Causality and Cope's Rule: evidence from the planktonic foraminifera

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Abstract
A literature-based compilation of phylogenetic relationships and biometric measurements of 342 Cenozoic planktonic foraminiferal species suggests that the group shows a net increase in size through the Cenozoic, thus appearing to follow Cope's Rule of phyletic size increase. However, when the data are corrected for size-related biases, they do not support the hypothesis that this apparent trend is driven by an organismal adaptive advantage of larger size.When the planktonic foraminifera return to their “primitive” globigerine morphology during the Eocene-Oligocene transition, there is no indication of size-dependent origination or extinction; however, when the extinction signal is decomposed into pseudoextinctions and true lineage terminations, a differential pulse of pseudoextinction is observed among the smaller forms. This observation suggests that smaller bodied species, rather than surviving stressful times with static morphologies, may evolve their way through times of crisis and go on to found lineages which, by virtue of their initial small size, are stochastically likely to increase in mean size during subsequent diversification. Thus, one general explanation for Cope's Rule might be that smaller bodied species are more adaptively responsive due to their tendency to have shorter generation times. During times of stress, this adaptive responsiveness may give them an advantage that is correlated with, but causally unrelated to, their size.