Diet

In comparison with the other subspecies, the diet of H. g. griseus is the best known and includes a diverse array of plants. In Analamazoatra Special Reserve (ASR), Wright (1986) reported that bamboo (Bambusa sp.) constitutes 90% of the animals' diet during the austral winter months. Like Pollock (1986), Wright also found H. g. griseus eating leafy parts of other plants and some fruit. Long-term studies conducted in RNP confirmed that bamboo is the mainstay of H. g. griseus (Grassi, 2001; Overdorff et al., 1997; Tan, 1999, 2000). In particular, at Talatakely, approximately 80% of the lemur's annual diet is comprised of bamboo (Cathariostachys madagascariensis, Cephalostachyum cf. perrieri, Cephalostachyum sp., Nastus elongatus, and Nastus sp.); parts ingested include the base of young leaves, immature pseudopetioles, branch shoots (i.e., emerged from the nodes), and ground shoots. The animals also consume a variety of nonbamboo foliage, fruit, and flowers (Tan, 1999, 2000). In fact, ongoing research (1996-2006) at Talatakely shows that the total number of food plants has exceeded 40 species for H. g. griseus (C. Tan, unpublished data).

By contrast, H. g. alaotrensis is strictly folivorous, relying solely on leaves and grasses around Lac Alaotra marshes. In a 15-month study, Mutschler (1999a,b) documented the animals eating 11 plant species; of these, 4 represented >95% of the total feeding records. Pith of papyrus stems ( Cyperus madagascariensis) as well as reed shoots (Phragmites communis), southern cut grass (Leersia hexandra), and millet (Echinochloa crusgalli) make up the principal diet (Mutschler, 1999a,b; Petter et al., 1977; Pollock, 1986).

Little is known about the feeding habits of the other subspecies of H. griseus. Petter et al. (1977) reported H. g. occidentalis eating fruits of Madagascar plum (Flacourtia ramontchi) and wild dates in ForĂȘt de Tsimembo. Additionally, in

Bemaraha and Sambirano, the animals were observed feeding on items such as liana flowers and pseudopetioles of a woody bamboo (Mutschler and Tan, 2003). There is no published dietary information on H. g. meridionalis to date.

The diet of H.. aureus is known only from Talatakely, RNP. Nearly 90% of the animals' diet is comprised of bamboo, primarily the giant bamboo (Cathariostachys madagascariensis) (Tan, 1999, 2000). Whereas young leaf bases and young pseudopetioles of bamboo are consumed throughout the year, branch shoots and ground shoots are seasonal food items. H. aureus also supplements its diet with leaves, stems, fruit, and flowers of other plants and mushrooms. There is extensive dietary overlap between H. aureus and H. g. griseus; however, the food diversity of H. aureus is relatively lower.

Bamboo culm pith and shoots represent the main components of the diet of H. simus (Petter et al., 1977; Tan, 1999, 2000; Wright and Randriamanantena, 1989). At Talatakely, RNP, H. simus specializes on the giant bamboo, which accounts for 95% of the annual diet (Tan, 1999, 2000). Unlike H. aureus and H. g. griseus that feed on only the tender sections of bamboo leaves (i.e., base of young leaves and immature pseudopetioles), H. simus consumes all parts of leaves, including mature leaf blades. Furthermore, H. simus differs from the two sym-patric congeners in having a strikingly seasonal diet—shifting from eating mainly ground shoots in the warm-wet season to culm pith in the cold-dry season (Tan, 1999, 2000). In highly disturbed forests, H. simus has been observed to survive on various plant materials. For example, in Kianjavato, animals were seen raiding rice crops and feeding on shoots of a woody bamboo (Valiha diffusa), young leaf bases of a viny bamboo, flowers of traveler's palm (Ravenala madagascariensis), jackfruit (Artocarpus integrifolius), mango (Mangifera indica), figs (Ficussp.), palm (Dypsis sp.), and leaves of African ginger (Aframomum sp.) and kikuyugrass (Pennisetum clandestinum) (Meier and Rumpler, 1987; C. Tan, unpublished data).

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