Population size, extinction, and speciation: the fission effect in Neogene Bivalvia

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Abstract
The extinction of a species represents reduction of both geographic range and population size to zero. Most workers have focused on geographic range as a variable strongly affecting the vulnerability of established species to extinction, but Lyellian percentages for Neogene bivalve faunas of California and Japan suggest that population size is a more important variable along continental shelves. The data employed to reach this conclusion are Lyellian percentages for latest Pliocene (∼2 ma old) bivalve faunas of California and Japan (N = 245 species). These regions did not suffer heavy extinction during the recent Ice Age, and for each region the Lyellian percentage is 70%–71%.Discrepancies in population size appear to explain the following differences in survivorship to the Recent (Lyellian percentage) for three pairs of subgroups: (1) burrowing nonsiphonate species (42%) versus burrowing siphonate species (84%), which suffer less heavy predation; (2) burrowing nonsiphonate species of small size (73%) versus burrowing nonsiphonate species of large body size (96%); (3) Pectinacea (30%) versus other epifauna (71%), which suffer less heavy predation. During the Mesozoic Era, when predation was less effective in benthic settings, mean species duration for the Pectinacea was much greater (∼20 ma).Along the west coast of North and Central America, mean geographic range is greater for siphonate species of large body size than for siphonate species of small body size and greater still for pectinacean species. These ranges are inversely related to mean species longevity for the three groups, which indicates that geographic range is not of first-order importance in influencing species longevity. Species with nonplanktotrophic development neither exhibit narrow geographic ranges along the west coast of North and Central America nor have experienced high rates of extinction in California and Japan.Rates of extinction are so high for Neogene pectinaceans and nonsiphonate burrowers that without enjoying high rates of speciation these groups could not exist at the diversities they have maintained during the Neogene Period. They are apparently speciating rapidly because of the fission effect: the relatively frequent generation of new species from populations that are fragmented by heavy predation. Thus, ironically, there may be a tendency for high rates of speciation to be approximately offset by high rates of extinction. Only if mean population size for species in a particular group becomes extremely small is it likely to result in a high rate of extinction and a low rate of speciation—and hence a dramatic decline of the group. The fission effect may contribute to the general correlation in the animal world between rate of speciation and rate of extinction.