The effect of positive interactions on community structure in a multi-species metacommunity model along an environmental gradient

Positive interactions are widely recognized as playing a major role in the organization of community structure and diversity. As such, recent theoretical and empirical works have revealed the significant contribution of positive interactions in shaping species’ geographical distributions, particularly in harsh abiotic conditions. In this report, we explore the joint influence of local dispersal and an environmental gradient on the spatial distribution, structure and function of communities containing positive interactions. While most previous theoretical efforts were limited to modelling the dynamics of single pairs of associated species being mutualist or competitor, here we employ a spatially explicit multi-species metacommunity model covering a rich range of interspecific interactions (mutualism, competition and exploitation) along an environmental gradient. We find that mutualistic interactions dominate in communities with low diversity characterized by limited species dispersal and poor habitat quality. On the other hand, the fraction of mutualistic interactions decreases at the expense of exploitation and competition with the increase in diversity caused by higher dispersal and/or habitat quality. Our multi-species model exemplifies the ubiquitous presence of mutualistic interactions and the role of mutualistic species as facilitators for the further establishment of species during ecosystem assembly. We therefore argue that mutualism is an essential component driving the origination of complex and diverse communities.     

Landscape constraints on functional diversity of birds and insects in tropical agroecosystems

In this paper, we analyzeatabases on birds and insects to assess patterns of functional diversity in human-dominated landscapes in the tropics. A perspective from developed landscapes is essential for understanding remnant natural ecosystems, because most species experience their surroundings at spatial scales beyond the plot level, and spillover between natural and managed ecosystems is common. Agricultural bird species have greater habitat and diet breadth than forest species. Based on a global data base, bird assemblages in tropical agroforest ecosystems were composed of disproportionately more frugivorous and nectarivorous, but fewer insectivorous bird species compared with forest. Similarly, insect predators of plant-feeding arthropods were more diverse in Ecuadorian agroforest and forest compared with rice and pasture, while, in Indonesia, bee diversity was also higher in forested habitats. Hence, diversity of insectivorous birds and insect predators as well as bee pollinators declined with agricultural transformation. In contrast, with increasing agricultural intensification, avian pollinators and seed dispersers initially increase then decrease in proportion. It is well established that the proximity of agricultural habitats to forests has a strong influence on the functional diversity of agroecosystems. Community similarity is higher among agricultural systems than in natural habitats and higher in simple than in complex landscapes for both birds and insects, so natural communities, low-intensity agriculture, and heterogeneous landscapes appear to be critical in the preservation of beta diversity. We require a better understanding of the relative role of landscape composition and the spatial configuration of landscape elements in affecting spillover of functionally important species across managed and natural habitats. This is important for data-based management of tropical human-dominated landscapes sustaining the capacity of communities to reorganize after disturbance and to ensure ecological functioning.     

Advances in pollination ecology from tropical plantation crops

Although ecologists traditionally focus on natural ecosystems, there is growing awareness that mixed landscapes of managed and unmanaged systems provide a research environment for understanding basic ecological relationships on a large scale. Here, we show how tropical agroforestry systems can be used to develop ideas about the mechanisms that influence species diversity and subsequent biotic interactions at different spatial scales. Our focus is on tropical plantation crops, mainly coffee and cacao, and their pollinators, which are of basic ecological interest as partners in an important mutualistic interaction. We review how insect-mediated pollination services depend on local agroforest and natural habitats in surrounding landscapes. Further, we evaluate the functional significance of pollinator diversity and the explanatory value of species traits, and we provide an intercontinental comparison of pollinator assemblages. We found that optimal pollination success might be best understood as a consequence of niche complementarities among pollinators in landscapes harboring various species. We further show that small cavity-nesting bees and small generalist beetles were especially affected by isolation from forest and that larger-bodied insects in the same landscapes were not similarly affected. We suggest that mixed tropical landscapes with agroforestry systems have great potential for future research on the interactions between plants and pollinators.     

Pyric Herbivory: Rewilding Landscapes through the Recoupling of Fire and Grazing

Abstract:  Our understanding of fire and grazing is largely based on small-scale experimental studies in which treatments are uniformly applied to experimental units that are considered homogenous. Any discussion of an interaction between fire and grazing is usually based on a statistical approach that ignores the spatial and temporal interactions on complex landscapes. We propose a new focus on the ecological interaction of fire and grazing in which each disturbance is spatially and temporally dependent on the other and results in a landscape where disturbance is best described as a shifting mosaic (a landscape with patches that vary with time since disturbance) that is critical to ecological structure and function of many ecosystems. We call this spatiotemporal interaction pyric herbivory (literal interpretation means grazing driven by fire). Pyric herbivory is the spatial and temporal interaction of fire and grazing, where positive and negative feedbacks promote a shifting pattern of disturbance across the landscape. We present data we collected from the Tallgrass Prairie Preserve in the southern Great Plains of North America that demonstrates that the interaction between free-roaming bison ( Bison bison ) and random fires promotes heterogeneity and provides the foundation for biological diversity and ecosystem function of North American and African grasslands. This study is different from other studies of fire and grazing because the fires we examined were random and grazing animals were free to roam and select from burned and unburned patches. For ecosystems across the globe with a long history of fire and grazing, pyric herbivory with any grazing herbivore is likely more effective at restoring evolutionary disturbance patterns than a focus on restoring any large vertebrate while ignoring the interaction with fire and other disturbances .     

Metapopulation Extinction Risk under Spatially Autocorrelated Disturbance

Abstract:  Recent extinction models generally show that spatial aggregation of habitat reduces overall extinction risk because sites emptied by local extinction are more rapidly recolonized. We extended such an investigation to include spatial structure in the disturbance regime. A spatially explicit metapopulation model was developed with a wide range of dispersal distances. The degree of aggregation of both habitat and disturbance pattern could be varied from a random distribution, through the intermediate case of a fractal distribution, all the way to complete aggregation (single block). Increasing spatial aggregation of disturbance generally increased extinction risk. The relative risk faced by populations in different landscapes varied greatly, depending on the disturbance regime. With random disturbance, the spatial aggregation of habitat reduced extinction risk, as in earlier studies. Where disturbance was spatially autocorrelated, however, this advantage was eliminated or reversed because populations in aggregated habitats are at risk of mass extinction from coarse-scale disturbance events. The effects of spatial patterns on extinction risk tended to be reduced by long-distance dispersal. Given the high levels of spatial correlation in natural and anthropogenic disturbance processes, population vulnerability may be greatly underestimated both by classical (nonspatial) models and by those that consider spatial structure in habitat alone.     

Consequences of dominance: a review of evenness effects on local and regional ecosystem processes

The composition of communities is strongly altered by anthropogenic manipulations of biogeochemical cycles, abiotic conditions, and trophic structure in all major ecosystems. Whereas the effects of species loss on ecosystem processes have received broad attention, the consequences of altered species dominance for emergent properties of communities and ecosystems are poorly investigated. Here we propose a framework guiding our understanding of how dominance affects species interactions within communities, processes within ecosystems, and dynamics on regional scales. Dominance (or the complementary term, evenness) reflects the distribution of traits in a community, which in turn affects the strength and sign of both intraspecifc and interspecific interactions. Consequently, dominance also mediates the effect of such interactions on species coexistence. We review the evidence for the fact that dominance directly affects ecosystem functions such as process rates via species identity (the dominant trait) and evenness (the frequency distribution of traits), and indirectly alters the relationship between process rates and species richness. Dominance also influences the temporal and spatial variability of aggregate community properties and compositional stability (invasibility). Finally, we propose that dominance affects regional species coexistence by altering metacommunity dynamics. Local dominance leads to high beta diversity, and rare species can persist because of source-sink dynamics, but anthropogenically induced environmental changes result in regional dominance and low beta diversity, reducing regional coexistence. Given the rapid anthropogenic alterations of dominance in many ecosystems and the strong implications of these changes, dominance should be considered explicitly in the analysis of consequences of altered biodiversity.     

Spatial clustering of habitat structure effects patterns of community composition and diversity

Natural ecosystems often show highly productive habitats that are clustered in space. Environmental disturbances are also often nonrandomly distributed in space and are either intrinsically linked to habitat quality or independent in occurrence. Theoretical studies predict that configuration and aggregation of habitat patch quality and disturbances can affect metacommunity composition and diversity, but experimental evidence is largely lacking. In a metacommunity experiment, we tested the effects of spatially autocorrelated disturbance and spatial aggregation of patch quality on regional and local richness, among-community dissimilarity, and community composition. We found that spatial aggregation of patch quality generally increased among-community dissimilarity (based on two measures of beta diversity) of communities containing protozoa and rotifers in microcosms. There were significant interacting effects of landscape structure and location of disturbances on beta diversity, which depended in part on the specific beta diversity measures used. Effects of disturbance on composition and richness in aggregated landscapes were generally dependent on distance and connectivity among habitat patches of different types. Our results also show that effects of disturbances in single patches cannot directly be extrapolated to the landscape scale: the predictions may be correct when only species richness is considered, but important changes in beta diversity may be overlooked. There is a need for biodiversity and conservation studies to consider the spatial aggregation of habitat quality and disturbance, as well as connectivity among spatial aggregations.     

Fire and forest landscapes in the Georgia Piedmont: an assessment of spatial modeling assumptions

Landscape simulation models are widely used to study the behavior of ecological systems. As computing power has increased, these models have become more complex and incorporated more realistic spatial representations of landscape patterns and ecological processes. The goal of this research was to examine the sensitivity of simulated landscape patterns to fundamental spatial modeling assumptions. The LANDIS simulator was parameterized for forests of the Georgia Piedmont and used to model landscape-scale community dynamics at fire return intervals from 20 to 100 years. A base scenario incorporating localized seed dispersal along with landform-related variation in species establishment rates and disturbance regimes was contrasted with three alternative scenarios. The uniform habitat scenario applied the same set of species establishment coefficients across all landforms. The uniform dispersal scenario removed the effects of seed source abundance and pattern on species establishment. The uniform disturbance scenario assumed identical disturbance regimes on all landforms.At the shortest fire return intervals, fire severities were low and the stand age distribution was dominated by older forests. At longer fire return intervals, fire severities were high and the stand age distribution was skewed toward younger forests. Species composition generally followed a gradient from fire-resistant species at short fire return intervals to fire-sensitive species at longer fire return intervals. However, some species exhibited bimodal distributions with high abundances at both short and long fire return intervals. Landscape responses to fire were similar in the uniform habitat scenario and the base scenario. Communities were less sensitive to fire return interval and had more fire-sensitive species in the uniform dispersal scenario than in the base scenario. Species composition in the uniform disturbance scenario was similar to the base scenario for the longest fire-intervals, but was more sensitive to changes in the fire regime at shorter fire return intervals. In models of Piedmont forest landscapes, accurate spatial representations of dispersal and fire regime heterogeneity are essential for predicting landscape-scale species composition under changing fire regimes. In contrast, the precise spatial representation of species–habitat relationships may be considerably less important.     

Measuring impacts on species with models and metrics of varying ecological and computational complexity

Approaches to assess the impacts of landscape disturbance scenarios on species range from metrics based on patterns of occurrence or habitat to comprehensive models that explicitly include ecological processes. The choice of metrics and models affects how impacts are interpreted and conservation decisions. We explored the impacts of 3 realistic disturbance scenarios on 4 species with different ecological and taxonomic traits. We used progressively more complex models and metrics to evaluate relative impact and rank of scenarios on the species. Models ranged from species distribution models that relied on implicit assumptions about environmental factors and species presence to highly parameterized spatially explicit population models that explicitly included ecological processes and stochasticity. Metrics performed consistently in ranking different scenarios in order of severity primarily when variation in impact was driven by habitat amount. However, they differed in rank for cases where dispersal dynamics were critical in influencing metapopulation persistence. Impacts of scenarios on species with low dispersal ability were better characterized using models that explicitly captured these processes. Metapopulation capacity provided rank orders that most consistently correlated with those from highly parameterized and data-rich models and incorporated information about dispersal with little additional computational and data cost. Our results highlight the importance of explicitly considering species’ ecology, spatial configuration of habitat, and disturbance when choosing indicators of species persistence. We suggest using hybrid approaches that are a mixture of simple and complex models to improve multispecies assessments.     

Interactive effects of disturbance and dispersal directionality on species richness and composition in metacommunities

Dispersal among ecological communities is usually assumed to be random in direction, or to vary in distance or frequency among species. However, a variety of natural systems and types of organisms may experience dispersal that is biased by directional currents or by gravity on hillslopes. We developed a general model for competing species in metacommunities to evaluate the role of directionally biased dispersal on species diversity, abundance, and traits. In parallel, we tested the role of directionally biased dispersal on communities in a microcosm experiment with protists and rotifers. Both the model and experiment independently demonstrated that diversity in local communities was reduced by directionally biased dispersal, especially dispersal that was biased away from disturbed patches. Abundance of species (and composition) in local communities was a product of disturbance intensity but not dispersal directionality. High disturbance selected for species with high intrinsic growth rates and low competitive abilities. Overall, our conclusions about the key role of dispersal directionality in (meta)communities seem robust and general, since they were supported both by the model, which was set in a general framework and not parameterized to fit to a specific system, and by a specific experimental test with microcosms.     

Mass effects mediate coexistence in competing shrews

Recent developments in metacommunity theory have raised awareness that processes occurring at regional scales might interfere with local dynamics and affect conditions for the local coexistence of competing species. Four main paradigms are recognized in this context (namely, neutral, patch-dynamics, species-sorting, and mass-effect), which differ according to the role assigned to ecological or life-history differences among competing species, as well as to the relative time scale of regional vs. local dynamics. We investigated the patterns of regional and local coexistence of two species of shrews (Crocidura russula and Sorex coronatus) sharing a similar diet (generalist insectivores) over four generations, in a spatially structured habitat at the altitudinal limit of their distributions. Local populations were small, and regional dynamics were strong, with high rates of extinction and recolonization. Niche analysis revealed significant habitat differentiation on a few important variables, including temperature and availability of winter resting sites. In sites suitable for both species, we found instances of local coexistence with no evidence of competitive exclusion. Patterns of temporal succession did not differ from random, with no suggestion of a colonization-competition trade-off. Altogether, our data provide support for the mass-effect paradigm, where regional coexistence is mediated by specialization on different habitat types, and local coexistence by rescue effects from source sites. The strong regional dynamics and demographic stochasticity, together with high dispersal rates, presumably contributed to mass effects by overriding local differences in specific competitive abilities.     

Landscape-level impact of tropical forest loss and fragmentation on bird occurrence in eastern Guatemala

Tropical forest destruction and fragmentation of habitat patches may reduce population persistence at the landscape level. Given the complex nature of simultaneously evaluating the effects of these factors on biotic populations, statistical presence/absence modelling has become an important tool in conservation biology. This study uses logistic regression to evaluate the independent effects of tropical forest cover and fragmentation on bird occurrence in eastern Guatemala. Logistic regression models were constructed for 10 species with varying response to habitat alteration. Predictive variables quantified forest cover, fragmentation and their interaction at three different radii (200, 500 and 1000 m scales) of 112 points where presence of target species was determined. Most species elicited a response to the 1000 m scale, which was greater than most species’ reported territory size. Thus, their presence at the landscape scale is probably regulated by extra-territorial phenomena, such as dispersal. Although proportion of forest cover was the most important predictor of species’ presence, there was strong evidence of area-independent and -dependent fragmentation effects on species presence, results that contrast with other studies from northernmost latitudes. Species’ habitat breadth was positively correlated with AIC model values, indicating a better fit for species more restricted to tropical forest. Species with a narrower habitat breadth also elicited stronger negative responses to forest loss. Habitat breadth is thus a simple measure that can be directly related to species’ vulnerability to landscape modification. Model predictive accuracy was acceptable for 4 of 10 species, which were in turn those with narrower habitat breadths.     

Spatially stochastic settlement and the coexistence of benthic marine animals

For sessile organisms, dispersal and recruitment are typically spatially stochastic, but there is little understanding of how this variability scales up to influence processes such as competitive coexistence. Here we argue that coexistence of benthic marine animals is enhanced by stochastic differences between species in the spatial distribution of larval settlement. Differentiation of settlement distributions among competitors results in intraspecifically aggregated settlement, which can reduce overall interspecific competition and increase overall intraspecific competition. We test for the components of this mechanism using a pair of subtidal invertebrates, and we find that the mean interspecific effect of the dominant competitor is substantially reduced by natural settlement variability. Using a simulation parameterized with experimental data, we find that variable settlement could play an important role in long-term coexistence between these species. This mechanism may apply broadly to benthic marine communities, which can be highly diverse and typically exhibit large settlement fluctuation over a range of scales.     

Spatial and temporal patterns of seed dispersal: an important determinant of grassland invasion.

We measured spatial and temporal patterns of seed dispersal and seedling recruitment for 58 species in a grassland community to test whether seed dispersal could predict patterns of invasion after disturbance. For the 12 most abundant grasses, recruitment of native species was dependent on the propagule supply of both native and exotic species. Variability in seed rain on small spatial (1-10 m) and temporal (within season) scales led to qualitative differences in the outcome of disturbance colonization such that native species dominated disturbances when exotic seed supply was low but failed to establish when exotic seed supply was high. Local dispersal and spatial heterogeneity in species composition promoted coexistence of native and exotic species by creating refuges from high exotic seed supply within native dominated patches. Despite this, copious exotic seed production strongly limited recruitment of native species in exotic dominated patches. Most grasslands in California are presently dominated by exotic species, suggesting that competition at the seedling stage is a major barrier to native species restoration.     

Linking life history to landscape for threatened species conservation in a multiuse region

Landscape-scale conservation that considers metapopulation dynamics will be essential for preventing declines of species facing multiple threats to their survival. Toward this end, we developed a novel approach that combines occurrence records, spatial–environmental data, and genetic information to model habitat, connectivity, and patterns of genetic structure and link spatial attributes to underlying ecological mechanisms. Using the threatened northern quoll (Dasyurus hallucatus) as a case study, we applied this approach to address the need for conservation decision-making tools that promote resilient metapopulations of this threatened species in the Pilbara, Western Australia, a multiuse landscape that is a hotspot for biodiversity and mining. Habitat and connectivity were predicted by different landscape characteristics. Whereas habitat suitability was overwhelmingly driven by terrain ruggedness, dispersal was facilitated by proximity to watercourses. Although there is limited evidence for major physical barriers in the Pilbara, areas with high silt and clay content (i.e., alluvial and hardpan plains) showed high resistance to dispersal. Climate subtlety shaped distributions and patterns of genetic turnover, suggesting the potential for local adaptation. By understanding these spatial–environmental associations and linking them to life-history and metapopulation dynamics, we highlight opportunities to provide targeted species management. To support this, we have created habitat, connectivity, and genetic uniqueness maps for conservation decision-making in the region. These tools have the potential to provide a more holistic approach to conservation in multiuse landscapes globally.     

What enables coexistence in plant communities? Weak versus strong species traits and the role of local processes

Explaining the coexistence of species that basically depend on the same resources has been a brainteaser for generations of ecologists. Different mechanisms have been proposed to facilitate coexistence in plant communities, where space is an important resource. Using a stochastic cellular automaton simulation model we analyze - separately and in combination - the influence of different species traits and processes which alter local competition on the coexistence of plant species over a fixed time horizon. We show that different species traits operate on different time scales in competition. We therefore suggest the concept of weak versus strong traits according to short- or long-term exclusion of species differing in these traits. As a consequence, highly non-linear trade-offs between weak and strong traits can result in communities. Furthermore, we found that trade-offs based on physiological species traits such as plant lifetime, dispersal range and plant growth, did not support broad and long-term coexistence—further processes such as density-dependent mortality and light-dependent colonization were necessary. This suggests that coexistence in plant communities requires (stabilizing) local processes to support the (equalizing) trade-offs in species traits.     

Natural and anthropogenic influences on the diversity structure of reef fish communities in the Tuamotu Archipelago (French Polynesia)

Despite the rapid rate of human-induced species losses, the relative influence of natural and anthropogenic factors on the functional diversity of species assemblages remains unknown for most ecosystems. A model was previously developed to predict the diversity structure of coral reef fish assemblages in 10 atolls of low human pressure and contrasting morphology of the Tuamotu Archipelago (French Polynesia). This existing model predicted smoothed histograms (spectra) of species richness according to size classes, diet classes and life-history classes of fish assemblages using a combination of environmental characteristics at different spatial scales. The present study applied the model to Tikehau, another atoll of the same archipelago where commercial fishing is practiced and where the same sampling strategy was reproduced. Significant differences appeared between predicted and observed species richness in several size, diet and life-history classes of fish assemblages in Tikehau. Two parameters which were not accounted for in the initial model, i.e. fishing pressure and atoll position within the archipelago, explained together 63% of variance in model residuals, >60% being explained by fishing pressure only. The respective effects of fishing and atoll position on the diversity of coral reef fish assemblages are discussed, with the potential of such modelling approach to assess the relative importance of factors affecting functional diversity within communities.     

The effects of habitat loss, fragmentation, and community homogenization on resilience in estuaries.

When changes in the frequency and extent of disturbance outstrip the recovery potential of resident communities, the selective removal of species contributes to habitat loss and fragmentation across landscapes. The degree to which habitat change is likely to influence community resilience will depend on metacommunity structure and connectivity. Thus ecological connectivity is central to understanding the potential for cumulative effects to impact upon diversity. The importance of these issues to coastal marine communities, where the prevailing concept of open communities composed of highly dispersive species is being challenged, indicates that these systems may be more sensitive to cumulative impacts than previously thought. We conducted a disturbance-recovery experiment across gradients of community type and environmental conditions to assess the roles of ecological connectivity and regional variations in community structure on the recovery of species richness, total abundance, and community composition in Mahurangi Harbour, New Zealand. After 394 days, significant differences in recovery between sites were apparent. Statistical models explaining a high proportion of the variability (R2 > 0.92) suggested that community recovery rates were controlled by a combination of physical and ecological features operating across spatial scales, affecting successional processes. The dynamic and complex interplay of ecological and environmental processes we observed driving patch recovery across the estuarine landscape are integral to recovery from disturbances in heterogeneous environments. This link between succession/recovery, disturbance, and heterogeneity confirms the utility of disturbance-recovery experiments as assays for cumulative change due to fragmentation and habitat change in estuaries.     

Integrating functional connectivity and fire management for better conservation outcomes

Globally, the mean abundance of terrestrial animals has fallen by 50% since 1970, and populations face ongoing threats associated with habitat loss, fragmentation, climate change, and disturbance. Climate change can influence the quality of remaining habitat directly and indirectly by precipitating increases in the extent, frequency, and severity of natural disturbances, such as fire. Species face the combined threats of habitat clearance, changing climates, and altered disturbance regimes, each of which may interact and have cascading impacts on animal populations. Typically, conservation agencies are limited in their capacity to mitigate rates of habitat clearance, habitat fragmentation, or climate change, yet fire management is increasingly used worldwide to reduce wildfire risk and achieve conservation outcomes. A popular approach to ecological fire management involves the creation of fire mosaics to promote animal diversity. However, this strategy has 2 fundamental limitations: the effect of fire on animal movement within or among habitat patches is not considered and the implications of the current fire regime for long-term population persistence are overlooked. Spatial and temporal patterns in fire history can influence animal movement, which is essential to the survival of individual animals, maintenance of genetic diversity, and persistence of populations, species, and ecosystems. We argue that there is rich potential for fire managers to manipulate animal movement patterns; enhance functional connectivity, gene flow, and genetic diversity; and increase the capacity of populations to persist under shifting environmental conditions. Recent methodological advances, such as spatiotemporal connectivity modeling, spatially explicit individual-based simulation, and fire-regime modeling can be integrated to achieve better outcomes for biodiversity in human-modified, fire-prone landscapes. Article impact statement: Land managers may conserve populations by using fire to sustain or enhance functional connectivity.     

Reconciling empirical ecology with neutral community models

Neutral community models embody the idea that individuals are ecologically equivalent, having equal fitness over all environmental conditions, and describe how the spatial dynamics and speciation of such individuals can produce a wide range of patterns of distribution, diversity, and abundance. Neutral models have been controversial, provoking a rush of tests and comments. The debate has been spurred by the suggestion that we should test mechanisms. However, the mechanisms and the spatial scales of interest have never clearly been described, and consequently, the tests have often been only peripherally relevant. At least two mechanisms are present in spatially structured neutral models. Dispersal limitation causes clumping of a species, which increases the strength of intraspecific competition and reduces the strength of interspecific competition. This may prolong coexistence and enhance local and regional diversity. Speciation is present in some neutral models and gives a donor-controlled input of new species, many of which remain rare or are short lived, but which directly add to species diversity. Spatial scale is an important consideration in neutral models. Ecological equivalence and equal fitness have implicit spatial scales because dispersal limitation and its emergent effects operate at population levels, and populations and communities are defined at a chosen spatial scale in recent neutral models; equality is measured relative to a metacommunity, and this necessitates defining the spatial scale of that metacommunity. Furthermore, dispersal has its own scales. Thorough empirical tests of neutral models will require both tests of mechanisms and pattern-producing ability, and will involve coupling theoretical models and experiments.