Varecia variegata (Kerr, 1792)

Russell A. Mittermeier, Anthony B. Rylands & Don E. Wilson, 2013, Lemuridae, Handbook of the Mammals of the World – Volume 3 Primates, Barcelona: Lynx Edicions, pp. 90-141 : 140-141

20.

Black-and-white Ruffed Lemur

Varecia variegata

French: Vari noir et blanc / German: Schwarzweil 3er Vari / Spanish: Lémur de collar blanco y negro

Other common names: Pied Black-and-white Ruffed Lemur; Northern Black-and-white Ruffed Lemur (subcincta), Southern Black-and-white Ruffed Lemur (editorum), Variegated Ruffed Lemur (variegata)

Taxonomy. Lemur macaco variegata Kerr, 1792 ,

Madagascar.

The color pattern in this species 1s extremely variable. Several major pattern morphs may be identified, a few of which maybe distinct enough to be considered valid subspecies. One such undescribed form (known only from asingle specimen collected in southern Madagascar) is said to be dark brown with two black lateral spots, while another (from south-eastern Madagascar) is almost completely white dorsally. The subspecies subcincta maypossibly intergrade with V. rubra : the confluence of the Vohimara and Antainambalana rivers has been investigatedas a possible contact or hybridization zone between the two, but results are inconclusive. Moreover, individuals with the coat pattern of the subspecies variegata are also found well inside the distribution of the subspecies editorum. This needs furtherinvestigation, but if thereis actual sympatry, these subspecies will require recognitionas full species. On the other hand, it maybethat thereis so muchvariation within this species that recognition of anysubspeciesis inappropriate. The subspecies would also appearto intergrade into one another, with intermediate patterns occurring even within otherwise “typical” populations. Three subspecies recognized.

Subspecies and Distribution.

V.v.variegataKerr,1792—CEMadagascar,fromtheAnoveRiver(betweenSoanieranaIvongoandMananara)StotheregionSofMananjaryasfarastheManampatranaorMananaraRivers;thedistributionisnowhighlyfragmented.

V.v.editorumHill,1953—CEMadagascar,extendingverypatchilyfromtheforestsofMantadiaNationalParkStojustSofMananjary;distributionsofeditorumandvariegataappeartooverlap,andintermediateformsmayexist,butsofar,nofieldsurveysorgeneticstudieshaveconfirmedthis.

V. v. subcincta A. Smith, 1833 — NE Madagascar, from the Antainambalana River (in the region of Maraontsetra) S to the Anove River (between Soanierana Ivongo and Mananara); formerly S as far as the region of Toamasina. Introduced into Nosy Mangabe (Antongil Bay) in the 1930s.

Descriptive notes. Head-body45 cm, tail 60-61 cm; weight 3-6 kg (males) and 3-7 kg (females). The three subspecies differ slightly but significantly in body weight andtail length; the “Northern Black-and-white Ruffed Lemur ” ( V. v. subcincta ) is the smallest in this respect, and the “Southern Black-and-white Ruffed Lemur ” (V. v. editorum ) is the largest. Apart from longertails in females, thereis no difference in size or coloration between sexes. Individuals fromthe introduced population on Nosy Mangabeare smaller than those from all other populations, which may be due to ecological constraints on this small island. As its common English namesuggests, Black-and-white Ruffed Lemurs are variably covered in black and white fur. In general, individuals are predominantly white to the south ofthe range (with black oftenrestricted to shoulders andflanks) and increasingly black to the north (with a white band aroundthe body and white forearms and flanks). In the “Variegated Black-and-white Ruffed Lemur ” (V. v. variegata ), furis black on the abdomen, tail, extremities, inner aspects oflimbs, forehead, circumorbital area, and top ofthe head. In contrast, the back, flanks, rump, and most ofthe hindlimbs are usually white. Shoulders are black. Thereis a thick white longitudinal band in the centerofthe back that distinguishes the Variegated Black-and-white Ruffed Lemurfrom Southern Black-and-white Ruffed Lemur . This pattern may vary according to locality, and intermediate forms mayexist. In the Southern Black-and-white Ruffed Lemur , the abdomen, tail, extremities, inner aspects of limbs, forehead, circumorbital area, top of the head, and anterior part ofthe back are black, whereas the posteriorpart ofthe back, flanks, rump, and outeraspects oflimbs are usually white. Like the Variegated Black-andwhite Ruffed Lemur , the coat of the Southern Black-and-white Ruffed Lemuris fluffy, the tail is long and bushy, and ears are ruffed with long thick white hair. The Northern Black-and-white Ruffed Lemurhas a distinct white belt aroundits otherwise black back. The abdomen, tail, extremities, inneraspects of limbs, forehead, circum-orbital area, and top ofthe headare also black. Outer aspects oflimbs andthe base ofthetail are white. In all subspecies eyes are yellow or amber-colored.

Habitat. The Variegated Black-and-white Ruffed Lemur inhabits remnant tracts of tropical moist lowland and montane forest from sea level to 1300 m. The Southern Black-and-white Ruffed Lemur inhabits lowland to mid-altitude primary and secondary rainforests from sea level to 1300 m. The Northern Black-and-white Ruffed Lemur prefers lowland rainforest.

Food and Feeding. Black-and-white Ruffed Lemurs eat mainly large ripe fruits, supplemented with young leaves, seeds, flowers, and nectar, depending on the season. It is one of the most frugivorous lemur species and relies heavily on large fruit trees. It may be an important disperser of the seeds of Canarium madagascariense ( Burseraceae ). It appears that only a small number oftree species are exploited. Individuals occasionally eat soil.

Breeding. The Black-and-white Ruffed Lemur is polygamous. Females have a 30day reproductive cycle. The vulva remains closed except during estrus. Reproduction varies considerably between years, with an average interbirth interval of one year. Mating takes place in May-July, with females only fertile for one day during each estrus. Births occur from September through early November, after gestation of only 99-106 days. In captivity, litters can be 1-5 in size but are usually 2-3. It is doubtful that female Blackand-white Ruffed Lemurs are as productive in the wild, given that litter size appears to be related to the mother’s non-pregnant body weight and captive females tend to be heavier. Newborns are rather poorly developed at first, each weighingjust under 100 g, but they are able to see and grow very rapidly. They have the same general fur pattern as adults, along with light blue eyes that change to golden by about the second week of life. Ruffed lemurs are probably the only primates that build nests exclusively for birth and the first couple of weeks of infant rearing. Nests are crudely constructed out of branches and leaves and lined with the mother’s fur, pulled from her flanks. At first, instead of bringing newborns with her while foraging, the mother will leave them on their own in the nest. Later, when she does begin to move them, they are carried in her mouth by the skin of their flanks. One reason for Varecia parking their infants in nests might be that females have to travel large distances between dispersed food resources and are thus unable to afford the energy-consuming practice of infant carrying. When suckling, a female lies directly on her infants. Multiple individuals participate in caring for the young. Young attain roughly three-quarters of their adult body weight by four months of age. Females reach sexual maturity at just under two years, but males take three years to mature. Captive individuals have lived for up to 35 years in zoos.

Activity patterns. The Black-and-white Ruffed Lemur is arboreal and diurnal. On Nosy Mangabe, their activity budgetis seasonally variable. In warm months, individuals are most active. Generally, activity peaks in the early morning and late afternoon.

Movements, Home range and Social organization. The Variegated Black-and-white Ruffed Lemur has been studied in Betampona. Long-term field research shows significant variation in home range size, group size, and territorial behavior. The Southern Black-and-white Ruffed Lemur has been studied in Ranomafana National Park and Manombo Special Reserve. One group lived in a 197ha home range and was monogamous. A long-term study of the Northern Black-and-white Ruffed Lemur was carried out on Nosy Mangabe Special Reserve, where it had a fission-fusion social system in which large groups of up to 16 individuals split into smaller core-groups when fruits were scarce—a behavior that may enhance foraging efficiency of each individual. Core groups were not cohesive and continued to split and reform. Density estimates are 0-4-2-5 ind/km? in Manombo, 10-15 ind/km? in Antanamalaza, and 29-43 ind/km? on Nosy Mangabe.

Status and Conservation. CITES Appendix I. Classified as Critically Endangered on The IUCN Red List (including all three subspecies). Main threats are habitat loss from slash-and-burn agriculture, logging, and mining, but hunting is also a major factor and can be severe in some areas. Studies ofvillages in the Makira Forest found that ruffed lemur meat is a desirable food item and that current levels of hunting are unsustainable. Stochastic events such as cyclones can severely affect small populations. The Blackand-white Ruffed Lemur is very patchily distributed—the result ofits dependency on lowto mid-altitude primary forest with large fruit trees and its susceptibility to hunting and trapping. The Variegated Black-and-white Ruffed Lemur is found in one national park (Zahamena), two strict nature reserves (Betampona and Zahamena), and two special reserves (Ambatovaky and Marotandrano). Individuals with the subspecies variegata coat pattern are also found in protected areas further south, well inside the range of editorum. In November 1997, individuals of this subspecies that had been born and raised in zoos in the USA were returned to Madagascar and released in the Betampona Strict Nature Reserve. A study of this reintroduction effort is ongoing. The Southern Black-and-white Ruffed Lemur occurs naturally in two national parks (Mantadia and Ranomafana), Manombo Special Reserve, Anosibe an’ Ala Classified Forest, and unprotected forests of Fandriana (proposed as a national park), Kianjavato, Lakia, Maromizaha, Tolongoina, and Vatovavy. Because distributions of editorum and variegata seem to overlap, some of the aforementioned localities might also hold populations of individuals with a variegata coat pattern. Past studies indicated its presence in Andringitra National Park, but recent surveys have failed to find it there. The original population in the Analamazaotra Special Reserve was extirpated by the 1980s, butit was successfully reintroduced in 2006 by E. Louis Jr. and a team of Malagasy researchers. The Northern Black-and-white Ruffed Lemuris found in Mananara-Nord National Park, Nosy Mangabe Special Reserve, and the soon-to-be protected forests of Makira.

Bibliography. Baden et al. (2008), Balko (1998), Balko & Underwood (2005), Britt (1997, 1998, 2000), Britt, lambana et al. (2003), Britt, Welck & Katz (2000, 2001), Brockman et al. (1987a, 1987b), Downman (1993), Foerg (1982a), Geissman & Mutschler (2006), Golden (2005), Goodman & Rasolonandrasana (2001), Hekkala et al. (2007), Irwin et al. (2005), Iwano (1989), Kaufman (1991), Lindsay & Simons (1986), Mallinson & Barker (1998), Mittermeier, Langrand et al. (2010), Mittermeier, Tattersall et al. (1994), Morland (1990, 1991a, 1991b, 1993a, 1993b), Nicoll & Langrand (1989), Overdorff et al. (2005), Pereira, Kaufman et al. (1990), Pereira, Klepper & Simons (1987), Pereira, Seeligson & Macedonia (1988), Petter et al. (1977), Rakotonirina, Rajaonson, Ratolojanahary, Missirli et al. (2011), Ralainasolo et al. (2005), Raps & White (1995), Rasmussen (1985), Ratsimbazafy (2002, 2006), Ratsimbazafy et al. (2002), Schwitzer & Kaumanns (2001a, 2009), Simons & Lindsay (1987), Tattersall (1982), Terranova & Coffman (1997), Tombomiadana & Rakotondravony (2000), Vasey (2003, 2005b), Vasey & Tattersall (2002), Weigler et al. (1994), White (1991), White, Balko & Fox (1993), White, Burton et al. (1992), Wyner et al. (1999a).

Figures

8. Brown (Lemur Fulemurfulvus), 9. Rufous Brown Lemur (Eulemur rufus), 10. Red-fronted Brown Lemur (FEulemur rufifrons), 11. White-fronted Brown Lemur (Eulemur albifrons), 12. Sanford’s Brown Lemur (Fulemur sanfordi), 13. White-collared Brown Lemur (Eulemur cinereiceps), 14. Red-collared Brown Lemur (Eulemur collars), 15. Black Lemur (Eulemur macaco), 16. Blue-eyed Black (Lemur Eulemurflavifrons), 17. Crowned Lemur (Eulemur coronatus), 18. Red-bellied Lemur (Eulemur rubriventer), 19. Mongoose Lemur (Eulemur mongoz)

Subspecies and Distribution. V.v.variegataKerr,1792—CEMadagascar,fromtheAnoveRiver(betweenSoanieranaIvongoandMananara)StotheregionSofMananjaryasfarastheManampatranaorMananaraRivers;thedistributionisnowhighlyfragmented. V.v.editorumHill,1953—CEMadagascar,extendingverypatchilyfromtheforestsofMantadiaNationalParkStojustSofMananjary;distributionsofeditorumandvariegataappeartooverlap,andintermediateformsmayexist,butsofar,nofieldsurveysorgeneticstudieshaveconfirmedthis. V. v. subcincta A. Smith, 1833 — NE Madagascar, from the Antainambalana River (in the region of Maraontsetra) S to the Anove River (between Soanierana Ivongo and Mananara); formerly S as far as the region of Toamasina. Introduced into Nosy Mangabe (Antongil Bay) in the 1930s.