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.

A recent increase in studies of b diversity has yielded a confusing array of concepts, measures and methods. Here, we provide a roadmap of the most widely used and ecologically relevant approaches for analysis through a series of mission statements. We distinguish two types of b diversity: directional turnover along a gradient vs. non-directional variation. Different measures emphasize different properties of ecological data. Such properties include the degree of emphasis on presence ⁄ absence vs. relative abundance information and the inclusion vs. exclusion of joint absences. Judicious use of multiple measures in concert can uncover the underlying nature of patterns in b diversity for a given dataset. A case study of Indonesian coral assemblages shows the utility of a multi-faceted approach. We advocate careful consideration of relevant questions, matched by appropriate analyses. The rigorous application of null models will also help to reveal potential processes driving observed patterns in b diversity.

Deterministic theories in community ecology suggest that local, niche-based processes, such as environmental filtering, biotic interactions and interspecific trade-offs largely determine patterns of species diversity and composition. In contrast, more stochastic theories emphasize the importance of chance colonization, random extinction and ecological drift. The schisms between deterministic and stochastic perspectives, which date back to the earliest days of ecology, continue to fuel contemporary debates (e.g. niches versus neutrality). As illustrated by the pioneering studies of Robert H. MacArthur and co-workers, resolution to these debates requires consideration of how the importance of local processes changes across scales. Here, we develop a framework for disentangling the relative importance of deterministic and stochastic processes in generating site-to-site variation in species composition (b-diversity) along ecological gradients (disturbance, productivity and biotic interactions) and among biogeographic regions that differ in the size of the regional species pool. We illustrate how to discern the importance of deterministic processes using nullmodel approaches that explicitly account for local and regional factors that inherently create stochastic turnover. By embracing processes across scales, we can build a more synthetic framework for understanding how niches structure patterns of biodiversity in the face of stochastic processes that emerge from local and biogeographic factors.

Community assembly provides a conceptual foundation for understanding the processes that determine which and how many species live in a particular locality. Evidence suggests that community assembly often leads to a single stable equilibrium, such that the conditions of the environment and interspecific interactions determine which species will exist there. In such cases, regions of local communities with similar environmental conditions should have similar community composition. Other evidence suggests that community assembly can lead to multiple stable equilibria. Thus, the resulting community depends on the assembly history, even when all species have access to the community. In these cases, a region of local communities with similar environmental conditions can be very dissimilar in their community composition. Both regional and local factors should determine the patterns by which communities assemble, and the resultant degree of similarity or dissimilarity among localities with similar environments. A single equilibrium in more likely to be realized in systems with small regional species pools, high rates of connectance, low productivity and high disturbance. Multiple stable equilibria are more likely in systems with large regional species pools, low rates of connectance, high productivity and low disturbance. I illustrate preliminary evidence for these predictions from an observational study of small pond communities, and show important effects on community similarity, as well as on local and regional species richness.

Historically, the biodiversity and composition of species in a locality was thought to be influenced primarily by deterministic factors. In such cases, species' niches create differential responses to environmental conditions and interspecific interactions, which combine to determine that locality's biodiversity and species composition. More recently, proponents of the neutral theory have placed a premium on how stochastic factors, such as birth, death, colonization, and extinction (termed ''ecological drift'') influence diversity and species composition in a locality independent of their niches. Here, I develop the hypothesis that the relative importance of stochastic ecological drift and/or priority effects depend on the harshness of the ecological filter in those habitats. I established long-term experimental ponds to explore the relative importance of community assembly history and drought on patterns of community compositional similarity among ponds that were otherwise similar in their environmental conditions. I show considerable site-to-site variation in pond community composition in the absence of drought that likely resulted from a combination of stochastic ecological drift and priority effects. However, in ponds that experienced drought, I found much higher similarity among communities that likely resulted from niche-selection filtering out species from the regional pool that could not tolerate such environmental harshness. These results implicate the critical role for understanding the processes of community assembly when examining patterns of biodiversity at different spatial scales.disturbance ͉ ecological drift ͉ niche ͉ similarity ͉ species composition