HMW 3 - Species accounts: Eastern Woolly Lemur (Avahi laniger)

Family INDRIDAE
Genus AVAHI

Jourdan, 1834

 

1. Eastern Woolly Lemur Avahi laniger

French: Avahi laineux / German: Östlicher Wollmaki / Spanish: Lémur lanudo oriental

Other common names: Eastern Avahi, Gmelin’s Woolly Lemur

Taxonomy. Lemur laniger Gmelin, 1788, Madagascar, Betanimena country, Antongil Bay.

This species is monotypic.

Distribution. NE & E Madagascar from the Bemarivo River in the N to the Mangoro/Nesivolo rivers in the S; prior to the recent taxonomic splitting of this species, it was thought to range almost the entire length of E Madagascar’s rainforests from the Ankarana Massif in the extreme N to the Tolagnaro (= Fort-Dauphin) region in the extreme S.

Descriptive notes. Head–body 27·7–32·2 cm, tail 30·4–36·6 cm; weight 1·1–1·3 kg. The dorsal coat of the Eastern Woolly Lemur is gray-brown to reddish, becoming paler toward the rusty-red tail. The chest and abdomen are gray. The face is brownish, with a lighter band or distinct patches above the eyes and lighter fur on the cheeks and throat. Small ears are largely hidden by thick fur. Dense, short fur is tightly curled on the back.

Habitat. Tropical moist lowland, montane, and secondary forest.

Food and Feeding. The Eastern Woolly Lemur eats mainly immature leaves and buds, supplemented with fruits, flowers, and bark.

Breeding. There is no information available for this species.

Activity patterns. There is no specific information available for this species, but the Eastern Woolly Lemur is nocturnal and arboreal.

Movements, Home range and Social organization. Groups of up to five Eastern Woolly Lemurs have been reported, and home ranges of 1–2 ha are defended aggressively. Small group size indicates monogamous groupings, but longitudinal and genetic data are needed to better understand this social behavior. Research at the Analamazaotra Special Reserve suggests a social organization based on monogamous pairs and their offspring. Research at the Analamazaotra provides estimates of densities reaching 72–100 ind/km2.

Status and Conservation. CITES Appendix I. Classified as Least Concern on The IUCN Red List. The conservation status of the Eastern Woolly Lemur is in need of reconsideration. It suffers mainly from habitat destruction due to logging and slash-and-burn agriculture. It is sometimes captured opportunistically at its daytime sleeping sites or in traps baited with fruit. In some places, it is actively pursued by hunters with slingshots (Mananara-Nord) or spears (Makira). Indeed, recent data indicate that there has been a considerable upsurge in hunting of the Eastern Woolly Lemur. It occurs in four national parks (Mananara-Nord, Mantadia, Marojejy, and Zahamena), two strict nature reserves (Betampona and Zahamena), five special reserves (Ambatovaky, Ambohitantely, Analamazaotra, Anjanaharibe-Sud, and Mangerivola), and the soon-to-be-protected forests of Makira. It is still unclear which species of woolly lemur occurs in Ankarana National Park; for now, it is tentatively consider to be the Western Woolly Lemur (A. occidentalis).

Bibliography. Albignac (1981b), Andriantompohavana et al. (2007), Britt et al. (1999), Fowler et al. (1989), Ganzhorn (1988), Ganzhorn et al. (1985), Glander et al. (1992), Goodman et al. (1998), Harcourt (1991), Hawkins et al. (1990), Lei et al. (2008), Mittermeier et al. (2010), Napier & Napier (1967), Nicoll & Langrand (1989), Petter et al. (1977), Razanahoera-Rakotomalala (1981), Rumpler et al. (2011), Schmid & Smolker (1998), Sterling & Ramaroson (1996), Tattersall (1982a, 1982b), Zaramody et al. (2006).

 

2. Masoala Woolly Lemur Avahi mooreorum

French: Avahi des Moore / German: Masoala-Wollmaki / Spanish: Lémur lanudo de Moore

Other common names: Moore’s Woolly Lemur

Taxonomy. Avahi mooreorum Lei et al., 2008, Madagascar, province of Antsiranana, Masoala National Park (approximately 15’ 40° S, 49’ 57° E).

This species is monotypic.

Distribution. NE Madagascar, currently known only from Masoala National Park.

Descriptive notes. Head–body 28·4–33 cm, tail 29·4–37·2 cm; weight 920 g. The Masoala Woolly Lemur is slightly lighter and slightly longer in body length than the Eastern Woolly Lemur (A. laniger). The overall pelage color is a mottled mixture of chocolate-brown and light brown on the dorsum, gradually lightening toward the base of the tail, which is cream-colored. A distinct whitish patch, characteristic of the genus, is present on the posterior surface of each hindlimb. The ventral surface, including the underside of limbs, is gray, and the tail is reddish-brown. The head is darker than the back, and a facemask is apparent, although not as pronounced as in other eastern species of Avahi species. There is no noticeable eyebrow, but a whitish patch is present under each mandible. Ears are not readily seen, blending in as they do with the rest of the head.

Habitat. Primary rainforest.

Food and Feeding. There is no information available for this species.

Breeding. There is no information available for this species.

Activity patterns. There is no specific information available for this species, but the Masoala Woolly Lemur is nocturnal and arboreal.

Movements, Home range and Social organization. There is no information available for this species.

Status and Conservation. CITES Appendix I. The conservation status of the Masoala Woolly Lemur has not been assessed on The IUCN Red List. It occurs in the Masoala National Park. Further surveys are required to confirm the northern extent of its distribution within this protected area and possibly in intervening forest fragments between Anjanaharibe-Sud Special Reserve (where the Eastern Woolly Lemur has been confirmed to occur) and the Masoala Peninsula.

Bibliography. Andriantompohavana et al. (2007), Lei et al. (2008), Mittermeier et al. (2010).

 

3. Peyrieras’ Woolly Lemur Avahi peyrierasi

French: Avahi de Peyrieras / German: Ranomafana-Wollmaki / Spanish: Lémur lanudo de Peyrieras

Taxonomy. Avahi peyrierasi Zaramody et al., 2006, Madagas 47° 26’ E, 21° 16’ S), Fianarantsoa Province.

Based on molecular sequence data, three “types” of A. peyrierasi were detected in 2007 and 2008. Their relationships or possible species status remains to be determined. Monotypic.

Distribution. SE Madagascar, currently known from S of the Mangoro/Nesivolo river systems in the forests of Manara, Vatoalatsaka, Sangalampona, Mahasoarivo, and Ranomafana; the S extent of the distribution remains unclear. The precise distributional limits of Peyrieras’ Woolly Lemur and in particular its relationship to the Southern Woolly Lemur (A. meridionalis) and the Manombo Woolly Lemur (A. ramanantsoavanai) in the south are undetermined.

Descriptive notes. Head–body 26–31·7 cm, tail 28·5–34·4 cm; weight 1–1·1 kg. Peyrieras’ Woolly Lemur is similar to the Eastern Woolly Lemur (A. laniger), but it is slightly smaller. The dorsal fur is gray-brown, with either a gray or a white underside and a red-brown tail. Outsides of the thighs are gray-brown, and insides of the thighs are white. There are small white bands visible along interior parts of the legs and, in some cases, along the upper parts. In some individuals, the face is completely encircled by a white border of fur, and white beards and cheeks are also present.

Habitat. Primary tropical moist lowland, montane, secondary, and disturbed forest. Peyrieras’ Woolly Lemur seems to prefer intact primary forest with tall trees and high understory visibility.

Food and Feeding. Peyrieras’ Woolly Lemur eats mainly leaves, along with some flowers and fruits. At Ranomafana, the most frequent eaten leaves are from Harungana madagascariensis (Hypericaceae). They eat flowers of Erythroxylum (Erythroxylaceae) and fruits of Rheedia (Clusiaceae).

Breeding. There is no information available for this species.

Activity patterns. Peyrieras’ Woolly Lemur is nocturnal and arboreal. One study at Ranomafana showed that they rested for nearly 60% of the night, fed for 22%, traveled for 13%, and groomed for 5%.

Movements, Home range and Social organization. Recent surveys of Peyrieras’ Woolly Lemur indicated that densities at Ranomafana were twice as high in lightly disturbed primary forest than in heavily logged forest (40 ind/km2 vs. 19 ind/km2).

Status and Conservation. CITES Appendix I. Classified as Data Deficient on The IUCN Red List. Peyrieras’ Woolly Lemur occurs in the national parks of Andringitra, Midongy du Sud, and Ranomafana.

Bibliography. Andriantompohavana et al. (2007), Ganzhorn, (1988), Harcourt (1991), Herrera et al. (2011), Lei et al. (2008), Markolf et al. (2011), Mittermeier et al. (2010), Rumpler et al. (2011), Zaramody et al. (2006).

 

4. Betsileo Woolly Lemur Avahi betsileo

French: Avahi des Betsileo / German: Betsileo-Wollmaki / Spanish: Lémur lanudo de Betsileo

Taxonomy. Avahi betsileo Andriantompohavana et al., 2007, Province of Fianarantsoa, region Amoron’i Mania, district of Fandriana, Bemosary Classified Forest, approx. 20° 20’ S, 47° 33’ E, and south of the Mangoro River.

This species is monotypic.

Distribution. EC Madagascar, currently known only from the Bemosary Classified Forest (Fandriana), but limits of the distribution may ultimately prove to extend from the Mangoro River in the N to the Mananjary River in the S.

Descriptive notes. Head–body 26–31 cm, tail 28·3–34·4 cm; weight 1–1·2 kg. The Betsileo Woolly Lemur differs from other eastern woolly lemurs in having primarily light reddish-brown pelage on the upper body and on the dorsal surface of extremities. The underside is dark gray toward the midline, diffusing to a light gray ventro-laterally. The tail is mainly reddish-brown and darker on the dorsal surface than on the ventral side, which is a lighter reddish-blonde. There is a distinct facial mask, with grayish pelage under the mandible and diffuse, cream-colored eyebrow markings. The fur is thicker on the head than in other eastern species of Avahi, giving it a more rounded or oval-like appearance.

Habitat. Primary tropical rainforest.

Food and Feeding. There is no information available for this species.

Breeding. There is no information available for this species.

Activity patterns. There is no specific information available for this species, but the Betsileo Woolly Lemur is nocturnal and arboreal.

Movements, Home range and Social organization. There is no information available for this species.

Status and Conservation. CITES Appendix I. Classified as Data Deficient on The IUCN Red List. The Betsileo Woolly Lemur is not known to occur in any official protected areas but is found in Bemosary Classified Forest, which provides some degree of protection.

Bibliography. Andriantompohavana et al. (2007), Lei et al. (2008), Markolf et al. (2011), Mittermeier et al. (2010), Rumpler et al. (2011).

 

5. Manombo Woolly Lemur Avahi ramanantsoavanai

French: Avahi de Manombo / German: Manombo-Wollmaki / Spanish: Lémur lanudo de Manombo

Other common names: Ramanantsoavana’s Southern Woolly Lemur, Ramanantsoavana’s Woolly Lemur

Taxonomy. Avahi meridionalis ramanantsoavani Zaramody et al., 2006, Madagascar, Reserve of Manombo (approx. 47° 41’ E, 23° 01’ S), Fianarantsoa Province.

This species is monotypic.

Distribution. SE coastal Madagascar, currently known only from the area of its type locality, the Manombo Special Reserve and Agnalazaha Forest.

Descriptive notes. Head–body 24–31 cm, tail 33–40 cm; weight 900–1000 g. The Manombo Woolly Lemur is slightly smaller than the Eastern Woolly Lemur (A.laniger) and Peyrieras’ Woolly Lemur (A. peyrierasi). The dorsal fur is gray-brown, and the ventrum is gray—the latter overtaking laterally from a white band on the posterior legs. The tail is red-brown. The facial mask differs slightly from that of the Eastern Woolly Lemur in that the fur of some individual Manombo Woolly Lemurs is lighter, while the outline of others may be more pronounced.

Habitat. Primary tropical rainforest.

Food and Feeding. There is no information available for this species.

Breeding. There is no information available for this species.

Activity patterns. There is no specific information available for this species, but the Manombo Woolly Lemur is nocturnal and arboreal.

Movements, Home range and Social organization. There is no information available for this species.

Status and Conservation. CITES Appendix I. Classified as Data Deficient on The IUCN Red List. The only protected areas in which the Manombo Woolly Lemur occurs are the Manombo Special Reserve and the nearby Agnalazaha Forest (currently managed by the Missouri Botanical Garden). Further studies are required to determine its exact distribution, and especially its distributional limits relative to those of the neighboring Peyrieras’ Woolly Lemur and the Southern Woolly Lemur (A. meridionalis).

Bibliography. Andriantompohavana et al. (2007), Lei et al. (2008), Markolf et al. (2011), Mittermeier et al. (2010), Rumpler et al. (2011), Zaramody et al. (2006).

 

6. Southern Woolly Lemur Avahi meridionalis

French: Avahi méridional / German: Südlicher Wollmaki / Spanish: Lémur lanudo meridional

Other common names: Southern Avahi

Taxonomy. Avahi meridionalismeridionalis Zaramody et al., 2006, Madagascar, Sainte-Luce (approx. 47° 11’ E, 24° 47’ S), Toliara Province.

This species is monotypic.

Distribution. SE Madagascar, apparently restricted to Andohahela National Park and the area of Sainte-Luce.

Descriptive notes. Head–body 23–29 cm, tail 30–33 cm; weight 1–1·1 kg. The Southern Woolly Lemur is similar in size to the Eastern Woolly Lemur (A.laniger) and Peyrieras’ Woolly Lemur (A. peyrierasi). The dorsal fur is gray-brown, toning down to light gray distally, and the ventrum is gray. The tail is red-brown and darkens distally.

Habitat. Primary tropical rainforest and coastal forest.

Food and Feeding. In Sainte Luce, the Southern Woolly Lemur feeds in at least 125 individuals trees of 43 species and 26 families. In the study there, two tree species, Cynometra cloiselii (Fabaceae) and Plectronia densiflora (Rubiaceae), accounted for 41% of total feeding time, and the top six preferred plant species made up 60% of total feeding time. Young leaves and flowers are eaten only from September to December when they are available. Flower feeding was limited mostly to green sepals. The Southern Woolly Lemur has never been seen eating fruits, despite their availability. When feeding on adult leaves, the Southern Woolly Lemur selects those with high protein content.

Breeding. Females give birth in August. Infants are carried by the mother for the first four months and become independent in December–January.

Activity patterns. The Southern Woolly Lemur is nocturnal and arboreal. In a study in Sainte Luce, individuals became active 17:34–17:46 h in June–August and 18:05–18:46 h in September–December. They spent 15% of their active time feeding, 67% resting, 14% moving, and 4% carrying out other activities such as grooming themselves and other individuals.

Movements, Home range and Social organization. Southern Woolly Lemurs live in pairs that sleep together in the same tree during most days and show some degree of cohesiveness during nightly travel and foraging. Home-range size is 2·5–3·5 ha. Home ranges of male and female pair partners overlap by 70–80%. Nightly path of movement is 500–800 m. Males have larger home ranges and travel longer distances than females. In a study in Sainte Luce, male and female pair partners spent an average of 19–35% of the night in close proximity to each other. Pair cohesion was higher after females gave birth. Adult densities were 0·6–2·6 ind/ha in a study carried out in forest fragments of Sainte Luce and Mandena.

Status and Conservation. CITES Appendix I. Classified as Data Deficient on The IUCN Red List. The Southern Woolly Lemur occurs in Andohahela National Park, Sainte-Luce Reserve, and Mandena Conservation Zone, east of Tolagnaro (= Fort-Dauphin). Further studies are required to determine its exact distribution, and especially its distributional limits relative to those of the neighboring species Peyrieras’ Woolly Lemur and the Manombo Woolly Lemur (A. ramanantsoavanai).

Bibliography. Andriantompohavana et al. (2007), Lei et al. (2008), Markolf et al. (2011), Mittermeier et al. (2010), Norscia (2008), Norscia & Borgognini-Tarli (2008), Norscia et al. (2012), Rumpler et al. (2011), Zaramody et al. (2006).

 

7. Western Woolly Lemur Avahi occidentalis

French: Avahi occidental / German: Mariarano-Wollmaki / Spanish: Lémur lanudo occidental

Other common names: Lorenz von Liburnau’s Woolly Lemur, Western Avahi

Taxonomy. Avahis laniger occidentalis von Lorenz-Liburnau, 1898, Madagascar, Ambondrobe, NE of Bombetoka Bay (15° 38’ S, 46° 24’ E).

This species is monotypic.

Distribution. NW Madagascar, the core distribution seems to be to the N and E of the Betsiboka River as far as the Bay of Narinda, and this is also the species present in Ankarafantsika National Park. The isolated population farther N in the Ankarana region may also be this species, but, in between, the Sambirano Woolly Lemur (A. unicolor) inhabits both the Ampasindava Peninsula and the Sambirano region, including the Manongarivo Special Reserve.

Descriptive notes. Head–body 26·9–30·3 cm, tail 30·7–37·7 cm; weight 830–1000 g. The Western Woolly Lemur is one of the smallest woolly lemurs and, consequently, among the smallest members of the family Indriidae. It is lighter in color than the Eastern Woolly Lemur (A. laniger), with dense, tightly curled fur of the back a light to medium gray, sometimes flecked with brown or olive, becoming paler toward the rear. Typically, the tail also is gray, but it is sometimes reddish. The face, throat, and cheeks are pale, not brown as in the Eastern Woolly Lemur.

Habitat. Tropical dry deciduous primary and secondary forests.

Food and Feeding. Young leaves and buds seem to be preferred food items of the Western Woolly Lemur, probably accounting for three-quarters of the dietary intake. More than 20 plant species have been documented as food items, many of which do not appear to be common in the surrounding forests.

Breeding. There is no information available for this species.

Activity patterns. The Western Woolly Lemur is nocturnal and arboreal. Feeding bouts occur more toward the beginning and end of the night, and individuals tend to be inactive in between.

Movements, Home range and Social organization. The Western Woolly Lemur lives in cohesive pairs. In the dry forests of Ankarafantsika, groups of up to five individuals, typically consisting of an adult pair and immature offspring, occupy home ranges of 1–2 ha. Territorial defense appears to be less vigorous and less vocal than in eastern rainforest species of Avahi, with greater overlap between neighboring groups. The Western Woolly Lemur exhibits female dominance over males. Density has been estimated at 67 ind/km2 in Ankarafantsika.

Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. The Western Woolly Lemur has a limited distribution, occurs in fragmented populations, and has experienced recent declines in habitat and overall numbers. It is threatened by forest destruction, mainly due to annual burning to create new pasture, and it is also hunted to some degree. Field surveys and genetic studies of populations of Avahi at Ankarana are needed to determine the taxonomic and conservation status of woolly lemurs in that area. The Western Woolly Lemur occurs in Ankarafantsika National Park, Bora Special Reserve, and Mariarano Classified Forest. It may perhaps be the species found in the Ankarana Special Reserve, although this has not been confirmed.

Bibliography. Albignac (1981b), Bauchot & Stephan (1966), Ganzhorn (1988), Lei et al. (2008), Mittermeier et al. (2010), Nicoll & Langrand (1989), Ramanankirahina et al. (2011), Randrianambinina et al. (2003c), Razanahoera-Rakotomalala (1981), Schmid & Smolker (1998), Sterling & McFadden (2000), Thalmann (2001), Thalmann & Geissmann (2000), Warren (1997), Warren & Crompton (1997a), Zaramody et al. (2006).

 

8. Bemaraha Woolly Lemur Avahi cleesei

French: Avahi de Cleese / German: Bemaraha-Wollmaki / Spanish: Lémur lanudo de Bemaraha

Other common names: Cleese’s Woolly Lemur

Taxonomy. Avahi cleesei Thalmann & Geissmann, 2005, 18° 59’ S, 44° 45’ E, approximately 3 km east-north-east of the village of Ambalarano, western Madagascar.

This species is monotypic.

Distribution. CW Madagascar, described from the forests of Ankindrodro, Ankinajao, and Ambalarano, it is known only from the Tsingy de Bemaraha region, N of the Manambolo River; the N limit of its distribution is unclear, and there is no evidence of its occurrence between the Sambao and Mahavavy rivers or between the Mahavavy and Betsiboka rivers.

Descriptive notes. Head–body 23–31 cm, tail 32–36 cm; weight 830–980 g. The fur of the upper body and head of the Bemaraha Woolly Lemur is predominantly brown-gray and woolly, with the ventral surface light gray and thin. The tail varies from brown-gray to beige, and it is slightly reddish on the dorsal surface near its base. Characteristic white patches are found on the dorsal surface of hindlimbs. The snout is black and hairless, although fur at the corners of the mouth is whitish. The face itself is slightly paler than the blackish forehead and crown, and the triangular upward extension of the facial area onto the crown contrasts with the triangular downward extension of the crown into the facial area that is seen in the Western Woolly Lemur (A. occidentalis) and the Sambirano Woolly Lemur (A. unicolor). The fur of the forehead immediately above the face is blackish, forming a dark chevron pattern. Eyes are maroon and have black hairless eyelids.

Habitat. Subhumid, dry deciduous forest characterized by a high proportion of evergreen trees, located close to the western Tsingy precipices, in the larger Tsingy crevasses or gorges, and along small seasonal rivulets and seasonal swamps near the Bemaraha Massif. Ironically, local densities of the Bemaraha Woolly Lemur appear to correlate positively with levels of habitat disturbance, meaning that higher densities seem to be found in disturbed habitats.

Food and Feeding. Diets of the Bemaraha Woolly Lemur are presumably similar to the Western Woolly Lemur. In a ten-day study using telemetry, a group of Bemaraha Woolly Lemurs fed on buds, sprouting buds, and young leaves.

Breeding. There is no information available for this species.

Activity patterns. The Bemaraha Woolly Lemur is nocturnal and arboreal. It exhibits three distinct activity peaks at 18:00–20:00 h, 22:00–24:00 h (but variable), and 03:00–05:00 h.

Movements, Home range and Social organization. There is no specific information available for this species, but home-range size is c.2 ha.

Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. The Bemaraha Woolly Lemur is reported to be generally rare. It evidently has a very restricted distribution and is threatened by continued habitat loss. Subhumid forests at the base of the escarpment of the Tsingy de Bemaraha in particular are under continuous pressure from annual bush fires, which reduce the forests in size in many places—some to only a few meters in width. Such forests are the only habitat where Bemaraha Woolly Lemurs have been found so far. It occurs in the contiguous Tsingy de Bemaraha National Park and Strict Nature Reserve.

Bibliography. Andriantompohavana et al. (2007), Lei et al. (2008), Mittermeier et al. (2010), Thalmann & Geissmann (2000, 2005, 2006), Thalmann & Rakotoarison (1994), Thalmann et al. (1999), Zaramody et al. (2006).

 

9. Sambirano Woolly Lemur Avahiunicolor

French: Avahi unicolore / German: Sambirano-Wollmaki / Spanish: Lémur lanudo de Sambirano

Other common names: Sambirano Avahi, Unicolor Woolly Lemur

Taxonomy. Avahi unicolor Thalmann & Geissmann, 2000, Madagascar, Cacamba (= Kakamba), Ampasindava Peninsula (13° 35’ E, 47° 57’ E).

Woolly lemurs found further to north in the Ankarana Special Reserve are generally lighter in color and may represent a distinct species or subspecies. Monotypic.

Distribution. NW Madagascar, probably restricted to the Sambirano region, including the Ampasindava Peninsula; the N extent of its distribution is possibly the Sambirano River and the S limit is the Andranomalaza (= Maetsamalaza) River. The woolly lemur reported from the western slopes of the Manongarivo Special Reserve was originally thought to be the Eastern Woolly Lemur (A. laniger), but subsequently determined to be this species.

Descriptive notes. Head–body 23–31 cm, tail 26·5–30·3 cm; weight 830–920 g. The Sambirano Woolly Lemur is one of the smallest woolly lemurs and, consequently, among the smallest members of the family Indriidae. The coat is woolly, and dorsal color is a sandy brownish-gray. The tail may be a darker gray-brown or more reddish-brown, although the base tends to be lighter, perhaps even cream-colored. There is a triangular beige or cream-colored pygal patch. The ventral coat is thinner than the dorsal coat and a lighter gray. The face is only slightly paler than the head and back, with short and straight hair, giving the impression of a facial ring or mask. The snout is hairless and black, and fur at the corners of the mouth is whitish. Eyes are maroon.

Habitat. Tropical moist lowland forest from sea level to elevations of 700 m.

Food and Feeding. There is no information available for this species.

Breeding. There is no information available for this species.

Activity patterns. There is no specific information available for this species, but the Sambirano Woolly Lemur is nocturnal and arboreal.

Movements, Home range and Social organization. There is no specific information available for this species, and it is known only from a few individuals.

Status and Conservation. CITES Appendix I. Classified as Data Deficient on The IUCN Red List. Very little is known about the Sambirano Woolly Lemur, which is evidently quite rare and difficult to find. Habitat destruction for slash-and-burn agriculture and charcoal production is the principal threat to its survival. The degree to which it is hunted is not known. The only official protected area in which it occurs is the Manongarivo Special Reserve, although it is also found in the Antafondro Classified Forest, which provides some protection. To enhance the conservation of the Sambirano Woolly Lemur, consideration should be given to establishing a protected area on the Ampasindava Peninsula and extending the boundaries of the Manongarivo reserve eastward.

Bibliography. Andriantompohavana et al. (2007), Groves (2001), Mittermeier et al. (2010), Raxworthy & Rakondroparany (1988), Thalmann & Geissmann (2000).

Genus PROPITHECUS

Bennett, 1832

 

10. Verreaux’s Sifaka Propithecusverreauxi

French: Sifaka de Verreaux / German: Larvensifaka / Spanish: Sifaca de Verreaux

Other common names: White Sifaka

Taxonomy. Propithecus verreauxiGrandidier, 1867, Madagascar, Tsifanihy (N. of Cap Sainte-Marie).

A distinctive color variant was described as a subspecies and given the name majori by W. Rothschild in 1894, but most experts now consider it a melanistic form of Verreaux’s Sifaka. Monotypic.

Distribution. SW & S Madagascar, in the W up to the Tsiribihina River, in the SE it is found near to (just N of) Tolagnaro (= Fort-Dauphin) in the Nahampoana Reserve, although it was probably introduced there; the distributional limit in the SE is the transitional and spiny forest patches of the Mangatsiaka Parcel of Andohahela National Park.

Descriptive notes. Head–body 40–48 cm, tail 50–60 cm; weight 2·9 kg. Verreaux’s Sifaka is one of the smaller species of Propithecus. Its pelage is long and thick, with the predominant coat color, including the tail, being generally a lustrous white, often with a silvery or golden tint on the back and flanks. This contrasts sharply with the black face, muzzle, hands, and feet. It also contrasts with the dark reddish-brown, chocolate-brown, or black crown that extends down the nape of the neck in both sexes. The underside is sparsely-furred, exposing the dark skin of the belly and giving the abdomen a grayish appearance. The snout is deep and narrow, and ears are slightly tufted with white. Males have a reddish-brown patch on the upper chest that is associated with a sternal gland, visible at the base of the throat. Juveniles are entirely white except for a dark brown spot on the crown and occasionally a rufous wash on the ventral side. A variant, originally considered a subspecies (majori), is also predominantly white, including its cheeks, ears, and forehead, but it has a chocolate-brown head cap and is brownish to brownish-black on the chest, back, inside of the limbs, and tail, except for its white tip. This variant is almost always found in groups of normally colored Verreaux’s Sifakas. Entirely white individuals also are occasionally observed with normally colored animals (e.g. in Berenty Reserve), and other variants may perhaps have maroon patches on the back, belly, or limbs.

Habitat. Typically tropical dry lowland and montane forest from sea level to elevations of 1300 m; also semi-arid spiny bush, brush-and-scrub thickets, deciduous gallery forest, riparian forest, and humid forests at low elevations.

Food and Feeding. Diets of Verreaux’s Sifaka are seasonally variable but consist mainly of young leaves, fruits, and flowers, supplemented with seeds, bark, dead wood, and termite soil. Leaves are the most important food item during the dry season, and fruits are most important during the wet season when parts of fewer plant species are eaten. Most seeds eaten are destroyed, meaning that Verreaux’s Sifaka is at least partly a seed predator. Survival of Verreaux’s Sifaka in Didiereaceae forest—including periods of severe drought—suggests that they do not need to drink. Water may be obtained indirectly during the dry season by eating bark and cambium of Operculicarya decaryi (Anacardiaceae).

Breeding. Reproduction of Verreaux’s Sifaka is seasonal and synchronized within groups and between them. Mating takes place in January–February. Females are receptive for only about a single day per year. A single dominant male monopolizes paternity in each group at Kirindy Forest, whereas paternity by extragroup males is common at Beza-Mahafaly Special Reserve. A single young is born in August–September after gestation of 162–170 days. The mother carries her infant on her belly for the first three months, at which point it shifts to her back. All group members interact with infants, with grooming, playing, carrying, and nursing being the most common alloparental behaviors. Infanticide has been reported. Young are almost completely independent at about six months. Age of sexual maturity varies by habitat; for example, in the spiny forests of Beza-Mahafaly, less than one-half of the females reproduced by six years of age, whereas three-year-old females at Berenty Reserve are routinely seen with newborns. Males are sexually mature at c.2·5 years old.

Activity patterns. Verreaux’s Sifakas are diurnal and mainly arboreal. They have the extraordinary ability to leap from one cactus-like Didiereaceae trunk to another—somehow managing to avoid the very hard, sharp spines of these tall, thin plants. Individuals regularly descend to the ground, where they proceed by a series of upright, usually slightly sideways, bipedal hops with arms raised above the head.

Movements, Home range and Social organization. Verreaux’s Sifakas tend to live in small to medium-sized, multimale–multifemale groups of 2–14 individuals (average 5–6). There are usually more males in a group than females. Home ranges may exceed 10 ha but are often much smaller. Rather than strict territorial boundaries, core areas and food resources of overlapping home ranges are defended against neighboring groups. Females appear to be dominant over males. Males fight with one another for dominance only during the mating season. Males disperse. Reported densities include 47 ind/km2 in the degraded forests of Belaoka Marovato, 150–200 ind/km2 at Berenty Reserve, and 400–500 ind/km2 at Antserananomby.

Status and Conservation. CITES Appendix I. Classified as Vulnerable on The IUCN Red List. Despite the large distribution of Verreaux’s Sifaka, the two principal habitats upon which it depends for survival—spiny forest and riparian or gallery forest—are under continual threat because of logging, slash-and-burn agriculture (especially for corn plantations), and charcoal and firewood collection. Although hunting is illegal and “fady” (taboo) to several of the tribes living within its distribution (e.g. Antandroy and Mahafaly), it is hunted for food by other tribes (e.g. Sakalava) and immigrants to the region. In the Isalo region, Verreaux’s Sifaka is known as “sifaka-bilany” (sifaka of the cooking pot), but it is unclear whether this is because of its popularity as a food item or because of the sooty black appearance of individuals from this part of its range. It occurs in four national parks (Andohahela, Isalo, Tsimanampetsotsa, and Zombitse-Vohibasia), two special reserves (Andranomena and Beza-Mahafaly), and two private reserves (Analabe and Berenty Reserve). Populations of the Verreaux’s Sifaka also are found in the Kirindy Forest (part of the Menabe-Antimena protected area) and a number of unprotected classified forests and forest reserves. Significant variation in densities has been noted in different forest types; in general, densities are lower in areas of degraded habitat but even very small forest patches can support sizeable numbers of Verreaux’s Sifaka.

Bibliography. Benadi et al. (2008), Brockman (1999, 2003), Brockman & Whitten (1996), Brockman et al. (1998), Carrai & Lunardini (1996), Carrai et al. (2003), Erkert & Kappeler (2004), Fenn et al. (1999), Fichtel (2004), Fichtel & Kappeler (2002), Fichtel & van Schaik (2006), Goodman (1999), Goodman & Raselimanana (2003), Goodman et al. (2004), Hawkins (1999), Howarth et al. (1986), Jolly (1966, 1972, 1977), Jolly et al. (1982b), Kappeler (1991), Kappeler & Schäffler (2008), Karpanty & Goodman (1999), Lawler et al. (2003, 2005), Lewis & Kappeler (2005a, 2000b), Lewis et al. (2003), Mertl-Millhollen (1979), Mittermeier et al. (1994, 2010), Nicoll & Langrand (1989), Norscia & Pelagi (2008), Norscia et al. (2005), O’Connor (1987), O’Connor et al. (1986, 1987), Oda (1998), Oda & Masataka (1996), Petter et al. (1977), Raharivololona & Ranaivosoa (2000), Ralisoamalala (1996), Rasoarimanana (2005), Rasoloarison et al. (1995), Richard (1974a, 1974b, 1976, 1977, 1978a, 1985, 1987, 1992, 2003), Richard & Nicoll (1987), Richard et al. (1991, 1993, 2000, 2002), Rumpler et al. (2011), Simmen et al. (2003), Sussman & Richard (1986), Sussman et al. (1987, 2003), Tattersall (1982a, 1982b), Trillmich et al. (2004).

 

11. Decken’s Sifaka Propithecusdeckenii

French: Sifaka de der Decken / German: Von-der-Decken-Sifaka / Spanish: Sifaca de Decken

Other common names: Von der Decken’s Sifaka

Taxonomy. Propithecus deckenii Peters, 1870, Madagascar, Kanatsy.

Some experts consider this species to be synonymous with P. coronatus. To the north in the coastal forests between the Mahavavy and Betsiboka rivers, the geographic separation between P. deckenii and P. coronatus appears clear, but in the upper reaches of the Mahavavy River, populations of the two species apparently hybridize, and there are individuals with intermediate coloration on islands in the middle of the Mahavavy. More confusing is the situation in the forests of the Bongolava Massif, far inland and to the south-east, where individuals with color patterns characteristic of both species have been observed. Furthermore, according to fairly recent surveys, populations representing both species can be found at a number of sites. Needless to say, the taxonomic status of these two forms and their relationship to one another require further investigation, especially in the field. Monotypic.

Distribution. CW Madagascar, found mainly between the Manambolo and Mahavavy rivers, the S limit of its distribution does not extend to the Tsiribihina River (which marks the N limit of Verreaux’s Sifaka, P. verreauxi). This species and the Crowned Sifaka (P. coronatus) are often found in the same areas, although the latter tends to be found further inland than the more coastal Decken’s Sifaka.

Descriptive notes. Head–body 42–48 cm, tail 50–60 cm; weight 2·6–2·9 kg. Decken’s Sifaka is a small species of Propithecus. The dorsal coat is creamy-white, often with faded silvery, golden, or pale brown tints on the neck, shoulders, back, and limbs. Some individuals have dark areas on the head and chest. The face is naked and black, although usually with a patch of white fur running across it. Bony pockets on either side of the muzzle give Decken’s Sifaka a rather blunt-nosed appearance, but not as extreme as in the Crowned Sifaka.

Habitat. Dry deciduous forest patches. Decken’s Sifaka seems to be fairly resilient to habitat degradation, and individuals have even been observed in Eucalyptus (Myrtaceae) trees in the middle of Soalala town.

Food and Feeding. There is no information available for this species.

Breeding. There is no information available for this species.

Activity patterns. There is no specific information available for this species, but Decken’s Sifaka is diurnal and mainly arboreal.

Movements, Home range and Social organization. There is no specific information available for this species, but it is believed to occur in groups of 2–10 individuals.

Status and Conservation. CITES Appendix I. Classified as Vulnerable on The IUCN Red List. Forests in the distribution of Decken’s Sifaka are already highly fragmented, and continued habitat loss is the greatest threat to its survival. Habitat is burned to provide pasture for livestock and cut for charcoal production. Hunting is rare because Decken’s Sifaka is protected by a very strong taboo over much of its range, leading them to become very tame. Nevertheless, if the taboo were to break down for whatever reason, Decken’s Sifaka could disappear very rapidly. It is occurs in three national parks (Baie de Baly, Tsingy de Bemaraha, and Tsingy de Namoroka), Tsingy de Bemaraha Strict Nature Reserve, and four special reserves (Ambohijanahary, Bemarivo, Kasijy, and Maningoza). It is also found in at least one classified forest (Tsiombikibo), which provides some degree of protection.

Bibliography. Curtis et al. (1998), Garbutt (2007), Hawkins et al. (1998), Mittermeier et al. (2010), Nicoll & Langrand (1989), Petter & Peyrieras (1972), Petter et al. (1977), Randrianarisoa et al. (2000, 2001a), Rumpler et al. (2011), Tattersall (1982a, 1982b, 1986), Thalmann & Rakotoarison (1994), Thalmann et al. (1999, 2002).

 

12. Crowned Sifaka Propithecus coronatus

French: Sifaka couronné / German: Kappensifaka / Spanish: Sifaca coronado

Taxonomy. Propithecus coronatus Milne-Edwards, 1871, Madagascar, Boueny Province.

Some experts consider this species to be synonymous with P. deckenii, with which it may occasionally hybridize along the upper reaches of the Mahavavy River and elsewhere. Collections made in the forests of Ambararatabe (to the west of the Mahavavy River) and sightings along the Bongolava Massif (west of Tsiroanomandidy) also appear to include individuals representing both species, as well as P. verreauxi, but reports of the co-occurrence of this species with P. deckenii along the lower course of the Mahavavy have never been confirmed. Monotypic.

Distribution. NW Madgascar, found discontinuously in the coast S of the Manambolo River, from where the distribution swings inland to the uplands before touching the coast again N and E of the Mahavavy River; reports of the Crowned Sifaka as far S as the Sakay River, as far E to Andanotongo, and SE beyond Tsiroanomandidy suggest that its distribution is more complicated than originally believed. Decken’s (P. deckenii) and Crowned sifakas are often found in the same areas, although the Crowned Sifaka tends to be found further inland than the more coastal Decken’s Sifaka. In general, the boundary between these two species is the Mahavavy River, while the Betsiboka River separates the Crowned Sifaka from Coquerel’s Sifaka (P. coquereli).

Descriptive notes.Head–body 39–45 cm, tail 48–57 cm; weight 3·2–3·7 kg. The Crowned Sifaka is a medium-sized species of Propithecus. The coat, including hindlimbs and tail, is creamy-white and variably tinted golden-yellow to golden-brown on the upper chest, shoulders, and upper forelimbs. It contrasts strongly with the chocolate-brown to black crown, forehead, cheeks, neck, and throat. The muzzle is blunt and rounded, even bulbous, in form, which readily distinguishes it from all others sifakas. The face is naked and black, but sometimes there is a patch of white fur across the bridge of the nose and very often slight white tufting around the ears.

Habitat. Tropical dry deciduous lowland forests from sea level to elevations of 700 m. The Crowned Sifaka occasionally enters mangrove forests.

Food and Feeding. Diets of the Crowned Sifaka consist mainly of leaves, supplemented with buds, unripe fruits, flowers, bark, and dead wood. In Badrala in the Antrema area, it consumes parts of at least 60 plant species in 32 families. In the wet season, only seven species made up 75% of the study group’s diet. Flower consumption increased in the wet season, whereas the bulk of the diet was mature leaves and fruits in the dry season.

Breeding. There is no information available for this species.

Activity patterns. The Crowned Sifaka is diurnal and mainly arboreal. In one study spanning the dry and wet seasons, individuals spent 36–39% of their active time feeding and foraging, 47–50% resting, and 5–7% traveling. They traveled more during the dry season than the wet season.

Movements, Home range and Social organization. Field studies of the Crowned Sifaka have been limited and short-term. At Anjamena, group size is 2–8 individuals, and home-range size is 1·2–1·5 ha, with territories defended aggressively against neighboring groups. Mean group size in Badrala is 4·3 individuals, with groups usually consisting of 1–3 breeding adult males, 1–4 breeding adult females, and 1–4 immature offspring. Densities of 48 groups/km2 and 173 ind/km2 have been recorded in Anjamena. Density of 300 ind/km2 was recorded at Badrala.

Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. Habitat loss is the principal threat faced by the Crowned Sifaka. Forest cover within its distribution has declined dramatically over the last few decades from burning for pasture and cutting for charcoal production. There is also some live capture for the illegal pet trade. It may occur in the Ambohijanahary and Kasijy special reserves and is found in the proposed Katsepy Reserve, which is already considered a sacred site by local people and thereby affords some level of protection. It also occurs in the Antrema Forest Station, which is the subject of a conservation program. A few classified forests within the distribution of the Crowned Sifaka may also harbor viable populations. It was present on the high plateau in the Ambohitantely Special Reserve until about the mid-20th century, but it is now absent from this small isolated locale.

Bibliography. Curtis et al. (1998), Groves (2001), Mittermeier et al. (2010), Muller et al. (2000), Petter & Andriatsafara (1987), Petter & Peyrieras (1972), Petter et al. (1977), Pichon et al. (2010), Randrianarisoa et al. (2001a, 2001b), Rumpler et al. (2011), Tattersall (1982a, 1982b, 1986), Thalmann et al. (2002).

 

13. Coquerel’s Sifaka Propithecuscoquereli

French: Sifaka de Coquerel / German: Coquerel-Sifaka / Spanish: Sifaca de Coquerel

Taxonomy. Cheirogalus [sic] coquereli Grandidier, 1867, Madagascar, Morondava.

This species is monotypic.

Distribution. NW Madagascar from near Bealalana in the N to the Betsiboka River, S limit is reportedly Ambato Boeni, and E boundary is near Antetemasy (just W of Befandriana Nord).

Descriptive notes. Head–body 42–50 cm, tail 50–60 cm; weight 3·7 kg. Coquerel’s Sifaka is a medium-sized species of Propithecus, with long dense fur and a low narrow snout. The dorsal color is mostly white, including the head and tail, with prominent chestnut-brown to maroon patches covering the chest and anterior and interior aspects of forelimbs and hindlimbs. There also are occasionally similarly colored or even silvery patches on the base of the back. The skin of the muzzle and face is bare and black, except for a white patch of fur extending across the bridge of the nose. Ears are small, black, and naked, and they are visible above the white fur of the head and cheeks. Eyes are yellow. An aberrant population has been reported in which individuals have the patterning described above but with a charcoal gray-black coloration instead of reddish-brown. The dark gray also extends somewhat on to the forehead of this variant.

Habitat. Commonly in mixed deciduous dry and semi-evergreen lowland forests from near sea level up to elevations of 300 m and often in brush-and-scrub and secondary forest immediately adjacent to primary forest. Coquerel’s Sifaka has also been seen in coastal mangroves in the Bay of Mahajamba.

Food and Feeding. Diets of Coquerel’s Sifaka consist mainly young leaves, flowers, fruits, bark, and dead wood in the wet season and mature leaves and buds in the dry season. As many as 98 different plant species have been found in its diet.

Breeding. Births of Coquerel’s Sifaka are clustered in June–July. Usually a single young is born after gestation of c.162 days. Infants cling to their mother’s chest for the first month or so and then transfer to her back. All group members interact with young; grooming, playing, carrying, and nursing are the most common alloparental behaviors. Complete independence occurs by about six months of age, and adult size is reached in one year. Sexual maturity in both sexes occurs at c.2·5 years.

Activity patterns. Coquerel’s Sifaka is diurnal and mainly arboreal. As with other western sifaka species, Coquerel’s Sifaka regularly descends to the ground. Interestingly, its ground locomotion is somewhat different from that of Verreaux’s Sifaka (P. verreauxi), further to the south. Whereas the latter usually bounds along the ground in a sideways position, Coquerel’s Sifaka leaps forward in a kangaroo-like fashion.

Movements, Home range and Social organization. In the forests of Ankarafantsika National Park, Coquerel’s Sifaka occurs in small mixed groups of 3–10 individuals, with home ranges of 4–9 ha, and with estimated densities approaching 60 ind/km2.

Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. Coquerel’s Sifaka is severely threatened by habitat destruction and hunting, and its restricted distribution makes it particularly susceptible to these threats. Slash-and-burn agriculture and annual burning to generate new pasture for livestock are the principle causes of forest loss, but trees in this part of Madagascar are also cut to produce charcoal. All of these practices are problematic even in officially protected areas. Hunting for food is an increasing; local traditions place taboos on the practice, but immigration to the region is changing these beliefs. The only protected areas in which Coquerel’s Sifaka is known to occur are the Ankarafantsika National Park and Bora Special Reserve. Unfortunately, hunting pressure on sifakas is significant in Ankarafantsika, and Bora has become seriously degraded. Populations have also been reported from the forests of Anjiamangirana, Anjajavy, the Narindra Peninsula, and Mariarano, which should be considered for protection.

Bibliography. Albignac (1981a), Eaglen & Boskoff (1978), Ganzhorn (1988), Garcia & Goodman (2003), Groves (2001), Kappeler (1991), Kubzdela et al. (1992), Mittermeier et al. (2006, 2010), Nicoll & Langrand (1989), Petter (1962), Poorman (1983), Randrianambinina et al. (2003c), Ravosa et al. (1993), Richard (1974b, 1976, 1978a, 1978b, 1987), Simons (1988), Tattersall (1982a, 1982b).

 

14. Tattersall’s Sifaka Propithecus tattersalli

French: Sifaka de Tattersall / German: Goldkronensifaka / Spanish: Sifaca de Tattersall

Other common names: Golden-crowned Sifaka

Taxonomy. Propithecus tattersalli Simons, 1988, Madagascar, dry forest about 6–7 km north-east of Daraina, Antseranana province (13° 9’ S, 49° 41’ E).

This species was first discovered in 1974, when I. Tattersall sighted groups of animals north of Vohémar and provisionally identified them as a variant of the Silky Sifaka (then P. diadema candidus). More than a decade passed before the species was finally described in 1988 when E. Simons named it in honor of Tattersall. Monotypic.

Distribution. Restricted range in NE Madagascar, limited to an extremely small area in the Ampandraha, Madirabe, and Daraina districts bounded by the Loky River in the N and the Manambato River in the S; its distribution is centered on the town of Daraina and covers c.245,000 ha.

Descriptive notes. Head–body 45–47 cm, tail 42–47 cm; weight 3·4–3·6 kg. Tattersall’s Sifaka is a medium-sized species, similar in size to the sifakas in the southern and western dry forest. The fur is notably short and sparse, being mainly creamy-white dorsally and on the shoulders, upper arms, and genital region, with a wash of golden-orange on the chest and rump. Forearms and tops of the legs often are pale orange. The crown is bright golden-orange, often extending to the shoulders and separated from the bare black skin of the face by a white ruff. Ears are prominent and black, sporting distinctive white tufts that give the head a somewhat triangular appearance. Eyes are yellow-orange.

Habitat. Primary dry deciduous, gallery, and semi-evergreen forest patches at 50–700 m above sea level and coastal/littoral forest. Forests in the distribution of Tattersall’s Sifaka are highly fragmented and isolated by extensive, human-altered degraded grasslands, dry scrub, and farmland. Most are deciduous secondary forsts similar in composition to transitional dry forests of western Madagascar. Forty-five of 75 forest tracts identified by researchers were inhabited by Tattersall’s Sifaka. Most of the year is dry, with the majority of the rainfall (200 cm annually) occurring in December–March.

Food and Feeding. Diets of Tattersall’s Sifaka are mainly composed of young leaves, flowers, seeds, and fruit. At Daraina, food availability varies during the year based on seasonal patterns of rainfall. Availability of ripe fruits, flowers, and immature leaves peaks during the rainy season (December–March). In the dry season, Tattersall’s Sifaka is forced to feed on less nutritious mature leaves, unripe fruits, and possibly tree bark. In one study population of Tattersall’s Sifaka, 60% of individuals had a larval parasite potentially detrimental to their health.

Breeding. The mating season of Tattersall’s Sifaka is in late January. Usually only one adult female per group breeds. Copulations only occur over a 24-hour period when the female is in estrus and fertile. Fertility is signaled by about a ten-hour long, externally visible pink genital swelling in females. Males’ testicles enlarge as the breeding season nears. Copulation is brief, often only 30–90 seconds and consists of a single mount. Single young are born from late June to August after gestation of 165–176 days. Newborns are sparsely furred. Weaning typically takes place in December. All group members interact with infants; grooming, playing, carrying, and nursing are the most common alloparental behaviors. Mothers wean their infants five months after birth, just before the subsequent breeding season. Sexual maturity in both sexes occurs at c.2·5 years of age.

Activity patterns. Tattersall’s Sifaka is arboreal and primarily diurnal, although individuals may become crepuscular during the rainy season. Groups sleep in high emergent trees at night.

Movements, Home range and Social organization. Tattersall’s Sifaka lives either in monogamous pairs or in multimale–multifemale groups of 3–10 individuals (average of five), the latter usually with two adults of each sex. Home-range size is 9–12 ha. Mean daily path of movement is 462–1077 m. Females are dominant over males. Males may change groups during the mating season. Recent density estimates are 10–23 ind/km2.

Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. Tattersall’s Sifaka is threatened by slash-and-burn agriculture, uncontrolled grass fires, collection of timber for housing and fuel, logging of precious hardwoods, and gold mining. In recent years, hunting for food has also become a serious problem, especially by itinerant gold miners who, unlike the local people, do not consider the animals “fady” (taboo). Tattersall’s Sifaka occurs in the 57,000-ha Loky-Manambato protected area in the Daraina region, which was established primarily for its benefit in 2005. The total world population is estimated to be over 18,000 individuals, and dispersal still seems to be possible among most forest fragments within its distribution. No Tattersall’s Sifakas are held in captivity.

Bibliography. Coffmann (1990), Garbutt (2007), Lehman et al. (2005a), Meyers (1993, 1996), Meyers & Ratsirarson (1989), Meyers & Wright (1993), Mittermeier et al. (2006, 2010), Pochron & Wright (2002, 2003), Quéméré et al. (2010a, 2010b), Randrianarisoa et al. (1999), Ravosa et al. (1993), Simons (1988), Tattersall (1982a, 1982b), Vargas et al. (2002).

 

15. Diademed Sifaka Propithecusdiadema

French: Sifaka à diadème / German: Diademsifaka / Spanish: Sifaca de diadema

Other common names: Diademed Simpona

Taxonomy. Propithecus diadema Bennett, 1832, Madagascar.

This species seems to form a clinal gradient with P. edwardsi at the southern limit of its range. A highly aberrant population, discovered in 1999 in the forests of Tsinjoarivo in south-central Madagascar, was originally believed to represent a different subspecies (marshi). It is somewhat smaller than typical P. diadema, with much longer canine teeth and different and quite variable markings (including at least one all-black individual), indicating that these animals may be a hybrid of P. diadema × P. edwardsi. A preliminary genetic study did not find sufficient evidence to warrant its recognition as a new taxon, but a more recent study indicated that it is distinct. Recent molecular analysis as part of a study on the phylogenies of the Indriidae revealed two well-supported mtDNA clades in P. diadema, one of which is likely to be more closely related to P. edwardsi than to the other P. diadema clade. Genetic distances between the two clades would warrant splitting the species into two taxa. Nevertheless, in the absence of data from nuclear genetic markers and with the two clades not showing any differences in their karyotypes, the authors of that study proposed male introgression as a possible cause for this paraphyly and did not delineate a new taxon. Monotypic.

Distribution. NE & E Madagascar from the Mananara River in the N to the Mangoro and Onive rivers in the S; although the precise distributional limits are unknown, the Diademed Sifaka is thought to be the most widely distributed of the sifakas, occurring throughout the Madagascar’s E rainforests. Historically, its distribution extended farther north to just south of the Antainambalana River, but it has not been found in recent years during fairly extensive studies of that region.

Descriptive notes. Head–body 50–55 cm, tail 44–50 cm; weight 5·7–6·8 kg. The Diademed Sifaka is the largest of the sifakas, and it rivals the slightly larger Indri (Indri indri) for being the largest living prosimian. It is also one of the most colorful and attractive, with its characteristic long silky fur and prominent “diadem” fringe across the forehead. Pelage of the forehead, cheeks, and throat is white with silvery or golden tints. Shoulders and upper back are slate-gray, and the lower back lightens to silver-gray. The ventral coat is typically white to pale gray. The crown is black, and this color sometimes extends to the nape. Arms and legs are orange to yellow-gold, hands and feet are black, and flanks and the tail are pale gray to white (the latter often with a golden-white base). The head is notably small and narrow. The muzzle is short, the bare face is dark gray to black, and eyes are reddish-brown. Males have a large reddish-brown cutaneous gland in the middle of their throat, and a perianal patch of similar color is also believed to be glandular. Juveniles resemble adults but they are paler, with a yellower frontal band and lighter yellow limbs.

Habitat. Primary highland rainforest at elevations of 200–1600 m. although elevations above 800 m are preferred. The Diademed Sifaka appears to require intact forest, but it can be found in small forest fragments.

Food and Feeding. Diets of the Diademed Sifaka consist mainly of ripe fruits, seeds, flowers, and young leaves; respective proportions of each vary according to seasonal abundance. Some bark also is consumed. The number of plant species in the diet each day often reaches 25 or more, including fruits of two preferred plants whose seeds contain significant levels of alkaloids. The Diademed Sifaka eats higher energy foods compared with sympatric Indri. Individuals are reported to search and smell large areas of leaves on the forest floor to locate hidden flowers of the subterranean parasitic plants Langsdorffia (Balanophoraceae) and Cytinus (Cytinaceae). This behavior is evidently learned by observing conspecifics.

Breeding. Females are sexually receptive for only one day per year, sometime in November–January. A single young is born in May–July, after gestation of 179 days. Infant mortality is high. Individuals may live up to 20 years in the wild.

Activity patterns. The Diademed Sifaka is diurnal and arboreal. In one study, individuals spent 49·4% of their time resting, 37·8% feeding, 2·4% in social behavior, 5·1% moving, and the remainder in other activities.

Movements, Home range and Social organization. The Diademed Sifaka has only been studied once for a significant period of time. It lives in female-dominated, multimale–multifemale groups of eight or more, consisting of 1–3 adult females and 1–2 adult males, with only one breeding pair in each group. Exclusive territories of 20–50 ha are maintained by scent marking, although there is little aggression over boundaries. Compared with the sympatric, similar-sized Indri, the Diademed Sifaka spends more time actively patrolling and defending its territories. Mean daily path of movement is 987–1629 m for the Diademed Sifaka compared to 774 m for the Indri. Males emigrate at age five to a neighboring group; females may either emigrate or remain in their natal group.

Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. The principal threat to the survival of the Diademed Sifaka is habitat loss from slash-and-burn agriculture, logging, and mining. Hunting for its meat and fur also has a very serious impact in many parts of its range, even within existing protected areas. As a result, it is now absent from many areas where it formerly occurred. It occurs in three national parks (Mananara-Nord, Mantadia, and Zahamena), two strict nature reserves (Betampona and Zahamena), three special reserves (Ambatovaky, Mangerivola, and Marotandrano), and Anjozorobe-Angavo protected area. It has also been successfully reintroduced into the Analamazaotra Special Reserve at Andasibe (= Périnet), where it was extirpated 30–40 years ago. The reintroduction, carried out by E. Louis Jr. and a team of Malagasy researchers, took place in 2007 with individuals rescued from the nearby Ambatovy mine site. Additional populations have been identified in two classified forests (Andriantantely and Tsinjoarivo), Marokitay Forest Reserve, and unprotected forests of Anosibe an’ala, Didy, Iofa, Maromiza, and Sandranantitra. Tsinjoarivo Classified Forest has already been recommended as a new protected area, based on the presence of the unusual population of Diademed Sifakas found there.

Bibliography. Andriaholinirina et al. (2004), Andriamasimanana et al. (2001), Britt et al. (1999), Garbutt (2007), Glander et al. (1992), Green & Sussman (1990), Groves (2001), Irwin (2006, 2007a, 2007b, 2008a, 2008b), Irwin & Ravelomanantsoa (2004), Irwin et al. (2007), Lehman & Wright (2000), Lehman et al. (2005a), Mayor et al. (2004), Mittermeier et al. (1992, 2006, 2010), Nicoll & Langrand (1989), Petter (1962), Petter & Petter-Rousseaux (1979), Petter et al. (1977), Powzyk (1997), Powzyk & Mowry (2003), Rumpler et al. (2011), Simons (1988), Smith & Jungers (1997), Tattersall (1982a, 1982b).

 

16. Milne-Edwards’ Sifaka Propithecus edwardsi

French: Sifaka de Milne-Edwards / German: Milne-Edwards-Sifaka / Spanish: Sifaca de Milne-Edwards

Other common names: Milne-Edwards’ Simpona

Taxonomy. Propithecus edwardsi Grandidier, 1871, Madagascar, west of Mananjary.

There appears to be a clinal gradient between this species and P. diadema. There are apparent intermediates between P. diadema and P. edwardsi at Anosiben’Ifody on the left bank of the upper Mangoro River, whereas typical P. edwardsi occurs further south along this river where it turns east toward the sea. The melanistic variant, (holomelas), long recognized as a subspecies of P. diadema, is now generally considered synonymous with this species. Almost entirely black except for a dark brown patch at the base of the tail, and perhaps slightly smaller in size on average than typical P. edwardsi, it was formerly found in the forests of Nandihizana, apparently in groups of normally colored P. edwardsi. Unfortunately, sifakas are now gone from this region, and no similarly colored animals have been found elsewhere, making it difficult to determine whether or not it was indeed a distinct taxon. Monotypic.

Distribution. CE Madagascar, the Mangoro and Onive rivers are the N limits of the present range, and the Rienana River in Andringitra National Park is the S boundary. The former distribution probably extended somewhat northward and further south to the Manampatrana River, but populations in those regions appear to have been extirpated.

Descriptive notes. Head–body 42–52 cm, tail 41–48 cm; weight 5·5 kg. Milne-Edwards’ Sifaka is a large, heavily-built species of Propithecus. The dorsal coat is dense and dark, varying from chocolate-brown to almost jet-black on the head, upper body, limbs, and tail. Bilateral whitish patches of varying extent grade into the darker surrounding fur on the back and flanks, sometimes meeting along the spine. The ventral coat is equally dark, sometimes paler around the upper part of the chest, but it is less dense than the dorsal coat. The face and ears are bare, and the skin is dark gray to black, with the ears barely discernible above the dark fur of the head. Eyes are orange-red. Juveniles resemble adults.

Habitat. Primary and slightly degraded secondary rainforest at middle to high elevations of 600–1600 m.

Food and Feeding. Diets of Milne-Edwards’ Sifaka consist mainly of young leaves, ripe fruits, seeds, and flowers, but they also eat some mature leaves, bark, subterranean fungus, and soil. About a dozen different plant species are sampled each day. Dietary composition appears to vary significantly, however, not only from month to month but also from year to year.

Breeding. Copulations of Milne-Edwards’ Sifaka only occur over a single 24-hour period when the female is in estrus and fertile, but females within a group are fertile on different days. Fertility is signaled by about a ten-hour long, externally visible pink genital swelling. Males’ testicles enlarge as the breeding season nears. Copulation is brief (often only 30–90 seconds) and consists of a single mount. A single young is born in June–July, after about a six-month gestation (average 179 days). Births in a group tend to be synchronous, with six of eight births in one study occurring during just one week. Average birth weight is 156 g. Infants transfer from their mother’s belly to her back after about 3–4 weeks, riding comfortably there at about two months of age. At night, mothers continue to sleep with their offspring until they are about two years old. Allogrooming and self-grooming commence in the fourth week of life. Play starts by week six and most often occurs between the infant and a juvenile. The infant spends more time on its mother than on other group member, and the mother is the main care provider. By one year, juveniles weigh 2·5 kg. Predation, especially by the Fosa (Cryptoprocta ferox), is a significant cause of death in infants. Infant mortality is high, with over one-half of female infants dying in their first year of life and only a one-quarter surviving to reproductive age. Infanticide has been well documented in Milne-Edwards’ Sifaka, particularly by unrelated adult male sifakas, which may serve as a male strategy to induce heat in females and shorten the interbirth interval from c.1·5 years (typical of mothers with infants surviving one year). Mothers wean their offspring six months after birth, which corresponds to just slightly before the subsequent breeding season. Age at first birth for females is about four years old, with an increase in fertility at six years. Males reach reproductive maturity at five years. About one-half of male and female Milne-Edwards’ Sifakas emigrate from their natal group, with females leaving before maturity and males both before and after maturity. One individual reached an age of 27 years.

Activity patterns. There is no specific information available for this species, but Milne-Edwards’ Sifaka is diurnal and arboreal.

Movements, Home range and Social organization. Field studies of Milne-Edwards’ Sifakas have been carried out at Ranomafana National Park. It lives in multimale–multifemale groups of 3–9 individuals, typically containing 1–3 adult females and 1–2 adult males, and only one breeding pair. Females are dominant to males. Home-range size is 40–250 ha. Exclusive territories are maintained, although there is little aggression over boundaries. Mean daily path of movement averages 670 m. Males emigrate at age five to a neighboring group; females may either emigrate or remain in their natal group. Milne-Edwards’ Sifaka populations can have polygynous, polygynandrous, polyandrous, and monogamous mating systems. Density at Ranomafana is relatively low at 7·6 ind/km2, with densities in healthy populations south of the park reportedly even lower at c.3 ind/km2.

Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. Habitat destruction for slash-and-burn agriculture, logging, and gold mining are the main threats to Milne-Edwards’ Sifakas. These activities sometimes take place even in protected areas. Hunting is also a problem, with shotguns, blowguns, and slingshots used as the primary weapons. Milne-Edwards’ Sifaka occurs in two national parks (Andringitra and Ranomafana), and it also may be found in Andohahela National Park and Pic d’Ivohibe Special Reserve. There are populations in the unprotected forests north of Ranomafana and a number of unsurveyed forest reserves in eastern Fianarantsoa Province. The total world population is estimated at 9000 individuals, with c.4500 in Ranomafana.

Bibliography. Andriaholinirina et al. (2004), Dunham et al. (2008), Erhart & Overdorff (1998), Feistner & Schmid (1999), Glander et al. (1992), Grieser (1992), Groves (2001), Groves & Helgen (2007), Hemingway (1995, 1996, 1998, 1999), Irwin et al. (2000, 2005), Lehman & Wright (2000), Lehman et al. (2005a, 2006b), Meyers (1993), Meyers & Wright (1993), Mittermeier et al. (1994, 2006a, 2010), Nicoll & Langrand (1989), O’Connor et al. (1986, 1987), Pochron & Wright (2002, 2003, 2005), Pochron et al. (2002, 2003, 2004, 2005a, 2005b), Sterling & Ramarason (1996), Tan & Wright (1995), Tattersall (1982a, 1982b, 1986), Wright (1995, 1998), Wright et al. (1987, 1997).

 

17. Silky Sifaka Propithecus candidus

French: Sifaka soyeux / German: Seidensifaka / Spanish: Sifaca sedoso

Other common names: Silky Simpona

Taxonomy. Propithecus candidus Grandidier, 1871, Madagascar, north of Bay of Antongil.

Variations in this taxon approach P. diadema. P.candidus may be the only member of its genus to show extreme individual variation in partial skin pigmentation loss, known as leucism. Although all infants are believed to be born with predominantly black faces, with age some individuals lose their pigmentation and show varying degrees of pink patches. The first western explorer to observe P.candidus was A. Grandidier in 1871, and he believed that it was an albino subspecies of P. diadema. All individuals have some skin pigment, and photo-phobic individuals have never been observed. Monotypic.

Distribution. Restricted and patchy range in NE Madagascar that includes the humid forest belt extending from the Marojejy Massif and the Andapa Basin to Maroantsetra; the Androranga River may be the NE distributional limit in the Tsaratanana Corridor, although further surveys are needed to confirm this, and the Antainambalana River in the Makira Forest protected area is currently regarded as the S boundary. Currently, Marojejy represents the N limit of its distribution, although historic range maps suggest that it once occurred as far north as the Bemarivo River near Sambava. The north-eastern distributional limit in Makira was only recently established, when a few groups were found in the Antohaka Lava Forest, but informal reports suggest that the unprotected Maherivaratra Forest, outside north-eastern Makira, may also contain Silky Sifakas.

Descriptive notes. Head–body 48–54 cm, tail 45–51 cm; weight 5–6 kg. The Silky Sifaka is a large white sifaka, with notably long and silky fur, which gives this species its common English name. Males and females are similar in size. Females are mainly creamy-white, with some individuals having silvery-gray tints on the crown, back, and limbs. There is usually a patch of rust-red (sometimes yellow) fur on the base of the tail. The muzzle and face are bare and normally slatey-gray-black (often with pinkish mottling around the mouth), but some individuals have all pink or all black faces. Tips of the naked black or pink ears protrude just beyond the white fur of the head and cheeks. Eyes are a deep orange-red. Adult males are readily distinguished from females by the pelage color on the upper chest; they have a large brown patch on the upper chest that results from scent marking with the sternal-gular gland. As rates of scent marking by males increase during the mating season, these chest patches become larger in size and can cover the entire front torso to the abdomen. Some Silky Sifakas in the Marojejy Massif have entirely pinkish faces and dark fur across their upper back and shoulders.

Habitat. Several types of elevation-specific habitats and, despite its extreme rarity, the greatest elevational range of any sifaka. In Marojejy and in Anjanaharibe-Sud, Silky Sifakas are most often encountered in undisturbed humid primary and montane rainforest at elevations of 700–1900 m. They are sometimes found in sclerophyllous forest and even low ericoid bush near their highest elevations. The southernmost Silky Sifakas in Makira adjacent to the Antainambalana River inhabit an unusually low-elevation forest fragment at 300–600 m.

Food and Feeding. The Silky Sifaka is a folivorous seed predator that consumes a wide variety of plant species. A three-month dietary study at Marojejy National Park documented use of 76 species in 42 families (mainly trees but also many lianas and epiphytes). The most important plant families in the diet were Moraceae, Fabaceae, Myrtaceae, Clusiaceae, and Apocynaceae. Fruit from Pachytrophe dimepate (Moraceae), seeds from Senna (Fabaceae), young leaves from Plectaneia thouarsii (Apocynaceae), and fruit from Eugenia (Myrtaceae) were the most preferred and accounted for c.37% of total feeding time. Overall, 52% of feeding time of Silky Sifakas was spent eating young and mature leaves, 34% ripe fruit, and 11% seeds. Flowers, bark, and soil were rarely eaten.

Breeding. Mating is believed to occur on a single day each year in December–January. Generally, female Silky Sifakas give birth to a single infant every two years and only occasionally in consecutive years. Births take place in June–July, after gestation of c.179 days. Infants initially grasp the fur of their mother’s belly and transfer to her back at about four weeks, riding jockey-style. Until they approach of maturity, offspring always sleep with their mothers. As is typical of Propithecus, all group members interact with infants. Grooming is the most frequent form of alloparental care, followed by playing and occasional carrying; nursing has been observed in a few remarkable instances. Mothers have been seen simultaneously nursing both newborn and older offspring. Interbirth interval averages 1·7 years, but it is affected by whether or not an infant survives its first year.

Activity patterns. The Silky Sifaka is diurnal and arboreal. Adults spend most of their day resting (44·4%) and foraging (21·9%), but they also devote a substantial amount of time to social behavior (16·8%). Long bouts of terrestrial play involving adults are not uncommon.

Movements, Home range and Social organization. The Silky Sifaka has been discussed in several survey reports and was the subject of two short-term studies. As with other eastern sifakas, it has a variable social structure, living in both pair-bonded and multimale–multifemale groups of up to nine individuals. Smaller groups of 3–4 individuals probably consist of an adult pair plus their offspring, while larger groups likely consist of mutually familiar foraging units that may contain more than one breeding pair and juveniles. Home-range size varies by site, ranging 34–47 ha. Daily movement averages 712 m. Exclusive territories are maintained, although there is little aggression over boundaries. Males emigrate at age five to a neighboring group. Although male and female dispersal is typical in eastern sifakas, female dispersal has not been observed. Rates of aggression are low and mainly occur during feeding. Females have feeding priority over males. Two populations from Marojejy National Park had densities of 40 ind/km2 and 90 ind/km2, each estimate from a different elevational range. Generally, Silky Sifakas occur at very low densities.

Status and Conservation. CITES Appendix I. Classified as Critically Endangered on The IUCN Red List. The Silky Sifaka is one of the rarest and most endangered sifakas, and it has been on the list of the World’s 25 Most Endangered Primates, produced jointly by the IUCN/SSC Primate Specialist Group, the International Primatological Society, and Conservation International, since its inception in 2000. Main threats are habitat destruction and hunting, which occur even within supposedly protected areas. There is no local “fady” (taboo) against eating Silky Sifakas, as exists for the Indri in at least some parts of its distribution. The situation has become even more severe since the political upheavals of early 2009. The Silky Sifaka is known to occur in the Anjanaharibe-Sud Special Reserve and Marojejy National Park. Unfortunately, Marojejy was one of the first protected areas to be hit by a wave of illegal rosewood logging shortly after the coup that ousted President M. Ravalomanana. It is very possible that hunting of Silky Sifakas accompanied these incursions, but this remains to be confirmed. Populations also are found in the forests of Makira (currently under temporary government protection), the Betaolana Corridor, and possibly some unsurveyed forest fragments south-east of Marojejy (e.g. Andranomenabe and Maherivaratra). Efforts should be made to stop the hunting of sifaka in Marojejy and elsewhere. The total world population is believed to number only a few hundred individuals, down from somewhat less than 1000 in 1992.

Bibliography. Cousins (2007), Duckworth et al. (1995), Groves (2001), Kelley & Mayor (2002), Lehman et al. (2005a), Mittermeier et al. (2010), Patel (2005, 2006a, 2006b, 2007a, 2007b, 2009a, 2009b, 2010), Patel & Andrianandrasana (2008), Patel et al. (2005), Pochron et al. (2004), Rasolofoson et al. (2007a, 2007b), Sterling & McFadden (2000), Tattersall (1982a, 1982b), Wilmé & Callmander (2006).

 

18. Perrier’s Sifaka Propithecus perrieri

French: Sifaka de Perrier / German: Perrier-Sifaka / Spanish: Sifaca de Perrier

Taxonomy. Propithecus perrieri Lavauden, 1931, Madagascar, Forest of Analamera, south-east of Diégo Suarez.

This species is monotypic.

Distribution. Restricted to extreme NE Madagascar (Analamerana and Andrafiamena massifs and the E edge of the Ankarana forests).

Descriptive notes. Head–body 43–47 cm, tail 42–45 cm; weight 4·4–4·5 kg. Perrier’s Sifaka is a medium-sized species with a relatively short tail. The dorsal coat is long, dense, and silky, being a deep lustrous black above with a shorter, rosy-brownish tinted chest and belly. The face is bare and black, and eyes are dark reddish-brown. Ears are naked in some individuals and furred in others, and they are largely concealed in the fur.

Habitat. Tropical dry deciduous and semi-humid lowland forest from sea level to elevations of 500 m. Perrier’s Sifaka is most abundant in semi-evergreen forests on sandstone that comprise only one-quarter of its remaining habitat; these forests are unfortunately also under the heaviest pressures from humans.

Food and Feeding. Diets of Perrier’s Sifaka consist mainly of leaves (both young and mature), unripe fruits, stems, and flowers. At least a dozen different plant species in nine families have been identified as food sources.

Breeding. There is no information available for this species.

Activity patterns. There is no specific information available for this species, but Perrier’s Sifaka is diurnal and arboreal.

Movements, Home range and Social organization. Perrier’s Sifaka lives in multimale–multifemale groups of 2–6 individuals, typically containing 1–3 adult females, 1–2 adult males, and only one breeding pair. Females are dominant over males. Home-range size is c.30 ha. Exclusive territories are maintained, although there is little aggression over boundaries. Males emigrate at age five to a neighboring group; females may either emigrate or remain in their natal group. Unlike related species, Perrier’s Sifakas routinely descend from the trees to cross open habitat and drink from riverbeds. Density within the Analamerana Special Reserve is estimated at 3–18 ind/km2.

Status and Conservation. CITES Appendix I. Classified as Critically Endangered on The IUCN Red List. Perrier’s Sifaka is one of the rarest and most endangered lemurs (and indeed of all the world’s primates). The total population could be as few as 500 individuals, with an effective breeding population of only 125 individuals. Slash-and-burn agriculture that destroys its very limited habitat is the greatest threat, and this is exacerbated by fires set to increase livestock pasture, cutting of trees for charcoal production, and forest destruction by itinerant miners. Local hunting also may be a problem. The only protected area in which it is known to occur is the Analamerana Special Reserve. A small number of Perrier’s Sifaka may still be found in the eastern section of Ankarana National Park, which is connected through a series of forest patches to populations at Andriafiamena and Analamerana. Perrier’s Sifaka has recently been seen in unprotected forest patches between Analamerana and Ankarana, and it is present in the Andavakoera Classified Forest, but probably in very low numbers. There is an urgent need for a full-time, long-term scientific presence in Analamerana and its expansion to include the forests of Andriafiamena and a connection to Ankarana. This effort should also include an educational campaign in the region and surveys in nearby forest patches to look for any other populations of Perrier’s Sifaka.

Bibliography. Banks et al. (2007), Fowler et al. (1989), Ganzhorn et al. (1996), Groves (2001), Harcourt & Thornback (1990), Hawkins et al. (1990), Lehman & Mayor (2004), Lehman et al. (2005a), Mayor & Lehman (1999), Meyers & Ratsirarson (1989), Mittermeier et al. (2010), Petter et al. (1977), Rasoloharijaona et al. (2005), Tattersall (1982a, 1982b).

Genus INDRI

É Geoffroy Saint-Hilaire & G. Cuvier, 1796

 

19. Indri Indri indri

French: Indri / German: Indri / Spanish: Indri

Other common names: Babakoto

Taxonomy. Lemur indri Gmelin, 1788, Madagascar.

There appears to be a regional trend regarding the amount of white and black fur on the coat, and as a consequence, two distinct subspecies were formerly recognized. This is now believed to constitute a cline, with darker individuals in the north of the distribution and lighter ones in the south. Monotypic.

Distribution. NE & CE Madagascar, found roughly from the Anjanaharibe-Sud Special Reserve in the N to the Anosibe an’ala Classified Forest in the S.

Descriptive notes. Head–body 64–72 cm, tail c.5 cm; weight 5·8–9 kg. The Indri is the largest living lemur, with only the Diademed Sifaka (Propithecus diadema) approaching it in size. The Indri’s body is thickly covered with long, silky fur above, which becomes shorter underneath, and the skin is black. The head is rounded with a prominent, nearly naked muzzle, large ears, and relatively large staring eyes that are directed more forward than laterally. Irises are yellow-brown, with circular pupils, and are accentuated by long eyelashes. Coat color varies considerably by region from predominantly black, contrasting with white or grayish-white patches on the crown, neck, flanks, forelimbs, and thighs (or any combination of these) to variegated black and white; the white areas correspond to the occipital cap, a broad collar extending up to and beyond the ears, and the outer surfaces of legs and lower arms. In some populations, heels and flanks may be tinged with red or gold. The northern form is predominantly black. The earliest Indris to be described had this darker color pattern, which is typical of museum specimens from Andapa and Maroantsetra and has also been observed in wild populations from Anjanaharibe-Sud, Ambatovaky, and Anjozorobe. Indris from central-eastern Madagascar are generally lighter in color. Further complicating the color variation among Indris, both melanistic and albino individuals have also been recorded. The classic pattern of the Giant Panda (Ailuropoda melanoleuca) appears to be less common among populations of Indris than the largely or entirely black pattern. In the Analamazaotra Special Reserve and in the Anjozorobe-Angavo protected area, male Indris are slightly larger than females. There is also some slight dimorphism in color pattern, with the chest of the male dark and that of the female much lighter. The male also has a white triangle at the base of its back, but this marking is absent in the female. Whether these differences hold true in other parts of the distribution of the Indri remains to be determined. Regardless of coat color or pattern, however, ears are always black, modestly tufted, and prominent.

Habitat. Primary and secondary tropical moist lowland and montane forest, also some disturbed habitats. The elevational range of the Indri is sea level to 1800 m. They are often found in mountainous habitats or steep terrain with numerous ridges and valleys. All levels of the canopy are used, although individuals tend to stay in the lower levels in October–December to avoid biting insects.

Food and Feeding. Indris feed mainly on immature leaves, although fruits, seeds, flowers, buds, and bark are also eaten; bark consumption varies seasonally. Individuals also descend to the ground to eat soil, which may help to detoxify seeds or perhaps facilitate the breakdown of their bulky diet.

Breeding. Reproduction of the Indri is highly seasonal. Mating takes place in December–March and is performed ventro/ventrally while hanging beneath a branch. Only one pair of adults in a group produces offspring. The female produces a single young about every 2–3 years—a very slow reproductive rate for a prosimian. Births usually occur in May–June (but can be as late as August), and gestation is 130–150 days. Indri young tend to be entirely black during the first 2–3 months of life, with white only on a small patch in the pygal region; they develop the characteristic gray and white patches of adults at 4–6 months of age. The mother carries her offspring across her abdomen for the first 4–5 months and then on her back. Young are reasonably independent at 8–10 months, but they stay close to the mother until well into their second year. They reach maturity at about four years of age, although females do not reach full sexual maturity until 7–9 years. Longevity is unknown.

Activity patterns. The Indri is diurnal and arboreal. Its signature vocalization is a loud, drawn out, wailing territorial call that is one of the best known and most characteristic sounds of the Malagasy rainforest. Territorial calls have been described as a plaintive bark, blending with lamenting cries that sound like a mixture of a human crying in pain and howling of a frightened dog. This call is heard most often during the morning hours but can be heard at any time of the day; it can carry across the forest for up to 3 km. It is also more frequently given in the warm rainy season than in colder months. The adult pair in a family social group typically leads the chorus, and neighboring groups often call sequentially in response. Spacing among groups is likely determined by these calls, which may explain the relatively small degree of home-range overlap.

Movements, Home range and Social organization. The Indri has been studied in the forests of Analamazaotra Special Reserve and nearby Mantadia National Park. It lives in small groups of 2–6 individuals, normally consisting of a monogamous adult pair (they evidently seek new partners only after a mate dies) and their offspring. Although groups in fragmented habitat tend to be larger than those in more extensive, undisturbed areas, this is not always the case. Changes in the composition of larger groups are quite frequent. Home-range size averages 18 ha in the fragmented forests of Analamazaotra, but it can be as large as 40 ha in the more pristine forests of Mantadia, where daily movements are 300–800 m. A large central part of each home range is defended, from which other groups are excluded. Individuals are extremely attached to their home range, and they always use the same trails within it. Adult males are responsible for territorial defense; the female moves to a safe location during encounters with intruders. There also is frequent agitation in the group and competition for food, and females and young are dominant in such situations, easily displacing an adult male. A female normally feeds higher in the trees than a male and has priority access to food resources. Before dusk, the group retires to a sleeping tree, bedding down on horizontal supports at heights of 10–30 m; the female is typically huddled with her offspring and separated from the male by 2–50 m. Reciprocal grooming has been reported, but not simultaneously. Densities typically are 9–16 ind/km2 but are as low as 5·2 ind/km2 in some areas. Indris can reach relatively high densities (c.22·9 ind/km2) if they are not hunted by local people.

Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. The principal threat to the Indri is habitat destruction for slash-and-burn agriculture, logging, and firewood gathering, affecting even protected areas. Contrary to what was believed in the past, illegal hunting of the Indri is also a major problem in certain areas. Although long thought to be protected by local “fady” (taboos), these do not appear to be universal, and Indris are now hunted even in places where such tribal taboos exist. In many areas, these taboos are breaking down with cultural erosion and immigration, and local people often find ways to circumvent taboos even if they are still in place. For example, a person forbidden to eat Indris can still hunt and sell them to others, and those forbidden to kill Indris can purchase them for food from someone else. Recent studies of villages in the Makira Forest indicate that Indris have been hunted in the past for their skins (worn as clothing), Indri meat is prized and fetches a premium price, and current levels of hunting are unsustainable. The Indri occur in three national parks (Mananara-Nord, Mantadia, and Zahamena), two strict nature reserves (Betampona and Zahamena), and five special reserves (Ambatovaky, Analamazaotra, Anjanaharibe-Sud, Mangerivola, and Marotandrano). It also is found in the Anjozorobe-Angavo Protected Area and the forests of Makira, which are currently under temporary government protection (although hunting pressure appears to be especially heavy at Makira). The corridor between Mantadia and Zahamena has been proposed as a new conservation site, and the Anosibe an’ala Classified Forest also should be considered for the creation of a new park or reserve. No population figures are available for the Indri, but a reasonable estimate would be 1000–10,000 individuals. The Indri does not occur on the Masoala Peninsula or in Marojejy National Park, despite the park being connected to forest less than 40 km away where the Indri occurs. Before large-scale deforestation, it was much more widely distributed, with a separate group said to occupy almost every ridge of Madagsacar’s eastern forests. Subfossil evidence indicates that they once occurred well into the interior of Madagascar at least as far west as the Itasy Massif, south-west to Ampoza-Ankazoabo, and north to the Ankarana Massif.

Bibliography. Britt et al. (1999, 2002), Garbutt (2007), Geissmann & Mutschler (2006), Glessner & Britt (2005), Godfrey et al. (1999), Golden (2005), Goodman & Ganzhorn (2004a, 2004b), Groves (2001), Jungers et al. (1995), Mittermeier et al. (2010), Nicoll & Langrand (1989), Oliver & O’Connor (1980), Petter (1962), Petter & Peyrieras (1972, 1974), Petter et al. (1977), Pollock (1975a, 1975b, 1977, 1979a, 1979b,1986a, 1986b), Powzyk. (1997), Powzyk & Mowry (2003, 2006), Powzyk & Thalmann (2003), Rand (1935), Rigamonti et al. (2005), Schmid & Smolker (1998), Tattersall (1982a, 1982b), Thalmann et al. (1993).