Ontogeny and the explanation of form: an allometric analysis

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Abstract
Significant allometry occurs during the ontogeny of every variable in Poecilozonites bermudensis, a Pleistocene-Recent land snail from Bermuda. The adaptive significance of shell allometry may lie in the necessity for preserving a high value of the foot surface/body volume ratio. In the absence of allometry, this ratio must decline as size increases. Three strategies could be used to keep this ratio sufficiently high: positive allometry of foot growth, structural strengthening of the foot, and the development of a foot initially large enough to withstand decline in the ratio during growth. As indicated by ontogenetic changes in apertural shape, a small amount of positive allometry occurs during ontogeny of the foot in P. bermudensis. This is not sufficient to prevent decline of the foot surface/body weight ratio, and I conclude that the strategy of possessing an initially large foot is also used. A simple model of doming, reflecting this latter strategy, is constructed (doming is a major allometric feature of P. bermudensis). In this model, the foot volume/body volume ratio is constant throughout ontogeny in each of two shells, but this value is higher in the more strongly domed shell.Knowledge of ontogenetic allometry is a prerequisite for understanding the phylogeny of P. bermudensis, for paedomorphosis has been the primary evolutionary event in this taxon. Paedomorphic samples are scaled-up replicas of juvenile shells of the central stock, P. bermudensis zonatus. The degree of ontogenetic retardation in development is the same for all variables (color, thickness, and external shape). Paedomorphosis has occurred several times during the Pleistocene, providing an example of iterative evolution at the infraspecific level. Four paedomorphic taxa are known: P. b. fasolti, P. b. siegmundi, P. b. sieglindae, all new; and P. b. bermudensis (Pfeiffer). They have the geographic distribution (small, peripheral isolates) expected of diverging populations and seem to be genetically distinct entities, not mere phenotypic variants. The most paedomorphic subspecies originated in red soils; paedomorphs did not evolve in times of carbonate-dune deposition. The thin shells of paedomorphs might have been adaptive in the low-calcium environment of red soils. The general significance of iterative evolution at the infraspecific level does not provide an adequate model for corresponding events at higher levels.

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