Avian Communal Breeding Systems

Communal breeding in birds may be the most critical test case for the application of Hamilton's (54, 55) ideas about inclusive fitness, altruism, and kin selection (73a) among vertebrates. Avian communal breeding systems (A CBS) feature precisely that phenomenon that seems most likely to require kin selection as a part of its evolutionary explanation, namely, the presence of a form of operational altruism [in which gains and losses are empirically determined ( 1 6)] known as helping behavior. A helper was defined by Skutch (99) as "a bird which assists in the nesting of an individual other than its mate, or feeds or otherwise attends a bird of whatever age which is neither its mate nor its dependent offspring." The obvious questions are then: (a) What selection pressures shape the kind and degree of helping behavior? (b) Is kin selection implicated in an evolutionary explanation of helping behavior, and, if so, to what extent? (c) Does helping behav­ ior qualify as altruism in the sense of Hamilton? As subjects for the study of such questions among vertebrates, communal birds have numerous practical advantages. The number of avian species having helpers runs in the hundreds. The giving of food and other kinds of aid by helpers to recipients is easily observable and quantifiable. The genetic relatedness of the partici­ pants can be determined by following successive generations of individually marked animals. Most species are diurnal, relatively conspicuous, and easily watched. Their small home ranges enable numerous social units to be monitored.

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