Abstract
The Darwin's finches on Isla Daphne Major, Galapagos, were studied between July 1975 and June 1978. Geospiza fortis and G. Scandens are residents, while G. fuliginosa and G. magnirostris are regular immigrants. The Daphne climate is unpredictably dry. The island has a simple plant community displaying marked annual and spatial variation in the foods which form finch diets. Breeding is stimulated by rain falling irregularly between January and April; G. scandens laid eggs with as little as 16 mm of rain, but G. fortis required 35 mm or more before laying. G. scandens consistently bred prior to the rains, associated with specialized exploitation of dry season Opuntia cactus flowers. The breeding system of both species was similar to that of other Geospiza species: monogamous matings on small, permanent, all—purpose territories. Reproductive output of both species varied. In 1976 single broods were produced by both species at high densities, with modes of three young. In 1977, only 24 mm of rain fell during the breeding season and G. scandens alone bred, with poor success. By 1978, G. scandens populations had declined by 66% and G. fortis by 85%. In 1978, both species laid an average of three clutches per pair, with a mode of four young per brood. During the 1977 drought, the sex ratio became skewed in favor of males in both species, and as a consequence some females bred successively with up to three different males in 1978. The skewed sex ratio retarded population recovery following the drought. The population decline was associated with a decline in seed abundance in the drought. As food disappeared, G. fortis diets widened to include a broad selection of food items, while G. scandens diets contracted to Opuntia seeds. Even in normal years, both species showed pronounced seasonal variation in feeding habits. Both fed heavily on Opuntia flowers in the late dry season, followed by a mixed diet of insects, fresh seed, and other green matter during the breeding season. After breeding, the birds fed primarily on seeds, the two species selecting different proportions of the available range of seed sizes. Overall, G. fortis displayed the feeding, breeding, and population dynamics characteristic of an ecologically generalized species, whereas G. scandens is a highly specialized species. Large variation in food supply, caused by large variation in rainfall, is responsible for the presence of only two species with breeding populations, for large fluctuations in their population sizes, and for their large clutch sizes and opportunistic breeding. Long—term studies are essential for an understanding of communities in such variable environments because some important events are rare.
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