Th e metacommunity framework is a powerful platform for evaluating patterns of species distribution in geographic or environmental space. Idealized patterns (checkerboard, Clementsian, evenly spaced, Gleasonian and nested distributions) give the framework shape. Each pattern represents an area in a multidimensional continuum of metacommunity structures; however, the current approach to analysis of spatial structure of metacommunities is incomplete. To address this, we describe additional non-random structures and illustrate how they may be discerned via objective criteria. First, we distinguish three distinct forms of species loss in nested structures, which should improve identifi cation of structuring mechanisms for nested patterns. Second, we defi ne six quasi-structures that are consistent with the conceptual underpinnings of Clementsian, Gleasonian, evenly spaced and nested distributions. Finally, we demonstrate how combinations of structures at smaller spatial extents may aggregate to form Clementsian structure at larger extents. Th ese refi nements should facilitate the identifi cation of best-fi t patterns, associated structuring mechanisms, and informative scales of analysis and interpretation. Th is conceptual and analytical framework may be applied to network properties within communities (i.e. structure of interspecifi c interactions) and has broad application in ecology and biogeography.

Techniques to evaluate elements of metacommunity structure (EMS; coherence, species turnover and range boundary clumping) have been available for several years. Such approaches are capable of determining which idealized pattern of species distribution best describes distributions in a metacommunity. Nonetheless, this approach rarely is employed and such aspects of metacommunity structure remain poorly understood. We expanded an extant method to better investigate metacommunity structure for systems that respond to multiple environmental gradients. We used data obtained from 26 sites throughout Paraguay as a model system to demonstrate application of this methodology. Using presence-absence data for bats, we evaluated coherence, species turnover and boundary clumping to distinguish among six idealized patterns of species distribution. Analyses were conducted for all bats as well as for each of three feeding ensembles (aerial insectivores, frugivores and molossid insectivores). For each group of bats, analyses were conducted separately for primary and secondary axes of ordination as defined by reciprocal averaging. The Paraguayan bat metacommunity evinced Clementsian distributions for primary and secondary ordination axes. Patterns of species distribution for aerial insectivores were dependent on ordination axis, showing Gleasonian distributions when ordinated according to the primary axis and Clementsian distributions when ordinated according to the secondary axis. Distribution patterns for frugivores and molossid insectivores were best described as random. Analysis of metacommunities using multiple ordination axes can provide a more complete picture of environmental variables that mold patterns of species distribution. Moreover, analysis of EMS along defined gradients (e.g., latitude, elevation and depth) or based on alternative ordination techniques may complement insights based on reciprocal averaging because the fundamental questions addressed in analyses are contingent on the ordination technique that is employed.

Distributions of species often exhibit nested structure, such that assemblages at species-poor sites are proper subsets of taxa at more species-rich sites. Traditionally, this has been viewed as a large-scale biogeographic pattern and treated implicitly as static from a temporal perspective. Nonetheless, recent work suggests that nestedness may arise at multiple spatio-temporal scales. A 13-year data set encompassing the effects of two largescale natural disturbances (hurricanes Hugo and Georges) on terrestrial gastropod assemblages was used to test the hypothesis that changes in species composition resulting from disturbance alter the degree of nestedness exhibited over time at two spatial scales. Gastropod assemblages were least nested immediately following disturbance, and nestedness increased thereafter. Although land-use history influenced the degree of nestedness, trajectories of nestedness following disturbance were similar irrespective of disturbance history or hurricane identity. The effects of hurricanes with respect to nestedness of terrestrial gastropods may be general and predictable, even though species respond to hurricanes in disparate fashions. By damaging some localities within the forest more severely than others, a hurricane dismantles extant patterns of species composition and severs connections among sites, as inhospitable microclimatic conditions limit dispersal of gastropods. As time passes and the forest canopy regenerates, conditions ameliorate, and movement among sites becomes more frequent. Thus, a conclusion based on a single time period may not characterize the study system in general. Consequently, explanations for nested structure that incorporate variability in ecological as well as evolutionary time will improve the applicability and comparability of nested subsets analysis across study systems.

Nestedness has received considerable attention in community ecology and conservation biology from both theoretical and empirical perspectives. This has lead to the creation of various metrics and null models to analyze nested subsets, all of which rely on the random placement of species to assess significance. However, if immigration and extinction are the processes that underlie species distributions on island systems, then null models might be better determined on the basis of randomly placed individuals. Consequently, we examined the effects of species–abundance distributions (uniform, dominance–decay, random–assortment, and dominance–preemption), island–size distributions (uniform and linear decrease), and total abundances (128, 256, 512, 1024, 2048, 4096 and 8192) on the degree of nestedness and its significance. Generally, matrices of species presence and absence created from the random placement of individuals were nested significantly according to null models based on the random placement of species. Island size and abundance had less of an effect on nestedness in systems dominated by only a few species than in systems in which abundances were distributed more evenly. Stochastic processes, such as the random placement of individuals, predispose systems to evince patterns of nestedness at the species level, which may account, in part, for the ubiquity of nestedness in nature.

Aim We tested the hypothesis that distributions of Mexican bats are defined by shared responses to environmental gradients for the entire Mexican bat metacommunity and for each of four metaensembles (frugivores, nectarivores, gleaning insectivores, and aerial insectivores). Further, we identified the main environmental factors to which bats respond for multiple spatial extents. Location Mexico.Methods Using bat presence-absence data, as well as vegetation composition for each of 31 sites, we analysed metacommunity structure via a comprehensive, hierarchical approach that uses reciprocal averaging (RA) to detect latent environmental gradients corresponding to each metacommunity structure (e.g. Clementsian, Gleasonian, nested, random). Canonical correspondence analysis (CCA) was used to relate such gradients to variation in vegetation composition.Results For all bat species and for each ensemble, the primary gradient of ordination from RA, which is based on species data only, recovered an axis of humidity that matched that obtained for the first axis of the CCA ordination, which is based both on vegetation attributes and on species composition of sites. For the complete assemblage as well as for aerial and gleaning insectivores, analyses revealed Clementsian or quasi-Clementsian structures with discrete compartments (distinctive groups of species along portions of an environmental gradient) coincident with the humidity gradient and with the NearcticNeotropical divide. Within-compartment analysis further revealed Clementsian or quasi-Clementsian structures corresponding to a gradient of elevational complexity that matched the second ordination axis in CCA. Frugivores had quasi-nested structure, whereas nectarivores had Gleasonian structure.Main conclusions Our hierarchical approach to metacommunity analysis detected complex metacommunity structures associated with multiple environmental gradients at different spatial extents. More importantly, the resulting structures and their extent along environmental gradients are determined by ensemble-specific characteristics and not by arbitrarily circumscribed study areas. This property renders compartment-level analyses particularly useful for large-scale ecological analyses in areas where more than one gradient may exist and species sorting may occur at multiple scales.