine "X",<br />
1901). In his taxonomic revision, Groves (2000) included as<br />
Cheirogaleus sibreei not only the holotype from Ankeramadinika<br />
(currently housed at the Natural History Museum in<br />
London) but also three additional specimens (3 skins and 1<br />
skull), two of which came from Ampasindava, northwestern<br />
Madagascar, and one from an unclear provenance (Imerina,<br />
which refers to a region of the central highlands around<br />
Antananarivo).<br />
The taxonomic shrinkage of Cheirogaleus<br />
The increase in the number of species within the genus<br />
Cheirogaleus was not surprising because dwarf lemurs occupy<br />
a wide variety of habitats in Madagascar,and their close relatives,the<br />
mouse lemurs (Microcebus), had undergone a taxonomic<br />
explosion of their own with more than 10 species<br />
described during the past <strong>15</strong> years (Louis et al.,2008;Olivieri<br />
et al., 2007; Radespiel et al., 2008). However, Groves’ 2000<br />
revision of dwarf lemur taxonomy did not escape criticism,<br />
not least of which had to do with the criteria that he used to<br />
define species, the lack of reliable locality information from<br />
museum specimens, and the absence of on-the-ground surveys<br />
to assess geographic boundaries and variation among<br />
species (Blanco et al., 2009; Tattersall, 2007). A recent and<br />
more comprehensive revision of dwarf lemur taxonomy was<br />
carried out by Groeneveld and colleagues, who compiled<br />
genetic and morphometric data from field as well as museum<br />
specimens from a variety of localities across Madagascar,<br />
including some of the specimens studied by Groves (Groeneveld<br />
et al., 2009; 2011). This research showed overall consistency<br />
between morphological and genetic data in recognizing<br />
only three Cheirogaleus species: C. medius, C. major<br />
and C. crossleyi. Individuals that previously had been assigned<br />
to C. adipicaudatus fell within the C. medius clade, and those<br />
named as C.ravus grouped with C.major.Results were inconclusive<br />
for C.minusculus and C.sibreei because holotype specimens<br />
were not available for sampling and their genetic affiliation<br />
could not be determined. Genetic data from one of the<br />
C. sibreei museum specimens from Ampasindava linked this<br />
specimen to C. medius. Nevertheless, the C. sibreei holotype<br />
from Ankeramadinika was larger and did not group morphologically<br />
with other C. medius. This suggested that the individuals<br />
from Ampasindava may have been misclassified by<br />
Groves as C. sibreei (Groeneveld et al., 2010). The status of<br />
this species remained equivocal.<br />
Second field discovery of C. sibreei, at last<br />
The story of a dwarf lemur named "May" told by Mitchell<br />
Irwin (2002) turned out to be rather prophetic. Irwin’s<br />
research team rescued this dwarf lemur badly burned in a
Page 68 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
human-induced fire nearby Andasivodihazo,one of the forest<br />
fragments at Tsinjoarivo (Fig. 1). At the time, "May" was believed<br />
to be, as were all eastern rainforest dwarf lemurs, C.<br />
major. Unfortunately, this female could not fully recover and<br />
died soon after the salvage, but her skeletal remains were<br />
carefully preserved and stored at the University of Antananarivo<br />
by Irwin’s team. Seven years later this specimen came<br />
to play a key role in our morphological analysis of dwarf<br />
lemurs at Tsinjoarivo.<br />
Fig.1: Map showing Tsinjoarivo and other localities associated<br />
with Sibree’s dwarf lemurs; see text for details.<br />
In 2006, with the logistic help of Mitchell Irwin and Jean-Luc<br />
Raharison, I began a survey of nocturnal lemurs at Tsinjoarivo.<br />
My assistants and I successfully trapped dwarf lemurs<br />
at two study sites: in one of the forest fragments (Andasivodihazo,<br />
19º41’<strong>15</strong>"S, 47º46’25"E, 1660 m) and within continuous<br />
forest (Vatateza, 19º43’<strong>15</strong>"S, 47º51’25"E, 1396 m)<br />
(Blanco et al., 2009). Even to an inexperienced eye, fragment<br />
dwarf lemurs looked different from continuous forest individuals,<br />
in that they were overall smaller, with grayer fur,<br />
marked eye rings and significantly larger female genitalia (Fig.<br />
2). Our morphological and dental analyses determined that<br />
of all the species described by Groves,C.sibreei was the most<br />
similar to forest fragment dwarf lemurs (Blanco et al., 2009).<br />
(Hopefully, sampling of C.sibreei’s holotype will be allowed in<br />
the near future to definitely determine whether or not there<br />
is a genetic match between this specimen and fragment<br />
dwarf lemurs from Tsinjoarivo.) Recent genetic analyses<br />
have confirmed not only that dwarf lemurs from Andasivodihazo<br />
constitute a different clade (and therefore an independent<br />
phylogenetic lineage), but also that the fragment<br />
dwarf lemur species had branched off first and was ancestral<br />
to the other dwarf lemur species (Groeneveld et al., 2010).<br />
To date, no other living population of C. sibreei has been<br />
reported in the wild and more intensive surveys around the<br />
Tsinjoarivo area (including possibly remaining forests nearby<br />
Ankeramadinika, ~100 km from Tsinjoarivo) are warranted<br />
to assess geographic boundaries and population density.<br />
Fig.2: Sibree’s dwarf lemur captured at Andasivodihazo, one<br />
of the forest fragments at Tsinjoarivo.<br />
Conservation concerns<br />
The genetic confirmation of three clades corresponding to C.<br />
medius, C. major and C. crossleyi, each of which has broad<br />
geographic distributions, implied that dwarf lemurs might be<br />
less threatened than previously thought (Groeneveld et al.,<br />
2009). However, the situation for C. sibreei is radically different.<br />
Tsinjoarivo’s unique geographic setting, continuous<br />
with the central plateau on the west and the steep escarpment<br />
of rainforest in the east, may harbor a unique array of<br />
animal communities. To date, C. sibreei has been captured in<br />
sympatry with C. crossleyi at one forest fragment, Andasivodihazo,and<br />
at one intermediate location,Ankadivory.Both of<br />
these areas are subjected to illicit logging and heavy deforestation<br />
(Fig.3).Furthermore,these forest sites are located towards<br />
Tsinjoarivo’s western boundary which reaches some<br />
of the highest altitudes (up to ~1650 m) known in eastern<br />
Fig. 3: Example of logging near Ankadivory, one of the forest<br />
sites where Sibree’s dwarf lemurs were captured.
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 69<br />
rainforests in Madagascar.So far,only C.crossleyi has been captured<br />
at Vatateza,a lower altitude site located within continuous<br />
and less disturbed forest.Virtually everything has to be<br />
learned about the "ancestral" C.sibreei,including how to prevent<br />
its extinction through habitat loss.Concerted efforts by<br />
organizations such as Sadabe (www.sadabe.org) are instrumental<br />
in raising awareness of endangered species and promoting<br />
research and educational programs in the Tsinjoarivo<br />
area.<br />
Acknowledgements<br />
I am particularly thankful to the research team in Madagascar:<br />
Malagasy students <strong>Vol</strong>olonirina Rahalinarivo and Mamihasimbola<br />
Rakotondratsima, and local assistants Noel Rakotoniaina,<br />
Edmond and Nirina Razanadrakoto. Additional<br />
thanks to Jean-Luc Raharison and Mitchell Irwin for providing<br />
logistical support and assistance in the field. I am also<br />
grateful to Laurie Godfrey, Stacy Gebo and Christoph<br />
Schwitzer for their comments. This research was supported<br />
by funds from the Rufford Foundation, MMBF/Conservation<br />
International Primate Action Fund and Primate Conservation<br />
Inc. I would like to thank the Ministère de l’Environnement<br />
et des forêts of the Malagasy government and the University<br />
of Antananarivo for permission to conduct this research.The<br />
project in Madagascar was facilitated by the Institute<br />
for the Conservation of Tropical Environments (ICTE,<br />
Patricia C. Wright) and the Madagascar Institute pour la<br />
Conservation des Ecosystèmes Tropicaux (MICET), especially<br />
Benjamin Andriamihaja.<br />
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B.; Ratsimbazafy, J. H.; Ratelolahy, F.; Randriamboavonjy,<br />
T.; Rasolofoharivelo, T.; Craul, M.; Rakotozafy, L.;<br />
Randrianarison,R.M.2008.Exceptional diversity of mouse<br />
lemurs (Microcebus spp.) in the Makira region with the<br />
description of one new species. Am. J. Primatol. 70: 1033-<br />
1046.<br />
Schwarz,E.1931.A revision of the genera and species of Madagascar<br />
Lemuridae.Proc.Zool.Soc.Lond.1931:399-426.<br />
Tattersall, I. 2007. Madagascar’s lemurs: Cryptic diversity or<br />
taxonomic inflation? Evol. Anthropol. 16: 12-23.<br />
Funding and Training<br />
AEECL Small Grants<br />
Since 2009,AEECL awards two small grants of up to € 1,000<br />
each year to graduate students, qualified conservationists<br />
and/or researchers to study lemurs in their natural habitat.<br />
Priority is given to proposals covering conservation-relevant<br />
research on those species red-listed as Vulnerable, Endangered,<br />
Critically Endangered or Data Deficient by the IUCN.<br />
We support original research<br />
that helps with establishing<br />
conservation action plans for<br />
the studied species. Grants<br />
are normally given to recent<br />
graduates from Malagasy universities to help building local<br />
capacity.<br />
We may also, in special circumstances, support studies on<br />
Malagasy species other than lemurs if the proposal provides<br />
satisfactory information as to how lemurs or the respective<br />
habitat/ecosystem as a whole will benefit from the research.<br />
All proposals will be assessed by the Board of Directors of<br />
AEECL and/or by external referees.The deadline for applications<br />
is February <strong>15</strong>th of each year. Successful applicants will<br />
be notified by June 1st.More information can be found on the<br />
AEECL website, www.aeecl.org.<br />
The Mohamed bin Zayed Species Conservation<br />
Fund<br />
Announced at the World Conservation Congress in Barcelona<br />
in 2008,The Mohamed bin Zayed Species Conservation<br />
Fund is a significant philanthropic endowment established to<br />
do the following:<br />
Provide targeted grants to individual species conservation<br />
initiatives;<br />
Recognize leaders in the field of species conservation;and Elevate<br />
the importance of species in the broader conservation<br />
debate.
Page 70 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
The fund’s reach is truly global,and its species interest is nondiscriminatory.<br />
It is open to applications for funding support<br />
from conservationists based in all parts of the world,and will<br />
potentially support projects focused on any and all kinds of<br />
plant and animal species, subject to the approval of an independent<br />
evaluation committee.<br />
Details on this important new source for species conservation<br />
initiatives and research can be found at<br />
www.mbzspeciesconservation.org<br />
CI Primate Action Fund<br />
The principal objective of Conservation International’s Primate<br />
Action Fund is to contribute to global biodiversity<br />
conservation by providing strategically targeted, catalytic<br />
support for the conservation of endangered nonhuman<br />
primates and their natural habitats.<br />
Projects submitted to the foundation should have one or<br />
more of the following characteristics:<br />
A focus on critically endangered and endangered nonhuman<br />
primates (and most especially those included in the<br />
biennial listing of the World’s 25 Most Endangered Primates)<br />
living in their natural habitats;<br />
Location in areas of high overall biodiversity and under great<br />
threat (e.g., "threatened hotspots", "megadiversity" countries)<br />
- to ensure maximum multiplier effect for each project;<br />
Direction and management by nationals from the tropical<br />
countries,to help increase local capacity for implementing<br />
biodiversity conservation;<br />
The ability to strengthen international networks of fieldbased<br />
primate specialists and enhance their capacity to be<br />
successful conservationists; and<br />
Projects that result in publication of information on endangered<br />
primate species in a format that is useful both to<br />
experts and the general public.<br />
Applications for support are considered throughout the year<br />
with no deadlines for submittal. Proposals should be sent by<br />
electronic mail to:<br />
Anthony B. Rylands, Primate Action Fund, Conservation International,<br />
2011 Crystal Drive, Suite 500, Arlington, VA<br />
22202, USA, a.rylands@conservation.org<br />
Recent Publications<br />
Lemurs of Madagascar, 3rd edition, by Russell A. Mittermeier,<br />
Edward E. Louis Jr., Matthew Richardson, Christoph<br />
Schwitzer,Olivier Langrand,Anthony B.Rylands,Frank Hawkins,<br />
Serge Rajaobelina, Jonah Ratsimbazafy, Rodin Rasoloarison,<br />
Christian Roos, Peter M. Kappeler and James Mackinnon.Illustrated<br />
by Stephen D.Nash.Conservation International,<br />
Tropical Field Guide Series, Arlington, VA, 2010. 762<br />
pp. ISBN: 978-1-934<strong>15</strong>1-23-5. US$55.00.<br />
In 2006,Madagascar was making significant progress towards<br />
conservation by expanding the protection of its natural treasures.At<br />
the same time,the second edition of Conservation<br />
International’s Tropical Field Guide Series, Lemurs of Madagascar,had<br />
just come off the press,a full twelve years after its<br />
much-celebrated predecessor. A lot has changed in four<br />
years. Political and economic instability has imperiled both<br />
the Malagasy people and their unique wildlife. Conservation<br />
has taken drastic steps backwards as the desperation of the<br />
masses and greed of a few elites and international profiteers<br />
has exacerbated the conflict between the domains of humans<br />
and wildlife. CI has answered the call to action by releasing<br />
a new third edition of its lemur field guide, dwarfing<br />
previous editions in both size and its depth of research and<br />
detail. With nearly 1,100 references to support it–up from<br />
approximately 500 references in the second edition–the<br />
third edition stands as more than just a complete compendium<br />
of our knowledge about lemurs, but the perfect guide<br />
for appreciating the history, diversity, uniqueness, and pure<br />
beauty of our strepsirrhine cousins.<br />
According to CI’s Jill Lucena,from early 2009,the 13 authors<br />
and dozens of contributors have worked tirelessly on the<br />
third edition,dedicating thousands of hours towards its production.<br />
Authors Matthew<br />
Richardson and Anthony B.<br />
Rylands, as well as illustrator<br />
Stephen D. Nash and<br />
graphic designer Paula K.<br />
Rylands,labored exclusively<br />
on the project for nine<br />
months. The end result is a<br />
field guide that will leave<br />
other academic fields envious!<br />
This new volume is 247<br />
pages longer than the previous<br />
edition, with 767 pages<br />
of carefully organized maps,<br />
photos, and colorful illustrations,<br />
in addition to all<br />
the details lemur enthusiasts<br />
and researchers have<br />
come to expect from this<br />
book.The content is so rich<br />
that the book’s dimensions<br />
have increased from 7.5" x<br />
4.5" x 1" to slightly more than 9.25" x 6.25" x 1.25" just to accommodate<br />
everything.And herein lies what may be the only<br />
problem with this new edition. Reviews of the previous two<br />
editions had lavished praise for not only the content and<br />
scope, but also the portability of the books. Although it will<br />
fit comfortably in a backpack, it will add more weight and<br />
consume more space than its predecessors. But given the<br />
content, that may be a small price to pay.<br />
The layout of the book has not changed much since the last<br />
edition. An enthralling chapter on Madagascar’s ancient geological<br />
history has been added, providing tantalizing details<br />
about the mysteries of Madagascar’s ancient past,while a few<br />
familiar chapters and appendices have been reordered. Each<br />
lemur family has now been assigned its own chapter.The section<br />
entitled "How to Use This Field Guide" still walks new<br />
readers through the layout of the book. The "Quick Visual<br />
Reference" and colored tabs facilitate speedy navigation and<br />
help to satiate an ecotourists’ spontaneous hunger for specific<br />
information. The "Lemur Life-list" returns in a more<br />
readable table format to help ecotourists record their first<br />
sightings of the numerous lemur species. Even the maps of<br />
the island have been revamped and are easier to read in this<br />
larger format.<br />
Introductory chapters discuss ancient geology,lemur origins,<br />
the extinct subfossil (giant) lemurs, the history of lemur re-
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 71<br />
search, and lemur conservation. Each chapter contains significant<br />
updates from the second edition. The ancient geology<br />
of Madagascar is covered in meticulous detail, while the theories<br />
of lemur origins are explored in depth, leaving both<br />
reader and researcher alike desperate for more definitive<br />
answers.The chapter on the extinct subfossil lemurs is beautifully<br />
illustrated with new peer-reviewed artwork from<br />
award-winning illustrator Stephen D. Nash, and has new details<br />
about their biology and extinction.The history of lemur<br />
research and discovery expands greatly upon the work from<br />
the last half century–a topic greatly underrepresented in the<br />
previous edition. Additionally, the chapter is loaded with<br />
newly added artwork from the 1700s and 1800s. The lemur<br />
conservation chapter provides a critical update on the newest<br />
emerging threats faced by lemurs and their habitat,<br />
namely the logging of precious hardwoods and bushmeat<br />
hunting. Additional detail is also provided about other important<br />
threats that received little mention previously, such<br />
as invasive species, cattle-raising, and mining.<br />
The bulk of the book details the description, geographic<br />
range,natural history,conservation status,and best locations<br />
to observe each of the 101 species and subspecies of lemur.<br />
(This total is up from the 71 taxa detailed in the second edition,yet<br />
the authors note that upcoming research may reveal<br />
as many as 110 to 125 lemur taxa!) Each species section<br />
sports a portrait photo, detailed range map, and other photos<br />
to enrich the lavish textual content.Once again,the third<br />
edition sets itself apart from the previous editions with its<br />
encyclopedic coverage of details from the lemur research literature.Species,such<br />
as the silky sifaka (Propithecus candidus),<br />
are discussed in significantly greater detail,bringing everyone<br />
from bright-eyed ecotourists to veteran lemur researcher<br />
up to speed on the latest findings.<br />
The second edition of Lemurs of Madagascar sold out quickly,<br />
leaving shelves empty as early as February 2008. The third<br />
edition is poised to do the same. With a print run of 10,000<br />
copies,nearly two-thirds are already spoken for according to<br />
Jill Lucena.To help promote conservation education in Madagascar,<br />
CI is generously donating 3,000 copies to its partner<br />
in the field,NGO Fanamby.CI hopes that the remaining copies<br />
will spark the public’s interest in Madagascar’s ecological<br />
gems and spur a new wave of ecotourism to bolster conservation<br />
efforts.<br />
Once again, CI has provided an invaluable tool for a diverse<br />
audience, which includes ecotourists, Malagasy tour guides,<br />
students,lemur researchers,and lemur enthusiasts.Although<br />
larger and not quite as portable as its predecessors, the increased<br />
size of the third edition hosts a wealth of enhanced<br />
encyclopedic detail,new and stunning artwork by Stephen D.<br />
Nash, and additional color photos and illustrations. With a<br />
copy of this printing in hand, the only things missing are a<br />
backpack full of supplies and an airline ticket to Madagascar.<br />
So what are you waiting for?<br />
Alex Dunkel<br />
Theses Completed<br />
Blanco,M.B.2010.Reproductive biology of mouse and dwarf<br />
lemurs of eastern Madagascar,with an emphasis on brown<br />
mouse lemurs (Microcebus rufus) at Ranomafana National<br />
Park,a southeastern rainforest.PhD Dissertation.University<br />
of Massachusetts, Amherst.<br />
This dissertation investigates reproductive schedules of<br />
brown mouse lemurs at Ranomafana,using intensive trap-<br />
ping techniques. The reproductive condition of female<br />
mouse lemurs was recorded on the basis of vaginal morphology,vaginal<br />
smears,body mass gain profiles and nipple<br />
development. Testis size was measured in males throughout<br />
the reproductive season. The timing of the first seasonal<br />
estrus was determined in frequently captured females<br />
over multiple years and it showed individual periodicities<br />
close to 365 days,consistent with endogenous regulation<br />
and entrainment by photoperiod. The timing of<br />
estrus did not correlate with female age or body mass.<br />
Males showed testicular regression during the rainy season,<br />
although there was high inter-individual variation in<br />
testes size at any given point during the reproductive season.<br />
Furthermore, some individuals completed testicular<br />
regression earlier than others. Implications for polyestry<br />
are discussed.<br />
For comparative purposes,mouse lemurs were also trapped<br />
at two study sites in the Tsinjoarivo area: one in a forest<br />
fragment and the other within continuous forest.<br />
These forests are higher in altitude than the main study<br />
area at Ranomafana. Trapping success for mouse lemurs<br />
was lower at Tsinjoarivo than Ranomafana.Albeit preliminary,<br />
data from Tsinjoarivo suggest that females have lower<br />
reproductive success than do females at Ranomafana.<br />
Nevertheless, mouse lemurs in the Tsinjoarivo forest<br />
fragment did not appear to be in "poorer" condition than<br />
those in the continuous forest.It had been reported in the<br />
literature that western gray mouse lemurs captured in secondary<br />
forests have lower body masses and lower recapture<br />
rates than those captured in primary forest; in fact,<br />
the opposite was true of the mouse lemurs at Tsinjoarivo.<br />
I additionally collected data on a larger member of the<br />
family Cheirogaleidae, the dwarf lemurs (Cheirogaleus),<br />
which live in sympatry with Microcebus at Ranomafana and<br />
Tsinjoarivo. I analyzed the patterns of growth, development<br />
and reproduction in Cheirogaleus and Microcebus and<br />
compared dwarf and mouse lemurs to other similarlysized<br />
prosimians which do not undergo torpor or hibernation.These<br />
comparisons draw attention to the unusual<br />
reproductive and metabolic strategies employed by cheirogaleids<br />
to cope with Madagascar’s unpredictable environments,<br />
which ultimately define their very unique life<br />
histories.<br />
Key words: Cheirogaleus, Madagascar, Microcebus, Mouse<br />
lemurs, Rainforest, Ranomafana, Reproduction, Tsinjoarivo.<br />
Bonaventure, R.T.A.R. 2010. Ecologie et comportement de<br />
Propithecus verrreauxi dans les zones d’extension de la<br />
Réserve Spéciale de Bezà Mahafaly.Engineer in agronomy,<br />
option Eaux et forêts.Eaux et forêts,Ecole Supérieure des<br />
Sciences Agronomiques de l’Université d’Antananarivo<br />
(ESSA), Madagascar.<br />
The population of Propithecus verreauxi in the special<br />
reserve of Bezà Mahafaly is one of the conservation targets<br />
of the site which is the subject of a long-term follow.<br />
Natural destruction of their habitat and the pressure of<br />
hunting which is exerted on this species outside of the<br />
current reserve are the origin of the sifakas’ decline.With<br />
the park extension project under way, furthering the<br />
knowledge on behavior and ecology of Propithecus verreauxi<br />
in disturbed areas outside the current reserve is essential<br />
for decision-making regarding conservation measures<br />
for this species. Thus,a study of behavior and ecology<br />
of the sifakas was carried out in the extension area of the<br />
special reserve of Bezà Mahafaly in gallery and transition<br />
forest at the end of the dry period. The study was centered<br />
on 3 focal groups including one in the gallery forest<br />
and two in the transition forest. The method of focal animal<br />
sampling was chosen to study their behavior.A floristic<br />
inventory according to the Gentry method, which<br />
includes transects of 2 x 50 m, was carried out to study<br />
the habitat. On the whole, 120h of observations of sifaka<br />
behavior were carried out and 12 transects were walked.<br />
In disturbed areas, the sifakas still consumed preferred<br />
plants.This resulted in a high intake of 2 or 3 easily digestible<br />
plant species while at the same time a large variety of<br />
other species was consumed.Thus,the disturbance of the<br />
sites did not influence food intake of the sifakas.Moreover,
Page 72 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
it was the group of sifakas in the gallery forest which was<br />
the most affected by resource scarcity in the dry period,<br />
necessitating a priorisation of the sifaka groups in this forest<br />
formation in the extension of the reserve.In addition,<br />
the availability of nutritional resources influenced spacial<br />
dispersion of group members as well as group size. With<br />
the current park extension project different access restrictions<br />
will be implemented, having as an objective the<br />
conservation of the sifaka groups as well as other conservation<br />
targets outside of the current reserve,while introducing<br />
a range of alternative income-generating activities<br />
and sustainable resource management practices for the<br />
local communities.<br />
Key words: Lemur, Propithecus verreauxi,Ecology, Behavior,<br />
Conservation, Special reserve, BezB Mahafaly, Madagascar.<br />
Delmore,K.2009.Maintenance of stability in the Andringitra<br />
brown lemur hybrid zone.M.A.thesis,University of Calgary,<br />
Calgary, Alberta, Canada.<br />
Two models of hybrid zone stability have been proposed:<br />
1) the tension zone model, which predicts that intrinsic<br />
selection acts against hybrids but is counteracted by dispersal<br />
of parentals into the zone and 2) the bounded superiority<br />
model,which predicts that exogenous selection<br />
favours hybrids within transitional habitats and parentals<br />
outside the zone.I used morphological,genetic and ecological<br />
data to evaluate these models in a stable hybrid zone<br />
between Eulemur rufifrons and E.cinereiceps in southeastern<br />
Madagascar. This zone appears to conform to the<br />
bounded superiority model: it was relatively wide and<br />
composed mostly of hybrids that were equally as fit as<br />
parentals.Gene flow between parental and hybrid populations<br />
was also limited, clines in multiple characters were<br />
non-coincident and significant ecological correlates were<br />
identified. Results suggest that hybridization can serve as<br />
an important evolutionary force and need not always be<br />
considered a conservation risk for endangered taxa.<br />
Hobinjatovo, T. 2009. Etude morphométrique et génétique<br />
de conservation d’Eulemur cinereiceps (Milne-Edwards et<br />
Grandidier, 1890) dans les forêts de Mahabo, de Manombo<br />
et de Vevembe, Madagascar. Mémoire de DEA en Biologie,Ecologie<br />
et Conservation Animale,Département de<br />
Biologie Animale,Faculté des Sciences,Université d’Antananarivo.<br />
Cette étude a pour but, d’une part, de connaître et de<br />
comparer la morphométrie d'’Eulemur cinereiceps de la forêt<br />
de Mahabo,de Manombo et de Vevembe - Madagascar,<br />
ainsi donc de voir le degré du dimorphisme sexuel et<br />
d’autre part, d’établir des données de bases génétiques<br />
pour avoir de plus amples informations en vue de la conservation<br />
de cette espèce en danger critique.La descente<br />
sur le terrain et la collecte des données ont été effectuées<br />
pendant deux semaines du mois de mai 2006 et le travail<br />
au laboratoire pour l’étude génétique a été fait du mois de<br />
Février au mois de Mai 2007.Ce lémurien pése en moyenne<br />
2,04 kg avec une longueur moyenne de la couronne de<br />
la tête de 10,84 cm,celle du corps de 33,31 cm et est doté<br />
d’une queue de 50,44 cm. La comparaison de la morphométrie<br />
des individus des sites d’étude, par le test U de<br />
Mann-Whitney montre que le Lémurien à Collier Blanc<br />
de Vevembe est de plus grande taille que ceux des 2 autres<br />
sites.La différence de taille entre le mâle et la femelle<br />
n’est pas significative. Chez cette espèce, le dimorphisme<br />
sexuel est très marqué concernant la couleur du pelage et<br />
la dimension de la canine supérieure,considérable chez le<br />
mâle.Le testicule droit du mâle est plus long,large et volumineux<br />
que le gauche.14 marqueurs génétiques polymorphiques<br />
ont été sélectionnés pour effectuer le génotypage<br />
par l’utilisation de la technologie moléculaire et à l’aide<br />
de la réaction en chaîne polymérasique. Les programmes<br />
de Cervus, GenePop, Fstat, Structure et Bottleneck ont<br />
été utilisés pour déterminer la structure et la nature génétique<br />
de la population d’Eulemur cinereiceps. Ces analyses<br />
ont permis de déterminer que la valeur de la diversité<br />
des gènes des populations est pareille et modérée, oscillant<br />
autour de 57 % ;la divergence génétique est comprise<br />
entre 0,05 à 0,<strong>15</strong> et est qualifiée de modérée; la richesse<br />
allélique varie de 2,928 à 3,632;une migration entre la population<br />
de Mahabo et de Vevembe,d’une part et celle de<br />
Manombo et de Vevembe d’autre part,a été identifiée;un<br />
certain degré de consanguinité a été constaté à Mahabo.<br />
La population de Manombo subit un goulot démographique<br />
et aucune structure distincte ni sous - structure n’a<br />
été observée au sein des populations de ces 3 sites.Même<br />
si cette espèce est en danger critique, sa santé génétique<br />
est modérée.Elle pourrait être bonne si les solutions adéquates<br />
sur la conservation génétique étaient appliquées.<br />
Dans le cas contraire,elle deviendrait désastreuse.La prise<br />
immédiate de mesures de conservation efficientes est<br />
donc nécessaire afin de préserver les populations pures<br />
d’E. cinereiceps et de protéger ses habitats.<br />
Mots-clés: Eulemur cinericeps, Mahabo, Manombo, Vevembe,<br />
Madagascar, Morphométrie, Mensuration, Génétique,<br />
Population, Conservation.<br />
Ingraldi, C. 2010. Forest fragmentation and edge effects on<br />
eight sympatric lemur species in southeast Madagascar.<br />
M.A. thesis, University of Calgary, Calgary, Alberta, Canada.<br />
Extensive slash-and-burn agriculture in southeastern Madagascar<br />
has led to the fragmentation of forests in this region,<br />
creating a constricted available habitat area and increasing<br />
the proportion of forest edge. I investigated the<br />
response to forest fragmentation and edge effects in eight<br />
lemur species through comparisons of species density<br />
and diversity between fragments, as well as correlation<br />
analyses including population distribution patterns,ecological<br />
variables,and distance from forest edge.I also include<br />
a more detailed focus on the behavioural response of Eulemur<br />
cinereiceps. Results were highly varied, with no species<br />
showing strong aversion to forest edge but with higher<br />
overall densities in larger,more connected fragments.<br />
Eulemur cinereiceps spent significantly more time near the<br />
forest edge while resting, but edge did not affect feeding<br />
patterns or food availability. These results suggest that<br />
conservation management should focus on maintaining<br />
large, complex fragments and improving connectivity<br />
through forest corridors.<br />
Mihaminekena,T.H.2010. Etude de la relation entre la dégradation<br />
de l’habitat et les activités de Propithecus edwardsi<br />
du Parc National de Ranomafana Ifanadiana, Madagascar.<br />
Mémoire de DEA en Paléontologie et évolution biologique,<br />
Biologie Evolutive, Primatologie, Département de<br />
Paléontologie et d’Anthropologie biologique, Faculté des<br />
Sciences, Université d’Antananarivo.<br />
Une étude sur le comportement et l’habitat de Propithecus<br />
edwardsi a été réalisée dans le Parc National de Ranomafana.Elle<br />
a été réalisée dans trois sites ayant chacun un<br />
degré de perturbation inégal: Talatakely (fortement perturbé);<br />
Sakaroa (moyennement perturbé) et Valohoaka<br />
(non perturbé). Sa finalité est d’analyser le type de comportement<br />
biologique adopté par l’espèce en réponse à la<br />
dégradation de son habitat et de le comparer entre les<br />
trois sites. La méthode adoptée est celle décrite par Altmann<br />
en 1974 qui consiste à déterminer l’activité du focal<br />
animal toutes les dix minutes et celle de tous les groupes<br />
toutes les cinq minutes. Pour toutes les activités, sauf le<br />
déplacement, la différence est toujours significative pour<br />
les trois sites. Le repos est plus élevé dans le site intact<br />
(36.6%) par rapport à l’alimentation.Inversement le repos<br />
est moins fréquent (28.9 %) que l’alimentation (53.6 %)<br />
dans le site fortement perturbé. La fréquence des activités<br />
de l’espèce dans le site moyennement perturbé est<br />
toujours comprise entre les deux sites perturbés et non<br />
perturbés. Pour ses activités, l’espèce utilise certains niveaux<br />
de strates, spécialement ceux compris entre 10 et<br />
<strong>15</strong> et <strong>15</strong> et 20 m. Néanmoins, l’espèce habitant le site intact<br />
se place à un niveau plus haut que celle de la forêt perturbée.<br />
Les grands arbres sont plus abondants à Valohoaka<br />
qu’à Sakaroa et à Talatakely:respectivement la hauteur<br />
varie de 10.80, 9.53 et 9.47 m; celui du DHP est de 13.59<br />
cm;12.17 et 11.09 cm.Les épaisseurs de la couronne sont<br />
respectivement 4.42,4.28 et 3.91 m.Les parties de plantes
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 73<br />
consommées pour chaque site sont significativement très<br />
différentes. La corrélation entre la consommation de jeunes<br />
feuilles et les activités exercées est toujours positive et significative<br />
quel que soit le site. Les femelles choisissent un niveau<br />
plus haut des arbres que les mâles à Valohoaka et à Talatakely<br />
par contre à Sakaroa c’est l’inverse. La cohésion du<br />
groupe est plus observée dans le site intact par rapport à celui<br />
dégradé.Bref,la perturbation influe les activités générales<br />
et la structure de l’habitat de Propithecus edwardsi.<br />
Mots-clés:Propithecus edwardsi, Lémuriens, Degré de perturbation,<br />
Habitat, Activités, Parc National de Ranomafana,<br />
Madagascar.<br />
Polowinsky, S.Y. 2009. Nutrition of captive Sclater’s lemurs<br />
(Eulemur macaco flavifrons GRAY, 1867) and crowned<br />
lemurs (Eulemur coronatus GRAY,1842),with special emphasis<br />
on the problem of obesity. PhD dissertation, Biology<br />
and Geography, University of Duisburg-Essen, Germany.<br />
This study was concerned with the obesity problem of blueeyed<br />
black lemurs and crowned lemurs in captivity. Its aims<br />
were to optimize the species’ diet in captivity by combining<br />
data obtained from individuals kept at different European<br />
zoos as well as from wild blue-eyed black lemurs to gain a<br />
better understanding of the ecological and nutritional needs<br />
of Eulemur macaco flavifrons in order to prevent individuals<br />
from becoming obese and to assist planned conservation<br />
measures.<br />
The captive part of the study was conducted in two European<br />
zoos:Cologne Zoo (Germany) and Parc Zoologique<br />
et Botanique de Mulhouse, Sud-Alsace (France). A longterm<br />
study with one group of blue-eyed black lemurs<br />
(1.3) and one group of crowned lemurs (1.2) was carried<br />
out at Cologne Zoo.In addition,three groups of blue-eyed<br />
black lemurs (2.1; 1.1; 1.1) and three groups of crowed<br />
lemurs (2.2; 2.1; 3.2) were studied at Mulhouse Zoo. The<br />
body weight development of captive individuals was registered<br />
and compared to body weight data of wild individuals.<br />
The obesity rate in captivity was recorded. An obese<br />
animal was identified as one weighing more than two standard<br />
deviations over the mean wild weight. Moreover,<br />
nutrient and energy intake of Eulemur macaco flavifrons<br />
and Eulemur coronatus at Cologne Zoo and Mulhouse Zoo<br />
were registered. In addition, digestibility trials were conducted.<br />
Samples of feeds and faeces were analyzed using<br />
Weende analysis and detergent analysis. In Madagascar,<br />
four groups of Eulemur macaco flavifrons in two forest fragments,one<br />
mainly primary forest and the other predominantly<br />
secondary forest,were observed.Samples of plants<br />
utilized by free-ranging blue-eyed black lemurs were collected.They<br />
were botanically classified and analyzed using<br />
Weende analysis and detergent analysis.<br />
The mean body weights of Eulemur macaco flavifrons as well<br />
as Eulemur coronatus in captivity were significantly higher<br />
than the mean body weight of free-ranging individuals.100<br />
% of the Eulemur macaco flavifrons sample and 33.3 % of<br />
the Eulemur coronatus sample were obese.Significant body<br />
weight differences were found between the groups studied<br />
at Cologne Zoo and Mulhouse Zoo, which could be<br />
explained by different feeding regimes. Comparing the<br />
diet of free-ranging blue-eyed black lemurs to the zoo<br />
diets that were based mainly on fruits and vegetables at<br />
Cologne Zoo and Mulhouse Zoo, considerable differences<br />
were found with respect to NDF,ADF,ADL and crude<br />
protein content, whereas ash and crude lipid content varied<br />
only slightly.The NFC and energy content in the zoo<br />
diets were almost twice as high as those in the diet of wild<br />
blue-eyed black lemurs. The high NFC,crude protein and<br />
metabolizable energy content and low fibre content of<br />
the zoo diets as compared to the wild diet,combined with<br />
a relatively high apparent digestibility of ~80 % for Eulemur<br />
macaco flavifrons and ~84 % for Eulemur coronatus, respectively,<br />
and in combination with lemurs’ typically low basal<br />
metabolic rates, all clearly contribute to the obesity problem<br />
of captive Eulemur macaco flavifrons.<br />
The presented data of food consumed by Eulemur macaco<br />
flavifrons in captivity and in the wild reveals elementary<br />
differences concerning nutrient and energy composition.<br />
Although a bright variety of fruits and vegetables could<br />
protect animals in captivity from stereotypic behaviour, a<br />
systematic reassessment of the zoo diet is suggested:<br />
increasing fibre content and decreasing energy density by<br />
feeding vegetables,and whenever possible,fresh plant material<br />
in appropriate quantities instead of energy-rich<br />
fruits, gruel or commercial feeds. Although the utilization<br />
of the food fibre content by a generalist frugivore like Eulemur<br />
macaco flavifrons or Eulemur coronatus is limited, fibre<br />
content plays an important role in the maintenance of<br />
physiological health. A zoo diet corresponding to the natural<br />
requirements of lemurs guarantees an optimization<br />
of breeding programmes and presents a valuable and necessary<br />
contribution to the preservation of these highly<br />
endangered species.<br />
Key words: Eulemur macaco flavifrons, Eulemur coronatus,<br />
Nutrition, Digestibility, Obesity, Captivity, Energy intake.<br />
Rafaliarison R.R. 2010. Activité générale du Prolemur simus:<br />
transition saison sèche - saison de pluies et activité de la<br />
femelle avant et après mise bas dans le Parc National<br />
Ranomafana.Département de Paléontologie et d’Anthropologie<br />
Biologique, Université d’Antananarivo, Madagascar.<br />
Cette étude a été réalisée dans la parcelle 3 du parc national<br />
Ranomafana qui abrite le seul groupe du parc.Elle nous<br />
aidera à mieux comprendre les variations de l’activité générale<br />
du Prolemur simus pendant la transition de la saison<br />
sèche à la saison de pluie ainsi que la variation de l’activité<br />
de la femelle avant et après mise bas. Les résultats ont<br />
montré que le Prolemur simus a dépensé la moitié de leur<br />
temps à l’alimentation suivi du repos. Les variations de la<br />
fréquence de l’activité sont en relation avec la partie consommée<br />
(tige, moelle ou jeunes pousses), la disponibilité<br />
alimentaire,la température et la pluie ainsi que la disponibilité<br />
en eau.La strate la plus utilisée est comprise entre 0<br />
à 5 m. La présence d’un nouveau né a une influence sur<br />
l’activité et la proximité des individus du groupe. Pour la<br />
femelle, il y a une diminution de la fréquence de l’alimentation<br />
après la mise bas.Il y a aussi une augmentation très<br />
marquée de la fréquence du repos après la mise bas.Le juvénile<br />
s’éloigne de la femelle après mise bas tandis que le<br />
mâle reste toujours près de la femelle.<br />
Mots clés: Prolemur simus, Activités, Mise bas, Parc National<br />
Ranomafana, Madagascar.<br />
This study was carried out in Parcel 3 of Ranomafana National<br />
Park,where the only group of Prolemur simus within<br />
the park is present.It concerns the variation in the general<br />
activities of P.simus during the transition from the dry season<br />
to the rainy season as well as the activity of the female<br />
before and after giving birth.The results showed that P. simus<br />
spent half of their time feeding, followed by resting.<br />
The variation in frequency of activities was related to the<br />
consumed plant parts (trunk,culm pith or bamboo shoot),<br />
availability of food,temperature,rain and the availability of<br />
water. The most frequently used forest stratum was between<br />
0 and 5 m of height. The presence of the new-born<br />
had an influence on the activity and the spacing of the individuals<br />
in the group.For the female,there was a reduction<br />
of the frequency of feeding after giving birth. There was<br />
also a very marked increase in the frequency of resting after<br />
birth. The juvenile stayed away from the female after<br />
she had given birth,but the male always remained close to<br />
the female.<br />
Key words:Prolemur simus,Activity,New born,Ranomafana<br />
National Park, Madagascar.<br />
Raharivololona,B.M.2010.Intestinal parasite infection of the<br />
gray mouse lemur (Microcebus murinus, J.F. Miller, 1777) in<br />
the south-eastern littoral forest of Madagascar. PhD Dissertation,<br />
Hamburg University, Hamburg, Germany.<br />
Madagascar’s plants and animals belong to one of the most<br />
unique and threatened biotas of the world. Lemurs are<br />
the flagship species associated with the biological crisis of<br />
the island and notably vulnerable to habitat degradation.<br />
While most studies on the effect of habitat destruction<br />
on species survival have focused on population reduction<br />
and forest degradation, indirect effects, such as altered<br />
parasite loads have received little attention. Parasitologi-
Page 74 Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
cal studies have concentrated on large primates, such as<br />
apes and monkeys.This is probably due to epidemiological<br />
interest in apes, which are genetically closer to humans<br />
and are known to be a reservoir of certain pests and<br />
diseases fatal to humans. Prosimians’ gastrointestinal parasites<br />
are less studied.<br />
The goal of this project was to assess and describe the<br />
gastrointestinal parasites of the lemur species Microcebus<br />
murinus (Family Cheirogaleidae), also known as the gray<br />
mouse lemur, from the littoral forest fragments of Mandena<br />
in extreme southeastern Madagascar. In addition, I<br />
wanted to evaluate the utility of determining gastrointestinal<br />
parasite loads based on fecal samples. From April<br />
2003 to October 2005, 427 fecal samples obtained<br />
from169 different individuals of M. murinus from five forest<br />
fragments were analyzed to assess the parasite species<br />
richness of this animal based on parasite egg morphology.Three<br />
individuals of M.murinus were also sacrified in<br />
order to look for adult worms for identification and confirmation<br />
of parasite species, and to localize their gastrointestinal<br />
parasites in the digestive tract. Screening all<br />
fecal samples by using the modified technique of the<br />
McMaster flotation, I noted that M. murinus harbored at<br />
least nine different intestinal parasites, which included 1)<br />
six Nemathelminthes:a member of the family Ascarididae,<br />
one species of the family Subuluridae represented by the<br />
genus Subulura,an unidentified Strongylida,a species of the<br />
genus Trichuris (Trichuridae), two species of the family<br />
Oxyuridae: the first belongs to the genus Lemuricola and<br />
the second is still unidentified;2) two Plathelminthes:two<br />
cestodes of the genus Hymenolepis (Hymenolepididae);3)<br />
one Protozoa: belonging to the order Coccidia.<br />
These gastrointesinal parasites of M.murinus from Mandena<br />
have not been previously described from this primate.<br />
The cestode infection deserves special attention,as these<br />
parasites have not been previously reprted from lemurs.<br />
Adult worms of Trichuris species were found in the caecum,<br />
as well as Lemuricola worms in the caecum and large<br />
intestine.Subulura worms were more abundant in the caecum<br />
than in the small and large intestine. A large number<br />
of Subulura larvae were observed in the caecum.As exemplified<br />
by the data on Subulura sp. worms in the digestive<br />
tract of M.murinus,the number of nematode parasite eggs<br />
and larvae found in the feces are correlated with the intensity<br />
of infection in the digestive tract.<br />
To assess effects of forest fragmentation and degradation,fecal<br />
samples from the first captureof 169 individuals of Microcebus<br />
murinus living in five littoral forest fragments were<br />
analyzed for gastrointestinal parasites. The fragments differed<br />
in size and forest quality. In good quality forest<br />
blocks, lemurs from a smaller fragment had higher prevalences<br />
and intensities of infection of gastrointestinal nematodes<br />
and protozoans than animals from a larger forest<br />
fragment. In larger forest blocks, excretion of eggs from<br />
Ascarididae and tapeworms was higher in a degraded forest<br />
fragment than in a better quality forest fragment.This<br />
situation was reversed in small forest fragments with fewer<br />
eggs of Subulura nematodes and protozoans shed by<br />
lemurs in the degraded fragment than by lemurs from the<br />
good quality fragment. The analyses are hampered by the<br />
fact that only one forest fragment was available per type<br />
of treatment. Keeping this limitation in mind, the results<br />
are consistent with other studies and indicate that forest<br />
degradation and fragmentation have marked effects on<br />
the level of parasitism of Madagascar’s lemurs.<br />
To assess seasonal effects on the excretion of gastrointetinal<br />
parasites I screened fecal samples from M. murinus<br />
caught during monthly trapping sessions for eggs and<br />
larvae of intestinal parasites. Parasite excretions changed<br />
seasonally when analyzed at the level of individual hosts.<br />
The number of parasite species and the abundance of parasite<br />
eggs and larvae in Microcebus feces were higher during<br />
the hot season than the cold season.Reduced parasite<br />
excretion during the cold season could be due to environmental<br />
factors or due to the ability of M. murinus to<br />
enter torpor and hibernation during the cold season<br />
which might lead to reduced metabolism of intestinal parasites<br />
and results in reduced shedding of parasite eggs.<br />
No such seasonal variation was found on the level of the<br />
lemur population when the analyses were based on samples<br />
of unknown origin.<br />
The study revealed noticeable effects of forest fragmentation<br />
on parasite loads as measured via the excretion of<br />
parasites. The disadvantageous consequences of increased<br />
parasite infections on the health of these animals is due<br />
to changes in habitat conditions and is a factor that needs<br />
to receive more attention when developing conservation<br />
plans.<br />
Key words: Microcebus murinus, Parasites, Habitat degradation,<br />
Fragmentation, Forest quality, Hymenolepis.<br />
Randrianarimanana, H.L. 2009. Etude comparative de l’alimentation<br />
et du comportement des deux espèces sympatriques<br />
d’Indriidés :Propitecus diadema et Indri indri dans<br />
le Réserve Naturelle Intégrale n°1 de Betampona (Tamatave).Mémoire<br />
de DEA en Paléontologie et évolution biologique,Biologie<br />
Evolutive,Primatologie,Département de<br />
Paléontologie et d’Anthropologie biologique, Faculté des<br />
Sciences, Université d’Antananarivo.<br />
Des études comportementales et nutritionnelles des<br />
deux espèces sympatriques d’Indriidés (Propithecus diadema<br />
et Indri indri) ont été réalisées pendant les mois de<br />
mars,avril,juin et juillet 2008 dans la Réserve Naturelle Intégrale<br />
numéro un de Betampona (Tamatave). Les données<br />
comportementales et nutritionnelles,la hauteur fréquentée<br />
et la nature des supports et des coordonnées<br />
géographiques ont été enregistrés toutes les 10 minutes.<br />
Des analyses statistiques ont été réalisées pour étudier<br />
comment ces deux plus grands lémuriens partagent leurs<br />
nourritures et habitats. Même si les deux espèces ont la<br />
même fréquence d’alimentation et sont toutes deux considérées<br />
folivores, Propithecus diadema consomme un peu<br />
plus de fruit qu’Indri indri (respectivement 33,6 et 9,4 %) et<br />
utilise beaucoup plus d’espèces végétales comme source<br />
de nourriture. Propithecus diadema fréquente des hauteurs<br />
beaucoup plus basses qu’Indri indri durant ses activités<br />
(8,3071et 10,208 m). De plus, cette dernière espèce<br />
utilise beaucoup de petits supports (respectivement 3,80<br />
et 5,38 cm) et peu inclinés (40,32° et 47,79°). Propithecus<br />
diadema se déplace beaucoup tandis qu’Indri indri se repose<br />
davantage.Malgré le chevauchement de leur territoire,<br />
ces deux espèces montrent une séparation de leur niche<br />
écologique.<br />
Mots-clés: Propithecus diadema, Indri indri, Alimentation,<br />
Comportement, Comparaison.<br />
Razafindratsima, O.H. 2009. Rôle écologique de Varecia rubra<br />
et d’Eulemur albifrons dans le Corridor Ambatolaidama du<br />
Parc National Masoala.Mémoire de DEA en Biologie,Ecologie<br />
et Conservation Animale, Département de Biologie<br />
Animale, Faculté des Sciences, Université d’Antananarivo.<br />
Une étude a été effectuée sur deux espèces de lémuriens<br />
sympatriques dans les forêts tropicales humides du corridor<br />
Ambatolaidama du Parc National Masoala - Varecia<br />
rubra (Geoffroy, 1812) et Eulemur albifrons (Geoffroy,<br />
1796). Le but est d’étudier les rôles écologiques de ces<br />
espèces dans la reforestation du corridor en tant que disséminatrices<br />
de graines. Ceci afin de mettre en évidence<br />
leur importance au niveau de ce site et par conséquent,<br />
d’élaborer une stratégie de conservation. Un groupe d’E.<br />
albifrons et trois groupes de V.rubra ont été suivis.Trois femelles<br />
de V.rubra,choisies comme animaux focaux,ont été<br />
munies de colliers à radio émetteur. Ces espèces ont fait<br />
l’objet de suivis quotidiens, de novembre 2006 à janvier<br />
2007, afin de collecter des informations sur leur régime<br />
alimentaire et leur défécation. Pour E. albifrons, aucune<br />
donnée sur son alimentation n’a été obtenue de part la<br />
difficulté de son suivi dû à l’absence de collier. Les fèces<br />
collectées sont analysées afin d’en extraire des graines qui<br />
ont été, ensuite, inventoriées, mesurées et identifiées. La<br />
viabilité de ces graines a été étudiée par un test d’immersion<br />
dans l’eau,puis par la mise en terre dans une pépinière<br />
des graines déféquées comparées avec celles extraites<br />
manuellement des fruits. Afin de comprendre le devenir<br />
de ces graines après leur dépôt, une étude de l’habitat où
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010 Page 75<br />
les fèces ont été déposées a été réalisée. Les résultats de<br />
cette étude ont démontré que V.rubra a une frugivore élevée<br />
(86.1 %). Aussi, ces deux espèces participent activement<br />
à la dissémination des graines de la majorité des<br />
espèces végétales du corridor, avec 16 graines par jour<br />
disséminées par V. rubra représentées par 34 espèces végétales<br />
appartenant à <strong>15</strong> familles, et 10 graines par jour<br />
pour E. albifrons réparties dans 8 familles avec 11 espèces.<br />
De plus, après leur passage au niveau du tube digestif de<br />
ces animaux,les graines sont viables (x2 = 107,283,ddl = 2,<br />
p = 0,0001) et ont une germination plus élevée que les<br />
graines témoin (x2 = 55,680, ddl = 1, p = 0,0001). Très peu<br />
d’entre elles ont subi des dommages (seulement de 3,8 %<br />
et 0,7 % respectivement pour les graines déféquées par V.<br />
rubra et E. albifons). L’étude de l’habitat démontre une<br />
réussite de germination malgré une forte pente.Ces deux<br />
espèces de lémuriens sont donc d’importantes disséminatrices<br />
de graines de la forêt humide du corridor Ambatolaidama.Elles<br />
jouent un rôle important dans le maintien<br />
de l’équilibre écologique et contribuent à la reforestation<br />
du corridor.<br />
Mots-clés: Varecia rubra, Eulemur albifrons, Primates, Régime<br />
alimentaire, Dispersion des graines, Parc National<br />
Masoala, Corridor, Ambatolaidama, Madagascar.<br />
Razakanirina H. 2010. Suivi phénologique global et statut de<br />
conservation de 4 espèces végétales (Strychnos decussata,<br />
Diospyros ferrea,Gardenia decaryi et Capurodendron gracilifolium)<br />
consommées par Propithecus verreauxi coronatus dans<br />
la forêt de Badrala (Antrema - Région Boeny). Mémoire<br />
de DEA en Biologie et Ecologie Végétale,option Ecologie<br />
végétale, Département de Biologie et Ecologie Végétale,<br />
Faculté des Sciences, Université d’Antananarivo.<br />
La Station Forestière à Usage Multiple d’Antrema constitue<br />
un des habitats de Propithecus verreauxi coronatus.Dans<br />
cette région, ces Lémuriens sont vénérés comme étant<br />
les représentants des ancêtres des "Sakalava". Cette culture<br />
leur offre donc un haut niveau de protection mais<br />
est- ce que la forêt peut leur fournir la nourriture dont ils<br />
ont besoin?<br />
Des études sur la phénologie et une évaluation du statut<br />
de conservation des quelques espèces consommées par<br />
ce lémurien ont été réalisées dans la forêt sèche sur dune<br />
de Badrala (partie Nord de la station),afin de faire ressortir<br />
les différents types phénologiques et la saisonnalité des<br />
différentes phénophases des arbres de cette forêt. Trois<br />
(3) plots de 2000 m2 ont été montés dans la forêt et dans<br />
chaque plot,tous les arbres à DHP > $10 cm sont numérotés<br />
et des paniers collecteurs de litières sont installés<br />
suivant un transect de 200 m. La phénologie des arbres a<br />
été suivie pendant une année grâce à des observations directes<br />
de chaque individu et à l’analyse des litières qui<br />
sont collectées tous les <strong>15</strong> jours. La période de floraison<br />
maximale des espèces se produit au début de la saison humide,<br />
avant l’apparition des feuilles et la défeuillaison est<br />
assez importante au milieu de la saison sèche. Strychnos<br />
decussata, Diospyros ferrea, Gardenia decaryi et Capurodendron<br />
gracilifolium sont classées en danger d’extinction<br />
(EN). Ainsi, des mesures de conservation sont à entreprendre<br />
afin de protéger ces espèces et les habitats de ce<br />
lémurien.<br />
Mots-clés: Suivi phénologique, Statut de conservation,<br />
Plantes consommées,Propithecus verreauxi coronatus,forêt<br />
sèche, Badrala, Station Forestière à Usage Multiple d’Antrema,<br />
Madagascar.<br />
Rued, A.C. 2009. Social structure and female foraging strategies<br />
in white-collared lemurs (Eulemur cinereiceps). M.A.<br />
thesis, University of Calgary, Calgary, Alberta, Canada.<br />
This thesis examines the nature of male-female affiliation<br />
in Eulemur cinereiceps, specifically whether it consists of<br />
special relationships or a central male social structure. A<br />
special relationship includes an unrelated male and female<br />
adult who preferentially associate and affiliate with each<br />
other over all other individuals within the group. I also<br />
examine the flexibility of female foraging strategies in<br />
response to changes in resource availability and energy<br />
requirements. I tested the resource defence hypothesis,<br />
which proposes that reproductive females form special<br />
relationships with males to improve foraging success and<br />
offset the energetic costs of reproduction.Data were collected<br />
on two small groups in Mahabo forest,on the southeastern<br />
coast of Madagascar.Analysis of social structure<br />
data suggested central male structure when resources<br />
were scarce and central female structure during the period<br />
of relative resource abundance.The resource defence<br />
hypothesis was not supported by foraging data.<br />
Solomon,S.K.2010.Living on the edge:a preliminary dry season<br />
study of crowned lemur (Eulemur coronatus, Gray<br />
1842) and Sanford’s lemur (E.sanfordi,Archbold 1932),responses<br />
to anthropogenic habitat changes in northern<br />
Madagascar. M.A., Anthropology (Environment & Sustainability),University<br />
of Western Ontario,London,Canada.<br />
Habitat fragmentation through anthropogenic disturbance<br />
is a significant threat to primates in all biogeographic<br />
areas. Recent research has shown that primates have<br />
non-patterned responses to this disturbance and that general<br />
models of changing primate behaviour are not effective<br />
conservation tools. Previous research on primates in<br />
fragments is concentrated in the Neotropics demonstrating<br />
a need to investigate species-specific responses in other<br />
areas of the world. This study examined two sympatric<br />
lemur species, the crowned lemur (Eulemur coronatus)<br />
and Sanford’s brown lemur (Eulemur sanfordi) and their responses<br />
to anthropogenic habitat fragmentation in northern<br />
Madagascar. Although habitat generalists, Sanford’s<br />
lemur was extirpated at the study site while crowned lemur<br />
density was low but viable; they were restricted to<br />
forest fragments on the periphery and top of limestone<br />
massifs. Conservation initiatives in these fragments are<br />
reliant on preserving fruit trees located in the remaining<br />
forest flatlands and the commitment of a community conservation<br />
group.<br />
Key words: Northern Madagascar, Crowned lemur, Sanford’s<br />
brown lemur,Forest fragmentation,Edge effects,Lemur<br />
density,Community conservation,Dry season,Deciduous<br />
forest.
Guidelines for authors<br />
Lemur News publishes manuscripts that deal largely or exclusively with lemurs and their habitats. The aims of Lemur News are: 1) to<br />
provide a forum for exchange of information about all aspects of lemur biology and conservation, and 2) to alert interested parties<br />
to particular threats to lemurs as they arise. Lemur News is distributed free of charge to all interested individuals and institutions. To<br />
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(*.doc or *.docx) or rich text format (*.rtf). They should generally be 1-8 pages long, including references and figures. Submissions to<br />
the “Articles” section should be divided into Introduction, Methods, Results and Discussion and should include a list of 4-6 key words.<br />
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Examples are given below.<br />
Journal article<br />
Ranaivoarisoa, J.F.; Ramanamahefa, R.; Louis, Jr., E.E.; Brenneman, R.A. 2006. Range extension of Perrier’s sifaka, Propithecus perrieri, in<br />
the Andrafiamena Classified Forest. Lemur News 11: 17-21.<br />
Book chapter<br />
Ganzhorn, J.U. 1994. Les lémuriens. Pp. 70-72. In: S.M. Goodman; O. Langrand (eds.). Inventaire biologique; Forêt de Zombitse.<br />
Recherches pour le Développement, Série Sciences Biologiques, n° Spécial. Centre d’Information et de Documentation Scientifique<br />
et Technique, Antananarivo, Madagascar.<br />
Book<br />
Mittermeier, R.A.; Konstant, W.R.; Hawkins, A.F.; Louis, E.E.; Langrand, O.; Ratsimbazafy, H.J.; Rasoloarison, M.R.; Ganzhorn, J.U.;<br />
Rajaobelina, S.; Tattersall, I.; Meyers, D.M. 2006. Lemurs of Madagascar. Second edition. Conservation International, Washington, DC,<br />
USA.<br />
Thesis<br />
Freed, B.Z. 1996. Co-occurrence among crowned lemurs<br />
(Lemur coronatus) and Sanford’s lemur (Lemur fulvus sanfordi)<br />
of Madagascar. Ph.D. thesis, Washington University, St. Louis, USA.<br />
Website<br />
IUCN. 2008. IUCN Red List of Threatened Species.<br />
. Downloaded on 21 April 2009.<br />
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Drawing by Stephen D. Nash
Editorial.................................................................................1<br />
Feature: Madagascar’s Environmental Crisis<br />
Madagascar’s illegal logging crisis: An update<br />
and discussion of possible solutions<br />
Erik R. Patel...............................................................................2<br />
Ongoing threats to lemurs and their habitat<br />
inside the Sahamalaza - Iles Radama National<br />
Park<br />
Melanie Seiler, Guy H. Randriatahina,<br />
Christoph Schwitzer.................................................................7<br />
News and Announcements..........................................9<br />
Short Communications<br />
Preliminary conservation status assessment for<br />
the Data Deficient northern giant mouse lemur<br />
Mirza zaza<br />
Eva Johanna Rode, K. Anne-Isola Nekaris,<br />
Christoph Schwitzer...............................................................11<br />
An observation of the hairy-eared dwarf lemur,<br />
Allocebus trichotis, in the Lakato region, eastern<br />
Madagascar<br />
Erwan Lagadec, Steven M. Goodman..................................12<br />
When big lemurs swallow up small ones:<br />
Coquerel’s dwarf lemur as a predator of grey<br />
mouse lemurs and endemic rodents<br />
Susanne Schliehe-Diecks, Matthias Markolf,<br />
Elise Huchard.........................................................................13<br />
Collective mobbing of a boa by a group of<br />
red-fronted lemurs (Eulemur fulvus rufus)<br />
Lennart Pyritz, Tianasoa Andrianjanahary..........................14<br />
Response of two nocturnal lemurs (Microcebus<br />
murinus and Lepilemur leucopus) to a potential<br />
boiidae (Sanzinia madagascariensis) predator<br />
Krista Fish...............................................................................16<br />
Effective predation defence in Cheirogaleus<br />
medius<br />
Kathrin H. Dausmann...........................................................18<br />
Lepilemur feeding observations from Northern<br />
Madagascar<br />
Andrew J. Lowin......................................................................20<br />
Hypotheses on ecological interactions between<br />
the aye-aye (Daubentonia madagascariensis)<br />
and microhylid frogs of the genus Platypelis in<br />
Tsaratanana bamboo forest<br />
Andolalao Rakotoarison, Solohery A. Rasamison,<br />
Emile Rajeriarison, David R. Vieites, Miguel Vences............21<br />
Discovery of crowned sifaka (Propithecus<br />
coronatus) in Dabolava, Miandrivazo, Menabe<br />
Region<br />
Josia Razafindramanana and Rija Rasamimanana............23<br />
Inferences about the distant past in Madagascar<br />
Elwyn L. Simons......................................................................25<br />
Husbandry guidelines for mouse lemurs at Paris<br />
Zoo<br />
Delphine Roullet....................................................................27<br />
Lemur News <strong>Vol</strong>. <strong>15</strong>, 2010<br />
ISSN 1608-1439<br />
Table of contents<br />
Articles<br />
Diurnal lemur density in the national park<br />
parcel Ivontaka Nord, UNESCO Biosphere<br />
Reserve of Mananara-Nord<br />
Marta Polasky Lyons..............................................................29<br />
Distribution of Prolemur simus north of the<br />
Mangoro-Nosivolo River - how far north do we<br />
really have to look?<br />
Rainer Dolch, Erik R. Patel, Jonah H. Ratsimbazafy,<br />
Christopher D. Golden, Tianasoa Ratolojanahary,<br />
Jean Rafalimandimby, Jonathan L. Fiely................................32<br />
Enquête préliminaire de la distribution des<br />
lémuriens de bambou dans et autour du<br />
Corridor forestier Fandriana-Vondrozo,<br />
Madagascar<br />
Andry Rajaonson, Maherisoa Ratolojanahary,<br />
Jonah Ratsimbazafy, Anna Feistner, Tony King....................34<br />
Effect of red ruffed lemur gut passage on the<br />
germination of native rainforest plant species<br />
Onja H. Razafindratsima, Emilienne Razafimahatratra.....39<br />
Feeding ecology of the crowned sifaka<br />
(Propithecus coronatus) in a coastal dry forest<br />
in northwest Madagascar (SFUM, Antrema)<br />
Claire Pichon, Rivo Ramanamisata, Laurent Tarnaud,<br />
Françoise Bayart, Annette Hladik, Claude Marcel<br />
Hladik, Bruno Simmen.......................................................... 42<br />
Effet de la dégradation de l'habitat sur la<br />
consommation alimentaire d'Eulemur<br />
rubriventer dans deux sites: Talatakely et<br />
Vatoharanana, du Parc National de Ranomafana<br />
Laingoniaina H. Rakotonirina, Germain J. Spiral,<br />
Jonah H. Ratsimbazafy, Soanorolalao Ravelonjanahary,<br />
Raharizelina Ralaiarison, Stacey Tecot, Alex Hall,<br />
Tricia Calhoon, Gisèle R. Randria..........................................47<br />
Observations of terrestrial latrine behaviour by<br />
the southern gentle lemur Hapalemur<br />
meridionalis in the Mandena littoral forest,<br />
southeast Madagascar<br />
Timothy M. Eppley, Giuseppe Donati....................................51<br />
Conservation des lémuriens via la protection de<br />
leurs habitats et le développement communautaire<br />
dans les corridors de Betaolana et Tsaratanana-<br />
Betaolana, région de SAVA<br />
Lala Razafy Fara, Iarilanto Andriamarosolo.........................54<br />
Genetic diversity in ten Indri (Indri indri)<br />
populations compared to other lemur species<br />
John Zaonarivelo, Rick Brenneman, Rambinintsoa<br />
Andriantompohavana, Edward E. Louis, Jr............................59<br />
Verreaux’s sifaka fur condition in the spiny forest<br />
of southern Androy<br />
Ivan Norscia, Jean Lambotsimihampy, Elisabetta Palagi.....64<br />
Rediscovery of Sibree’s dwarf lemur in the<br />
fragmented forests of Tsinjoarivo, central-eastern<br />
Madagascar<br />
Marina B. Blanco....................................................................67<br />
Funding and Training.....................................................69<br />
Recent Publications........................................................70<br />
Theses Completed..........................................................71