Ecological, taxonomic, and taphonomic components of the post-Paleozoic increase in sample-level species diversity of marine benthos

Abstract

Biological veracity of the sharp diversity increase observed in many analyses of the post-Paleozoic marine fossil record has been debated vigorously in recent years. To assess this question for sample-level (“alpha”) diversity, we used bulk samples of shelly invertebrates, representing three major fossil groups (brachiopods, bivalves, and gastropods), to compare the Jurassic and late Cenozoic sample-level diversity of marine benthos. After restricting the data set to single-bed, whole-fauna, bulk samples (n≥ 30 specimens) from comparable open marine siliciclastic facies, we were able to retain 427 samples (255 Jurassic and 172 late Cenozoic), with most of those samples originating from our own empirical work.Regardless of the diversity metric applied, the initial results suggest that standardized sample-level species (or genus) diversity, driven by evenness and/or richness of the most common taxa, increased between the Jurassic and late Cenozoic by at least a factor of 1.6. When the data are partitioned into the three dominant higher taxa, it becomes clear that (1) the bivalves, which dominated the samples for both time intervals, increased in sample-level diversity between the Jurassic and the late Cenozoic by a much smaller factor than the total fauna; (2) the removal of brachiopods, which were a noticeable component of the Jurassic samples, did not significantly affect standardized sample-level diversity estimates; and (3) the gastropods, which were rare in the Jurassic but common in many late Cenozoic samples, contributed notably to the increase in sample-level diversity observed between the two time intervals. Parallel to these changes, the samples revealed secular trends in ecological structure, including Jurassic to late Cenozoic increases in proportion of (1) infauna, (2) mobile forms, and (3) non-suspension-feeding organisms. These trends mostly persist when data are restricted to bivalves.Supplementary analyses indicate that these patterns cannot be attributed to sampling heterogeneities in paleolatitudinal range, lithology, or paleoenvironment of deposition. Likewise, when data are restricted to samples dominated by species with originally aragonitic shells, the observed temporal changes persist at a comparable magnitude, suggesting that the pervasive loss of aragonite in the older fossil record is unlikely to have been the primary cause of the observed patterns. The comparable ratio of identified to unidentified species and genera, observed when comparing the Jurassic and late Cenozoic samples, indicates that the relatively poorer (mold/cast) preservation of Jurassic aragonite species also is unlikely to have been responsible for the observed patterns. However, the diagenesis-related taphonomic and methodological artifacts cannot be ruled out as an at least partial contributor to the observed post-Paleozoic changes in diversity, taxonomic composition, and ecology (the outcomes of the three tests of the diagenetic bias available to us are incongruent).The study demonstrates that the post-Paleozoic trends in the sample-level diversity, ecology, and taxonomic structure of common taxa can be replicated across multiple studies. However, the diversity increase estimated here is much less prominent than suggested by many previous analyses. The results also narrow the list of causative explanations down to two testable hypotheses. The first isdiagenetic bias—a spurious trend driven by either (a) increasing taphonomic loss of small specimens in the older fossil record or (b) a shift in sampling procedures between predominantly lithified rocks of the Mesozoic and predominately unlithified, and therefore sievable, sediments of the late Cenozoic. The second hypothesis isgenuine biological changes—macroevolutionary trends in the structure of marine benthic associations through time, consistent with predictions of several related models such as evolutionary escalation, increased ecospace utilization, and the Mesozoic marine revolution. Future studies should focus on testing these two rival models, a key remaining challenge for identifying the primary causative mechanism for the long-term changes in sample-level diversity, ecology, and taxonomic structure observed in the Phanerozoic marine fossil record.