The metacommunity concept is an important way to think about linkages between different spatial scales in ecology. Here we review current understanding about this concept. We first investigate issues related to its definition as a set of local communities that are linked by dispersal of multiple potentially interacting species. We then identify four paradigms for metacommunities: the patch-dynamic view, the species-sorting view, the mass effects view and the neutral view, that each emphasizes different processes of potential importance in metacommunities. These have somewhat distinct intellectual histories and we discuss elements related to their potential future synthesis. We then use this framework to discuss why the concept is useful in modifying existing ecological thinking and illustrate this with a number of both theoretical and empirical examples. As ecologists strive to understand increasingly complex mechanisms and strive to work across multiple scales of spatio-temporal organization, concepts like the metacommunity can provide important insights that frequently contrast with those that would be obtained with more conventional approaches based on local communities alone.

Ecologists have identified several kinds of pattern in the distribution of species among sites, including a) nested subsets, b) checkerboards, c) Clementsian gradients, d) Gleasonian gradients, and e) evenly spaced gradients. Most past efforts to diagnose such patterns have focused on only one at a time, often contrasted with a sixth type of pattern, f) “randomness”. While there are statistical tests to distinguish each of the first five patterns from randomness, there are currently no established methods for discriminating among these first five patterns in a given data set. Here we propose a method that will identify which of these possibilities is most prevalent in a site‐by‐species incidence matrix based on three basic aspects of meta‐community structure. Our method is based on first ordinating the incidence matrix to identify the dominant axis of variation and identifying three aspects variation along this dominant axis. The first aspect, “coherence”, is the degree to which pattern can be collapsed into a single dimension. The second, “species turnover”, describes the number of species replacements along this dimension. The third aspect, “boundary clumping”, has to do with how the edges of species boundaries are distributed along this dimension. We present methods for analyzing these three aspects of meta‐community structure, use them to identify the six different patterns, and illustrate them with a representative set of cases drawn from previously published data.

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The diversity of life is heterogeneously distributed across the Earth. A primary cause for this pattern is the heterogeneity in the amount of energy, or primary productivity (the rate of carbon fixed through photosynthesis), available to the biota in a given location. But the shape of the relationship between productivity and species diversity is highly variable. In many cases, the relationship is 'hump-shaped', where diversity peaks at intermediate productivity. In other cases, diversity increases linearly with productivity. A possible reason for this discrepancy is that data are often collected at different spatial scales. If the mechanisms that determine species diversity vary with spatial scale, then so would the shape of the productivity-diversity relationship. Here, we present evidence for scale-dependent productivity-diversity patterns in ponds. When the data were viewed at a local scale (among ponds), the relationship was hump-shaped, whereas when the same data were viewed at a regional scale (among watersheds), the relationship was positively linear. This dependence on scale results because dissimilarity in local species composition within regions increased with productivity.

The neutral theory for community structure and biodiversity is dependent on the assumption that species are equivalent to each other in all important ecological respects. We explore what this concept of equivalence means in ecological communities, how such species may arise evolutionarily, and how the possibility of ecological equivalents relates to previous ideas about niche differentiation. We also show that the co-occurrence of ecologically similar or equivalent species is not incompatible with niche theory as has been supposed, because niche relations can sometimes favor coexistence of similar species. We argue that both evolutionary and ecological processes operate to promote the introduction and to sustain the persistence of ecologically similar and in many cases nearly equivalent species embedded in highly structured food webs. Future work should focus on synthesizing niche and neutral perspectives rather than dichotomously debating whether neutral or niche models provide better explanations for community structure and biodiversity.

Ecophylogenetics can be viewed as an emerging fusion of ecology, biogeography and macroevolution. This new and fastgrowing field is promoting the incorporation of evolution and historical contingencies into the ecological research agenda through the widespread use of phylogenetic data. Including phylogeny into ecological thinking represents an opportunity for biologists from different fields to collaborate and has provided promising avenues of research in both theoretical and empirical ecology, towards a better understanding of the assembly of communities, the functioning of ecosystems and their responses to environmental changes. The time is ripe to assess critically the extent to which the integration of phylogeny into these different fields of ecology has delivered on its promise. Here we review how phylogenetic information has been used to identify better the key components of species interactions with their biotic and abiotic environments, to determine the relationships between diversity and ecosystem functioning and ultimately to establish good management practices to protect overall biodiversity in the face of global change. We evaluate the relevance of information provided by phylogenies to ecologists, highlighting current potential weaknesses and needs for future developments. We suggest * Address for correspondence (E-mail: nmouquet@univ-montp2.fr). † These authors contributed equally to this work. that despite the strong progress that has been made, a consistent unified framework is still missing to link local ecological dynamics to macroevolution. This is a necessary step in order to interpret observed phylogenetic patterns in a wider ecological context. Beyond the fundamental question of how evolutionary history contributes to shape communities, ecophylogenetics will help ecology to become a better integrative and predictive science.

Trophic structure, the partitioning of biomass among trophic levels, is a major characteristic of ecosystems. Most studies of the forces that shape trophic structure emphasize either "bottom-up" or "top-down" regulation of populations and communities. Recent work has shown that these two forces are not mutually exclusive alternatives, but efforts to model their interaction still often yield unrealistic predictions. We focus on the problems involved with modeling situations in which community composition, including both the number of trophic levels and the species composition within a trophic level, can change. We review the development of these ideas, emphasizing in particular how compositional change can alter theoretical expectations about the regulation of trophic structure. A comparison of studies on the effects of predators and resource productivity in limnetic ecosystems reveals an intriguing disparity between the results of manipulative experiments and those of correlational studies. We suggest that this contrast is a result of the difference in the temporal scales operating in the two types of studies. Ecosystem-level variables may appear to approach an equilibrium in short-term press experiments; however, processes such as invasion and extinction of species will not have time to play out in most such experiments. We found that the responses of ecosystems to short-term experimental treatments involve less change in species composition than is found in natural communities that have diverged in response to local conditions over longer periods. We argue that the results of short-term experiments support the predictions of models in which 467 0066-4162/97/1120-0467$08.00 Annu. Rev. Ecol. Syst. 1997.28:467-494. Downloaded from www.annualreviews.org Access provided by University of Utah -Marriot Library on 12/03/14. For personal use only. 468LEIBOLD ET AL the species pool does not change, whereas correlational studies among systems support theories that incorporate compositional change.

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