Bamboo lemur
Hapalemur

TAXONOMY

Suborder: Strepsirrhini
Infraorder: Lemuriformes
Superfamily: Lemuroidea
Family: Lemuridae
Genus: Hapalemur
Species: H. alaotrensis, H. aureus, H. griseus, H. meridionalis, H. occidentalis

Other names: H. alaotrensis: H. griseus alaotrensis, Alaotra reed lemur, Alaotran bamboo lemur, Alaotran gentle lemur, Lac Alaotra bamboo lemur, Lake Alaotra gentle lemur; hapalemur d'Alaotra (French); Alaotra halbmaki (German); bandro (Malagasy); H. aureus: golden bamboo lemur, golden lemur; gouden halfmaki (Dutch); hapalémur doré (French); goldener bambuslemur (German); bokombolomena, varibolomena (Malagasy); lemur cariancho (Spanish); gyllenbambulemur, gyllenhalvmaki (Swedish); H. griseus: H. ranomafanensis, grey gentle lemur, eastern lesser bamboo lemur, eastern gray bamboo lemur, bamboo lemur, lesser bamboo lemur; kleine halfmaki (Dutch); hapalémur gris, petit hapalémur (French); ostlicher grauer halbmaki (German); bokombolo, kotrika, kontè (Malagasy); lemur cariancho (Spanish); grå bambulemur, grå halvmaki, grå lemurträskhalvmaki, västlig halvmaki (Swedish); H. meridionalis: rusty-gray lesser bamboo lemur, southern grey bamboo lemur, southern lesser bamboo lemur, southern gentle lemur; petit hapalémur méridional (French); halo (Malagasy); H. occidentalis: H. griseus occidentalis, Sambirano bamboo lemur, Sambirano lesser bamboo lemur, western gentle lemur, western grey bamboo lemur, western lesser bamboo lemur; hapalémur occidental (French); westlicher grauer halbmaki (German); bokombolo, akomba-valiha, bekola, kofiy, kitronytrony (Malagasy).

The taxonomy of the genus Hapalemur is debated. Groves (2005) recognizes four full species of Hapalemur: H. alaotrensis, H. aureus, H. griseus, and H. occidentalis. As with other animals, the data used to differentiate species or elevate types to species are different depending on the researcher. In the Hapalemur case, these lines of data are in conflict with one another, and result in confusion. For example, H. meridionalis is sometimes considered a full species based on cytogenetic data and H. alaotrensis which is morphologically specifically distinct but not so cytogenetically (Fausser et al. 2002; Rabarivola et al. 2007). To avoid confusion, the approach taken by Mittermeier et al. (2006) will be followed here in which five full species are recognized, but also a need for further clarification of the relationships between the species is acknowledged. Several subspecies of H. griseus have recently been described but are not included here as such (Rabarivola et al. 2007). Finally, while recognizing significant recent alterations in the distributions of H. griseus, H. meridionalis, and H. occidentalis due to new cytogenetic data (Rabarivola et al. 2007), information about these species is included under the taxon given by the researcher, even though in light of new data this may not be correct.

MORPHOLOGY

Bamboo lemurs are generally small to medium lemurs and are mostly gray (Mittermeier et al. 2006; 2008). They are approximately the same size as smallish housecats (Pollock 1986). The tail is usually longer than the body (Warter et al. 1987). The head is lighter in color than the rest of the body (Warter et al. 1987; Mittermeier et al. 2006). The ears of H. griseus, H. meridionalis, and H. occidentalis are larger than those possessed by H. alaotrensis and H. aureus, where they almost cannot be seen in the fur of the head (Groves 2001). The noses and faces of bamboo lemurs are shorter than in other types of lemur (Warter et al. 1987; Mittermeier et al. 2006). Bamboo lemurs possess scent glands on their forearms and on their underside near their armpits (Meier et al. 1987; Warter et al. 1987; Groves 2001).

H. aureus dorsal surfaces are grey-brown with underlying orange pelage and their ventrum is yellow. Their face is black with a gold-yellow ring around the face encompassing their cheeks, eyebrows and their throat (Meier et al. 1987). The nose of H. aureus is pink (Mittermeier et al. 2006). H. griseus are generally gray with some olive coloration (Groves 2001). H. occidentalis are gray-brown overall with a pale face (Groves 2001). H. meridionalis are generally gray as well but have some red coloration, particularly on the head and body (Groves 2001). H. alaotrensis have dense pelage which is gray-brown dorsally and lighter ventrally. The head and neck are chestnut-brown (Mittermeier et al. 2006).

H. griseus females average 66.94 cm (26.4 in) in length while males average 67.69 cm (26.7 in). H. meridionalis females average 66.78 cm (26.3 in) in length and males average 67.80 cm (26.3 in). H. occidentalis females average 66.5 cm (26.2 in) in length while males average 67.5 cm (26.6 in). H. alaotrensis have been recorded between 77.0 and 81.0 cm (30.3 and 31.9 in) in length (Rabarivola et al. 2007).

H. alaotrensis and H. aureus are significantly larger than the other species of bamboo lemur (Groves 2001; Rabarivola et al. 2007). H. griseus weigh on average 0.935 kg (2.1 lb) and H. aureus weigh on average 1.548 kg (3.4 lb) (Glander et al. 1992 compiled by Tan 1999; Tan 1999). H. alaotrensis weigh on average 1.24 kg (2.7 lb) (Mutschler 1998; 2002). H. occidentalis is similar in size to H. griseus, with females weighing on average 1.188 kg (2.6 lb) and males weighing 0.847 kg (1.9 lb) (Mittermeier et al. 2006; Rabarivola et al. 2007). H. meridionalis females weigh on average 0.870 kg (1.9 lb) while males weigh 0.839 kg (1.9 in) (Rabarivola et al. 2007). There is not significant body mass dimorphism between male and female bamboo lemurs (Kappeler 1991; Mittermeier et al. 2006).

Bamboo lemurs leap from support to support in a vertical posture, landing feet-first and are capable of running quickly on the ground (Petter & Peyrieras 1975; Wright 1988). They will also run quadrupedally along supports (Haring & Davis 1998). H. griseus are normally found around 5.9 m (19.4 ft) above the ground and travel through leaping (48%) and quadrupedal walking (38%) (Blanchard et al. 2007). Anecdotal evidence indicates that H. alaotrensis can swim (Petter & Peyrieras 1975).

In captivity, H. griseus have lived over 23 years (Weigl 2005).

RANGE

CURRENT RANGE MAPS (IUCN REDLIST):
Hapalemur alaotrensis | Hapalemur aureus | Hapalemur griseus | Hapalemur meridionalis | Hapalemur occidentalis

Like all lemurs, the genus Hapalemur only lives on Madagascar. Distribution accounts of H. griseus, H. occidentalis, and H. meridiontalis have been significantly and recently altered recently in light of new cytogenetic data (http://www.redlist.org; Rabarivola et al. 2007). Previously, the H. griseus distribution was considered to extend from the Anjanaharibe-Sud Special Reserve and the Marojejy National Park in northeastern Madagascar south to the region north of Andohahela, in the southeastern parts of the island (Mittermeier et al. 2006). The distribution was interrupted in the region around Lake Alaotra and met the distribution of H. meridionalis in the southeastern parts of the island (Mittermeier et al. 2006). H. occidentalis was thought to live in western and northern Madagascar. The species was known from the northern edges of the island, from the Ankarana and Analamerana forests and the Sambirano region west and south to a number of areas between the Mahavavy River and the Tsiribihina River in west-central Madagascar (Mittermeier et al. 2006). Reflecting data obtained in a recent study however (Rabarivola et al. 2007), the distribution of these three species has changed considerably.

The distribution of H. griseus has been significantly altered, with Hapalemur populations in western Madagascar between the Mahavavy and Tsiribihina Rivers previously assigned to H. occidentalis now attributed to H. griseus and with the southern extent of the H. griseus species now reduced, only extending as far south as Ranamofana (http://www.redlist.org; Rabarivola et al. 2007). Similarly, the H. occidentalis distribution has changed, with the west-central populations of the species now attributed to H. griseus. The distribution of H. occidentalis is now recorded as encompassing the previously recognized northern populations but now extending south in eastern Madagascar to Zahamena, east of Lake Alaotra (http://www.redlist.org; Rabarivola et al. 2007).

The type locality of H. meridionalis is the Mandena forestry station, approximately ten kilometers (6.2 miles) north of Tolagnaro (Ft. Dauphin) and at a minimum, the species is found between the Mananara River in the north and the Andohahela Nature Reserve although the distribution may reach as far north as Atsimo due to a hybrid form recorded at that locality (http://www.redlist.org; Warter & Tattersall 1994; Rabarivola et al. 2007).

The Alaotran bamboo lemur (H. alaotrensis) is found exclusively in reed beds surrounding Lake Alaotra, the largest lake in Madagascar, which is located on the eastern part of the island. The species distribution around the lake is divided into two subpopulations (Mutschler & Feistner 1995). H. aureus is found in eastern Madagascar, from Ranomafana National Park in the north (and possibly further northeast) and south to the Andringitra National Park, inhabiting a forest corridor between the two (http://www.redlist.org; Mittermeier et al. 2006).

HABITAT

In general, H. griseus and H. aureus are found in humid eastern lowland and montane forests (Mutschler & Tan 2003; Mittermeier et al. 2006). H. griseus are found in an altitudinal range between sea-level up to 2050 meters (6725.72 feet), while H. aureus range between 625 and 1625 meters (2050.5 and 5331.4 feet) (reviewed in Goodman & Ganzhorn 2004). H. meridionalis is known to reside in lowland and montane subtropical rain forests and littoral forests with marshy areas (Mutschler & Tan 2003; Mittermeier et al. 2006). This species exists at altitudes of up to 1600 m (5249.3 ft) (Mittermeier et al. 2006). H. occidentalis is found in dry to sub-humid primary and degraded deciduous forests as well as bamboo scrub, also sometimes occurring in areas under a degree of agricultural use and other degraded habitats (Hawkins et al. 1998; Mutschler & Tan 2003; Mittermeier et al. 2006; Martinez 2008). While characterized by their consumption of bamboo, bamboo lemurs are sometimes found in forests without any bamboo, as is the case in the Malahelo Forest, southeastern Madagascar, an intermediate forest type (Ramanamanjato et al. 2002). The same may be true of bamboo lemurs found in some bamboo-free areas of northwestern Madagascar (Hawkins et al. 1998).

Among the bamboo lemurs, H. alaotrensis inhabit a habitat unlike the other members of the genus. They are restricted to reed and papyrus marshland vegetation surrounding Lake Alaotra in eastern Madagascar, a habitat in which stands of reeds and papyrus are separated by water channels (Mutschler & Feistner 1995; Nievergelt et al. 1998). H. alaotrensis is unique among the primates in that it lives only in this marshy habitat (Nievergelt et al. 1998).

At one study site of H. griseus and H. aureus at Ranamafana National Park in eastern Madagascar, the average temperature is 21°C (69.8°F), but seasonally varies from 4-6° C (39.2-42.8°F) between June-August and 28-30°C (82.4-86.0°F) between November-January (see Tan 1999 and references therein). At this study site, a dry season occurs between April-November and a wet season between December-March (Atsalis 1998).

ECOLOGY

The feeding ecology of bamboo lemurs is not completely known. As is implied by their name however, most of the genus specializes in the consumption of bamboo. Bamboo lemurs can consume far more bamboo than should be lethal to animals of their size due to its cyanide content, although the way they manage to avoid poisoning remains unclear (Glander et al. 1989; Yamashita et al. 2010).

At the Ranomafana National Park, H. griseus consumed over 40 species of plant, but the diet consisted mostly of giant bamboo (Cathariostachys madagascariensis) (72%) (Tan 1999; 2006). The rest of the foods were other types of bamboo and grasses (16%), leaves from other plants (particularly lianas) (4%), fruits (5%), and other foods (3%) (Tan 1999). Most of the bamboo parts that are eaten are the young parts of the plants, and the lemurs avoid mature leaves (Tan 1999). From month to month, the diversity of the H. griseus diet remains relatively low, and they tend to rely on resources that are not seasonal (Tan 1999). However, only during the wet season are other foods such as fruit, fungus, mushrooms, leaf petioles, flowers and new leaves included in the diet, and even then at low levels (Overdorff et al. 1997). Geophagy in wild H. griseus is documented (Irwin et al. 2004).

Also at Ranomafana National Park, and similarly to H. griseus, H. aureus eats mostly giant bamboo (78%), other types of bamboo and grasses (10%), fruits (4%), and other foods (5%) (Tan 1999). H. alaotrensis are entirely folivorous, eating the leaves and stems of only 11 plant species (Mutschler 1998; 2002). However, over 95% of the diet was restricted to four species (Cyperus madagascariensis, Phragmites communis, Leersia hexandra, and Echinochloa crusgalli) (Mutschler 1998). In addition, only very specific parts of each species are eaten (Mutschler et al. 1998). In captivity, this species has been seen eating soil (Beattie & Feistner 1998). Little is known about the diet of H. occidentalis, but individuals have been observed eating fruits, bamboo and liana flowers (Petter et al. 1977 cited in Tan 2006; Mutschler & Tan 2003). The diet of H. meridionalis is completely unknown (Mutschler & Tan 2003; Tan 2006).

Activity patterns appear to vary by species among the bamboo lemurs. H. aureus is diurnal, as is H. griseus and preliminary data indicates that H. meridionalis is diurnal also (Mutschler & Tan 2003; Tan 2006; reviewed in Tan 2006). H. aureus spends its days resting (50%), feeding (41%), and traveling (8%) and has a discrete rest period between mid-morning and early afternoon (Tan 2000 cited in Tan 2006; Tan 2006). Activities are similar in H. griseus, who spend their days resting (41%), feeding (48%), and traveling (9%) (Tan 2000 cited in Tan 2006). Around dusk, H. griseus move to sleeping sites (Overdorff et al. 1997).

H. alaotrensis on the other hand, is cathemeral in its activity and exhibits substantial amounts of activity both during the day and during the night (Mutschler et al. 1998; Mutschler 2002). However, the species spends more time active during daylight hours than during the nighttime (Mutschler & Tan 2003; Tan 2006). During daylight, the H. alaotrensis activity is usually separated into an early morning activity period and a late afternoon activity period, but the day-to-day reality of this pattern is highly variable (Mutschler et al. 1998). However, nighttime activity is never less than 30 minutes (Mutschler et al. 1998). H. occidentalis show some evidence for substantial nighttime activity, with dry season (July-September) activity predominantly occurring during hours of darkness (Mutschler 2000 cited in Mutschler & Tan 2003). H. griseus activity changes seasonally as well, with more resting and less traveling and feeding during the warm wet season (Overdorff et al. 1997).

H. griseus home ranges average 0.15 km² (0.06 mi²) (although in a different study ranged between 0.06-0.1 km²), while H. aureus home ranges average 0.26 km² (0.1 mi²) (Wright 1986; Tan 1999). Recorded H. occidentalis home ranges are around 0.26 km² (0.1 mi²) (Gould & Sauther 2007). In their unique habitat around Lake Aloatra, H. alaotrensis home ranges average only 0.02 km² (0.008 mi²) (Mutschler et al. 1994 cited in Mutschler & Feistner 1995). The daily path of H. aureus averages 365 m (1197.5 ft) while the daily path of one H. griesus group averaged 425 m (1394.4 ft) (Wright 1986; Wright et al. 1987).

H. griseus at Kalambatritra Special Reserve in south-central Madagascar are known to exhibit latrine behavior (repeated group defecation at specific sites), and groups defecate in turn, with adults going first followed by juveniles (Irwin et al. 2004).

The Malagasy tree boa (Boa manditra) has been observed killing and consuming adult H. griseus in the wild (Rakotondravony et al. 1998). Henst's Goshawk (Accipiter henstii) and the Madagascar Harrier-hawk (Polyboroides radiatus) are both predators of H. griseus (Karpanty 2006).

Bamboo lemurs are often found in sympatry with other species of primates. For example, at Ranomafana National Park, H. griseus and H. aureus are sympatric with brown mouse lemurs (Microcebus rufus), greater dwarf lemurs (Cheirogaleus major), sportive lemurs (Lepilemur sp.), eastern avahis (Avahi laniger), aye-ayes (Daubentonia madagascariensis), greater bamboo lemurs (Prolemur simus), red-fronted lemurs (Eulemur rufus), red-bellied lemurs (Eulemur rubriventer), and Milne-Edwards' sifakas (Propithecus edwardsi) (Tan 1999).

While both H. griseus and H. aureus can live together in the same habitat and consume generally the same foods, they consume different parts of the same foods, avoiding some competition by eating parts of the same plants with different physical properties (Yamashita et al. 2009).

Content last modified: July 22, 2010

Written by Kurt Gron.

Cite this page as:
Gron KJ. 2010 July 22. Primate Factsheets: Bamboo lemur (Hapalemur) Taxonomy, Morphology, & Ecology . <http://pin.primate.wisc.edu/factsheets/entry/bamboo_lemur/taxon>. Accessed 2013 May 18.