Consequences of varying regional heterogeneity in source–sink metacommunities

Although the influence of dispersal on coexistence mechanisms in metacommunities has received great emphasis, few studies have addressed how such influence is affected varying regional heterogeneity. We present a mechanistic model of resource competition in a metacommunity based on classical models of plant competition for limiting resources. We defined regional heterogeneity as the differences in resource supply rates (or resource availabilities) across local communities. As suggested by previous work, the highest diversify occurred at intermediate levels of dispersal among local communities. However our model shows how the effects of dispersal depend on the amount of heterogeneity among local communities and vice versa. Both regional and local species richness were the highest when heterogeneity was intermediate. We suggest that empirical studies that found no evidence for source–sink or mass effects at the community level may have examined communities with limited ranges of dispersal and regional heterogeneity. This model of species coexistence contributes to a broader understanding of patterns in real communities.     

Environmental gradients determine the potential for ecosystem engineering effects

Understanding processes that determine biodiversity is a fundamental challenge in ecology. At the landscape scale, physical alteration of ecosystems by organisms, called ecosystem engineering, enhances biodiversity worldwide by increasing heterogeneity in resource conditions and enhancing species coexistence across engineered and non‐engineered habitats. Engineering–diversity relationships can vary along environmental gradients due to changes in the amount of physical structuring created by ecosystem engineering, but it is unclear how this variation is influenced by the responsiveness of non‐structural abiotic properties to engineering. Here we show that environmental gradients determine the capacity for engineering to alter resource availability and species diversity, independent of the magnitude of structural change produced by engineering. We created an experimental rainfall gradient in an arid grassland where rodents restructure soils by constructing large, long‐lasting burrows. We found that greater rainfall increased water availability and productivity in both burrow and inter‐burrow habitats, causing a decline in local (alpha) plant diversity within both of these habitats. However, increased rainfall also resulted in greater differences in soil resources between burrow and inter‐burrow habitats, which increased species turnover (beta diversity) across habitats and stabilized landscape‐level (gamma) diversity. These responses occurred regardless of rodent presence and without changes in the extent of physical alteration of soils by rodents. Our results suggest that environmental gradients can influence the effects of ecosystem engineering in maintaining biodiversity via resource heterogeneity and species turnover. In an era of rapid environmental change, accounting for this interaction may be critical to conservation and management.     

Regional enrichment of local assemblages is robust to variation in local productivity, abiotic gradients, and heterogeneity

Theory predicts that the effects of regional richness on the richness of local communities may depend on the productivity, resource availability, and/or heterogeneity of local sites. Using the wetland plant communities of 50 independent streams as 'regions', we tested whether: (1) local richness in 1-m² quadrats and 50-m stream segments was positively related to regional richness, even after environmental influences were considered; and (2) the effect of regional richness would interact with the effects of biomass, soil moisture, and/or heterogeneity on local richness. In models that explained up to 88% of variation in local richness, we found that richness at both local scales was positively related to regional richness, and that regional richness did not interact with any of the environmental gradients that also shaped local richness. We conclude that species availability from the regional pool may consistently enrich local communities, even while other constraints on local richness operate.     

Stoichiometrically explicit competition between grazers: species replacement, coexistence, and priority effects along resource supply gradients

Assuming key trade-offs among interactors, several models (resource ratio, keystone predation, intraguild predation) predict changes in species composition over resource supply gradients. Ecological stoichiometry could also predict compositional shifts of grazers over gradients of nutrient and light supply through a mechanism involving (mis)matches between elemental body composition of grazers and plants. This hypothesis is explored here using a suite of two-grazer, one-plant models that incorporate three key components: plant production depends on light and nutrients, nutrient content of plants can vary, and homeostatic grazers can be carbon or nutrient limited. The results from this suite closely resemble the classical resource ratio model describing plant competition for two resources. Here, the models predict shifts of grazer composition along resource supply gradients if species trade off competitive abilities for plant carbon and nutrients. Given this trade-off, superior nutrient competitors should dominate low nutrient environments, and superior carbon competitors should dominate high nutrient environments. At intermediate nutrient supply, species can coexist at a stable equilibrium, or alternative stable states emerge, depending on how grazers impact their resources. These results depend on food web architecture, however. For instance, predators can alter or reduce possibilities for stoichiometry-mediated coexistence of grazers.     

Niche-habitat mechanisms and biotic interactions explain the coexistence and abundance of congeneric sandgrouse species

Ascertaining which niche processes allow coexistence between closely related species is of special interest in ecology. We quantified variations in the environmental niches and densities of two congeneric species, the pin-tailed and the black-bellied sandgrouse (Pterocles alchata and Pterocles orientalis) in allopatry and sympatry under similar abiotic, habitat and dispersal contexts to understand their coexistence. Using principal component analysis, we defined environmental gradients (niche dimensions) including abiotic, habitat and anthropogenic variables, and calculated niche breadth, position and overlap of both species in sympatry and allopatry. Additionally, sandgrouse density was modelled as a function of the niche dimensions and the density of the other species. We found evidence that each species occupies distinct environmental niches in sympatry and in allopatry. The black-bellied sandgrouse exploits a broader range of environmental conditions (wider niche breadth) while the pin-tailed sandgrouse reaches high densities where conditions seem to match its optimum. In sympatry, both species shift their niches to intermediate positions, indicating the importance of abiotic factors in setting coexistence areas. Environmental conditions determine regional densities of pin-tailed sandgrouse whereas biotic interactions explain the density of the black-bellied sandgrouse in areas with abiotic conditions similarly conducive for both species. Highly suitable areas for the pin-tailed sandgrouse fall beyond the upper thermal limit of the black-bellied sandgrouse, leading to niche segregation and low densities for the latter. Finally, local niche shift and expansion plus possible heterospecific aggregation allow the pin-tailed sandgrouse to thrive in a priori less favourable environments. This work provides insight into how different mechanisms allow species coexistence and how species densities vary in sympatry compared to allopatry as a result of environmental filtering and biotic interactions.     

Spatial heterogeneity, source-sink dynamics, and the local coexistence of competing species

Patch occupancy theory predicts that a trade-off between competition and dispersal should lead to regional coexistence of competing species. Empirical investigations, however, find local coexistence of superior and inferior competitors, an outcome that cannot be explained within the patch occupancy framework because of the decoupling of local and spatial dynamics. We develop two-patch metapopulation models that explicitly consider the interaction between competition and dispersal. We show that a dispersal-competition trade-off can lead to local coexistence provided the inferior competitor is superior at colonizing empty patches as well as immigrating among occupied patches. Immigration from patches that the superior competitor cannot colonize rescues the inferior competitor from extinction in patches that both species colonize. Too much immigration, however, can be detrimental to coexistence. When competitive asymmetry between species is high, local coexistence is possible only if the dispersal rate of the inferior competitor occurs below a critical threshold. If competing species have comparable colonization abilities and the environment is otherwise spatially homogeneous, a superior ability to immigrate among occupied patches cannot prevent exclusion of the inferior competitor. If, however, biotic or abiotic factors create spatial heterogeneity in competitive rankings across the landscape, local coexistence can occur even in the absence of a dispersal-competition trade-off. In fact, coexistence requires that the dispersal rate of the overall inferior competitor not exceed a critical threshold. Explicit consideration of how dispersal modifies local competitive interactions shifts the focus from the patch occupancy approach with its emphasis on extinction-colonization dynamics to the realm of source-sink dynamics. The key to coexistence in this framework is spatial variance in fitness. Unlike in the patch occupancy framework, high rates of dispersal can undermine coexistence, and hence diversity, by reducing spatial variance in fitness.     

Spatial dynamics in model plant communities: what do we really know?

A variety of models have shown that spatial dynamics and small-scale endogenous heterogeneity (e.g., forest gaps or local resource depletion zones) can change the rate and outcome of competition in communities of plants or other sessile organisms. However, the theory appears complicated and hard to connect to real systems. We synthesize results from three different kinds of models: interacting particle systems, moment equations for spatial point processes, and metapopulation or patch models. Studies using all three frameworks agree that spatial dynamics need not enhance coexistence nor slow down dynamics; their effects depend on the underlying competitive interactions in the community. When similar species would coexist in a nonspatial habitat, endogenous spatial structure inhibits coexistence and slows dynamics. When a dominant species disperses poorly and the weaker species has higher fecundity or better dispersal, competition-colonization trade-offs enhance coexistence. Even when species have equal dispersal and per-generation fecundity, spatial successional niches where the weaker and faster-growing species can rapidly exploit ephemeral local resources can enhance coexistence. When interspecific competition is strong, spatial dynamics reduce founder control at large scales and short dispersal becomes advantageous. We describe a series of empirical tests to detect and distinguish among the suggested scenarios.     

Seed arrival, ecological filters, and plant species richness: a meta-analysis

Theoretical models predict that effects of dispersal on local biodiversity are influenced by the size and composition of the species pool, as well as ecological filters that limit local species membership. We tested these predictions by conducting a meta-analysis of 28 studies encompassing 62 experiments examining effects of propagule supply (seed arrival) on plant species richness under contrasting intensities of ecological filters (owing to disturbance and resource availability). Seed arrival increased local species richness in a wide range of communities (forest, grassland, montane, savanna, wetland), resulting in a positive mean effect size across experiments. Mean effect size was 70% higher in disturbed relative to undisturbed communities, suggesting that disturbance increases recruitment opportunities for immigrating species. In contrast, effect size was not significantly influenced by nutrient or water availability. Among seed-addition experiments, effect size was positively correlated with species and functional diversity within the pool of added seeds (species evenness and seed-size diversity), primarily in disturbed communities. Our analysis provides experimental support for the general hypothesis that species pools and local environmental heterogeneity interactively structure plant communities. We highlight empirical gaps that can be addressed by future experiments and discuss implications for community assembly, species coexistence, and the maintenance of biodiversity.     

Resource quantity, not resource heterogeneity, maintains plant diversity

Resource heterogeneity has often been proposed to explain the maintenance of plant species diversity and patterns of species diversity along productivity gradients. Resource heterogeneity should maintain biodiversity by preventing competitive exclusion because different species are superior competitors in different parts of a heterogeneous environment. In natural systems, however, resource heterogeneity covaries with average resource supply rate, making the effect of heterogeneity difficult to isolate. Using a novel experimental approach, we tested the independent effects of resource heterogeneity and average supply rate on plant species diversity. We show that the average supply rate of the most limiting resource controlled species diversity, whereas heterogeneity of this resource had virtually no effect. These findings also suggest that biodiversity declines with increasing productivity because at high enough levels of productivity one resource may always be driven to sufficiently short supply to exclude many species.     

Aggregation, intraguild interactions and the coexistence of competitors on small ephemeral patches

It is well established that intraspecific aggregation has the potential to promote coexistence in communities of species competing for patchy ephemeral resources. We developed a simulation model to explore the influence of aggregation on coexistence in such communities when an important assumption of previous studies – that interspecific interactions have only negative effects on the species involved – is relaxed. The model describes a community of competing insect larvae in which an interaction that is equivalent to intraguild predation (IGP) can occur, and is unusual in that it considers species exploiting very small resource patches (carrying capacity=1). Model simulations show that, in the absence of any intraspecific aggregation, variation between species in the way that resource heterogeneity affects survival increases the likelihood of species coexistence. Simulations also show that intraspecific aggregation of the dominant competitor's eggs across resource patches can promote coexistence by reducing the importance of interspecific competition relative to that of intraspecific competition. Crucially, however, this effect is altered if one competitor indulges in IGP. In general, coexistence is only possible when the species that is capable of IGP is less effective at exploiting the shared resource than its competitor. Because it reduces the relative importance of interspecific interactions, intraspecific aggregation of the eggs of a species that is the victim of IGP actually reduces the likelihood of coexistence in parts of parameter space in which the persistence of the other species is dependent on its ability to exploit its competitor. Since resource heterogeneity, intraspecific aggregation and IGP are all common phenomena, these findings shed light on mechanisms that are likely to influence diversity in communities exploiting patchy resources.     

Vegetation heterogeneity and diversity in flat and mountain landscapes of Patagonia (Argentina)

Abstract. We studied floristic and diversity patterns and their environmental controls in two landscapes of contrasting topography in the Patagonian steppe. The analyses were focused on the effects of water availability gradients and landscape configuration on plant species distribution and coexistence. Floristic variation was investigated using Correspondence Analysis. The relationship between floristic and environmental variation was analyzed using Canonical Correspondence Analysis and correlation tests. We explored diversity patterns by relating spatial distance to floristic dissimilarities. The floristic gradient was determined by shrub and grass species and was related to precipitation in the flat area, and to precipitation, elevation and potential radiation in the mountain area. Site species richness increased with water availability in both areas. Mean site species richness and species turnover in space was higher in the mountain than in the flat area. Landscape species richness and floristic gradients were more concentrated in the mountain than in the flat area. In contrast to shrubs and grasses, forb species distributions were uncoordinated and probably independent of any environmental gradient. Our results suggest (1) that landscape configuration affects species composition and diversity through its direct effect on abiotic environmental heterogeneity, and (2) that the environmental controls of the community composition vary depending on the plant functional type considered.     

Host specialist clownfishes are environmental niche generalists

Why generalist and specialist species coexist in nature is a question that has interested evolutionary biologists for a long time. While the coexistence of specialists and generalists exploiting resources on a single ecological dimension has been theoretically and empirically explored, biological systems with multiple resource dimensions (e.g. trophic, ecological) are less well understood. Yet, such systems may provide an alternative to the classical theory of stable evolutionary coexistence of generalist and specialist species on a single resource dimension. We explore such systems and the potential trade-offs between different resource dimensions in clownfishes. All species of this iconic clade are obligate mutualists with sea anemones yet show interspecific variation in anemone host specificity. Moreover, clownfishes developed variable environmental specialization across their distribution. In this study, we test for the existence of a relationship between host-specificity (number of anemones associated with a clownfish species) and environmental-specificity (expressed as the size of the ecological niche breadth across climatic gradients). We find a negative correlation between host range and environmental specificities in temperature, salinity and pH, probably indicating a trade-off between both types of specialization forcing species to specialize only in a single direction. Trade-offs in a multi-dimensional resource space could be a novel way of explaining the coexistence of generalist and specialists.     

The biogeography and filtering of woody plant functional diversity in North and South America

Aim In recent years evidence has accumulated that plant species are differentially sorted from regional assemblages into local assemblages along local‐scale environmental gradients on the basis of their function and abiotic filtering. The favourability hypothesis in biogeography proposes that in climatically difficult regions abiotic filtering should produce a regional assemblage that is less functionally diverse than that expected given the species richness and the global pool of traits. Thus it seems likely that differential filtering of plant traits along local‐scale gradients may scale up to explain the distribution, diversity and filtering of plant traits in regional‐scale assemblages across continents. The present work aims to address this prediction. Location North and South America. Methods We combine a dataset comprising over 5.5 million georeferenced plant occurrence records with several large plant functional trait databases in order to: (1) quantify how several critical traits associated with plant performance and ecology vary across environmental gradients; and (2) provide the first test of whether the woody plants found within 1° and 5° map grid cells are more or less functionally diverse than expected, given their species richness, across broad gradients. Results The results show that, for many of the traits studied, the overall distribution of functional traits in tropical regions often exceeds the expectations of random sampling given the species richness. Conversely, temperate regions often had narrower functional trait distributions than their smaller species pools would suggest. Main conclusion The results show that the overall distribution of function does increase towards the equator, but the functional diversity within regional‐scale tropical assemblages is higher than that expected given their species richness. These results are consistent with the hypothesis that abiotic filtering constrains the overall distribution of function in temperate assemblages, but tropical assemblages are not as tightly constrained.     

Fusing spatial resource heterogeneity with a competition–colonization trade-off in model communities

Two commonly cited mechanisms of multispecies coexistence in patchy environments are spatial heterogeneity in competitive abilities caused by variation in resources and a competition–colonization trade-off. In this paper, a model that fuses these mechanisms together is presented and analyzed. The model suggests that spatial variation in resource ratios can lead to multispecies coexistence, but this mechanism by itself is weak when the number of resources for which species compete is small. However, spatial resource heterogeneity is a powerful mechanism for multispecies coexistence when it acts synergistically with a competition–colonization trade-off. The model also shows how resource supply can control the competitive balance between species that are weak competitors but superior colonizers and strong competitors/inferior colonizers. This provides additional theoretical support for a possible explanation of empirically observed hump-shaped relationships between species diversity and ecological productivity.     

From plant neighbourhood to landscape scales: how grazing modifies native and exotic plant species richness in grassland

The interactive effect of grazing and soil resources on plant species richness and coexistence has been predicted to vary across spatial scales. When resources are not limiting, grazing should reduce competitive effects and increase colonisation and richness at fine scales. However, at broad scales richness is predicted to decline due to loss of grazing intolerant species. We examined these hypotheses in grasslands of southern Australia that varied in resources and ungulate grazing intensity since farming commenced 170 years ago. Fine-scale species richness was slightly greater in more intensively grazed upper slope sites with high nutrients but low water supply compared to those that were moderately grazed, largely due to a greater abundance of exotic species. At broader scales, exotic species richness declined with increasing grazing intensity whether nutrients or water supply were low or high. Native species richness declined at all scales in response to increasing grazing intensity and greater resource supply. Grazing also reduced fine-scale heterogeneity in native species richness and although exotics were also characterised by greater heterogeneity at fine scales, grazing effects varied across scales. In these grasslands patterns of plant species richness did not match predictions at all scales and this is likely to be due to differing responses of native and exotic species and their relative abundance in the regional species pool. Over the past 170 years intolerant native species have been eliminated from areas that are continually and heavily grazed, whereas transient, light grazing increases richness of both exotics and natives. The results support the observation that the processes and scales at which they operate differ between coevolved ungulate—grassland systems and those in transition due to recent invasion of herbivores and associated plant species.     

Large-scale multi-trophic co-response models and environmental control of pelagic food webs in Québec lakes

Environmental heterogeneity plays a fundamental role in driving species distributions by, for one, fostering niche dimensionality. Within lake ecosystems, species distributions and concordance patterns are driven by both local and regional heterogeneity, though their relative importance across trophic levels has rarely been explored. We developed a statistical framework to compare responses of taxa from different trophic levels to abiotic factors and determine how this affected multi-trophic network structures. In particular, we used multi-species concordance modelling (concordance analysis and RV coefficient) to determine species associations and correlations within and among three trophic levels (phytoplankton, zooplankton and fish communities sampled across 49 southern Québec lakes, covering eight hydrological regions). We then used multiple factor analysis, latent variable modelling and local contributions of sites to beta diversity to assess the relative importance of major environmental gradients in structuring species co-responses and species interaction turnover across the landscape. Our analyses confirmed that concordant species within each trophic level varied jointly or segregated into different pelagic food webs in Québec lakes where important acidification and eutrophication took place. Some keynote species were indicators of different food web compartments and distinguished groups of lakes along multiple environmental niche dimensions. Among the three trophic levels examined, zooplankton depicted the highest proportion of species concordance and appeared to act as a trophic linkage between phytoplankton and fish. Ultimately, the losses or gains in species richness and species interactions were strongly driven by environmental gradients. This study provides for the first time a combined analysis of the effects of environmental heterogeneity on ecological communities belonging to three trophic levels sampled near simultaneously across an 800 km broad lacustrine landscape. The new framework developed in this study has a great potential to better understand the complex response of aquatic ecosystems in a world increasingly affected by multiple, cumulative stressors.     

The relationship between species richness and ecosystem variability is shaped by the mechanism of coexistence

Theory relating species richness to ecosystem variability typically ignores the potential for environmental variability to promote species coexistence. Failure to account for fluctuation‐dependent coexistence may explain deviations from the expected negative diversity–ecosystem variability relationship, and limits our ability to predict the consequences of increases in environmental variability. We use a consumer‐resource model to explore how coexistence via the temporal storage effect and relative nonlinearity affects ecosystem variability. We show that a positive, rather than negative, diversity–ecosystem variability relationship is possible when ecosystem function is sampled across a natural gradient in environmental variability and diversity. We also show how fluctuation‐dependent coexistence can buffer ecosystem functioning against increasing environmental variability by promoting species richness and portfolio effects. Our work provides a general explanation for variation in observed diversity–ecosystem variability relationships and highlights the importance of conserving regional species pools to help buffer ecosystems against predicted increases in environmental variability.     

Positive interactions among competitors can produce species-rich communities

Although positive interactions between species are well documented, most ecological theory for investigating multispecies coexistence remains rooted in antagonistic interactions such as competition and predation. Standard resource-competition models from this theory predict that the number of coexisting species should not exceed the number of factors that limit population growth. Here I show that positive interactions among resource competitors can produce species-rich model communities supported by a single limiting resource. Simulations show that when resource competitors reduce each others' per capita mortality rate (e.g. by ameliorating an abiotic stress), stable multispecies coexistence with a single resource may be common, even while the net interspecific interaction remains negative. These results demonstrate that positive interactions may provide an important mechanism for generating species-rich communities in nature. They also show that focusing on the net interaction between species may conceal important coexistence mechanisms when species simultaneously engage in both antagonistic and positive interactions.     

Taxonomic scale-dependence of habitat niche partitioning and biotic neighbourhood on survival of tropical tree seedlings

In order to differentiate between mechanisms of species coexistence, we examined the relative importance of local biotic neighbourhood, abiotic habitat factors and species differences as factors influencing the survival of 2330 spatially mapped tropical tree seedlings of 15 species of Myristicaceae in two separate analyses in which individuals were identified first to species and then to genus. Using likelihood methods, we selected the most parsimonious candidate models as predictors of 3 year seedling survival in both sets of analyses. We found evidence for differential effects of abiotic niche and neighbourhood processes on individual survival between analyses at the genus and species levels. Niche partitioning (defined as an interaction of taxonomic identity and abiotic neighbourhood) was significant in analyses at the genus level, but did not differentiate among species in models of individual seedling survival. By contrast, conspecific and congeneric seedling and adult density were retained in the minimum adequate models of seedling survival at species and genus levels, respectively. We conclude that abiotic niche effects express differences in seedling survival among genera but not among species, and that, within genera, community and/or local variation in adult and seedling abundance drives variation in seedling survival. These data suggest that different mechanisms of coexistence among tropical tree taxa may function at different taxonomic or phylogenetic scales. This perspective helps to reconcile perceived differences of importance in the various non-mutually exclusive mechanisms of species coexistence in hyper-diverse tropical forests.     

Competition and coexistence in spatially subdivided habitats

While non-spatial models predict that like species cannot stably coexist, empirical studies suggest that similar species have similar distributions due to shared habitat requirements. A model is developed to discuss competition and coexistence in subdivided but locally stable habitats. The model predicts that in some cases it is possible for one species to exclude the other species from a geographic region, while in other cases two competing species can stably coexist. The equilibrium level and the fraction of doubly occupied patches, if there is coexistence, are determined by the strength of competition on colonization and exclusion in such a system. Also, it is possible for two ecologically identical species to stably coexist, and two asymmetrically competing species can coexist when there is a trade-off between local competition ability and invasion ability. When rescue effects are considered, the stable region at internal equilibrium point would be reduced, but the fraction of doubly occupied patches would be enlarged.