Maximum species richness at intermediate frequencies of disturbance: consistency among levels of productivity

Development of a mechanistic understanding and predictions of patterns of biodiversity is a central theme in ecology. One of the most influential theories, the intermediate disturbance hypothesis (IDH), predicts maximum diversity at intermediate levels of disturbance frequency. The dynamic equilibrium model (DEM), an extension of the IDH, predicts that the level of productivity determines at what frequency of disturbance maximum diversity occurs. To test, and contrast, the predictions of these two models, a field experiment on marine hard-substratum assemblages was conducted with seven levels of disturbance frequency and three levels of nutrient availability. Consistent with the IDH, maximum diversity, measured as species richness, was observed at an intermediate frequency of disturbance. Despite documented effects on productivity, the relationship between disturbance and diversity was not altered by the nutrient treatments. Thus, in this system the DEM did not improve the understanding of patterns of diversity compared to the IDH. Furthermore, it is suggested that careful consideration of measurements and practical definitions of productivity in natural assemblages is necessary for a rigorous test of the DEM.     

Trade-offs, temporal variation, and species coexistence in communities with intraguild predation

Intraguild predation/parasitism (IGP: competing species preying on or parasitizing each other) is widespread in nature, but the mechanisms by which intraguild prey and predators coexist remain elusive. Theory predicts that a trade-off between resource competition and IGP should allow local niche partitioning, but such trade-offs are expressed only at intermediate resource productivity and cannot explain observations of stable coexistence at high productivity. Coexistence must therefore involve additional mechanisms beside the trade-off, but very little is known about the operation of such mechanisms in nature. Here I present the first experimental test of multiple coexistence mechanisms in a natural community exhibiting IGP. The results suggest that, when resource productivity constrains the competition-IGP trade-off, a temporal refuge for the intraguild prey can not only promote coexistence, but also change species abundances to a pattern qualitatively different from that expected based on the trade-off or a refuge alone. This is the first empirical study to demonstrate a mechanism for why communities with IGP do not lose species diversity in highly productive environments. These results have implications for diversity maintenance in multi-trophic communities, and the use of multiple natural enemies in biological control.     

Partitioning the factors of spatial variation in regeneration density of shade-tolerant tree species

Understanding coexistence of highly shade-tolerant tree species is a longstanding challenge for forest ecologists. A conceptual model for the coexistence of sugar maple (Acer saccharum) and American beech (Fagus grandibfolia) has been proposed, based on a low-light survival/high-light growth trade-off, which interacts with soil fertility and small-scale spatiotemporal variation in the environment. In this study, we first tested whether the spatial distribution of seedlings and saplings can be predicted by the spatiotemporal variability of light availability and soil fertility, and second, the manner in which the process of environmental filtering changes with regeneration size. We evaluate the support for this hypothesis relative to the one for a neutral model, i.e., for seed rain density predicted from the distribution of adult trees. To do so, we performed intensive sampling over 86 quadrats (5 x 5 m) in a 0.24-ha plot in a mature maple-beech community in Quebec, Canada. Maple and beech abundance, soil characteristics, light availability, and growth history (used as a proxy for spatiotemporal variation in light availability) were finely measured to model variation in sapling composition across different size classes. Results indicate that the variables selected to model species distribution do effectively change with size, but not as predicted by the conceptual model. Our results show that variability in the environment is not sufficient to differentiate these species' distributions in space. Although species differ in their spatial distribution in the small size classes, they tend to correlate at the larger size class in which recruitment occurs. Overall, the results are not supportive of a model of coexistence based on small-scale variations in the environment. We propose that, at the scale of a local stand, the lack of fit of the model could result from the high similarity of species in the range of environmental conditions encountered, and we suggest that coexistence would be stable only at larger spatial scales at which variability in the environment is greater.     

Scale-dependent response diversity of seabirds to prey in the North Sea

Functional response diversity is defined as the diversity of responses to environmental change among species that contribute to the same ecosystem function. Because different ecological processes dominate on different spatial and temporal scales, response diversity is likely to be scale dependent. Using three extensive data sets on seabirds, pelagic fish, and zooplankton, we investigate the strength and diversity in the response of seabirds to prey in the North Sea over three scales of ecological organization. Two-stage analyses were used to partition the variance in the abundance of predators and prey among the different scales of investigation: variation from year to year, variation among habitats, and variation on the local patch scale. On the year-to-year scale, we found a strong and synchronous response of seabirds to the abundance of prey, resulting in low response diversity. Conversely, as different seabird species were found in habitats dominated by different prey species, we found a high diversity in the response of seabirds to prey on the habitat scale. Finally, on the local patch scale, seabirds were organized in multispecies patches. These patches were weakly associated with patches of prey, resulting in a weak response strength and a low response diversity. We suggest that ecological similarities among seabird species resulted in low response diversity on the year-to-year scale. On the habitat scale, we suggest that high response diversity was due to interspecific competition and niche segregation among seabird species. On the local patch scale, we suggest that facilitation with respect to the detection and accessibility of prey patches resulted in overlapping distribution of seabirds but weak associations with prey. The observed scale dependencies in response strength and diversity have implications for how the seabird community will respond to different environmental disturbances.     

Application of an ecological framework linking scales based on self-thinning.

Barnes and Roderick developed a generic, theoretical framework for vegetation modeling across scales. Inclusion of a self-thinning mechanism connects the individual to the larger-scale population and, being based on the conservation of mass, all mass flux processes are integral to the formulation. Significantly, disturbance (both regular and stochastic) and its impact at larger scales are included in the formulation. The purpose of this paper is to illustrate how this model can be used to predict patch and ecosystem dry mass, and consequently system carbon. Examples from pine plantations and mixed forests are considered, with these applications requiring estimates of system carrying capacity and the growth rates of individual plants. The results indicate that the model is relatively simple and straightforward to apply, and its predictions compare well with the data. A significant feature of this approach is that the impact of local scale data on the dynamics of larger patch and ecosystem scales can be determined explicitly, as we show by example. Further, the general formulation has an analytic solution based on characteristics of the individual, facilitating practical and predictive application.     

Scale-dependent responses of plant biodiversity to nitrogen enrichment

Experimental studies demonstrating that nitrogen (N) enrichment reduces plant diversity within individual plots have led to the conclusion that anthropogenic N enrichment is a threat to global biodiversity. These conclusions overlook the influence of spatial scale, however, as N enrichment may alter beta diversity (i.e., how similar plots are in their species composition), which would likely alter the degree to which N-induced changes in diversity within localities translate to changes in diversity at larger scales that are relevant to policy and management. Currently, it is unclear how N enrichment affects biodiversity at scales larger than a small plot. We synthesized data from 18 N-enrichment experiments across North America to examine the effects of N enrichment on plant species diversity at three spatial scales: small (within plots), intermediate (among plots), and large (within and among plots). We found that N enrichment reduced plant diversity within plots by an average of 25% (ranging from a reduction of 61% to an increase of 5%) and frequently enhanced beta diversity. The extent to which N enrichment altered beta diversity, however, varied substantially among sites (from a 22% increase to an 18% reduction) and was contingent on site productivity. Specifically, N enrichment enhanced beta diversity at low-productivity sites but reduced beta diversity at high-productivity sites. N-induced changes in beta diversity generally reduced the extent of species loss at larger scales to an average of 22% (ranging from a reduction of 54% to an increase of 18%). Our results demonstrate that N enrichment often reduces biodiversity at both local and regional scales, but that a focus on the effects of N enrichment on biodiversity at small spatial scales may often overestimate (and sometimes underestimate) declines in regional biodiversity by failing to recognize the effects of N on beta diversity.     

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.     

Alternative strategies in avian scavengers: how subordinate species foil the despotic distribution

Trade-offs in species’ traits can mediate competition and enable coexistence. A key challenge in ecology is understanding the role of species’ trade-offs in maintaining diversity, and evolutionary trade-offs between the abilities of competing species are best understood by considering how competitive advantages change along an environmental gradient. Previous studies of such trade-offs are generally limited to two-species systems and a single trade-off. In this study, I consider the effect of trade-offs in search efficiency and competitive abilities on habitat use patterns among a diverse avian scavenger guild. I hypothesize that species’ dominance status and search efficiency will both be correlated with patch quality. Using counts of searching birds in areas that vary in habitat quality in terms of both wildlife and human settlement density and observations at experimental carcasses, I assess the competitive ability, search efficiency, and habitat use of seven avian scavenger species in Masai Mara National Reserve, Kenya. Findings support the hypothesis with Bateleurs, a species with high search efficiency, and Ruppell’s, Lappet-faced, and White-backed vultures, species with high individual or social dominance, preferentially exploiting habitats of high quality, while Tawny eagles and Hooded vultures, species with low search efficiency and competitive ability, prefer habitats of low quality. This paper demonstrates the importance of considering multiple strategies for assessing the effect of competition on habitat use within complex communities.     

Does local competition increase the coexistence of species in intransitive networks?

Competitive intransitivity, a situation in which species' competitive ranks cannot be listed in a strict hierarchy, promotes species coexistence through "enemy's enemy indirect facilitation." Theory suggests that intransitivity-mediated coexistence is enhanced when competitive interactions occur at local spatial scales, although this hypothesis has not been thoroughly tested. Here, we use a lattice model to investigate the effect of local vs. global competition on intransitivity-mediated coexistence across a range of species richness values and levels of intransitivity. Our simulations show that local competition can enhance intransitivity-mediated coexistence in the short-term, yet hinder it in the long-term, when compared to global competition. This occurs because local competition slows species disaggregation, allowing weaker competitors to persist longer in the shifting spatial refuges of intransitive networks, enhancing short-term coexistence. Conversely, our simulations show that, in the long-term, local competition traps disaggregated species in unfavorable areas of the competitive arena, where they are excluded by superior competitors. As a result, in the long-term, global intransitive competition allows a greater number of species to coexist than local intransitive competition.     

Interacting effects of habitat destruction and changing disturbance rates on biodiversity: Who is going to survive?

Changes in disturbance rates due to climate change may increase or decrease diversity, whereas permanent loss of habitat is generally believed to decrease diversity. It is, however, very likely that the effects of disturbances and habitat destruction interact. Understanding such combined effects is essential to predict the response of communities to global changes and in particular which functional types of species are most endangered. Using an individual-based spatially explicit community model, we investigate (1) whether diversity-disturbance curves alter when spatially uncorrelated or autocorrelated habitat destruction is added, and (2) which functional types of species are able to survive under these altered conditions. Model communities consisted of four functional types of species trading off between colonisation ability and competition strength. We found that habitat destruction may alter both height and shape of diversity-disturbance curves: maximum diversity at intermediate disturbance rates may shift to other disturbance rates or even split into two peaks giving rise to bimodal diversity-disturbance relationships with different sub-communities persisting at low and high disturbance rates. Diversity responded differentially depending on how the colonisation-competition trade-off was represented. Our results suggest that, for trade-offs in seed production rate, generally the best coloniser will better withstand the interacting effects of habitat destruction and changing disturbance rates; however, for trade-offs in mean dispersal distances, functional types characterized by intermediate abilities will perform best. We conclude that predictions of the impacts of changing disturbance rates on biodiversity depend on community structure and cannot be made without knowledge of concurrent permanent habitat destruction.     

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.     

Physical and biological disturbances interact differently with productivity: effects on floral and faunal richness

Physical and biological disturbances are ecological processes affecting patterns in biodiversity at a range of scales in a variety of terrestrial and aquatic systems. Theoretical and empirical evidence suggest that effects of disturbance on diversity differ qualitatively and quantitatively, depending on levels of productivity (e.g., the dynamic equilibrium model). In this study we contrasted the interactive effects between physical disturbance and productivity to those between biological disturbance and productivity. Furthermore, to evaluate how these effects varied among different components of marine hard-substratum assemblages, analyses were done separately on algal and invertebrate richness, as well as richness of the whole assemblage. Physical disturbance (wave action) was simulated at five distinct frequencies, while biological disturbance (grazing periwinkles) was manipulated as present or absent, and productivity was manipulated as high or ambient. Uni- and multivariate analyses both showed significant effects of physical disturbance and interactive effects between biological disturbance and productivity on the composition of assemblages and total species richness. Algal richness was significantly affected by productivity and biological disturbance, whereas invertebrate richness was affected by physical disturbance only. Thus, we show, for the first time, that biological disturbance and physical disturbance interact differently with productivity, because these two types of disturbances affect different components of assemblages. These patterns might be explained by differences in the distribution (i.e., press vs. pulse) and degree of selectivity between disturbances. Because different types of disturbance can affect different components of assemblages, general ecological models will benefit from using natural diverse communities, and studies concerned with particular subsets of assemblages may be misleading. In conclusion, this study shows that the outcome of experiments on effects of disturbance and productivity on diversity is greatly influenced by the composition of the assemblage under study, as well as on the type of disturbance that is used as an experimental treatment.     

Scale-Dependent Effects of Habitat Disturbance on Species Richness in Tropical Forests

Abstract: Despite growing concern, no consensus has emerged over the effects of habitat modification on species diversity in tropical forests. Even for comparatively well-studied taxa such as Lepidoptera, disturbance has been reported to increase and decrease diversity with approximately equal frequency. Species diversity within landscapes depends on the spatial scale at which communities are sampled, and the effects of disturbance in tropical forests have been studied at a wide range of spatial scales. Yet the question of how disturbance affects diversity at different spatial scales has not been addressed. We reanalyzed data from previous studies to examine the relationship between spatial scale and effects of disturbance on tropical-forest Lepidoptera. Disturbance had opposite effects on diversity at large and small scales: as scale decreased, the probability of a positive effect of disturbance on diversity increased. We also explicitly examined the relationship between spatial scale and the diversity of butterflies in selectively logged and unlogged forest in Maluku Province, Indonesia. Species richness increased with spatial scale in both logged and unlogged forest, but at a significantly faster rate in unlogged forest, whereas species evenness increased with scale in unlogged forest but did not increase with scale in logged forest. These data indicate that the effects of habitat modification on species diversity are heavily scale-dependent. As a result, recorded effects of disturbance were strongly influenced by the spatial scale at which species assemblages were sampled. Future studies need to account for this by explicitly examining the effects of disturbance at a number of different spatial scales. A further problem arises because the relationship between scale and diversity is likely to differ among taxa in relation to mobility. This may explain to some extent why the measured effects of disturbance have differed between relatively mobile and immobile taxa.     

The influence of environmental spatial structure on the life-history traits and diversity of species in a metacommunity

Several models have been proposed to understand how so many species can coexist in ecosystems. Despite evidence showing that natural habitats are often patchy and fragmented, these models rarely take into account environmental spatial structure. In this study we investigated the influence of spatial structure in habitat and disturbance regime upon species’ traits and species’ coexistence in a metacommunity. We used a population-based model to simulate competing species in spatially explicit landscapes. The species traits we focused on were dispersal ability, competitiveness, reproductive investment and survival rate. Communities were characterized by their species richness and by the four life-history traits averaged over all the surviving species. Our results show that spatial structure and disturbance have a strong influence on the equilibrium life-history traits within a metacommunity. In the absence of disturbance, spatially structured landscapes favour species investing more in reproduction, but less in dispersal and survival. However, this influence is strongly dependent on the disturbance rate, pointing to an important interaction between spatial structure and disturbance. This interaction also plays a role in species coexistence. While spatial structure tends to reduce diversity in the absence of disturbance, the tendency is reversed when disturbance occurs. In conclusion, the spatial structure of communities is an important determinant of their diversity and characteristic traits. These traits are likely to influence important ecological properties such as resistance to invasion or response to climate change, which in turn will determine the fate of ecosystems facing the current global ecological crisis.     

Coexistence of intraguild predators and prey in resource-rich environments

The prevalence of intraguild predation (IGP) in productive environments has long puzzled ecologists. Theory predicts the exclusion of intraguild prey from such environments, but data consistently defy this expectation. This suggests that coexistence mechanisms at high resource productivity may differ from those at lower productivity. Here I present a mathematical model that investigates multiple coexistence mechanisms. I incorporate two biological features widely observed in IGP communities: intraspecific interference via cannibalism or superparasitism, and temporal refuges arising from differential sensitivities to abiotic variation. I develop predictions based on three aspects of the IG prey-IG predator interaction: mutual invasibility, transient dynamics, and long-term abundances. These predictions specify the conditions under which coexistence mechanisms reinforce vs. deter one another: when a competition-IGP trade-off allows coexistence at intermediate productivity a temporal refuge for the intraguild prey always allows coexistence at high productivity, but intraspecific interference does so only at a net fitness cost to the intraguild predator. Intraspecific interference that benefits the intraguild predator not only reduces tradeoff-mediated coexistence at intermediate productivity, but also undermines the refuge's coexistence-enhancing effect at high productivity. Different mechanism combinations yield characteristic signatures in time series data during transient dynamics. By judicious measurement of parameters and examining time series for critical signatures, one can elucidate the mechanisms that allow IGP to prevail in resource-rich environments.     

Evolving metacommunities: toward an evolutionary perspective on metacommunities

The metacommunity framework predicts that local coexistence depends on the outcome of local species interactions and regional migration. In analogous fashion, spatial structure among populations can shape species interactions through evolutionary mechanisms. Yet, most metacommunity theories assume that populations do not evolve. Here, we evaluate how evolution shapes local species coexistence and exclusion within the multiscale and multispecies context embodied by the metacommunity framework. In general, coexistence in joint ecological-evolutionary models requires low to intermediate dispersal rates that can promote maintenance of both regional species and genetic diversity. These conditions support a set of key mechanisms that modify patterns of species coexistence including local adaptation, gene storage effects, genetic rescue effects, spatial genetic subsidies, and metacommunity evolution. Multispecies extensions indicate that correlated selection can further alter the outcome of interspecific interactions depending on the magnitude and direction of correlations and shape of fitness trade-offs. We suggest that an evolving metacommunity perspective has the potential to generate novel predictions about community structure and function by incorporating the genetic and species diversity that characterize natural communities. In adopting such a perspective, we seek to facilitate understanding about the interactions between evolutionary and metacommunity dynamics.     

Productivity alters the scale dependence of the diversity-invasibility relationship

At small scales, areas with high native diversity are often resistant to invasion, while at large scales, areas with more native species harbor more exotic species, suggesting that different processes control the relationship between native and exotic species diversity at different spatial scales. Although the small-scale negative relationship between native and exotic diversity has a satisfactory explanation, we lack a mechanistic explanation for the change in relationship to positive at large scales. We investigated the native-exotic diversity relationship at three scales (range: 1-4000 km2) in California serpentine, a system with a wide range in the productivity of sites from harsh to lush. Native and exotic diversity were positively correlated at all three scales; it is rarer to detect a positive relationship at the small scales within which interactions between individuals occur. However, although positively correlated on average, the small-scale relationship between native and exotic diversity was positive at low-productivity sites and negative at high-productivity sites. Thus, the change in the relationship between native and exotic diversity does not depend on spatial scale per se, but occurs whenever environmental conditions change to promote species coexistence rather than competitive exclusion. This occurred within a single spatial scale when the environment shifted from being locally unproductive to productive.     

Elephants as agents of habitat creation for small vertebrates at the patch scale

Ecologists increasingly recognize the ability of certain species to influence ecological processes by engineering the physical environment, but efforts to develop a predictive understanding of this phenomenon are in their early stages. While many believe that the landscape-scale effects of ecosystem engineers will be to increase habitat diversity and therefore the abundance and richness of other species, few generalities exist about the effects of engineering at the scale of the engineered patch. According to one hypothesis, activities that increase structural habitat complexity within engineered patches will have positive effects on the abundance or diversity of other organisms. Here I show that, by damaging trees and increasing their structural complexity, browsing elephants create refuges used by a common arboreal lizard. Observational surveys and a lizard transplant experiment revealed that lizards preferentially occupy trees with real or simulated elephant damage. A second experiment showed that lizards vacate trees when elephant-engineered refuges are removed. Furthermore, local lizard densities increased with (and may be constrained by) local densities of elephant-damaged trees. This facilitative effect of elephants upon lizards via patch-scale habitat modification runs contrary to previously documented negative effects of the entire ungulate guild on lizards at the landscape scale, suggesting that net indirect effects of large herbivores comprise opposing trophic and engineering interactions operating at different spatial scales. Such powerful megaherbivore-initiated interactions suggest that anthropogenic changes in large-mammal densities will have important cascading consequences for ecological communities.     

Compositional stability and diversity of vascular plant communities following logging disturbance in Appalachian forests

Human-caused changes in disturbance regimes and introductions of nonnative species have the potential to result in widespread, directional changes in forest community structure. The degree that plant community composition persists or changes following disturbances depends on the balance between local extirpation and colonization by new species, including nonnatives. In this study, we examined species losses and gains, and entry of native vs. exotic species to determine how oak forests in the Appalachian Mountains might shift in species composition following a gradient of pulse disturbances (timber harvesting). We asked (1) how compositional stability of the plant community (resistance and resilience) was influenced by disturbance intensity, (2) whether community responses were driven by extirpation or colonization of species, and (3) how disturbance intensity influenced total and functional group diversity, including the nonnative proportion of the flora through time. We collected data at three spatial scales and three times, including just before, one year post-disturbance, and 10 years post-disturbance. Resistance was estimated using community distance measures between pre- and one year post-disturbance, and resilience using community distance between pre- and 10-year post-disturbance conditions. The number of colonizing and extirpated species between sampling times was analyzed for all species combined and for six functional groups. Resistance and resilience decreased with increasing timber-harvesting disturbance; compositional stability was lower in the most disturbed plots, which was driven by colonization, but not extirpation, of species. Colonization of species also led to increases in diversity after disturbance that was typically maintained after 10 years following disturbance. Most of the community-level responses were driven by post-disturbance colonization of native forbs and graminoids. The nonnative proportion of plant species tended to increase following disturbance, especially at large spatial scales in the most disturbed treatments, but tended to decrease through time following disturbance due to canopy development. The results of this study are consistent with the theory that resources released by disturbance have strong influences on species colonization and community composition. The effects of management activities tested in this study, which span a gradient of timber-harvesting disturbance, shift species composition largely via an increase in species colonization and diversity.     

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.