Large-scale heterogeneity of the fossil record: implications for Phanerozoic biodiversity studies.

Patterns of origination, extinction and standing diversity through time have been inferred from tallies of taxa preserved in the fossil record. This approach assumes that sampling of the fossil record is effectively uniform over time. Although recent evidence suggests that our sampling of the available rock record has indeed been very thorough and effective, there is also overwhelming evidence that the rock record available for sampling is itself distorted by major systematic biases. Data on rock outcrop area compiled for post-Palaeozoic sediments from Western Europe at stage level are presented. These show a strongly cyclical pattern corresponding to first- and second-order sequence stratigraphical depositional cycles. Standing diversity increases over time and, at the coarsest scale, is decoupled from surface outcrop area. This increasing trend can therefore be considered a real pattern. Changes in standing diversity and origination rates over time-scales measured in tens of millions of years, however, are strongly correlated with surface outcrop area. Extinction peaks conform to a random-walk model, but larger peaks occur at just two positions with respect to second-order stratigraphical sequences, towards the culmination of stacked transgressive system tracts and close to system bases, precisely the positions where taxonomic last occurrences are predicted to cluster under a random distribution model. Many of the taxonomic patterns that have been described from the fossil record conform to a species-area effect. Whether this arises primarily from sampling bias, or from changing surface area of marine shelf seas through time and its effect on biodiversity, remains problematic.