Stabilizing Niche Differences Research Articles

Frequency-dependent tree growth depends on climate.

Climate and competition interact to affect species' performance, such as growth and survival, and help determine species distributions and coexistence. However, it is unclear how climatic conditions modulate frequency-dependent performance, that is, how performance changes as a species becomes locally rare or common. This is critical because declines in performance as a species becomes more common (negative frequency dependence) is a signature of niche differences among species that stabilize coexistence, whereas positive frequency dependence leads to priority effects and hampers species coexistence. Here, we used dendrochronology and hierarchical models to test whether frequency-dependent growth of sugar pine (Pinus lambertiana) depends on climatic conditions. We found that growth rates were strongly dependent on annual precipitation, but no frequency dependence was evident across all years. However, there was a strong interaction between precipitation and frequency dependence, revealing stabilizing niche differences in dry years but positive frequency dependence in wet years. These differences emerged because of precipitation-driven changes in the direction and strength of both con- and heterospecific competition. Overall, these results show how stabilizing and destabilizing effects can be temporally dynamic for long-lived species and interact with climate variation.

A native annual forb locally excludes a closely related introduced species that co-occurs in oak-savanna habitat remnants.

Despite the ubiquity of introduced species, their long-term impacts on native plant abundance and diversity remain poorly understood. Coexistence theory offers a tool for advancing this understanding by providing a framework to link short-term individual measurements with long-term population dynamics by directly quantifying the niche and average fitness differences between species. We observed that a pair of closely related and functionally similar annual plants with different origins—native Plectritis congesta and introduced Valerianella locusta—co-occur at the community scale but rarely at the local scale of direct interaction. To test whether niche and/or fitness differences preclude local-scale long-term coexistence, we parameterized models of competitor dynamics with results from a controlled outdoor pot experiment, where we manipulated densities of each species. To evaluate the hypothesis that niche and fitness differences exhibit environmental dependency, leading to community-scale coexistence despite local competitive exclusion, we replicated this experiment with a water availability treatment to determine if this key limiting resource alters the long-term prediction. Water availability impacted population vital rates and intensities of intraspecific versus interspecific competition between P. congesta and V. locusta. Despite environmental influence on competition our model predicts that native P. congesta competitively excludes introduced V. locusta in direct competition across water availability conditions because of an absence of stabilizing niche differences combined with a difference in average fitness, although this advantage weakens in drier conditions. Further, field data demonstrated that P. congesta densities have a negative effect on V. locusta seed prediction. We conclude that native P. congesta limits abundances of introduced V. locusta at the direct-interaction scale, and we posit that V. locusta may rely on spatially dependent coexistence mechanisms to maintain coexistence at the site scale. In quantifying this competitive outcome our study demonstrates mechanistically how a native species may limit the abundance of an introduced invader.

Natural enemies do not contribute to negative frequency-dependence in native and exotic grassland plants

Species’ responses to antagonists, such as fungal pathogens and insect herbivores, are part of a species’ niche and are thought to contribute to species coexistence. In theory, coexistence is supposed to be maintained by the effects of equalizing mechanisms that reduce fitness differences among species, and stabilizing mechanisms that are the result of niche differences. The presence of stabilizing niche differences is hallmarked by negative frequency-dependent per capita rates of increase. However, rigorous experimental testing of how insect herbivores and fungal pathogens contribute to negative frequency dependence among co-occurring plant species, in particular in the context of biological invasions, is scarce. We present an experimental study in a grassland in central Germany, in which we established native and exotic-dominated communities on 1 m²-plots, and reduced above- and belowground fungal pathogens and insect herbivores by applying pesticides. We manipulated the frequencies of six native and six exotic focal species, in order to assemble one dominant and five rare species per community. We quantified productivity and diversity for each community, and recorded data on per capita rates of increase and other fitness components for two native and two exotic focal species over three years. We found that native and exotic-dominated communities did not differ in their productivity and diversity. Reduction of fungi and insects increased productivity but the effect was similar for both species origins, suggesting that effects of enemy release are transitory or not existent. Negative frequency-dependence was common in our species, but could not be linked to interactions with antagonists. Although focal species responded to the reduction of antagonists, these effects were only present at low frequency, suggesting a greater importance of intraspecific competition, e.g. for resources, in explaining the observed stabilizing niche differences. Offsetting responses of the two exotic focal species to both antagonist groups highlight the significance of equalizing mechanisms among native and exotic species at the community level. Our study demonstrates the importance of both equalizing and stabilizing mechanisms in the context of biological invasions and differential contributions of antagonistic interactions to these mechanisms.

Cxr: A toolbox for modelling species coexistence in R

Abstract Recent developments in modern coexistence theory (MCT) have advanced our understanding of how species interactions among themselves and with the environment influence community dynamics. Although the formulation of MCT is mathematically clear, its application to empirical cases is still challenging, which precludes its adoption by a large range of ecologists and evolutionary biologists interested in broad questions related to community assembly and the maintenance of species diversity. We developed cxr, an r package that provides a complete toolbox for calculating species vital rates and interaction parameters, from which the user can obtain estimates of coexistence outcomes based on stabilizing niche differences and average fitness differences. Our aim is to offer a highly versatile package to accommodate different research needs. This means that the user can define population models, use different optimization algorithms and include the effect of external covariates on species interactions, which may include environmental variables (e.g. temperature, precipitation, salinity) and biotic controls (e.g. predation, pollination, mycorrhizae). To illustrate the functionality and versatility of cxr, we provide a complete set of population dynamic models and a dataset from a highly diverse grassland community. By building bridges between MCT formulation and its implementation, we provide tools to obtain a deeper mechanistic understanding of how species interactions determine basic patterns such as species abundances and dominance, which are core information for many applied fields, such as conservation, restoration and invasion biology. Finally, the package is not limited taxonomically to any particular group. The application of tools derived from MCT to a wide range of different systems can create feedbacks between empirical and theoretical studies in a way that stimulates a better understanding of the processes maintaining biodiversity.

Functional traits and phenotypic plasticity modulate species coexistence across contrasting climatic conditions

Functional traits are expected to modulate plant competitive dynamics. However, how traits and their plasticity in response to contrasting environments connect with the mechanisms determining species coexistence remains poorly understood. Here, we couple field experiments under two contrasting climatic conditions to a plant population model describing competitive dynamics between 10 annual plant species in order to evaluate how 19 functional traits, covering physiological, morphological and reproductive characteristics, are associated with species’ niche and fitness differences. We find a rich diversity of univariate and multidimensional associations, which highlight the primary role of traits related to water- and light-use-efficiency for modulating the determinants of competitive outcomes. Importantly, such traits and their plasticity promote species coexistence across climatic conditions by enhancing stabilizing niche differences and by generating competitive trade-offs between species. Our study represents a significant advance showing how leading dimensions of plant function connect to the mechanisms determining the maintenance of biodiversity.

Mutualists Stabilize the Coexistence of Congeneric Legumes.

Coexistence requires that stabilizing niche differences, which cause species to limit themselves more than others, outweigh relative fitness differences, which cause competitive exclusion. Interactions with shared mutualists, which can differentially affect host fitness and change in magnitude with host frequency, can satisfy these conditions for coexistence, yet empirical tests of mutualist effects on relative fitness and stabilizing niche differences are largely lacking within the framework of coexistence theory. Here, we show that N-fixing rhizobial mutualists mediate coexistence in four naturally co-occurring congeneric legume (Trifolium) species. Using experimental greenhouse communities, we quantified relative fitness and stabilizing niche differences for each species in the presence of rhizobia originating from conspecific or congeneric hosts. Rhizobia stabilized coexistence by increasing the self-limitation of Trifolium species grown with rhizobia isolated from conspecifics, thus allowing congeners to increase when rare. Greenhouse-measured invasion growth rates predicted natural, unmanipulated coexistence dynamics of Trifolium species over 2 years at our field sites. Our results demonstrate that interactions with shared mutualists can stabilize the coexistence of closely related species.

Distinct responses of niche and fitness differences to water availability underlie variable coexistence outcomes in semi‐arid annual plant communities

Abstract Climate change is predicted to have profound consequences for multispecies coexistence, and thus, patterns of biological diversity. These consequences will be mediated by direct and indirect impacts of environmental change on species’ vital rates and interactions. While the impacts of environmental change on individual species has received much attention to date, the consequences for coexistence mediated by changes in the strength and direction of multispecies interactions are not as well understood. To investigate how coexistence dynamics may be sensitive to environmental change, we conducted a field experiment in a diverse semi‐arid annual plant system. We imposed a water manipulation treatment in two sites that vary in aridity and associated rainfall. Focusing on four common annual plant species in these sites, we quantified the fecundity (seed production) of individuals in response to a gradient of intra‐ and interspecific competitor densities and aridity. We then used these fecundities to parameterize an annual plant population model and examine the influence of aridity and species identity on resultant coexistence dynamics (as a function of stabilizing niche differences and fitness inequalities). While the responses of some vital rates and competitive impacts to watering varied somewhat predictably across sites, coexistence metrics encapsulating changes in these vital rates and interaction strengths did not. Fitness inequalities among our focal species were driven largely by differences in sensitivity to competition, which were almost always much greater than the magnitude of stabilizing niche differences. These findings were surprising given observational evidence suggesting that these species do coexist at local scales in these natural communities. Synthesis. Our study is one of the first to explicitly consider the influence of environmental variation on the individual components of coexistence outcomes. We show that environmental change has the ability to influence coexistence not only through direct pathways (i.e., vital rates), but also indirect pathways (i.e., species interactions). Despite the consistency of many of the responses of these individual components to environmental variation, their combined influence on predictions of both current and future coexistence remains unclear.

A structural approach for understanding multispecies coexistence

AbstractAlthough observations of species‐rich communities have long served as a primary motivation for research on the coexistence of competitors, the majority of our empirical and theoretical understanding comes from two‐species systems. How much of the coexistence observed in species‐rich communities results from indirect effects among competitors that only emerge in diverse systems remains poorly understood. Resolving this issue requires simple, scalable, and intuitive metrics for quantifying the conditions for coexistence in multispecies systems, and how these conditions differ from those expected based solely on pairwise interactions. To achieve these aims, we develop a structural approach for studying the set of parameter values compatible withn‐species coexistence given the geometric constraints imposed by the matrix of competition coefficients. We derive novel mathematical metrics analogous to stabilizing niche differences and fitness differences that measure the range of conditions compatible with multispecies coexistence, incorporating the effects of indirect interactions emerging in diverse systems. We show how our measures can be used to quantify the extent to which the conditions for coexistence in multispecies systems differ from those that allow pairwise coexistence, and apply the method to a field system of annual plants. We conclude by presenting new challenges and empirical opportunities emerging from our structural metrics of multispecies coexistence.

Ecological drift and the distribution of species diversity.

Ecological drift causes species abundances to fluctuate randomly, lowering diversity within communities and increasing differences among otherwise equivalent communities. Despite broad interest in ecological drift, ecologists have little experimental evidence of its consequences in nature, where competitive forces modulate species abundances. We manipulated drift by imposing 40-fold variation in the size of experimentally assembled annual plant communities and holding their edge-to-interior ratios comparable. Drift over three generations was greater than predicted by neutral models, causing high extinction rates and fast divergence in composition among smaller communities. Competitive asymmetries drove populations of most species to small enough sizes that demographic stochasticity could markedly influence dynamics, increasing the importance of drift in communities. The strong effects of drift occurred despite stabilizing niche differences, which cause species to have greater population growth rates when at low local abundance. Overall, the importance of ecological drift appears greater in non-neutral communities than previously recognized, and varies with community size and the type and strength of density dependence.

Ecological and Evolutionary Diversification Within the Genus <I>Carex</I> (Cyperaceae): Consequences for Community Assembly in Subarctic Fens

The concept of limiting similarity predicts that closely related taxa are less likely to co-occur than expected by chance. The degree to which the phylogenetic relatedness in plant communities is in accord with limiting similarity has been little tested at the scale where the consequences of adaptive differentiation during speciation should be most evident: the scale of neighboring, congeneric plants within a community. To quantify species co-occurrence patterns in relation to environment, we sampled sedge species, their rooting level relative to the water table, and the water pH in 2,124 0.25 m2 quadrats distributed across 29 subarctic fens in the central Labrador Peninsula. We estimated phylogenetic relationships using four DNA regions (ETS, ITS, matK, trnL-trnF) for all species of Carex (42), Eriophorum (6), and Trichophorum (2) in the region, of which 21, four, and two, respectively, occurred in the sampled fens. We demonstrate that closely related species of Carex are less likely to co-occur than expected by chance using 1) a probabilistic method to test the significance of pairwise co-occurrence patterns of species, and 2) linear mixed modeling to relate these patterns to phylogenetic relationships and ecological tolerances along gradients of substrate pH and rooting level in relation to the water table. The results also indicate that suites of species with significant mutual pairwise co-occurrence belong to distant lineages within the Cariceae-Dulichieae-Scirpeae clade of Cyperaceae and have stabilizing niche differences. We suggest that niche differentiation during the evolution and diversification of a clade of wetland Carex species over the past few million years, especially during the dynamic glacial cycles of the Pleistocene, has resulted in diverse sedge communities that share space and resources in harsh northern peatland habitats.

Trait adaptation promotes species coexistence in diverse predator and prey communities.

Species can adjust their traits in response to selection which may strongly influence species coexistence. Nevertheless, current theory mainly assumes distinct and time‐invariant trait values. We examined the combined effects of the range and the speed of trait adaptation on species coexistence using an innovative multispecies predator–prey model. It allows for temporal trait changes of all predator and prey species and thus simultaneous coadaptation within and among trophic levels. We show that very small or slow trait adaptation did not facilitate coexistence because the stabilizing niche differences were not sufficient to offset the fitness differences. In contrast, sufficiently large and fast trait adaptation jointly promoted stable or neutrally stable species coexistence. Continuous trait adjustments in response to selection enabled a temporally variable convergence and divergence of species traits; that is, species became temporally more similar (neutral theory) or dissimilar (niche theory) depending on the selection pressure, resulting over time in a balance between niche differences stabilizing coexistence and fitness differences promoting competitive exclusion. Furthermore, coadaptation allowed prey and predator species to cluster into different functional groups. This equalized the fitness of similar species while maintaining sufficient niche differences among functionally different species delaying or preventing competitive exclusion. In contrast to previous studies, the emergent feedback between biomass and trait dynamics enabled supersaturated coexistence for a broad range of potential trait adaptation and parameters. We conclude that accounting for trait adaptation may explain stable and supersaturated species coexistence for a broad range of environmental conditions in natural systems when the absence of such adaptive changes would preclude it. Small trait changes, coincident with those that may occur within many natural populations, greatly enlarged the number of coexisting species.

Potential mechanisms of coexistence in closely related forbs

The stable coexistence of very similar species has perplexed ecologists for decades and has been central to the development of coexistence theory. According to modern coexistence theory, species can coexist stably (i.e. persist indefinitely with no long‐term density trends) as long as species' niche differences exceed competitive ability differences, even if these differences are very small. Recent studies have directly quantified niche and competitive ability differences in experimental communities at small spatial scales, but provide limited information about stable coexistence across spatial scales in heterogeneous natural communities. In this study, we use experimental and observational approaches to explore evidence for niche and competitive ability differences between two closely related, ecologically similar and widely coexisting annual forbs: Trachymene cyanopetala and T. ornata. We experimentally tested for stabilizing niche differences and competitive ability differences between these species by manipulating species' frequencies, under both well‐watered and water‐stressed conditions. We considered these experimental results in light of extensive field observations to explore evidence of niche segregation at a range of spatial scales. We found little evidence of intra‐specific stabilization or competitive ability differences in laboratory experiments while observational studies suggested niche segregation across pollinator assemblages and small‐scale microclimate heterogeneity. Though we did not quantify long‐term stabilization of coexisting populations of these species, results are consistent with expectations for stable coexistence of similar species via a spatial storage effect allowing niche differences to overcome even small (to absent) competitive ability differences.

Within-Host Niche Differences and Fitness Trade-offs Promote Coexistence of Plant Viruses.

Pathogens live in diverse, competitive communities, yet the processes that maintain pathogen diversity remain elusive. Here, we use a species-rich, well-studied plant virus system, the barley yellow dwarf viruses, to examine the mechanisms that regulate pathogen diversity. We empirically parameterized models of three viruses, their two aphid vectors, and one perennial grass host. We found that high densities of both aphids maximized virus diversity and that competition limited the coexistence of two closely related viruses. Even limited ability to simultaneously infect (coinfect) host individuals strongly promoted virus coexistence; preventing coinfection led to priority effects. Coinfection generated stabilizing niche differences by allowing viruses to share hosts. However, coexistence also required trade-offs between vector generalist and specialist life-history strategies. Our predicted outcomes broadly concur with previous field observations. These results show how competition within individual hosts and vectors may lead to unexpected population-level outcomes between pathogens, including coexistence, competitive exclusion, and priority effects, and how contemporary coexistence theory can help to predict these outcomes.

Large niche differences emerge at the recruitment stage to stabilize grassland coexistence

Niche differences and average fitness differences jointly determine coexistence. However, little empirical information about the magnitude of these two mechanisms is available. Using multispecies population models fit to long‐term demographic data for common, co‐occurring species in five grassland and shrubland plant communities in western North America, we estimated the strength of stabilizing niche differences and average fitness differences. In all five communities, both pairwise and full community comparisons showed evidence for strong stabilizing mechanisms and relatively small average fitness differences. For a total of 17 species pairs, a measure of niche differences based on simulations of invasion growth rates ranged from 0.59 to 0.93 with a mean of 0.81, where 0 indicates complete niche overlap and 1 indicates zero niche overlap. A corresponding measure of average fitness differences ranged from 1.02 to 2.54 with a mean of 1.53, where 1 indicates identical fitness and a value of 2 indicates a fourfold difference in sensitivity to competition. Comparisons of full communities displayed similar patterns: niche differences ranged from 0.58 to 0.69 with a mean of 0.64, and the average fitness differences ranged from 1.42 to 1.63 with a mean of 1.47. In almost every case, the stabilizing mechanisms were much stronger than minimally necessary to prevent competitive exclusion. Considering that all but one of the species we studied are perennial grasses, which are often grouped in the same functional type, the magnitude of these niche differences is surprising. In all five communities, differences between intra‐ and interspecific effects at the recruitment stage contributed far more to stabilization than interactions affecting growth and survival. Our results indicate that for these abundant, co‐occurring species (1) dynamics are far from neutral, with strong niche differences and weak fitness differences combining to stabilize coexistence, and (2) processes operating at early life stages account for a large proportion of the stabilizing effect. Given the limitations of our inductive approach, both these findings represent hypotheses in need of experimental testing.

Plant functional traits and the multidimensional nature of species coexistence.

Understanding the processes maintaining species diversity is a central problem in ecology, with implications for the conservation and management of ecosystems. Although biologists often assume that trait differences between competitors promote diversity, empirical evidence connecting functional traits to the niche differences that stabilize species coexistence is rare. Obtaining such evidence is critical because traits also underlie the average fitness differences driving competitive exclusion, and this complicates efforts to infer community dynamics from phenotypic patterns. We coupled field-parameterized mathematical models of competition between 102 pairs of annual plants with detailed sampling of leaf, seed, root, and whole-plant functional traits to relate phenotypic differences to stabilizing niche and average fitness differences. Single functional traits were often well correlated with average fitness differences between species, indicating that competitive dominance was associated with late phenology, deep rooting, and several other traits. In contrast, single functional traits were poorly correlated with the stabilizing niche differences that promote coexistence. Niche differences could only be described by combinations of traits, corresponding to differentiation between species in multiple ecological dimensions. In addition, several traits were associated with both fitness differences and stabilizing niche differences. These complex relationships between phenotypic differences and the dynamics of competing species argue against the simple use of single functional traits to infer community assembly processes but lay the groundwork for a theoretically justified trait-based community ecology.

Functional equivalence, competitive hierarchy and facilitation determine species coexistence in highly invaded grasslands.

Alien and native plant species often differ in functional traits. Trait differences could lead to niche differences that minimize competitive interactions and stabilize coexistence. However, trait differences could also translate into average fitness differences, leading to a competitive hierarchy that prevents coexistence. We tested whether trait differences between alien and native species translated into average fitness or stabilizing niche differences, and whether competition could explain observed coexistence within invaded grassland communities (New Zealand). Trait differences reflected marked competitive hierarchy, suggesting average fitness differences. Species coexistence was determined by a trade-off between species susceptibility to herbivory vs competitive hierarchy and facilitation. Importantly, although aliens and natives differed in their trait values, they did not differ in their competitive response, highlighting the importance of equalizing mechanisms in structuring invaded communities. Only a few alien species with a particular set of traits were able to jeopardize species coexistence when grazing was ceased. Our study explains why some alien species coexist with natives, whereas others have strongimpacts on native communities. It highlights that trait differences can underlie several coexistence processes and that the demonstration of trait differences between aliens and natives is only a first step to understanding the role of biotic interactions in structuring invaded communities.

Trait-mediated assembly processes predict successional changes in community diversity of tropical forests

Interspecific differences in relative fitness can cause local dominance by a single species. However, stabilizing interspecific niche differences can promote local diversity. Understanding these mechanisms requires that we simultaneously quantify their effects on demography and link these effects to community dynamics. Successional forests are ideal systems for testing assembly theory because they exhibit rapid community assembly. Here, we leverage functional trait and long-term demographic data to build spatially explicit models of successional community dynamics of lowland rainforests in Costa Rica. First, we ask what the effects and relative importance of four trait-mediated community assembly processes are on tree survival, a major component of fitness. We model trait correlations with relative fitness differences that are both density-independent and -dependent in addition to trait correlations with stabilizing niche differences. Second, we ask how the relative importance of these trait-mediated processes relates to successional changes in functional diversity. Tree dynamics were more strongly influenced by trait-related interspecific variation in average survival than trait-related responses to neighbors, with wood specific gravity (WSG) positively correlated with greater survival. Our findings also suggest that competition was mediated by stabilizing niche differences associated with specific leaf area (SLA) and leaf dry matter content (LDMC). These drivers of individual-level survival were reflected in successional shifts to higher SLA and LDMC diversity but lower WSG diversity. Our study makes significant advances to identifying the links between individual tree performance, species functional traits, and mechanisms of tropical forest succession.

Phenology effects on invasion success: insights from coupling field experiments to coexistence theory.

Ecologists have identified a growing number of functional traits that promote invasion. However, whether trait differences between exotic and native species promote invasion success by enhancing niche differences or giving invaders competitive advantages is poorly understood. We explored the mechanisms by which phenology determines invasion success in a California annual plant community by quantifying how the seasonal timing of growth relates to niche differences that stabilize coexistence, and the competitive ability differences that drive dominance and exclusion. We parameterized models of community dynamics from experimentally assembled annual communities in which exotic plants displayed earlier, coincident, or later phenology than native residents. Using recent theoretical advances from the coexistence literature, we found that differences in phenology promote stabilizing niche differences between exotic and native species. However, phenology was more strongly related to competitive ability differences, allowing later invaders to outcompete earlier native competitors and native residents to outcompete earlier invaders in field experiments. Few of these insights could be inferred by comparing the competitive outcomes across invaders, highlighting the need to quantify niche and competitive ability differences when disentangling how species differences drive invasion success.

Rethinking Community Assembly through the Lens of Coexistence Theory

Although research on the role of competitive interactions during community assembly began decades ago, a recent revival of interest has led to new discoveries and research opportunities. Using contemporary coexistence theory that emphasizes stabilizing niche differences and relative fitness differences, we evaluate three empirical approaches for studying community assembly. We show that experimental manipulations of the abiotic or biotic environment, assessments of trait-phylogeny-environment relationships, and investigations of frequency-dependent population growth all suggest strong influences of stabilizing niche differences and fitness differences on the outcome of plant community assembly. Nonetheless, due to the limitations of these approaches applied in isolation, we still have a poor understanding of which niche axes and which traits determine the outcome of competition and community structure. Combining current approaches represents our best chance of achieving this goal, which is fundamental to conceptual ecology and to the management of plant communities under global change.

Forecasting plant community impacts of climate variability and change: when do competitive interactions matter?

Summary1. A change in a climate variable may alter a species’ abundance not only through a direct effect on that species’ vital rates, but also through ‘indirect’ effects mediated by species interactions. While recent work has highlighted cases in which indirect effects overwhelm the direct effects of climate, we lack robust generalizations to predict the strength of indirect effects.2. For communities dominated by non‐trophic interactions, we propose that the potential for indirect effects of climate change declines with the strength of stabilizing niche differences.3. We tested this hypothesis by analysing an empirically parameterized four species population model. We quantified negative frequency dependence in population growth rates as a measure of stabilizing niche differences and projected the sensitivity of each species to direct and indirect effects of climate perturbations.4. Consistent with our hypothesis, species’ sensitivities to indirect effects decreased rapidly with increasing stabilization by niche differences.5. Synthesis. Information about niche differences can identify species sensitive to indirect effects of climate change and determine when multispecies forecasting approaches are necessary. However, practical application of this generalization will require methods to predict niche differences from easily collected data.