Difficulties with estimating and interpreting species pools and the implications for understanding patterns of diversity

Evidence has been accumulating that species pools play a major role in regulating variations in small-scale diversity. However, our ability to unambiguously estimate and interpret species pools remains a major impediment to understanding the processes that control patterns of diversity. Two main approaches have been employed to evaluate the relationships between species pools and species diversity. The direct approach has been to estimate the actual sizes of species pools by sampling discrete areas at larger spatial scales and then relating these estimates to samples taken at smaller scales. The indirect approach has been to search for correlations between abiotic environmental factors and patterns of diversity that are indicative of gradients in species pools. Both of these approaches have substantial predictive capability but also have limitations that impair our ability to draw unambiguous interpretations about causal factors. A primary difficulty for the direct approach is in deciding which species in the larger pool of potential species are actually capable of living in a sample. In this regard, the indirect approach requires fewer assumptions and has the ability to detect previously unsuspected gradients in species pools. As with the direct approach, assessing the causes for observed gradients in species pools remains a limitation for the indirect approach. Consideration of experimental studies of potential niches suggests that it may be valuable to distinguish between potential and observed species pools if the role of competitive exclusion is to be fully assessed. This paper concludes by arguing for (1) an increased use of multivariate studies that examine the effects of species pools indirectly and (2) further experimental studies designed to determine potential species pools.