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Vol. 15 - Deutsches Primatenzentrum

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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 />

References<br />

Asa, C.S. 2008. Scent and the single male: ring-tailed lemurs<br />

produce honest signals.Molecular Ecology 17:3223-3224.<br />

Bollen,A.;Donati,G.2006.Conservation status of the littoral<br />

forest of south-eastern Madagascar: a review. Oryx 40:<br />

57-66.<br />

Boonstra,R.;Krebs,C.J.;Kenney,A.1996.Why lemmings have<br />

indoor plumbing in summer.Canadian Journal of Zoology<br />

74: 1947-1949.<br />

Charles-Dominique,P.;Hladik,C.M.1971.Le Lepilemur de sud<br />

de Madagascar: ecologie, alimentation, et vie sociale.<br />

Revue d’Écologie (La Terre et la Vie) 25: 3-66.<br />

Curtis, D.; Donati, G.; Rasmussen, M. 2006. Cathemerality.<br />

Folia Primatologica 77: 5-6.<br />

Epple, G. 1986. Communication by chemical signals. Pp.<br />

531-580. In: G. Mitchell; J. Erwin (eds.), Comparative<br />

Primate Biology, <strong>Vol</strong>ume 2A: Behavior, Conservation and<br />

Ecology. Alan R. Liss, New York.<br />

Feeley,K. 2005.The role of clumped defecation in the spatial<br />

distribution of soil nutrients and the availability of nutrients<br />

for plant uptake. Journal of Tropical Ecology 21:<br />

99-102.<br />

Gilbert, K.A. 1997. Red howling monkey use of specific defecation<br />

sites as a parasite avoidance strategy.Animal Behaviour<br />

54: 451-455.<br />

Goodman, S.M.; Pidgeon, M. 1999. Carnivora of the Réserve<br />

Naturelle Intégrale d’Andohahela, Madagascar. Fieldiana:<br />

Zoology 94: 256-68.<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<br />

and their Ecological Basis. Plenum Press, New York.<br />

Irwin, M.T.; Samonds, K.E.; Raharison, J.-L.; Wright, P.C. 2004<br />

Lemur latrines: observations of latrine behavior in wild

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