Seed dispersal by long-tailed macaques.

We review here the methods by which long-tailed macaques (Macaca fascicularis fascicularis) process seeds in Bukit Timah Nature Reserve, Singapore, and the factors that influence this. Feeding observations have revealed that these macaques either destroy seeds in their mouth with their teeth, spit them out whole from the mouth after removing much of the flesh, or else clean the flesh off them at the front of the mouth and then drop them. Absence of these observed behaviors is associated with the presence of intact seeds in the feces, indicating that macaques swallow some seeds whole. All these options were exhibited by one group of macaques and were not random alternatives; evidence links their frequency to the type of fruit (fleshy vs. dry) and the size of the seed(s). Adaptations of the mouth of long-tailed macaques, such as gape and the presence of well-innervated cheek pouches and relatively large incisor and postcanine teeth, are predicted to influence these thresholds strongly. In a faunally depleted reserve like Bukit Timah, this dispersal is effective, but we see no evidence in this of coevolution between macaques and fruits. Many seeds in the macaque diet are probably dispersed more effectively by other vertebrates, such as birds, bats, gibbons, and civets, when these are present. However, in sites where large vertebrates have been eliminated, macaques may become important for dispersing large, large-seeded fruits.