The Red Queen revisited: reevaluating the age selectivity of Phanerozoic marine genus extinctions

Extinction risk is inversely related to genus age (time since first appearance) in most intervals of the Phanerozoic marine fossil record, in apparent contradiction to the macroevolu- tionary Red Queen's Hypothesis, which posits that extinction risk is independent of taxon age. Age- dependent increases in the mean species richness and geographic range of genera have been in- voked to reconcile this genus-level observation with the presumed prevalence of Red Queen dy- namics at the species level. Here we test these explanations with data from the Paleobiology Da- tabase. Multiple logistic regression demonstrates that the association of extinction risk with genus age is not adequately explained by species richness or geographic range: there is a residual asso- ciation between age and extinction risk even when range and richness effects are accounted for. Throughout most of the Phanerozoic the age selectivity gradient is highest among the youngest age cohorts, whereas there is no association between age and extinction risk among older age co- horts. Some of the apparent age selectivity of extinction in the global fauna is attributable to dif- ferences in extinction rate among taxonomic groups, but extinction risk declines with genus age even within most taxonomic orders. Notable exceptions to this pattern include the Cambrian- Ordovician, latest Permian, Triassic, and Paleocene intervals. The association of age with extinction risk could reflect sampling heterogeneity or taxonomic practice more than biological reality, but at present it is difficult to evaluate or correct for such biases. Alternatively, the pattern may reflect consistent extinction selectivity on some as-yet unidentified covariate of genus age. Although this latter explanation is not compatible with a Red Queen model if most genus extinctions have re- sulted from biological interactions, it may be applicable if most genus extinctions have instead been caused by recurrent physical disturbances that repeatedly impose similar selective pressures.