|Scientific Name:||Varecia variegata|
|Species Authority:||(Kerr, 1792)|
|Infra-specific Taxa Assessed:|
|Taxonomic Notes:||There are three recognized subspecies: the nominate form, V. v. variegata; V. v. editorum; and V. v. subcincta.|
|Red List Category & Criteria:||Critically Endangered A2cd ver 3.1|
|Assessor(s):||Andriaholinirina, N., Baden, A., Blanco, M., Chikhi, L., Cooke, A., Davies, N., Dolch, R., Donati, G., Ganzhorn, J., Golden, C., Groeneveld, L.F., Hapke, A., Irwin, M., Johnson, S., Kappeler, P., King, T., Lewis, R., Louis, E.E., Markolf, M., Mass, V., Mittermeier, R.A., Nichols, R., Patel, E., Rabarivola, C.J., Raharivololona, B., Rajaobelina, S., Rakotoarisoa, G., Rakotomanga, B., Rakotonanahary, J., Rakotondrainibe, H., Rakotondratsimba, G., Rakotondratsimba, M., Rakotonirina, L., Ralainasolo, F.B., Ralison, J., Ramahaleo, T., Ranaivoarisoa, J.F., Randrianahaleo, S.I., Randrianambinina, B., Randrianarimanana, L., Randrianasolo, H., Randriatahina, G., Rasamimananana, H., Rasolofoharivelo, T., Rasoloharijaona, S., Ratelolahy, F., Ratsimbazafy, J., Ratsimbazafy, N., Razafindraibe, H., Razafindramanana, J., Rowe, N., Salmona, J., Seiler, M., Volampeno, S., Wright, P., Youssouf, J., Zaonarivelo, J. & Zaramody, A.|
|Reviewer(s):||Schwitzer, C. & Molur, S.|
|Facilitator/Compiler(s):||Chiozza, F. & Clark, F.|
Listed as Critically Endangered as the species is suspected to have undergone a population decline of ≥80% over a period of 21 years (three generations), primarily due to observed and inferred continuing decline in area, extent and quality of habitat from slash-and-burn agriculture, logging and mining, in addition to exploitation through unsustainable hunting pressure. These causes have not ceased, and will to a large extent not be easily reversible.
|Range Description:||This species inhabits lowland to mid-altitude rain forests in eastern Madagascar. There are three subspecies:
The subspecies V. v. subcincta is assumed to represent the northernmost subspecies, the northern limit of its range being the Antainambalana River (east of which occurs the Red Ruffed Lemur). Its range extends southwards to the Anova River, including part of Makira, Mananara-Nord, Atialanankorendrina, and Marotandrano. This subspecies was introduced to the island of Nosy Mangabe in the Bay of Antongil back in the 1930s and still occurs there (Kuhn 1972). The distribution of this lemur is very patchy throughout its range, except for Nosy Mangabe, where it lives at a relatively high density (Morland 1991).
The nominate subspecies V. v. variegata occurs south of the Anove River, from about Ambatovaky south to about Betampona and Zahamena National Park (including Ambatovaky), although the southern limit is not yet clearly defined.
The subspecies V. v. editorum is the southernmost subspecies and is known with certainty only from Mantadia southwards to Manombo Special Reserve. According to Groves (2001), the ranges of V. v. editorum and V. v. variegata overlap, and intermediate forms exist. The form occurring in Mangerivola Special Reserve is unknown.
|Range Map:||Click here to open the map viewer and explore range.|
Population densities recorded range from 0.4-2.5/km² in Manombo, to 10-15 individuals/km² in Antanamalaza, to 25 individuals/km² in Mangevo (Ranomafana National Park) (Baden 2011) and 29-43 individuals/km² on Nosy Mangabe (Vasey 2003). No Varecia were recorded during 2007 surveys at Manombo (also completely absent from nearby Agnalazaha and closest interior forests) suggesting that the Manombo population is small and likely not recovered from Cyclone Gretelle in 1997.
|Habitat and Ecology:||
This species is very patchily distributed in lowland to mid-altitude rain forests. Black-and-white Ruffed Lemurs maintain large home ranges (e.g., 87.8 – 90.5 ha; Baden 2011) consisting of low-to-mid elevation primary forest with tall trees with broad crowns. They are almost exclusively frugivorous, and as they are very selective feeders, which makes them especially susceptible to disturbance (e.g. see White et al. 1995, Ratsimbazafy 2002). In Vatovavy and Sangasanga, Varecia favor areas far from the forest edge, and areas with a large basal area of known food species. However, preferences did not result in total avoidance areas within fragments; they were fairly ubiquitous across limited forests available (Note: this landscape is at lower elevation than RNP (100-600 m above sea level). See Mittermeier et al. 2008 and references therein.
Group size and structure appear to vary considerably between study sites (e.g., pairs: White 1989; cohesive MM-MF: Balko 1998; fission-fusion: Morland 1991, Vasey 2003, Baden 2011). Females usually give birth to two to three young, which are left in a nest when young and afterwards carried in the mother’s mouth (Baden 2011). Boom-bust reproductive strategies [e.g., one reproductive event documented in 6 years of observation (Baden 2011); ‘reproductive collapse’ observed (Vasey 2009); long IBI during this same time in Kianjavato (Holmes, unpublished data)], possibly mitigated by climatic variation/resource availability/abundance. Ruffed lemurs are probably the only primates that build nests exclusively for the birth and the first days of rearing infants (Mittermeier et al. 2008, and references therein).
Varecia variegata among the highest genetic diversity of primates (3.1x higher than the genetic diversity present in humans) (Perry et al. 2012). Comparison of population genetics between fragmented and continuous landscapes in southeast suggest that small geographic separation may not be sufficient to disrupt gene flow among V. variegata populations. Rather, fragment isolation or age may play a greater role than size in the genetic consequences of habitat fragmentation (Holmes et al. submitted).
|Use and Trade:||This species is hunted.|
|Major Threat(s):||The principal threat to its survival is habitat loss due to slash-and-burn agriculture, logging and mining. They are large bodied and diurnal, and therefore also among the most heavily hunted of all lemur species. The seasonality of their vocalizations (due to increased food availability) has been tied to increased levels of hunting (Ratsimbazafy 2002, Vasey 2003). In Makira, where they are one of the more expensive and desired meats, hunting is largely unsustainable (Golden 2005). It is one of the first lemurs to disappear where humans encroach upon rain forest habitats.|
This species is listed on Appendix I of CITES.
V. v. subcincta is recorded from Mananara-Nord National Park and Nosy Mangabe Special Reserve. V. v. variegata is recorded from Zahamena National Park, and two nature reserves (Betampona and Zahamena) and from Ambatovaky and Marotandrano Special Reserves. V. v. editorum is recorded from Mantadia National Park, Ranomafana National Park and Manombo Special Reserve. Now extirpated from Analamazaotra Special Reserve, and no longer reported from Andringitra National Park.
Manongarivo, to which it is not possible to assign a subspecies, is a Special Reserve. Fandriana (in the range of V. v. editorum) is in the process of becoming a national park. Unprotected forests such as Tolodoina, Vatovavy, Atialanankorendrina and Makira should be included in protected areas.
In November 1997, V. v. editorum born and raised in US zoological institutions were returned to Madagascar and released in the Betampona Reserve (Britt et al. 2003). The study of this reintroduction effort is ongoing. An education campaign against hunting, using this species as a flagship, is recommended.
Baden A. 2011. Communal infant care in black-and-white ruffed lemurs (Varecia variegata)". Stony Brook University.
Balko EA. 1998. A behaviorally plastic response to forest composition and logging disturbance by Varecia variegata in Ranomafana National Park, Madagascar. State University of New York.
Britt, A., Iambana, B. R., Welch, C. R. and Katz, A. S. 2003. Restocking of Varecia variegata variegata in the Réserve Naturelle Intégrale de Betampona. In: S. M. Goodman and J. P. Benstead (eds), The Natural History of Madagascar, pp. 1545–1551. The University of Chicago Press, Chicago, USA.
Golden, C. D. 2005. Eaten to endangerment: Mammal hunting and the bushmeat trade in Madagascar's Makira Forest. Undergraduate Thesis, Harvard University.
Groves, C. P. 2001. Primate taxonomy. Smithsonian Institution Press, Washington, DC, USA.
IUCN. 2014. The IUCN Red List of Threatened Species. Version 2014.1. Available at: www.iucnredlist.org. (Accessed: 12 June 2014).
Kuhn, H. J. 1972. Die Geschichte der Säugetiere Madagaskars. Zeitschrift des Kölner Zoo 15(1): 28–42.
Mittermeier, R., Louis, E., Hawkins, F., Langrand, O., Ganzhorn, J., Konstant, W., Rasoloarison, R., Rajaobelina, S. and Richardson, M. 2008. Lemurs of Madagascar, 3rd edition. Conservation International.
Morland, H. S. 1991a. Social organization and ecology of black and white ruffed lemurs (Varecia variegata variegata) in lowland rain forest, Nosy Mangabe, Madagascar. Ph.D. Thesis, Yale University.
Rakotonirina, L.H.F., Rajaonson, A., Ratolojanahary, J.H., Missirli, J.M., Razafy Fara, L., Raholijaona, Andriamanajaranirina, M. & King, T. 2013. Southern range extensions for the critically endangered black-and-white ruffed lemur Varecia variegata and greater bamboo lemur Prolemur simus. Primate Conservation 26: 49-55.
Ratsimbazafy, J. 2002. On the brink of extinction and the process of recovery: responses of black-and-white ruffed lemurs (Varecia variegata variegata) to disturbance in Manombo Forest, Madagascar. Ph.D. Thesis, Stony Brook University.
Vasey, N. 2003. Varecia, ruffed lemurs. In: S. M. Goodman andJ. P. Benstead (ed.), The Natural History of Madagascar, pp. 1332-1336. University of Chicago Press, Chicago, USA.
Vasey, N., Borgerson, C. 2009. Social and ecological factors associated with temporary reproductive collapse in cooperatively breeding red variegated lemurs (Varecia rubra) on the Masoala Peninsula, Madagascar. American Journal of Physical Anthropology 48: 261.
White, F.J. 1989. Diet, ranging behavior and social organisation of the black-and-white ruffed lemur, Varecia variegata variegata, in southern Madagascar. American Journal of Physical Anthropology 78(2): 23.
White, F. J., Overdorff, D. J., Balko, E. A. and Wright, P. C. 1995. Distribution of ruffed lemurs (Varecia variegata) in Ranomafana National Park, Madagascar. Folia Primatologica 64: 124–131.
|Citation:||Andriaholinirina, N., Baden, A., Blanco, M., Chikhi, L., Cooke, A., Davies, N., Dolch, R., Donati, G., Ganzhorn, J., Golden, C., Groeneveld, L.F., Hapke, A., Irwin, M., Johnson, S., Kappeler, P., King, T., Lewis, R., Louis, E.E., Markolf, M., Mass, V., Mittermeier, R.A., Nichols, R., Patel, E., Rabarivola, C.J., Raharivololona, B., Rajaobelina, S., Rakotoarisoa, G., Rakotomanga, B., Rakotonanahary, J., Rakotondrainibe, H., Rakotondratsimba, G., Rakotondratsimba, M., Rakotonirina, L., Ralainasolo, F.B., Ralison, J., Ramahaleo, T., Ranaivoarisoa, J.F., Randrianahaleo, S.I., Randrianambinina, B., Randrianarimanana, L., Randrianasolo, H., Randriatahina, G., Rasamimananana, H., Rasolofoharivelo, T., Rasoloharijaona, S., Ratelolahy, F., Ratsimbazafy, J., Ratsimbazafy, N., Razafindraibe, H., Razafindramanana, J., Rowe, N., Salmona, J., Seiler, M., Volampeno, S., Wright, P., Youssouf, J., Zaonarivelo, J. & Zaramody, A. 2014. Varecia variegata. The IUCN Red List of Threatened Species. Version 2015.2. <www.iucnredlist.org>. Downloaded on 30 July 2015.|
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