Allometry and heterochrony in an Eocene echinoid lineage: morphological change as a by-product of size selection

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Abstract
Three closely related species of the irregular echinoid Oligopygus succeed one another in the late Eocene deposits of Florida. This apparent lineage, where each species is similar but larger than the preceding species, offers an excellent opportunity to examine relationships between heterochrony and environmental change. Bivariate and multivariate techniques illustrate patterns of simple and complex allometry and indicate that simple size increase related to changes in the timing of maturation account for most of the morphological differences among species. Those differences that are not the by-products of size increase are due to heterochronic changes dissociated from the otherwise global (i.e., whole organism) hypermorphosis and were probably needed to increase the relative food intake of the larger forms. Unlike the vast majority of allometric studies, which show constant growth ratios, the ontogenies analyzed here often exhibit curvilinear trajectories. Thus, heterochronic events are seen as systematic alterations of already changing growth ratios. It is suggested that these heterochronic changes provided quick, efficient responses to selective pressures on a plurality of characters and behaviors. As the Florida carbonate environment became more stable, responses associated with K-type life history strategies were favored, resulting in “size selection.” Thus, the morphological differentia among these species may not have resulted from selection directed at those traits, but were either simple sequelae of size increase or of changes which occurred to permit the size increase.