Biotic Response in Aquatic Reptiles (Testudines) during Earliest Eocene Climatic Warming

The earliest Eocene is marked by significant events of global warming: the Paleocene-Eocene Thermal Maximum (PETM) at ~55.8 Ma and two short-lived events (ETM2 or Elmo and H2) approximately 2 Ma later. These environmental changes induced strong responses in the continental biota. Noteworthy changes in North American mid-latitude faunas and floras that are temporally correlated with earliest Eocene warming events include: increased diversity; turnover; and significant range changes, comprising both northward shifts in ranges of North American taxa as well as intercontinental dispersal across Holarctica. Evidence for these biotic changes comes directly from the fossil record and indirectly from phylogeographic analyses of molecular phylogenies of extant biota. To date, the stratigraphic record of biotic change has only been examined for the flora and terrestrial mammals. Data on reptiles and for continental aquatic systems are particularly lacking. In order to assess the impact of climate-mediated faunal change in aquatic systems during early Paleogene warming, we have focused on developing a detailed record of fossil turtles (Testudines) from the Bighorn Basin of Wyoming, where these records can be directly compared to similarly studied mammalian and floral data and to isotopic studies that provide independent proxies of climate change. Using genus-level occurrence data from more than 450 stratigraphically-constrained localities spanning ~2.5 Ma, we calculated first and last appearances, taxonomic richness, and relative abundance as measured by presence-absence (site occupancy). Among turtles, taxonomic richness increased episodically through the earliest Eocene with two new taxa appearing at the PETM, two immediately following it, and two at Biohorizon B, an interval associated with the younger hyperthermals. These new, immigrant taxa eventually comprised 40% of known generic richness. Phylogenetically, the inferred biogeographic source regions are southern North America and Asia, with an equal number of taxa originating in each area. Although immigrant taxa comprised less than half of the known earliest Eocene diversity, their relative dominance in these assemblages varied markedly. Within the PETM interval, immigrant taxa comprise nearly 70% of occurrences. Post-PETM, as temperatures cooled, immigrant taxa and taxa persisting from the Paleocene showed greater evenness, but immigrant taxa again became dominant with renewed warming. Among immigrant taxa, intercontinental dispersers are much more common than those that that dispersed from southern North America. These data are consistent with and stratigraphically correlative with significant changes in the mammalian fauna and flora of the Bighorn Basin and underline the importance of climatic change as a driver in these events. However, the magnitude and relative importance of intra- vs. intercontinental dispersal has not yet been fully examined in other taxonomic groups. The asymmetry of response following immigration that we observe in turtles may be taxon-specific, unique to aquatic systems, or may illustrate a more general pattern of how biotas respond to significant climate change.