The causes of extinction

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Abstract
A species may go extinct either because it is unable to evolve rapidly enough to meet changing circumstances, or because its niche disappears and no capacity for rapid evolution could have saved it. Although recent extinctions can usually be interpreted as resulting from niche disappearance, the taxonomic distribution of parthenogens suggests that inability to evolve may also be important. A second distinction is between physical and biotic causes of extinction. Fossil evidence for constant taxonomic diversity, combined with species turnover, implies that biotic factors have been important. A similar conclusion emerges from studies of recent introductions of predators, competitors and parasites into new areas. The term 'species selection' should be confined to cases in which the outcome of selection is determined by properties of the population as a whole, rather than of individuals. The process has been of only trivial importance in producing complex adaptations, but of major importance in determining the distribution of different types of organisms. An adequate interpretation of the fossil record requires a theory of the coevolution of many interacting species. Such a theory is at present lacking, but various approaches to it are discussed.