Increases in the frequency, severity and duration of temperature extremes are anticipated in the near future. Although recent work suggests that changes in temperature variation will have disproportionately greater effects on species than changes to the mean, much of climate change research in ecology has focused on the impacts of mean temperature change. Here, we couple finegrained climate projections (2050-2059) to thermal performance data from 38 ectothermic invertebrate species and contrast projections with those of a simple model. We show that projections based on mean temperature change alone differ substantially from those incorporating changes to the variation, and to the mean and variation in concert. Although most species show increases in performance at greater mean temperatures, the effect of mean and variance change together yields a range of responses, with temperate species at greatest risk of performance declines. Our work highlights the importance of using fine-grained temporal data to incorporate the full extent of temperature variation when assessing and projecting performance.

A fundamental goal of ecology is to understand what controls the distribution and abundance of species. Both environmental niches and trade-offs among species in dispersal and competitive ability have traditionally been cited as determinants of plant community composition. More recently, neutral models have shown that communities of species with identical life-history characteristics and no adaptation to environmental niches can form spatial distribution patterns similar to those found in nature, so long as the species have a limited dispersal distance. If there is a strong correlation between geographic distance and change in environmental conditions, however, such spatial patterns can arise through either neutral or niche-based processes. To test these competing theories, we developed a sampling design that decoupled distance and environment in the understory plant communities of an old-growth, temperate forest. We found strong evidence of niche-structuring but almost no support for neutral predictions. Dispersal limitation acted in conjunction with environmental gradients to determine species' distributions, and both functional and phylogenetic constraints appear to contribute to the niche differentiation that structures community assembly. Our results indicate that testing a neutral hypothesis without accounting for environmental gradients will at best cause unexplained variation in plant distributions and may well provide misleading support for neutrality because of a correlation between geographic distance and environment. N eutral theory (1-3) demonstrates that the compounded effects of dispersal limitation, speciation, and the role of chance through time can cause ecologically identical species to form patterns of distribution and abundance similar to those found in nature. The assumption of ecological equivalence among species in neutral theory challenges contemporary views on the importance of evolution and ecological adaptation in determining patterns of distribution and abundance, and thus current approaches to species conservation (2, 4). Although some assumptions of neutral theory have been questioned (5, 6), the value of the theory lies in the degree to which it can predict or refute the importance of species-specific traits in determining distribution and abundance. Aspects of neutral theory can be tested by model fitting (6), but a stronger test lies in directly assessing the degree to which species distributions are explained by environmental heterogeneity versus neutral processes operating on natural landscapes (4, 6, 7).Because of the spatial effects of dispersal limitation, neutral theory predicts that the compositional similarity between plant communities will decrease as the distance between two points increases (3,8). In contrast, niche theory predicts that community composition will change as a result of species-specific differences in evolved adaptive responses along environmental gradients (9, 10). These hypotheses are hard to distinguish in natural ecosystems, because a change in envi...

Summary1. For plant invaders, being different is often equated with being successful, yet the mechanistic connection remains unclear. 2. Classic niche theory predicts that invaders with niches distinct from the native flora should coexist with little interaction with native species, yet such invaders often have substantial impacts. Meanwhile, invaders that overlap in niche space with native species should either be repelled or dominate, yet these invaders often naturalize with little effect. Such discrepancies between theory and observation raise questions about how species differences influence invader establishment and impact. 3. Here, we review these issues in light of recent work on coexistence theory, which shows how niche and fitness differences between natives and invaders interact to determine invasion outcomes. We show how successful invader establishment depends on either a fitness advantage or niche difference from resident species, but that only the former allows invaders to become dominant. 4. By identifying the role of niche and fitness differences in leading invasion hypotheses, we unify their predictions for invasion success while highlighting new approaches for evaluating the importance of species differences for invasion. 5. Synthesis. Situating the invasion process within a recent coexistence framework broadens our understanding of invasion mechanisms and more tightly links problems in invasion ecology with our more general understanding of community dynamics.

Changing temperature can substantially shift ecological communities by altering the strength and stability of trophic interactions. Because many ecological rates are constrained by temperature, new approaches are required to understand how simultaneous changes in multiple rates alter the relative performance of species and their trophic interactions. We develop an energetic approach to identify the relationship between biomass fluxes and standing biomass across trophic levels. Our approach links ecological rates and trophic dynamics to measure temperature-dependent changes to the strength of trophic interactions and determine how these changes alter food web stability. It accomplishes this by using biomass as a common energetic currency and isolating three temperature-dependent processes that are common to all consumer-resource interactions: biomass accumulation of the resource, resource consumption and consumer mortality. Using this framework, we clarify when and how temperature alters consumer to resource biomass ratios, equilibrium resilience, consumer variability, extinction risk and transient vs. equilibrium dynamics. Finally, we characterise key asymmetries in species responses to temperature that produce these distinct dynamic behaviours and identify when they are likely to emerge. Overall, our framework provides a mechanistic and more unified understanding of the temperature dependence of trophic dynamics in terms of ecological rates, biomass ratios and stability.

1. Statistical tests partitioning community variation into environmental and spatial components have been widely used to test ecological theories and explore the determinants of community structure for applied conservation questions. Despite the wide use of these tests, there is considerable debate about their relative effectiveness. 2. We used simulated communities to evaluate the most commonly employed tests that partition community variation: regression on distance matrices and canonical ordination using a third-order polynomial, principal components of neighbour matrices (PCNM) or Moran's eigenvector maps (MEM) to model spatial components. Each test was evaluated under a variety of realistic sampling scenarios. 3. All tests failed to correctly model spatial and environmental components of variation, and in some cases produced biased estimates of the relative importance of components. Regression on distance matrices under-fit the spatial component, and ordination models consistently under-fit the environmental component. The PCNM and MEM approaches often produced inflated R 2 statistics, apparently as a result of statistical artefacts involving selection of superfluous axes. This problem occurred regardless of the forward-selection technique used. 4. Both sample configuration and the underlying linear model used to analyse species-environment relationships also revealed strong potential to bias results. 5. Synthesis and applications. Several common applications of variation partitioning in ecology now appear inappropriate. These potentially include decisions for community conservation based on inferred relative strengths of niche and dispersal processes, inferred community responses to climate change, and numerous additional analyses that depend on precise results from multivariate variation-partitioning techniques. We clarify the appropriate uses of these analyses in research programmes, and outline potential steps to improve them.