Prolemur simus (Gray, 1870)

Russell A. Mittermeier, Anthony B. Rylands & Don E. Wilson, 2013, Lemuridae, Handbook of the Mammals of the World – Volume 3 Primates, Barcelona: Lynx Edicions, pp. 90-141 : 127-128

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Greater Bamboo Lemur

Prolemur simus

French: Lémur a nez large / German: Gro 3er Bambuslemur / Spanish: Lémur del bambu grande

Other common names: Broad-nosed Bamboo Lemur, Broad-nosed Gentle Lemur

Taxonomy. Hapalemur simus Gray, 1871 ,

Madagascar.

Subfossil specimens from Ampasambazimba and Andrafiabe, despite the considerable geographic gap separating them from extant (Ranomafana) and historic (Antongil Bay) populations of Prolemur simus , show no sign of being different from them. Monotypic.

Distribution. CE & SE Madagascar; until recently restricted to the SC portion of the country’s rainforests including Ranomafana and Andringitra national parks, Ambositra-Vondrozo Corridor and isolated forests between and to the E ofthese localities (e.g. Ambolomavo, Ifanadiana, Kianjavato), and one locality N of the Manampatrana River (Evendra); recent surveys have now confirmed its presence in the forests of Torotorofotsy, in the region of Andasibe-Mantadia, and 18 sites in and around the Ankeniheny-Zahamena Corridor, and at five additional sites around the Marolambo forest corridor and as far N as Zahamena National Park, and it extends to near the Midongy du Sud National Park in the S, 670 km S of Zahamena. Reports of large bamboo lemursfitting this species’ description have been filtering in from various other remote sites (e.g. Mananara region).

Descriptive notes. Head-body 40-42 cm, tail 45-48 cm; weight 2.2-2.5 kg (males possibly heavier than females). The Greater Bamboo Lemur is the largest of the bamboo lemurs and so distinct that it has been placed in its own genus. Its pelage is somewhat drab, being typically grayish-brown with a slight reddish tinge on the dorsum and tail. The crown, neck, shoulders, and upper arms tend to be more olive-brown. The ventral coat is a lighter creamy-brown. There is a rusty-brown pygal patch. The muzzle is blunt and dark gray, and ears are large with (usually) prominent white tufts. In 2007, P. Garbutt mentioned a population of the Greater Bamboo Lemur discovered on the Andringitra Massif that might be a distinct color morph. Although generally similar to typical Greater Bamboo Lemur ,its head, upper parts, throat, and underparts are uniformly deep golden-red, and the tip of the tail is gray. The face and muzzle are dark, and ears are large and prominent but lack the characteristic white tufts of the species. The taxonomic identity of this morph requires further investigation.

Habitat. Humid primary and secondary rainforest associated with giant bamboo ( Cathariostachys madagascariensis, Poaceae ), and marshland, at elevations of 28-1600 m. The Greater Bamboo Lemur also occurs in degraded habitat and plantations. It is sympatric with the Gray Bamboo Lemur ( H. griseus ranomafanensis) in Ranomafana National Park, the Southern Bamboo Lemur ( H. meridionalis ) in Andringitra National Park, and the Golden Bamboo Lemur ( H. aureus ) in both these protected areas.

Food and Feeding. The Greater Bamboo Lemur subsists predominantly on bamboo. Giant bamboo can account for as much as 95% of the diet, with shoots, young and mature leaves, and pith consumed. Presence of giant bamboo significantly affects the distribution of the Greater Bamboo Lemur . In the dry season, inner pith of the culm is most commonly consumed, whereas in the rainy season, shoots are mainly eaten. Other food items include the bamboo species Valiha diffusa and Bambusa vulgaris, flowers of the traveler's palm ( Ravenala madagascariensis, Strelitziaceae ), figs ( Ficus , Moraceae ), palm fruits ( Dypsis spp. , Arecaceae ), leaves of Pennisetum clandestinum ( Poaceae ), Aframomum angustifolium ( Zingiberaceae ), and Rubus moluccanus ( Rosaceae ). Greater Bamboo Lemurs also eat various agricultural crops including Mangifera indica ( Anacardiaceae ; mango), Litchi sinensis ( Sapindaceae ; lychee), Annona squamosa ( Annonaceae ; custard apple), Artocarpus heterophyllus ( Moraceae ; jackfruit), A. altilis ( Moraceae ; breadfruit), Musa sp. ( Musaceae ; banana), Saccharum sp. ( Poaceae ; sugar cane), and Coffea spp. ( Rubiaceae ; coffee). Overall, their diet is high in fiber, cellulose, and lignin and low in protein. When eating bamboo, individuals strip the outside ofthe live stalk and tear apart the bamboo pole to get at the soft pith, an impressive accomplishment for such a small animal. Damage to bamboo is a signature of the Greater Bamboo Lemur , allowing detection of its presence in an area.

Breeding. The vulva of the female Greater Bamboo Lemursis closed during anestrus. Mating begins in late May or June. Females typically give birth to a single young in October-November, after gestation of ¢.150 days. Birth weight in captivity is ¢.80 g. Interbirth interval is one year. Most care of the infant is provided by the mother, who carriesit all the time during the first four months of life. Unlike species of Hapalemur , female Greater Bamboo Lemurs do not park their infants on branches while they forage. Instead, infants are kept in close contact until they are ready to explore independently. They are weaned by 8:5—-10 months but will eat solid bamboo at eight weeks. Copulatory play begins at two years of age. Females become sexually mature at 1-5 years and males at 2-5 years old. Individuals have lived more than 17 years in captivity.

Activity patterns. The Greater Bamboo Lemur is arboreal and cathemeral. Observations of wild populations and captive animals suggest that it is active both day and night throughout the year.

Movements, Home range and Social organization. The Greater Bamboo Lemur lives in small monogamous groups or larger polygamous groups of up to 28 animals. Home ranges are large and vary from 60 ha at Ranomafana to several hundred hectares at Torotorofotsy. Home ranges may overlap those of other sympatric bamboo lemur species, and Greater Bamboo Lemurs frequently form mixed-species associations with species of Hapalemur and Eulemur . Males disperse from their natal groups as adults.

Status and Conservation. CITES Appendix I. Classified as Critically Endangered on The IUCN Red List. The Greater Bamboo Lemur has been on the list of the World's 25 Most Endangered Primates , prepared every two years by the IUCN/SSC Primate Specialist Group, the International Primatological Society, and Conservation International since 2002. It is threatened mainly by slash-and-burn agriculture, mining, illegal logging, and cutting of bamboo. It is also hunted for food with slingshots, spears, and snares. Recent surveys have shown that the Greater Bamboo Lemur occurs in at least 36 sites from Zahamena National Park in the north to Midongy du Sud National Park in the south. Although recent surveys have tripled the number of known sites of occurrence of the Greater Bamboo Lemur and quadrupled total population estimates, and it appears safer than just a few years ago,it is probably still one of the most endangered of all lemurs. Historical records and subfossil remains confirm a much more widespread distribution in the past, covering the northern, north-western, central, and eastern parts of Madagascar, including Ampasambazimba in the Itasy Basin, Grotte d’Andrafiabe on the Ankarana Massif, Grottes d’Anjohibe near Mahajanga, and Tsingy de Bemaraha. The Greater Bamboo Lemur may still have been present in the Ankarana region, at the northern tip of Madagascar, up until the mid-20" century at least, and it may still exist there. It is quite possible that future field surveys will discover additional populations. It is known to occur in three national parks ( Andringitra, Ranomafana, and Zahamena ) and Ambositra-Vondrozo Corridor that is under temporary protection and Ankeniheny-Zahamena Corridor that will soon receive protected area status.

Bibliography. Albrecht et al. (1990), Andriaholinirina et al. (2003), Arrigo-Nelson & Wright (2004), Dolch, Fiely et al. (2008), Dolch, Hilgartner et al. (2004), Fleagle (1999), Garbutt (2007), Gauthier et al. (1998), Godfrey & Vuillaume-Randriamanantena (1986), Godfrey, Jungers et al. (1999), Godfrey, Simons et al. (2004), Goodman et al. (2001), Grassi (1998), Irwin et al. (2005), Kappeler (1991), Klopfer & Boskoff (1979), Leclercq & Santini-Palka (1995), Macedonia & Stanger (1994), Meier (1987), Meier & Rumpler (1987), Meier et al. (1987), Mittermeier et al. (2010), Mutschler & Tan (2003), Petter & Charles-Dominique (1979), Rakotonirina, Rajaonson, Ratolojanahary, Missirli et al. (2011), Rakotonirina, Rajaonson, Ratolojanahary, Rafalimandimby et al. (2011), Rakotosamimanana et al. (2004), Ravaloharimanitra et al. (2011), Rumpler et al. (1989), Santini-Palka (1994), Schmid & Alonso (2005), Schwarz (1931a), Stanger-Hall (1997), Sterling & Ramaroson (1996), Tan (1999, 2000, 2006), Tattersall (1982), Vuillaume-Randriamanantena et al. (1985), Weigl (2005), Wilson et al. (1989), Wright & Randrimanantena (1989), Wright, Daniels et al. (1987), Wright, Johnson et al. (2008), Wright, Larney et al. (2009).

Figures

0. Greater Bamboo Lemur (Prolemur simus), 1. Gray Bamboo Lemur (Hapalemur griseus), 2. Southern Bamboo Lemur (Hapalemur meridionalis), 3. Northern Bamboo Lemur (Hapalemur occidentalis), 4. Lac Alaotra Bamboo Lemur (Hapalemur alaotrensis), 5. Golden Bamboo Lemur (Hapalemur aureus)

Distribution. CE & SE Madagascar; until recently restricted to the SC portion of the country’s rainforests including Ranomafana and Andringitra national parks, Ambositra-Vondrozo Corridor and isolated forests between and to the E ofthese localities (e.g. Ambolomavo, Ifanadiana, Kianjavato), and one locality N of the Manampatrana River (Evendra); recent surveys have now confirmed its presence in the forests of Torotorofotsy, in the region of Andasibe-Mantadia, and 18 sites in and around the Ankeniheny-Zahamena Corridor, and at five additional sites around the Marolambo forest corridor and as far N as Zahamena National Park, and it extends to near the Midongy du Sud National Park in the S, 670 km S of Zahamena. Reports of large bamboo lemursfitting this species’ description have been filtering in from various other remote sites (e.g. Mananara region).