Like many concepts in ecology (metacommunity, community), the idea of a regional species pool is useful, makes conceptual sense, and is incredibly difficult to apply to real data. Originally, the idea of a species pool came from the theory of island biogeography (MacArthur and Wilson, 1967), where it referred to all the species that could disperse to an island. Today, the regional species pool appears frequently, across null models, studies of community assembly both empirical and theoretical, and metacommunity theory.
Understanding how particular processes shape community membership—whether the environmental, competition, or dispersal limitation—depends on knowing the identity of all the species that could have potentially assembled there. The species pool as defined by the research provides the frame of reference against which to consider a community's composition. Most null models of community assembly rely on correctly identifying this set of species, and worse, tend to be very sensitive to bias in how the regional pool is defined. If you include all species physically present in a region, in your species pool, environmental filtering may appear to be particularly important simply because many of those species can’t actually survive in your community (the narcissus effect). Given the importance of null models to community ecology, defining the species pool appropriately is an ongoing concern.
There are many decisions that can be made when asking 'which species could potentially be members of a community'? You could include all species that can physically arrive at a site (so only dispersal or geographic distance limits membership), or only include those species that can both arrive and establish (both dispersal and environmental conditions limit membership). Further, the availability of data is key: if you use observational data used to determine the environmental limitations, you may also incorporate the outcome of biotic interactions indirectly. If some species are rare and have low observation likelihoods, they will be under-represented. Abundances may be useful but inaccurate depending on how they are measured. Finally, it is common to define species as either present or not present for a species pool; this binary approach may conceal ecologically important information.
|The 'filtering' heuristic for understanding community membership. Species groups 1-3 could each be defined as a regional species pool, depending on the definition applied.|
Lessard et al. (2016) illustrates how to produce this kind of process-based species pool with various constraints (figure below). Their three-step approach is to:
- Define absolutely all possible members of regional pool. This is determined by identifying all assemblages in the region containing at least one species also found in the focal community (creating a 'dispersion field') (figure below, section A). This delineates a large region and identifies all species within it.
- Calculate the probability of resampling a species from the focal community elsewhere in the dispersion field. This is done in the context of the process of interest. For example, the probability of observing a species in the focal community and another community might be determined based on the geographical or environmental distance between those sites. Every site in the dispersion field would now have a probability (or distance really) associated with it, representing some similarity with the focal site.
- Finally, apply constraints to the calculated probabilities. You might choose to consider only the species within communities that are at least 50% similar to the focal community, for example. Such constraints would reflect the strength or importance of filtering by the process of interest.
Both comparative and probabilistic approaches to defining species are logical advances, and one way of dealing with the untidy concept of the species pool. If this topic is of interest, a few other papers, albeit slightly less recent, are definitely worth reading: Pigot and Etienne 2015; Lessard et al. 2012, Carsten et al., 2013.
|From Lessard et al., 2016. The three steps to build a species pool.|