Review
Phylogenetic analysis of community assembly and structure over space and time

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Evolutionary ecologists are increasingly combining phylogenetic data with distributional and ecological data to assess how and why communities of species differ from random expectations for evolutionary and ecological relatedness. Of particular interest have been the roles of environmental filtering and competitive interactions, or alternatively neutral effects, in dictating community composition. Our goal is to place current research within a dynamic framework, specifically using recent phylogenetic studies from insular environments to provide an explicit spatial and temporal context. We compare communities over a range of evolutionary, ecological and geographic scales that differ in the extent to which speciation and adaptation contribute to community assembly and structure. This perspective allows insights into the processes that can generate community structure, as well as the evolutionary dynamics of community assembly.

Section snippets

Bringing phylogeny into community ecology

Recent years have seen a growing interest in incorporating phylogenetic data into studies of community assembly and structure, ranging from the scale of individual guilds 1, 2 to whole biomes [3]. The basic premise is that phylogenetic data provide a historical framework to quantify evolutionary and ecological patterns and infer evolutionary and ecological processes. There have now been multiple studies examining the tendency of species to either maintain or shift their niches, and the

When are immigration and speciation important for community assembly?

The phylogenetic structure of a community is strongly dependent on the number and identity of available colonists (Figure 1). An unpopulated area in close proximity to a source of colonists will be colonized rapidly from that source. Under this scenario where immigration is important, assembly patterns are largely stochastic and dictated by propagule pressure, although turnover eventually leads to a more deterministic set of members within a given community [12]. All else being equal, it is

How are communities phylogenetically structured?

Many studies have now shown that patterns of co-occurrence can deviate from random expectations with regard to phylogenetic relatedness, either as phylogenetic overdispersion or phylogenetic clustering (Table 1). When species enter a community through immigration from a regional pool of species, community assembly can be largely neutral in that species have an equal chance of establishment or, alternatively, can be shaped by ecological processes. Here again, islands – and the temporal framework

Why are communities phylogenetically structured?

Depending upon the spatial and temporal scale, patterns of both phylogenetic overdispersion and clustering can be consistent with either environmental filtering or competitive interaction selecting for species composition within a community (Figure 3). To understand the relative importance of environmental filtering and competitive interaction requires an evolutionary analysis of traits that are important for niche occupancy. If traits associated with niche exhibit phylogenetic conservatism,

Is community assembly a predictable process?

The theory of island biogeography 21, 22 assumes that immigration is essentially neutral with respect to the identity of the colonists. However, experiments have shown that, although a community can support more species in the early phase of assembly while most species are rare (termed the noninteractive phase) 12, 61, as population sizes increase, and competition likely becomes stronger, the species number drops (interactive equilibrium), with more highly coadapted sets of species tending to

Conclusions

The incorporation of phylogenetics into community ecology has allowed key insights into the assembly and structure of communities. Here we have used insular environments to provide a temporal context for understanding the role of spatial and temporal scale in dictating pattern. Although much remains to be understood, we can draw some preliminary conclusions. Spatial (isolation) and time (for adaptation) interact to dictate first whether a niche is more likely to be filled by immigrants or

Acknowledgements

We thank G.K. Roderick and three anonymous reviewers for their suggestions. Our research is funded by the Natural Environment Research Council (B.C.E.) and the National Science Foundation (R.G.G.).

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