Revised perspectives on Devonian biozonation and environmental volatility in the wake of recent time-scale revisions

Revised time-scales for the Devonian Period have highlighted major discrepancies in the frequency of evolutionary, eustatic, carbon cycle, and biotic events. The number of conodont biozones per million years shows strong variation among stages. Using two alternative time scales, the lowest values are in the Emsian and highest in the Givetian, Frasnian, and Famennian stages. A similar result is obtained by dividing the Devonian time scale into 12 equal 5-million year bins and determining the number of conodont zonal subdivisions in each bin. The record of ammonoid biozones also shows interesting parallels. The acuity of biostratigraphic zonation and zonal subdivisions appears to be correlative with the frequency of environmental perturbations, including eustatic, positive carbon isotopic excursions, and hypoxic/anoxic related bioevents, and these are similarly unevenly distributed. All calculations of environmental volatility metrics show the late Eifelian through early Frasnian to be the most volatile interval of the Devonian, followed by the mid-late Famennian, while the Lochkovian through Emsian and mid-Frasnian through early Famennian are the least volatile. Furthermore, the majority of major bioevents occur in the Eifelian through early Frasnian, and late Famennian. The strong evidence for higher biotic and environmental volatility in the late Eifelian through early Frasnian and late Famennian rock record corresponds to the coolest paleoclimate conditions in the Devonian according to recent conodont apatite delta O-18 compilations, and the transition from an overall cooling to warming paleotemperature trend during the mid-Givetian. Regardless of mechanism, strong variations in volatility occurred during the Devonian Period. Intervals with greater biostratigraphic resolution also show much stronger environmental volatility including a series of relatively quasi-stable conditions punctuated by major turnovers with abrupt sea-level rise, widespread hypoxia, and changes in the carbon cycle. It was during these intervals that much net evolutionary and ecological change appears to have taken place.