The importance of phylogenetic analysis for the assessment of species turnover: a case history of Paleocene mammals in North America

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Abstract
During the latest Cretaceous and the Paleocene in western North America, disappearance rates for mammalian genera track appearance rates, both reaching their peak in the early Paleocene (Puercan) following the extinction of non-avian dinosaurs. Some of the disappearances during this time were pseudoextinctions that resulted when ancestral species disappeared during speciation.Species-level cladistic analyses and a well-constrained biostratigraphic framework are required to study this form of pseudoextinction. Cladistic analyses show that monophyly cannot be established or rejected for some species because these species lack autapomorphies (uniquely derived character states) that unite their constituent members. Such taxa, termed metaspecies, are potential ancestors to species and higher clades with which they share a node in the cladogram.A hypothetical species-level cladistic analysis coupled with three different hypothetical biostratigraphies shows how different models of speciation (bifurcation, budding, or anagenesis) result in very different patterns of true versus pseudoextinction. Depending on the speciation model, true extinction can be overestimated by as much as a factor of four, raising the specter of mass extinction. Species-level studies for three early Tertiary mammalian taxa—taeniodont eutherians, taeniolabidid multituberculates, and periptychid ungulates—use the same procedures. They show that almost 25% of disappearances during the early Paleocene (Puercan) for species in the analysis were pseudoextinctions of metaspecies. Budding and anagenetic-like peripatric speciation, but not bifurcation, are seen in the three examples.Equating disappearance to true extinction can profoundly affect interpretations of faunal turnover, especially during mass extinctions or major faunal reorganizations. Some authors use pseudoextinction to describe the taxonomic rather than evolutionary disappearance of nonmonophyletic groups. Pseudoextinction, as used here refers only to the evolutionary disappearance of metaspecies via speciation. Both usages seem appropriate but should not be confounded.