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210 FELDMAN et al. observing another animal interacting with that stimulus. The behavior itself is then acquired by individual learning which is the result of the animal's own experience of the consequences of its actions. If, as in this case, primitive forms of social learning can be regarded as fairly simple extensions of individual learning, the principal precursor of social learning and cultural transmission may well have been a capacity for individual learning (Laland et al., 1993). Hence it is pertinent to ask what conditions might favor the evolution of social learning in its earliest phases from individual learning.

This question has been addressed theoretically by Boyd and Richerson (1985, 1988) and Rogers (1988). Their basic premises are as follows. The environment may change between generations, and there is an optimal behavior appropriate to each environmental state. Individual learners achieve this optimal behavior on their own. Social learners, on the other hand, must copy an animal of the immediately preceding generation. The term" copy" is used broadly to include all means by