Paleoenvironmental basis of cognitive evolution in great apes.

A bias favoring tree-dominated habitats and ripe-fruit frugivory has persisted in great ape evolution since the early Miocene. This bias is indicated by fossil ape paleoenvironments, molar morphology, dental microwear, the geographic pattern of extinctions, and extant apes' reliance on wooded settings. The ephemeral aspect of high-quality fruit has placed a premium on cognitive and social means of finding and defending food sources, and appears related to great apes' affinity since the Miocene for wooded, fruit-rich environments. These habitats have, however, undergone a severe withdrawal toward the low latitudes of Africa and Southeast Asia since the late Miocene, corresponding to a decline in the diversity of great apes beginning 9.5 million years ago. Plio-Pleistocene records imply that wooded settings of Africa and SE Asia were prone to substantial fragmentation and coalescence. Once apes were confined to equatorial settings, therefore, habitat instability heightened the spatial/temporal uncertainty of ripe-fruit sources. Prolonged learning, the assignment of attributes to distant places, mental representation, and reliance on fallback foods were all favored in this dynamic environmental context. These abilities helped sustain forest frugivory in most lineages. Fluid social grouping afforded the animals opportunities to locate ephemeral foods in continuous and fragmented forests. Fission-fusion grouping also magnified the problems of object impermanence (of individuals) and dispersion manifested by food sources in the ecological realm. Thus the spatial and temporal dynamics of fruit and wooded habitats since the Miocene are reflected in important components of great ape cognition, foraging, and sociality. In contrast to great apes, cercopithecoid monkeys have increased their plant dietary options and diversified in seasonal environments since the late Miocene. Early hominins eventually severed the habitat bias that characterized the evolution of great apes, and later expanded into diverse environments.