Section 4: Lemur Taxa of Conservation Concern
Aye-Aye
Daubentonia madagascariensis
The aye-aye is the only living representative of the family Daubentonidae. It is an elusive nocturnal lemur that appears to be widely, though sparsely, distributed throughout eastern, northern and northwestern Madagascar (Fig. 4). The aye-aye seems very adaptable and is found in areas of primary and secondary rain forest, deciduous forest, secondary growth, dry scrub forest and cultivated areas (Tattersall, 1982).
Daubentonia spends the day in a nest, usually built in the fork of a tree or in vine tangles. Single individuals are most commonly observed. The aye-aye's diet consists largely of fruits, principally coconuts, and insect larvae (Petter, 1977), but they are reported to eat a wide variety of items including plant galls, bamboo tree bark, adult insects and small vertebrates (Petter and Petter-Rousseaux, 1967; Pollock et al., 1985; Iwano and Iwakaw, 1988). It appears that the aye-aye may give birth once every two to three years (Petter and Peyrieras, 1970a).
The aye-aye is found in both the Eastern and Western Regions of Madagascar, and has a larger distribution than was originally believed. It has been reported from the following protected areas: the Verezanantsoro, Mantady, Montagne d'Ambre and Ranomafana National Parks; the Andohahela, Bemaraha, Betampona, Marojejy and Zahamena Nature Reserves; the Ambatovaky, Analamazaotra, Analamera, Ankarana, Foret d'Ambre, Manombo, and Manongarivo Special Reserves, and it was introduced to the Nosy Mangabe Special Reserve in 1966 (Ganzhorn, 1986; O'Connor et al, 1986; Albignac, 1987; Petter and Andriatsarafara, 1987; Raxworthy and Rakotondraparany, 1988; Wright, 1988: Nicoll and Langrand, 1989; Safford et al., 1989; Thompson and Evans, 1991; Hawkins et al., in press). Aye-aye have recently been sighted in the forests surrounding Daraina (D. Meyers, pers. comm.) Eleanor Sterling of Yale University has recently completed two years of field research on the aye-ayes of Nosy Mangabe.
Captive aye-aye (Daubentonia madagascariensis) at the Duke University Primate Center.
David Haring
Estimates of aye-aye populations are needed for each of the protected areas listed above. In addition, most of the above-mentioned areas require better protection, as well as public awareness campaigns to discourage killing of aye-ayes. Albignac (1988) suggests that such programs compensate local people for damage done by aye-ayes to their crops. Searches for new aye-aye populations should be conducted in the Marotandrano and Tampoketsa-Analamaitso Special Reserves and the Midongy-Sud Classified Forest. Creation of new protected areas around the city of Maroantsetra and on the Masoala Peninsula should also help safeguard the aye-aye's future.
The captive breeding effort for this species should be expanded beyond the small colonies that currently exist at the Jersey Wildlife Preservation Trust, Vincennes Zoo and the Duke University Primate Center, which now total about 10 animals.
Figure 4. Distribution of Daubentonia madagascariensis.
The recently rediscovered hairy-eared dwarf lemur (Allocebus trichotis).
Bernhard Meier
Hairy-Eared Dwarf Lemur
Allocebus trichotis
Until its rediscovery in 1989 in northeast Madagascar, the tiny, hairy-eared dwarf lemur, a monotypic genus, was known from only five museum specimens. Meier and Albignac (1989) consider that the distribution of Allocebus trichotis may be restricted and patchy. Its numbers are almost certainly declining as the eastern rain forests continue to be cut (Richard and Sussman, 1975, 1987).
Little is known about this nocturnal species, but it appears to occur only in lowland rain forest. Local people reported that they did not see active hairy-eared dwarf lemurs between May and September, and it appears that they are in some kind of hibernation during that time (Meier and Albignac, 1989). The animals are usually found sleeping in tree holes.
The main threat to this species is slash-and-burn agriculture. An area around the Verezanantsoro National Park (Fig. 5), the only known locality for Allocebus, has been designated as the Mananara-Nord Biosphere Reserve (Greve, 1991). Surveys are needed to locate other remaining populations of this species. Four Allocebus have been collected for captive breeding in conjunction with the Mananara-Nord Biosphere Reserve project (Meier and Albignac, 1989). This program should be expanded to include colonies abroad.
Figure 5. Distribution of Allocebus trichotis.
Sclater's Lemur
Eulemur macaco flavifrons
The black lemur (Eulemur macaco) occurs in the evergreen forests of the Sambirano region in northwestern Madagascar, as well as on the islands of Nosy Be and Nosy Komba and several smaller islands in the Nose Be area. Two subspecies are recognized, Eulemur m. macaco and E. m. flavifrons, the latter regarded as endangered.
Male Sclater's lemurs (E. m. flavifrons) are uniformly black, perhaps with a brown tint to the fur, while the females have a reddish-tan coat (Koenders et al., 1985; Birkel, 1987). The subspecies is reported to occur from the Andranomalaza River in the north, south to the Sandrakota River, but is not known to occur in any protected area (Fig. 6).
Meyers et al. (1989) recently studied the distribution and status of Lemur macaco and found evidence of a geographical cline between E. m. macaco and E. m. flavifrons. Populations of what they considered pure E. m. flavifrons were small, patchy and found entirely outside the Sambirano region. Sclater's lemur is threatened by hunting, trapping and forest destruction for agricultural development (Koenders et al., 1985).
In 1988, France's Strasbourg University and the Mulhouse Zoo proposed a conservation program for Sclater's lemur to the Malagasy Government, which included captive breeding efforts, field studies, creating a special reserve within its range and the training of a Malagasy student to work on the project (Lernould and Rumpler, 1988). This program is also supported by the Cologne Zoo, Saarbruken Zoo and the Duke University Primate Center (Harcourt and Thornback, 1990). Subsequent recommendations by Meyers et al. (1989) focus heavily on conducting a socioeconomic survey of the Andranomalaza region, and changing current land-use practices to conserve forest habitat. They point out the difficulty in creating a special reserve that will protect a significant population of E. m. flavifrons, but suggest that increased protection of the Manongarivo Special Reserve will protect significant populations of E. m. flavifrons and E. m. macaco hybrids. Sclater's lemur has bred successfully at France's Mulhouse Zoo, and this captive breeding program should be expanded.
Female Sclater's lemur (Eulemur macaco flavifrons).
David Mevers
Male and (possibly hybrid) female Sclater's lemur.
Russell A. Mittermeier
Figure 6. Distribution of Eulemur macaco subspecies.
Red Ruffed Lemur
Varecia variegata rubra
The ruffed lemur (Varecia variegata) is an eastern rain forest species which does not appear to be common anywhere. Two subspecies are generally recognized; both are considered endangered. The coat of Varecia variegata rubra is primarily a deep rusty red with black extremities, forehead, crown, ventrum and tail. A patch of white fur occurs on the back of the neck.
The red ruffed lemur is restricted to the forests of the Masoala Peninsula (Figs. 7,17) (Petter et al., 1977; Petter and Petter-Rousseaux, 1979; Tattersall, 1977a, 1982), parts of which have been heavily degraded for agricultural development (Simons and Lindsay, 1987). Very little information is available regarding the red ruffed lemur's conservation status, ecology or behavior. The only protected area in which it was known to occur, the former Masoala Nature Reserve, was degazetted in 1964 to permit timbering. Hunting and trapping of all lemurs is common in this area (Tattersall, 1977a; Constable et al., 1985; Simons and Lindsay, 1987).
The Missouri Botanical Garden is actively involved in the creation of a national park on the Masoala Peninsula, with funding provided by USAID. A detailed survey of this region is needed to determine the distribution of the red ruffed lemur. A successful international captive breeding program is already underway which could possibly result in a reintroduction effort.
Figure 7. Distribution of Varecia variegata rubra and Varecia variegata variegata in the area of the Masoala Peninsula.
The red ruffed lemur (Varecia variegata rubra).
Photo courtesy of the San Diego Zoo
Golden Bamboo Lemur
Hapalemur aureus
The golden bamboo lemur was discovered only in 1987. A medium-sized gray-brown lemur with a black face and golden-yellow eyebrows, cheeks and throat, it is patchily distributed over a small area of rain forest in southeastern Madagascar. Hapalemur aureus is known to occur in only one protected area, the newly-created Ranomafana National Park (Figs. 8,9), where it is sympatric with both of the other members of the genus Hapalemur, the larger H. simus and the smaller H. griseus (Meier et al., 1987). Its numbers in Ranomafana are estimated at 1,000 (P. Wright, pers. comm.).
The species feeds almost exclusively on plants of the family Graminae, particularly the giant bamboo (Cephalostachium viguieri), but also on bamboo creeper and bamboo grass (Meier et al., 1987; Meier and Rumpler, 1987). It has been observed in groups of between two to six individuals (Meier et al., 1987; Wright et al., 1987) and appears to be active in the early morning and evening and is probably also active for part of the night.
Slash-and-burn agriculture is the main threat to this species. Surveys are needed to determine if the species survives in areas other than Ranomafana. Two Hapalemur aureus were taken into captivity at Madagascar's Parc Tsimbazaza, representing the “type” and the “paratype” of this newly-described species. This pair has successfully bred four times; three young survive. This captive breeding program should be expanded.
Golden bamboo lemur (Hapalemur aureus).
Russell A. Mittermeier
Greater bamboo lemur (Hapalemur simus).
Russell A. Mittermeier
Greater Bamboo Lemur or Broad-Nosed Gentle Lemur
Hapalemur simus
The greater bamboo lemur is now known from only the humid forest east of Fianarantsoa (Figs. 8,9) (Tattersall, 1982; Meier and Rumpler, 1987; Wright et al., 1987), but the species used to be distributed throughout northern, northwestern, central and eastern Madagascar (Godfrey and Vuillaume-Randriamanantena, 1986). It is charcoal-gray in color with paler underparts and white ear tufts (Tattersall, 1982). This species may also occur in the forest of Ampasinambo, 80 km east of Ambositra, and there is a possibility that it might be discovered at Ankarana. Bamboo is plentiful in that area and recent feeding damage similar to that produced by the greater bamboo lemur was seen there in 1986 (Fowler et al., 1989; Wilson et al., 1989). The species is found in rain forest areas where there are considerable quantities of the giant bamboo (Cephalostachium viguieri). It has been seen in groups of up to seven individuals.
Habitat destruction by slash and burn agriculture and the cutting of bamboo are the major threats to H. simus. The population at Kianjavato is also threatened by hunting (Meier and Rumpler, 1987).
Currently, Ranomafana National Park is the only protected area in which Hapalemur simus is known to occur. P. Wright (pers. comm.) estimates its numbers in the 41,600 ha area to be about 1,000. Increased protection for Ranomafana is needed to reverse a 15-year trend of habitat destruction along its borders. Searches for new populations should be conducted in other areas, including Ankarana and Ampasinambo. Several greater bamboo lemurs have been taken into captivity in Europe. A captive breeding program should be established in Madagascar as well.
Lac Alaotra Bamboo Lemur
Hapalemur griseus alaotrensis
There are three subspecies of bamboo or gentle lemur (Hapalemur griseus), one of which, H. g. griseus, is widely distributed throughout the eastern rainforest. Another, H. g. occidentalis, occurs in two isolated populations in the west. The third, H. g. alaotrensis, is restricted to the reed beds of Lac Alaotra and the surrounding marshes (Figs. 8,10) (Petter et al., 1977; Petter and Petter-Rousseaux, 1979; Tattersall, 1982).
The Lac Alaotra bamboo lemur is gray-brown in color and weighs about 1 kg. It is known to feed on the leaves and young shoots of the reed Phragmites and on the buds and pith of Papyrus in the wild and on bamboo in captivity (L. Durrell, pers. comm.). Although it normally moves about by clinging and leaping, it is also reported to swim very well. Group sizes range from three to four in certain seasons to congregations as large as 30–40 individuals at other times of the year (Petter and Peyrieras, 1970b, 1975).
This subspecies is severely threatened by habitat loss due to the annual burning of reed beds and the capture of these lemurs as food as they flee the fires (Petter and Peyrieras, 1970b; Jolly et al., 1984). In addition, the lake is continually drained for rice irrigation and the reeds are cut to fashion mats, fish traps, screens, barriers and fencing (Pollock, 1986). The Lac Alaotra bamboo lemur is not found in any protected area.
Recommendations for the creation of a national park have been submitted to the Malagasy authorities. Two areas have been targeted for protection, 2,000 ha along the northern part and 3,000 ha along the central part of Lac Alaotra. A long-term field study should be initiated as soon as possible to determine this species needs. A small number of individuals were recently taken into captivity by the Jersey Wildlife Preservation Trust. This captive breeding program should be expanded.
Lac Alaotra bamboo lemur (Hapalemur griseus alaotrensis).
James Morgan
Figure 8. Distribution of Hapalemur aureus, Hapalemur simus and Hapalemur griseus alaotrensis.
Figure 9. Ranomafana National Park and the region in which Hapalemur aureus and Hapalemur simus are known to occur.
Figure 10. The location of Lac Alaotra in relation to the Zahamena and Betampona Nature Reserves.
Indri
Indri indri
The indri is the largest living lemur, weighing 7–8 kg or more (Pollock, 1984), and is a monotypic genus. Its coloration is variable, mostly black with some white, gray or brown (Tattersall, 1982; Jenkins, 1987), and it is the only lemur species with only a stump of a tail. The indri is now confined to the eastern rain forest from the Mangoro River northwards to near the latitude of Sambava (Fig. 11), but excluding the Masoala Peninsula (Petter et al., 1977; Tattersall, 1982). Population densities vary widely (Pollock, 1975).
Much of what we know about the ecology and behavior of this species comes from research conducted by Pollock (1975, 1977, 1979, 1984) mainly on one group at Perinet. Indri are strictly diurnal and live in groups of 2–5 individuals; loud morning calls advertise the presence of groups within their ranges. Indris feed on leaves, flowers and fruit. Females give birth to young every two to three years.
The main threat to Indri indri is slash-and-burn agriculture, a practice that continues even in protected areas. Although hunting remains a problem for many lemur species, it is taboo to hunt the indri in many areas. Nonetheless, the animal is sometimes killed for food anyway, as was the case with Chinese workers on the Antananarivo-Tamatave road in the mid-1980s. The indri is reported to occur in the following protected areas: the newly-created Verezanantsoro and Mantady National Parks, the Betampona and Zahamena Nature Reserves, and the Ambatovaky, Analamazaotra and Anjanharibe-Sud Special Reserves (Nicoll and Langrand, 1989; Thompson and Evans, 1991). Based upon its geographic range, the indri may also be present in the Mangerivola and Marotandrano Special Reserves.
Figure 11. Distribution of the indri (Indri indri).
The indri (Indri indri) at the Perinet-Analamazaotra Reserve.
Russell A. Mittermeier
Surveys are needed to determine the size and status of indri populations in protected areas, and searches should also be conducted in areas not currently protected in order to determine the true distribution of this species. The indri has never been kept successfully in captivity, but a carefully planned program might be established in Madagascar.
Silky Sifaka
Propithecus diadema candidus
Diademed sifakas (Propithecus diadema sspp.) are large, diurnal lemurs found in Madagascar's eastern rain forest.
Propithecus diadema candidus is found north of Maroantsetra to the Andapa Basin and the Marojejy Massif (Fig. 12). Its numbers are unknown, but it appears to be rare throughout its range (Tattersall, 1982). The common English name for this subspecies comes from its dense silky white pelage. There have been no detailed studies of the silky sifaka, but brief observations suggest that groups are small and contain a dominant pair and their offspring. Habitat destruction is the major threat to this subspecies; it is also hunted even in protected areas (Safford et al., 1989). Propithecus diadema candidus is known to inhabit the Marojejy Nature Reserve and the Anjanaharibe-Sud Special Reserve (Simona, 1988; Nicoll and Langrand, 1989; Safford et al., 1989).
It is important to determine the status of the silky sifaka in the two protected areas in which it is known to occur, and to increase protection of these populations. In addition, searches for this subspecies should be made in the area northeast of Maroantsetra. Its presence would strengthen the proposal that has already been made for a new special reserve in that area. None are in captivity at present, but the Duke University Primate Center has developed protocols for Propithecus husbandry that could be useful in establishing captive colonies.
The silky sifaka (Propithecus diadema candidus).
Stephen Nash
Perrier's sifaka (Propithecus diadema perrieri).
Stephen Nash
Perrier's Sifaka
Propithecus diadema perrieri
Perrier's sifaka has long, dense and predominantly black fur. It is the rarest of the diademed sifaka subspecies and is believed to be restricted to the drier forests south and east of Anivorano Nord (Figs. 12,13) (Petter et al., 1977; Tattersall, 1982; Meyers and Ratsirarson, 1989).
Propithecus diadema perrieri is known to occur in the Analamera and Ankarana Special Reserves, but is apparently rare in the latter (Meyers and Ratsirarson, 1989; Hawkins et al., in press). Based upon studies conducted in Analamera, group size ranges from one to six individuals; the diet includes mature leaves, unripe fruit, petioles, young leaves, stems and flowers. The total population of Perrier's sifaka is probably close to 2,000 (Meyers and Ratsirarson, 1989).
Threats to Perrier's sifaka are habitat destruction for agricultural development, fires, livestock grazing and the cutting of trees to produce charcoal. It is also hunted for food in the Analamera region. Meyers (pers. comm.) reports that the 100 km2 area of this sifaka's range is only 50% forested, and that hunting appears to be on the rise in the southern part of its range. There is a local taboo against hunting sifakas in the Ankarana region (Fowler et al., 1989) and in the northern part of their range (Meyers and Ratsirarson, 1989).
Better protection for Perrier's sifaka is needed in both the Analamera and Ankarana Reserves (Nicoll and Langrand, 1989). In addition, a well-protected forest corridor should be created between these two reserves (Harcourt and Thornback, 1990). Ecological studies should begin as soon as possible, and the feasibility of a captive breeding program should be evaluated.
Tattersall's Sifaka or Golden-Crowned Sifaka
Propithecus tattersalli
Tattersall's sifaka is a newly-described species with a very limited distribution in northeast Madagascar near Daraina (Figs. 12,13). Although original estimates of its population were in the low hundreds (Simons, 1988), Meyers and Ratsirarson (1989) have shown that it inhabits a much larger area than was originally thought. D. Meyers (pers. comm.) estimates that Tattersall's sifaka inhabits a region of about 1,500 km2 (approximately 20% forested) at relatively high densities, and that the total population may be as high as 8,000.
Propithecus tattersalli is found primarily in dry forests, but it was recently observed in humid evergreen forests near Binara (Meyers and Ratsirarson, 1989). It has been observed in groups of three to six individuals and at densities of 60–70 individuals/km2.
This species does not occur in any protected area at the present time. It is threatened by brush fires and loss of habitat to agriculture. Despite the fact that local custom in Daraina forbids the hunting or eating of lemurs, some gold miners attracted to this region do not respect this tradition.
Meyers and Ratsirarson (1989) have recommended that a national park of approximately 20,000 ha be created to protect Propithecus tattersalli, and that the park include the forests of Binara as well as those northeast of Daraina. Methods of sustainable land use must be employed around this protected area and a public education program launched. A long-term ecological study should begin as soon as possible.
A small captive colony was established several years ago at the Duke University Primate Center. This colony should be expanded and others created.
Crowned Sifaka
Propithecus verreauxi coronatus
Propithecus verreauxi is the most common and best-studied species in the genus, but it is still threatened by habitat destruction throughout its range.
The crowned sifaka (P. v. coronatus) is a uniformly light-colored animal with dark brown head, cheeks, throat, and inner arms. It is known from a small area of northwestern Madagascar just south of Mahajanga where its range adjoins that of P. v. deckeni to the southwest and P. v. coquereli to the northeast (Fig. 12). P. v. coronatus appears to be restricted to forests on the eastern side of the Mahavavy River (Rabemazava, 1990). As suggested by Tattersall (1982), the relationship between coronatus and deckeni remains to be clarified.
Very little information exists about this subspecies and it is not known to occur in any protected areas. Surveys of the Kasijy Special Reserve may identify new populations. Genetic analyses should be performed on sifakas inhabiting these protected areas, as well as on known populations of P. v. deckeni, to clarify the relationship between these two taxa. Several crowned sifakas are kept at the Vincennes Zoo in France.
Tattersall's sifaka (Propithecus tattersalli).
David Haring
Crowned sifaka (Propithecus verreauxi coronatus).
Russell A. Mittermeier
Figure 12. Distribution of Propithecus, Propithecus diadema perrieri, Propithecus tattersalli and Propithecus verreauxi deckeni/coronatus.
Figure 13. Occurrence of Propithecus diadema perrieri and Propithecus tattersalli in northern Madagascar.
Coquerel's Dwarf Lemur
Mirza coquereli
Coquerel's dwarf lemur is a nocturnal species with a discontinuous distribution in areas of dry deciduous forest and the more humid Sambirano region in western Madagascar (Fig. 14). It weighs about 300g and has a grayish-brown coat. Its diet includes insects, spiders, frogs, chameleons, small birds, fruits, flowers, buds, gums and insect secretions (Petter et al., 1971; Hladik et al., 1980; Andrianarivo, 1981). Mirza coquereli spends the day in a nest made of vines, branches and leaves; in some areas these nests appear to be clustered in communities. Both sexes are territorial (Pages 1978, 1980).
Like other lemurs, Mirza coquereli is threatened by habitat destruction due to cultivation and fires. However, it appears to adapt well to secondary forest. It has been reported to occur in the Tsingy de Bemahara Nature Reserve, the Andranomena Special Reserve and the Analabe Private Reserve (Nicoll and Langrand, 1989).
Surveys of Mirza coquereli populations in the above-mentioned reserves are required, as are searches for other populations, beginning perhaps with the Kirindy and Tsimembo Classified Forests. Captive breeding programs have been established at a number of zoos and related institutions. These efforts should be better coordinated and a studbook established.
Coquerel's dwarf lemur (Mirza coquereli) at the Duke University Primate Center.
Benjamin Freed
Figure 14. Distribution of Mirza coquereli.
Fork-Marked Dwarf Lemur
Phaner furcifer
The fork-marked dwarf lemur is a small nocturnal species with a wide but discontinuous distribution (Fig. 15). It inhabits both humid and dry deciduous forest as well as secondary forest (Harcourt and Thornback, 1990). Its color is gray-brown above with lighter underparts and a broad dark dorsal stripe that bifurcates on the crown, the two stripes continuing to the eyes (Tattersall, 1982).
Originally considered a monotypic species, Phaner furcifer was recently divided into four subspecies by Groves and Tattersall (1991). P. f. furcifer is found on the Masoala Peninsula, P. f. electromontis in the area of Mt. d'Ambre, P. f. pallescens in western Madagascar from south of the Fiherenana River to the region of Soalala, and P. f. parienti in the Sambirano region. Disjunct populations have also been reported from the Tsaratanana Massif in the north and the Andohahela Nature Reserve in the south (Andriampianina and Peyrieras, 1972; Russell and McGeorge, 1977; Petter et al., 1977; Petter and Petter-Rousseaux, 1979; Tattersall, 1982).
The fork-marked dwarf lemur eats tree gums, sap, bud exudates, insects and larvae; it spends the day in tree holes or abandoned nests of other lemurs and is reported to be a very vocal species (Petter et al., 1971, 1975; Charles-Dominique and Petter, 1980).
All four subspecies of Phaner furcifer are threatened by habitat destruction due to fires and the clearing of forest land for pasture and crops (Harcourt and Thornback, 1990).
Based upon a presumed distribution limited to the Masoala Peninsula, Phaner f. furcifer would not occur in any protected area. However, fork-marked dwarf lemurs have been reported from the Betampona and Zahamena Nature Reserves, and this probably represents P. f. furcifer (Jenkins, 1987; Katz, 1990; Daniels, 1991). P. f. electromontis is reported from the Montagne d'Ambre National Park, the Ankarana and Foret d'Ambre Special Reserves, and from the forests near Daraina (D. Meyers, pers. comm.) P. f. pallescens is reported from the Tsingy de Bemaraha Nature Reserve, the Andranomena and Analamera Special Reserves the Analabe Private Reserve and the Kirindy, Tsimembo and Zombitse Classified Forests (Nicoll and Langrand, 1989). P. f. parienti is reported from the Manongarivo and Tsaratanana Special Reserves (Raxworthy and Rakotondraparany, 1988; Nicoll and Langrand, 1989; Hawkins et al., in press). At present, it is not clear which subspecies is represented by the fork-marked dwarf lemur population reported from the Andohahela Nature Reserve in southeastern Madagascar.
Surveys of little-studied protected areas throughout Madagascar could very well identify new populations of Phaner furcifer, and these should be undertaken in conjunction with searches for other priority lemur taxa. In addition, genetic research is needed to clarify the taxonomy of Phaner.
The fork-marked dwarf lemur (Phaner furcifer pallescens), this one from the vicinity of Analabe, southwest Madagascar.
Russell A. Mittermeier
Figure 15. Distribution of Phaner furcifer subspecies.
Ring-Tailed Lemur
Lemur catta
The diurnal ring-tailed lemur is found in dry brush, scrub and closed canopy forests of southern and southwestern Madagascar (Fig. 16), and is probably the most terrestrial and best-studied of all lemur taxa. It is also the lemur most commonly seen in zoos. It has a very distinctive gray coat on its body, arms and legs, white underparts, a white face with black muzzle and eye rings, and a characteristic black-and-white striped tail (Tattersall, 1982).
Based upon the revised taxonomy of Simons and Rumpler (1988), Lemur catta is now a monotypic genus. The northern limit of its range appears to be the forests south of Morondava on Madagascar's west coast (Sussman, 1977), and it is said to range into the interior highlands further than any other lemur (Tattersall, 1982). Long-term studies at the Berenty Private Reserve and Beza-Mahafaly Special Reserve show that ring-tailed lemurs feed on fruit, leaves, flowers, bark and sap from more than 30 plant species, but the mainstay of their diet is the kily tree, Tamarindus indica. Group sizes range from three to two dozen individuals, with a female-dominated social hierarchy (Jolly, 1966; Sussman, 1974).
Recent satellite surveys of southern Madagascar indicate that Lemur catta habitat is disappearing at an alarming rate. This is due largely to fires, overgrazing by livestock and the cutting of trees to produce charcoal (Harocourt and Thornback, 1990). Ring-tailed lemurs are also hunted with dogs in some areas (O'Connor, 1987) and often kept as pets.
Lemur catta is found in the Isalo National Park, Tsimanampetsotsa, Andohahela and Andringitra Nature Reserves, the Beza-Mahafaly Special Reserve and Berenty Private Reserve (Nicoll and Langrand, 1989). Surveys are needed to determine the distribution and sizes of remaining populations. Efforts should also be made to link captive breeding programs — there are probably more than 1,000 ring-tailed lemurs in over 100 zoos worldwide — with conservation programs in the field.
The ring-tailed lemur (Lemur catta).
Russell A. Mittermeier
Figure 16. Distribution of the ring-tailed lemur (Lemur catta).
Black-and-White Ruffed Lemur
Varecia variegata variegata
The black-and-white ruffed lemur occurs in the eastern rain forest (Fig. 17), extending southward from the Antainambalana River (where its range meets that of Varecia variegata rubra) to just north of the Mananara River (Petter et al., 1977; Tattersall, 1982; Petter and Petter-Rousseaux, 1979); it was also introduced to the island of Nosy Mangabe in the 1930s (Constable et al., 1985). Considerable variation in the coat color and pattern of the black-and-white ruffed lemur suggests that several varieties, ranging from predominantly white to predominantly black in color, might be recognized as subspecies (Petter et al., 1977; Tattersall, 1982).
Varecia variegata variegata appears to occur at low densities throughout its range, with the exception of the population on Nosy Mangabe (Pollock, 1984). It eats mostly fruit, supplemented with nectar, seeds and leaves (White, 1989; Simons Morland, in prep). Groups range in size between two and five individuals and they are very territorial (Petter et al., 1977; Pollock, 1979; Jolly et al., 1984). A great deal has been learned about the reproductive behavior of this species through studies of captive animals (Bogart et al., 1977; Boskoff, 1977; Foerg, 1982).
Black-and-white ruffed lemurs are found in the Verezanantsoro, Mantady and Ranomafana National Parks, the Andringitra, Betampona and Zahamena Nature Reserves and in the Ambatovaky, Nosy Mangabe and Analamazaotra Special Reserves (Pollock, 1984; Nicoll and Langrand, 1989; Harcourt and Thornback, 1990; Thompson and Evans, 1991). Better protection of wildlife is needed in most of these areas, where local villagers commonly keep Varecia and other lemurs as pets.
Searches for Varecia variegata variegata should be conducted in the Mangerivola, Marotandrano and Tampoketsa-Analamaitso Special Reserves, and creation of a new protected area to the west or southwest of Maroantsetra should be considered. A survey is being conducted by Hilary Simons-Morland to identify distinguishable varieties of this species. Genetic studies are needed to clarify the status of these varieties, and long-term field studies of known populations should be carried out in several protected areas. A successful captive breeding program makes reintroduction a possibility, but this should take place only after the genetic questions have been answered (St. Catherine's Workshop, 1986).
Figure 17. Distribution of ruffed lemurs (Varecia variegata).
Black-and-white ruffed lemur (Varecia variegata variegata).
Russell A. Mittermeier
A black-and-white ruffed lemur at Zoo Ivoloina, Madagascar, showing darker pattern of coloration on back and flanks.
William R. Konstant
Simpona or Diademed Sifaka
Propithecus diadema diadema
This is the most widely distributed of the P. diadema subspecies, and is found in the eastern rainforests (Fig. 18) from the Mangoro River in the south to near Maroantsera in the north (Petter et al., 1977; Petter and Petter-Rousseaux, 1979; Tattersall, 1982). The diademed sifaka's hindquarters and hindlimbs are a light golden color; its tail, forelimbs, forehead, throat and cheeks are white; and its face crown and extremities are black (Tattersall, 1982).
Propithecus d. diadema is found in the Verezanantsoro and Mantady National Parks, the Betampona and Zahamena Nature Reserves and the Ambatovaky and Analamazaotra Special Reserves (Pollock, 1975, 1984; Simons, 1984; Raxworthy, 1986, 1988; Nicoll and Langrand, 1989; Katz, 1991; Thompson and Evans, 1991). It is threatened by habitat destruction due to agricultural encroachment and logging, and by hunting (Simons, 1984). There are currently no individuals in captivity.
All of the above-mentioned parks and reserves are in need of better protection, and studies of the diademed sifakas are necessary to determine population sizes, limits of its distribution and ecological requirements (Harcourt and Thornback, 1990). Searches for the diademed sifaka should be conducted in the Mangerivola, Marotandrano and Tampoketsa-Analamaitso Special Reserves and in the region southwest of Maroantsera, where a new special reserve is proposed. A captive breeding program should also be developed.
Diademed sifaka (Propithecus diadema diadema).
Alison Richard
Milne-Edwards' sifaka (Propithecus diadema edwardsi) in the Ranomafana National Park.
Russell A. Mittermeier
Simpona or Milne-Edward's Sifaka
Propithecus diadema edwardsi
Milne-Edward's sifaka is reported to occur in the eastern rainforest (Fig. 18) southward from the Mangoro River to Manakara (Tattersall, 1982). Its color is almost entirely black or chocolate brown, with variable white patches on the back and flanks.
Studies of P. d. edwardsi have been conducted at Ranomafana National Park, where group size ranges from four to eight animals; no territorial behavior is displayed and the sifakas eat leaves, fruits and flowers (Wright et al., 1987). This subspecies is also reported from the Andohahela Nature Reserve (O'Connor et al., 1986, 1987), the Pic d'Ivohibe Special Reserve and the Midongy-Sud Classified Forest (Nicoll and Langrand, 1989). The primary threat to these and other populations is habitat destruction. There are none in captivity.
Surveys and better protection are needed for the protected areas listed above. Wright (1988) describes plans for increasing protection at Ranomafana National Park, where long-term studies should be continued. Searches for new populations should be conducted at the Andringitra Nature Reserve, which is very close to Pic d'Ivohibe. Genetic studies and fieldwork should be conducted to clarify the relationship with P. d. holomelas, and a captive breeding program should begin as soon as possible.
Coquerel's Sifaka
Propithecus verreauxi coquereli
Coquerel's sifaka inhabits deciduous and evergreen forests in northwestern Madagascar (Fig. 19), north of the Betsiboka River to Antsohihy and east to Antetemazy (Tattersall, 1982). Its color is white on the dorsal surface and maroon on the abdomen, inner thighs and forelimbs.
The average family group is four to five individuals; there is considerable overlap between home ranges and no evidence of territoriality (Petter, 1962; Richard, 1974; Albignac, 1981). Coquerel's sifaka eats leaves, buds, flowers, fruit and bark (Richard, 1974).
This animal is known to inhabit the Ankarafantsika Nature Reserve and may be present in the Bora Special Reserve as well, but habitat destruction within these protected areas is a major threat. Hunting is not yet a serious problem due to local taboos.
Increased protection is needed in both reserves in which P. v. coquereli appears to occur, as are surveys to determine its status elsewhere. Searches for other populations should be conducted in the Bongolava region between Ankarafantsika and Bora. The Duke University Primate Center maintains a small colony of this subspecies; this captive breeding program should be expanded.
Coquerel's sifaka (Propithecus verreauxi coquereli) at the Ampijoroa Station, Ankarafantsika Nature Reserve.
Russell A. Mittermeier
Decken's Sifaka
Propithecus verreauxi deckeni
Decken's sifaka is found in dry deciduous forest southwest of the Betsiboka River to near Antsalova (Tattersall, 1982), although there is some question about the boundaries of its range and that of P. v. coronatus (Figs. 12,19). Decken's sifaka is almost completely white in color (Harcourt and Thornback, 1990). There have been no studies of its behavior and ecology.
This animal is found in the Tsingy de Bemaraha and Namoroka Nature Reserves and is reportedly abundant in the Ambohijanahary Special Reserve (Nicoll and Langrand, 1989). The principal threat to its existence, even within these protected areas, is habitat destruction due to fires set annually to clear land for pasture.
All of the reserves mentioned above require better protection. Searches and surveys to confirm the existence of Decken's or crowned sifakas should be conducted in the Bemarivo, Maningozo and Kasijy Special Reserves, in the Lac Kinkony and Tsimembo Classified Forests, and in the region surrounding Soalala. Genetic research is necessary to clarify the relationship between P. v. coronatus and P. v. deckeni.
Figure 18. Distribution of Propithecus diadema diadema and Propithecus diadema edwardsi.
Figure 19. Distribution of Propithecus verreauxi coquereli and Propithecus verreauxi deckeni.
Crowned Lemur
Eulemur coronatus
The crowned lemur is diurnal and is found in dry and humid forests of northern Madagascar. Its range extends west as far as the Ankarana Massif and east to the Fanambana River (Fig. 20). Male crowned lemurs are brown with a black crown between the ears; females are gray with a light brown crown (Tattersall, 1982).
Population densities for this species vary considerably throughout its range (Arbelot-Tracqui, 1983; Fowler et al., 1989; Wilson et al., 1989; Hawkins et al., in press). It is primarily frugivorous, leaves being taken only rarely. Typical groups number five or more individuals and do not appear to be territorial towards conspecifics. They do act aggressively, however, toward other lemur species (Wilson et al., 1989).
Eulemur coronatus is reported from Montagne d'Ambre National Park and the Analamera, Ankarana and Foret d'Ambre Special Reserves (Nicoll and Langrand, 1989), but it remains threatened within these protected areas due to brush fires, logging, grazing and hunting (Wilson et al., 1988). A small number breed in captivity in European and North American zoos, as well as in Madagascar's Parc Tsimbazaza.
Better protection for crowned lemurs and other species is needed in Madagascar's northern reserves, as are surveys to determine their numbers and conservation status. The captive breeding effort for Eulemur coronatus should be expanded.
Female crowned lemur (Eulemur coronatus).
Russell A. Mittermeier
Figure 20. Distribution of crowned (E. coronatus) and mongoose lemurs (E. mongoz)
Mongoose Lemur
Eulemur mongoz
The precise limits of the mongoose lemur's range in western Madagascar remain to be determined (Fig. 20); it is also found in the Comores on the islands of Moheli and Anjouan, where it was probably introduced (Petter, 1977; Tattersall, 1976, 1982). The species is sexually dichromatic; females are grayish-brown with a dark face and white cheeks and beard, while males are gray with pale faces and reddish-brown cheeks and beard.
Both diurnal and nocturnal activity have been reported for this species (Tattersall and Sussman, 1975; Harrington, 1978). Mongoose lemurs are reported to eat flowers, fruit and leaves (Andriatsarafara, 1988) and to live in family groups averaging five individuals.
Eulemur mongoz is known to occur in only one protected area in Madagascar, the Ankarafantsika Nature Reserve, which is threatened by clearance for pastures and crops, and burning to produce charcoal (Nicoll and Langrand, 1989). In the Comores, although protected by law, mongoose lemurs are commonly captured for pets; habitat destruction by man and cyclones has also decreased the amount of forest habitat available to Eulemur mongoz (Tattersall, 1977, 1983). Captive colonies of mongoose lemurs exist in European and North American zoos. Particular success has been achieved at France's Mulhouse Zoo and at the Duke University Primare Center.
Mongoose lemur (Eulemur mongoz) family. Adult male on left and adult female at lower right.
Courtesy of the Zoological Society of Philadelphia
Better protection is needed for the lemurs of Ankarafantsika, the only reserve in which Eulemur mongoz is found. The creation of officially protected areas is probably necessary to safeguard this species in the Comores. An increased effort should be made to breed this species in captivity.
White-Collared Brown Lemur
Eulemur fulvus albocollaris
The brown lemur, Eulemur fulvus, is the most widespread of Madagascar's diurnal lemurs and is represented by six subspecies. The white-collared brown lemur (Eulemur f. albocollaris) appears to have a limited distribution between the Mananara and Faraony Rivers in southeastern Madagascar (Fig. 21). It is a rich brown-colored lemur with a white or orange beard (Tattersall, 1982).
No detailed studies have been conducted on this subspecies to date. The only protected area in which it is known to occur is the Manombo Special Reserve (Nicoll and Langrand, 1989; S. Goodman and T. Schulenberg, pers. comm.), where it is common in both parcels but hunted throughout. The reserve itself suffers from deforestation for fuel wood production. Only a few white-collared brown lemurs are found in captivity.
A survey of the Manombo Reserve is needed (St. Catherine's Workshop, 1986), and it will likely require better protection. A captive breeding program should begin as soon as possible.
Figure 21. Distribution of Eulemur fulvus albocollaris.
White-collared brown lemur (Eulemur fulvus albocollaris).
Russell A. Mittermeier
Mayotte Brown Lemur
Eulemur fulvus mayottensis
This subspecies is found in secondary forest, the only forest remaining on the island of Mayotte in the Comores (Fig. 22), and may be the result of the introduction of Eulemur f. fulvus several hundred years ago (Tattersall, 1977). The Mayotte lemur eats mainly fruit, leaves and flowers, and appears to live in loosely associated groups averaging nine to ten individuals.
Tattersall (1983, 1989) has reported an accelerating decline in the numbers of lemurs on Mayotte and an increasing exploitation of their habitat. He estimated that the population stood at 50,000 in 1975 but, due to uncontrolled development in the 1980s, it is very likely to be half that number today. Although lemurs are sometimes hunted, this is less a threat. Unfortunately, no protected areas exist on Mayotte (Harcourt and Thornback, 1990). A significant captive population exists in North American and European institutions (Wilde et al., 1988).
A study should be undertaken to determine whether the Mayotte lemur is genetically distinct from Eulemur f. fulvus. In any case, a protected area of some kind should be established on Mayotte to protect this Comores lemur population. Efforts to breed it in captivity should continue.
Figure 22. Distribution of Eulemur fulvus mayottensis and Hapalemur griseus occidentalis.
Mayotte Brown Lemur (Eulemur fulvus mayottensis).
Ian Tattersall
Western gentle lemur (Hapalemur griseus occidentalis).
Stephen Nash
Western Gentle Lemur
Hapalemur griseus occidentalis
The western gentle lemur is a slightly smaller animal, rarer and more sparsely distributed than the nominate subspecies. It is gray-brown in color and occurs in several isolated populations in western Madagascar (Fig. 22) near Lake Bemamba and in the Sambirano region (Petter and Andriatsarafara, 1987). There are also recent reports of its occurrence in the Ankarana Special Reserve in northwest Madagascar (Hawkins et al., in press). There are apparently no western gentle lemurs in captivity.
Hapalemur griseus occidentalis is a diurnal lemur living in groups of up to four individuals, and appears to be confined to forests that contain bamboo (Tattersall, 1982). It is threatened primarily by destruction of its forest habitat, in the form of fires set to promote grass growth for livestock grazing.
This subspecies is found in three protected areas: the Tsingy de Bemaraha Nature Reserve and the Ankarana and Manongarivo Special Reserves. Surveys are needed to determine the size and status of its populations in these three reserves, as are searches for new populations in surrounding areas.
Avahi or Woolly Lemur
Avahi laniger
The avahi is a small, nocturnal, folivorous lemur. There are two subspecies, Avahi l. laniger, widely distributed in the eastern rainforests, and A. l. occidentalis, with a more restricted distribution in northwestern Madagascar (Fig. 23) (Tattersall, 1982). Avahi is the smallest member of the family Indridae, averaging about 1 kg in weight, and is grayish-brown in color. Although there have been no long-term studies of this species, it is reported to be monogamous, adult females giving birth to one young each year. Woolly lemurs are also reported to be territorial; family groups of 3–5 animals aggressively defend home ranges of 1–2 hectares.
Avahi l. laniger was originally reported to occur from the latitude of Sambava in the north to that of Fort Dauphin in the south (Petter et al., 1977; Petter and Petter-Rousseaux, 1979; Tattersall, 1982). The other protected areas in which it can be found include: the Mantady and Ranomafana National Parks; the Andohahela, Andringitra, Betampona, Marojejy and Zahamena Nature Reserves; and the Ambatovaky, Ambohitantely, Analamazaotra and Anjanaharibe-Sud Special Reserves (Pollock, 1984; Simons, 1984; O'Connor et al., 1986; Rax-worthy, 1986; Nicoll and Langrand, 1989; Safford et al., 1989; Thompson and Evans, 1991). The major threat to the eastern avahi is habitat destruction for timber and agricultural land.
Eastern avahi (Avahi laniger laniger) photographed in the Zahamena Nature Reserve.
Chris Raxworthy
The western avahi (A. l. occidentalis) appears to have a disjunct distribution north and east of the Betsiboka River. It has been reported from the Ankarafantsika and Tsingy de Bemaraha Nature Reserves and the Manongarivo Special Reserve (Rax-worthy and Rakotondraparany, 1988; Thalmann, 1990). It is not clear which subspecies of A. laniger occurs in the Ankarana Special Reserve (Fowler et al., 1989; Hawkins et al., in press). The principal threat to this and other lemur species in these protected areas is habitat destruction due to fires set to promote growth of new grass.
Better protection is needed for all of the national parks and reserves listed above. A management plan for the Ankarafantsika Nature Reserve is being developed with financial assistance from the World Bank (Nicoll and Langrand, 1989). Creation of a new protected area on the Masoala Peninsula would help safeguard populations of Avahi l. laniger). There are no Avahi in captivity, and no recommendations to establish a captive breeding program at this time.
