Varecia variegata variegata (Kerr, 1792)
The distribution of V. v. variegata, is poorly known (Tattersall, 1977,1982). It is found in the eastern rain forest, extending southwards from the Antainambalana River (which is the boundary between the two subspecies) as far as Manakara (Petter and Petter, 1971) or to just north of the Mananara River (Petter et al, 1977; Tattersall, 1982; Petter and Petter-Rousseaux, 1979). This subspecies is also found on the small island of Nosy Mangabe where it was introduced in the 1930s (J. Petter pers. comm. to Constable et al, 1985).
Numbers are not known. The Black and White Ruffed Lemur does not appear to occur at high densities anywhere other than on Nosy Mangabe Pollock, 1984). It is estimated (Simons Morland, in prep) that there may be as many as 100-150 individuals on the 520 ha island (i.e. approximately 20-30 individuals per sq. km). In 1984, Pollock estimated between 56 and 84 animals on the island. Density on the island in 1983 was estimated at 175 animals per sq. km (Iwano, 1989), i.e. a total of 910 individuals, which is a much higher estimate than that of Simons Morland or Pollock.
A three month (June-August 1988) study of the Black and White Ruffed Lemur has been carried out near Ranomafana in the south-east of Madagascar (White, 1989). The results of a longer term study (1600 observation hours between July 1987 and January 1989) of the subspecies on Nosy Mangabe are currently being written up (Simons Morland, in prep). At Ranomafana, the study group consisted of an adult male and adult female which traveled through a large home range, of 197 ha, as a cohesive pair (White, 1989). A subadult was observed in the area but, though it exchanged calls with the adult pair it did not associate with them. The pair frequently ranged more than I km each day, usually feeding, traveling and resting high (20-25 m) in the canopy. Locomotion was principally quadrupedal, with frequent leaping (White, 1989; Pereira et al, 1988). White et al (1989) suggest that the larger groups of Varecia with smaller home ranges that are found on islands and in isolated forest blocks may be a consequence of the limited space available there for dispersion. The diet of V. variegata in both Ranomafana and Nosy Mangabe was mostly fruit, supplemented with small amounts of nectar, seeds and leaves; on Nosy Mangabe, the diet varied seasonally (White, 1989; Simons Morland, in prep). Some earth was also eaten. Chorusing loud calls were used as territorial advertisement and for coordination of movement within the territory (White, 1989). Simons Morland (in prep) reports that it is female V. v. variegata which defend the territories. Other reports of group size are of between two and five individuals (Petter et al, 1977; Pollock, 1979; Jolly et al, 1984). On Nosy Mangabe, there was seasonal variation in activity levels and patterns; levels were highest during the summer months and some nocturnal activity may have occurred (Simons Morland, in prep.). In general, the Ruffed Lemurs on the island were most active in the early morning and late afternoon/evening (Simons Morland, in prep). Pollock (1979) describes Varecia as crepuscular. In captivity, the animals were more active in the morning and evening and there was no sign of nocturnal activity (Klopfer and Dugard, 1976; Kress et al, 1978).
Most details of reproduction come from studies in captivity. Gestation period is 90-102 days (Hick, 1976; Bogart et al, 1977; Boskoff, 1977). Up to six offspring may be produced in a litter (Anon, 1984), although two or three is the most common number and primiparous females frequently have singletons (Boskoff, 1977; Foerg, 1982). On Nosy Mangabe, most females had twins, these were born in October and November (Simons Morland, in prep). When the infants are born, they do not cling to their mothers' fur, as happens in most other lemur species, but are left in nests (Petter et al, 1977; Klopfer and Dugard, 1976; Jolly et al, 1984). These may be constructed by the female, but are frequently just bundles of epiphytes (Jolly et al, 1984). On Nosy Mangabe, infants were kept in nests constructed by their mother, 15-20 m high in large trees; they were never seen in thick tangles of epiphytes but were parked in trees once they were one to two weeks old (Simons Morland, 1989, in prep). In a forest enclosure at Duke, nests were built by the female Varecia on the ground; infants remained in these until they were approximately three weeks old after which their mothers frequently left them parked high up in trees (Pereira et al, 1987). When they are carried, it is in their mothers' mouth (Klopfer and Dugard, 1976; Petter et al, 1977). The infants begin to follow their mother at three weeks of age and are as fully mobile and active as the adults at seven weeks old (Klopfer and Boskoff, 1979). In the wild, infants were close to adult size at six months of age (Simons Morland, in prep). Females can conceive at 20 months of age (Boskoff, 1977), but, in captivity, average age at first reproduction is 3.4 years (SSP Masterplan, 1988). Simons Morland (in prep.) suggests that the high rate of population increase seen in captive Ruffed Lemurs is not typical of wild populations.
There is considerable destruction of the eastern forests, these are being cleared principally for growing crops. The lemurs are heavily hunted for food, both trapping and shooting occurs (Nicoll and Langrand, 1989; Constable et al, 1985; Lindsay and Simons, 1986). Iwano (1989) implies that there was a considerable decline in the number of Varecia present on Nosy Mangabe between 1983 and 1984 because of the poaching of this species on the island.
The Black and White Ruffed Lemur is present in Betampona Nature Reserve, it is reported to be common in Zahamena Nature Reserve (Pollock, 1984; Nicoll and Langrand, 1989; Simons Morland, in litt.) and occurs in Andringitra Nature Reserve (Nicoll and Langrand, 1989). Nicoll and Langrand (1989) were informed that it was in Marojejy Nature Reserve but an expedition there in 1988 failed to see or hear them (Stafford et al, 1989; W. Duckworth, pers. comm.). The subspecies is also found in the Special Reserve of Nosy Mangabe and seems to be reappearing in Analamazaotra Special Reserve (Nicoll and Langrand, 1989).
A number of new protected areas, in which Varecia is present, have been proposed (Nicoll and Langrand, 1989). These are Ranomafana, Mantady (both proposed as National Parks) and Mananara (proposed as a Biosphere Reserve). Surveys to discover the distribution and numbers of existing populations of V. v. variegata are essential. Special attention should be paid to determining whether there are several subspecies within the Black and White Ruffed Lemur population, or whether the variation in color is nothing more than individual variation (see Remarks below). If there are, in fact, distinct forms, the conservation problems for this species will be much greater than currently recognized (St Catherine's Workshop, 1986).
ISIS (1989) lists 243 Black and White Ruffed Lemurs (Lemur variegatus variegatus and L. v. subcinctus) in 49 institutes; about 95% of these are reported to be captive born. They also report a further 134 individuals of Lemur variegatus for which they give no subspecific name (ISIS, 1989). Wilde et al (1988) list a further 20 animals in four institutes that are not in the ISIS lists. The Species Survival Masterplan (1988) for the Ruffed Lemur records 265 individuals of this subspecies in North America. Duke Primate Center has the largest collection with 22 individuals held in February 1989 (Katz, in litt.). In Madagascar, there are six individuals held at Ivoloina, near Toamasina (Katz, in litt.). Two pairs of these were captive bred animals returned to Madagascar by Duke Primate Center in November 1988 (Katz, in litt.). The animals were supplied by San Diego and San Antonio Zoo as well as DPC. Brockman (1989) recommends equalization of founder representation and controlled reproduction for the captive population of this subspecies.
There is considerable variation in coat color and pattern within V. v. variegata, and it is possible that better knowledge of the distribution of the varieties may ultimately suggest their recognition as subspecies (Petter et al, 1977; Tattersall, 1982). Tattersall (1982) recognizes four distinct and consistent coat patterns within the Black and White Ruffed Lemur: Type a) Face black except for short white hairs on muzzle below eyes; black forehead and crown; ears, cheeks and throat tufted white; otherwise white except for ventrum, tail, lateral aspect of thighs and shoulders, proximal part of forelimbs and extremities, all of which are black. Type b1) Resembles type a, except that the black shoulder patches extend posteriorly onto the flanks and medially to meet in the midline. Type b2) Pattern as in b1, except that a narrow white stripe runs forward in the dorsal midline, invading the back forequarters but not reaching the neck area. Type c) Entirely black except for white cheeks, ears and throat, a white transverse band extending across the back and sides just below the shoulders and another across the rump extending down the posterior aspect of the thighs onto the lateral surface of the lower leg. White patches also occur laterally on the lower arm. It is this dark type, often referred to as V. v. subcinctus that is found on Nosy Mangabe and in the surrounding mainland forests (Simons Morland, in litt.). Adult sized animals weighed on Nosy Mangabe were between 2400 and 3700g; weights were seasonally variable (Simons Morland, in prep.). The Malagasy names of this subspecies are varikandana and varikandra (Tattersall, 1982).