Patchiness of local species richness and its implication for large-scale diversity patterns: an example from the middle Miocene of the Paratethys

The best hope for understanding global diversity patterns is to compare local assemblages, which are mostly preserved in taphonomically-complex shell beds. The present study investigates the variability in faunal composition and diversity at the scale of a single outcrop. A total of 152 species (3315 shells) occurred in 25 samples from 5 tempestitic shell beds. Although sampling intensity was high, total species richness was not captured by far at the hierarchical levels present (outcrop, shell beds, samples) because the majority of species is rare. In contrast, sampling intensity was sufficient to cover the most abundant species, as indicated by stable evenness values. Four taxa dominate the assemblage, but their rank order differs strongly between individual shell beds and individual samples; significant differences between some shell beds are evident for faunal composition, and one shell bed differs from all others with respect to species accumulation curves. Within shell beds, rarefaction curves are generally characterized by strongly overlapping confidence intervals, but outliers occur in three of five shell beds. Patchiness is additionally indicated by a wide scatter of diversity indices in some shell beds and by a wide scatter of samples of one shell bed in an ordination on faunal composition. Most of the outcrop-scale variability in faunal composition and diversity can be related to differences between shell beds. This suggests that sampling a single shell bed of the outcrop is insufficient to characterize the local fauna and its diversity, even when sampling intensity (i.e. the number of samples and shells) within the shell bed was high. Similarly, a single sample from such a shell bed may not be sufficient to characterize its diversity, even when the number of counted shells was high. It is therefore confirmed that sampling strategy and sampling intensity are crucial to confidently characterize the shelly assemblages at such a small spatial scale and that dispersed sampling effort with many small replicate samples will characterize a local assemblage and its diversity better than a few large samples. Diversity comparisons of individual samples between localities must account for the high variability present at the smaller spatial scale, as observed in our study.