Abstract

Estimates of taxonomic richness and abundance are complicated by sampling biases. The failure to sample rare taxa is most often attributed to inadequate sampling and to removal during the process of sample-size standardization. Here I present two methods for unveiling rare diversity by integrating species presence/absence data from museum collections and the literature with quantitative estimates of species richness and abundance gathered from field-based bulk samples. Combining museum, literature, and field data can provide a more comprehensive estimate of taxonomic richness and abundance without substantial increase in current sampling effort. First, in a given bulk sample, the lowest proportional abundance value observed can be used to estimate the maximum abundance of rare species known to have occurred at the locality at least once but not recorded in the current sample. Second, a model-selection approach can be used, in which a set of relative abundance distribution models are fit to the bulk-sample abundance data and the parameter estimates for the best model used to calculate the abundance distribution for all species known from the locality. The Paleogene marine fossil record of the U.S. Gulf Coastal Plain is suitable for applying these methods, because (1) the molluscan fauna is well represented in museum collections and the literature, (2) the molluscan fauna has been taxonomically standardized, and (3) many classic localities remain accessible for standardized bulk sampling. I introduce these methods by applying them to a single locality and then, using the faunas of the Gosport, Moodys Branch, and Red Bluff Formations, I demonstrate how the model-fitting approach can be used to compare taxonomic richness among multiple localities. A substantial fraction of the molluscan richness known from each locality is not captured in bulk samples and much of this unobserved richness may be attributed to the rarity of species. The multiple-locality comparison suggests that the greatest Paleogene decline in standing richness occurred in the middle Eocene and that the recovery of richness following the Eocene-Oligocene extinction was quite rapid despite substantial loss of taxa. These analyses underscore the magnitude of veiled diversity in marine fossil assemblages and the potential of existing sources of data to unveil rare taxa, allowing them to be incorporated into quantitative diversity studies.

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