Escalation and Extinction Selectivity: Morphology Versus Isotopic Reconstruction of Bivalve Metabolism

Abstract

Abstract. Studies that have tested and failed to support the hypothesis that escalated species (e.g., those with predationresistant adaptations) are more susceptible to elimination during mass extinctions have concentrated on the distribution and degree of morphological defenses in molluscan species. This morphological approach to determining level of escalation in bivalves may be oversimplified because it does not account for metabolic rate, which is an important measure of escalation that is less readily accessible for fossils. Shell growth rates in living bivalves are positively correlated with metabolic rate and thus are potential indicators of level of escalation. To evaluate this approach, we used oxygen isotopes to reconstruct shell growth rates for two bivalve species (Macrocallista marylandica and Glossus markoei) from Miocene-aged sediments of Maryland. Although both species are classified as non-escalated based on morphology, the isotopic data indicate that M. marylandica was a faster-growing species with a higher metabolic rate and G. markoei was a slower-growing species with a lower metabolic rate. Based on these results, we predict that some morphologically non-escalated species in previous tests of extinction selectivity should be reclassified as escalated because of their fast shell growth rates (i.e., high metabolic rates). Studies that evaluate the level of escalation of a fauna should take into account the energetic physiology of taxa to avoid misleading results.Key words. Escalation, extinction selectivity, isotopes, metabolism, predation, shell growth rates. A fundamental question in macroevolution is whether there are long-term, predictable patterns or trends in the history of life (Gould 1985(Gould , 1990Vermeij 1987 Vermeij , 1999. Implicit in this question are two significant and related issues: the role of biotic and abiotic factors in evolution, and the effect of mass extinctions, including their selectivity, on evolution. Two end-member points of view on these questions have been articulated most forcefully by