Geophagy as a correlate of folivory in red-handed howler monkeys (Alouatta belzebul) from eastern Brazilian Amazonia

Howler monkeys, Alouatta spp., are the most folivorous of neotropical primates (Platyrrhini), although Amazonian red-handed howlers (Alouatta belzebul) are relatively frugivorous. The feeding ecology of a free-ranging group of A. belzebul was monitored at a site in eastern Brazilian Amazonia over a ten-month period (November 1997 to August 1998). The first half of the study period encompassed the peak of the wet season, during which the study group's diet was predominantly frugivorous (scan sample records: 53.5% fruit, 40.8% leaves), and the second half, the transition from wet to dry season, characterized by a marked shift to folivory (18.9% fruit, 77.9% leaves). This shift was accompanied by a marked increase in mature, as opposed to flush leaves, which are relatively rich in secondary compounds. Ingestion of soil from termitaria was recorded on a total of 26 occasions, all of which occurred during the second half of the study period. Soil from termitaria was relatively rich in elements such as Ca and Na and in organic carbon, in comparison with that from the forest floor. The extent to which the monkeys ingested soil for their mineral supplements, or as an aid for the digestion of leaves, in particular the absorption of secondary compounds, remains unclear. The marked correlation with the observed patterns of folivory suggests that the latter function may have been the primary motive for geophagy in this species.