Evolutionary paleontologists seek explanations of form and pattern in a diversity of contexts. A review of this diversity is presented to underscore the difficulty of producing a general model of the factors controlling the morphology of real organisms. A model specific to the marine prosobranch gastropod radula identifies seven factors contributing to form and pattern: (1) phylogenetic, (2) mechanical, (3) ecological, (4) programmatic, (5) maturational, (6) degenerative, and (7) constructional. Aspects of radular morphology attributable to each factor are illustrated in scanning electron micrographs of radulae of marine prosobranch gastropods. Applications of and extrapolations from the radular model are made to promote recognition of a broader range of interacting factors in other systems. Some aspects of form and pattern convey important information about non-evolutionary processes and phenomena and are best examined outside of the evolutionary framework.
Six distinctive types of fossiliferous marine deposits occur on the Galádpagos Islands that provide evidence for the age of emergence of the islands above sea level and hence a maximum age for the islands' terrestrial biota. These subtidal to supratidal deposits include (i) volcanic tuffs with fossils, (ii) limestones and sandstones interbedded with basalt, (iii) terrace deposits, (iv) beach rock, (v) supratidal talus deposits, and (vi) recently uplifted tidal and subtidal rocks and sand. With the exception of (vi), the deposits were previously assigned ages varying from Miocene to Pleistocene, but all are less than about 2 million years old. This age, together with independently determined geologic ages, indicate that the islands emerged from the sea relatively recently and that all evolution of the islands' unique terrestrial biota occurred within the past 3 to 4 million years.
The structural diversity of gastropod veliger larvae offers an instructive counterpoint to the view of larval forms as conservative archetypes. Larval structure, function, and development are fine-tuned for survival in the plankton. Accordingly, the study of larval adaptation provides an important perspective for evolutionary-developmental biology as an integrated science. Patterns of breakage and repair in the field, as well as patterns of breakage in arranged encounters with zooplankton under laboratory conditions, are two powerful sources of data on the adaptive significance of morphological and microsculptural features of the gastropod larval shell. Shells of the planktonic veliger larvae of the caenogastropod Nassarius paupertus [GOULD] preserve multiple repaired breaks, attributed to unsuccessful zooplankton predators. In culture, larvae isolated from concentrated zooplankton samples rapidly repaired broken apertural margins and restored the "ideal" apertural form, in which an elaborate projection or "beak" covers the head of the swimming veliger. When individuals with repaired apertures were reintroduced to a concentrated mixture of potential zooplankton predators, the repaired margins were rapidly chipped and broken back. The projecting beak of the larval shell is the first line of mechanical defense, covering the larval head and mouth and potentially the most vulnerable part of the shell to breakage. Patterns of mechanical failure show that spiral ridges do reinforce the beak and retard breakage. The capacity for rapid shell repair and regeneration, and the evolution of features that resist or retard mechanical damage, may play a more prominent role than previously thought in enhancing the ability of larvae to survive in the plankton.
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