The fitness value of ecological information in a variable world.

Information processing is increasingly recognized as a fundamental component of life in variable environments, including the evolved use of environmental cues, biomolecular networks, and social learning. Despite this, ecology lacks a quantitative framework for understanding how population, community, and ecosystem dynamics depend on information processing. Here, we review the rationale and evidence for 'fitness value of information' (FVOI), and synthesize theoretical work in ecology, information theory, and probability behind this general mathematical framework. The FVOI quantifies how species' per capita population growth rates can depend on the use of information in their environment. FVOI is a breakthrough approach to linking information processing and ecological and evolutionary outcomes in a changing environment, addressing longstanding questions about how information mediates the effects of environmental change and species interactions.

Climate-driven range shifts reduce persistence of competitors in a perennial plant community.

Forecasting the impacts of climate change on species persistence in diverse natural communities requires a way to account for indirect effects mediated through species interactions. In particular, we expect species to experience major changes in competition as they track favorable climates. Here, we combine experimental data with a recently developed theoretical framework based on coexistence theory to measure the impact of climate-driven range shifts on alpine plant persistence under climate change. We transplanted three co-dominant alpine perennial species to five elevations, creating a maximum of 5°C increase in average growing-season temperature. We statistically modeled species' demographic rates in response to the environment and interpolated species' intrinsic ranges-the environmental mapping of reproduction in the absence of competition. We used low-density population growth rates-species' initial rate of invasion into an established community-as a metric of persistence. Further analysis of low-density growth rates (LGRs) allowed us to parse the direct impacts of climate change from indirect impacts mediated by shifting competition. Our models predict qualitatively different range shifts for each species based on the climate conditions under which growth rates are maximized and where they are zero. Overall, climate change is predicted to increase the intrinsic (competition free) growth rates of all species, as warmer and wetter conditions increase the favorability of alpine habitat. However, these benefits are entirely negated by increased competition arising from greater overlap between competitors in their intrinsic ranges. Species were highly dispersal limited, which can prevent species from tracking shifting intrinsic ranges by reducing population spread rates. Yet dispersal limitation also promoted species' persistence because it promotes persistence mechanisms. Our study demonstrates the complex pathways by which climate change impacts species' persistence by altering their competitive environment, and highlights how a persistence framework based on LGRs can help disentangle impacts.

Species persistence under climate change: a geographical scale coexistence problem.

Forecasting the impacts of climate change on biological diversity requires better ways to incorporate competitive interactions into predictions of species' range dynamics and persistence. This problem has been studied extensively in a different context by theoreticians evaluating the coexistence of species in spatially heterogeneous environments. Here, we show how spatial coexistence theory can be adapted to provide a mathematical framework for understanding species persistence in competitive communities under climate change. We first show how the spatial low-density growth rate provides the relevant metric of species persistence along a climate gradient. We then analyse a model of multiple migrating competitors to show how mechanisms contributing to low-density growth rates quantify the effect of different competitive processes on persistence, and how these processes change in strength with species' asynchronous migration under climate change. Finally, we outline the empirical utility of the framework, showing how the theory can scale up from local measurements of species performance and competitive interactions to range-scale metrics of persistence. Treating species' range dynamics as a geographical-scale coexistence problem presents its own set of challenges, but building from a well-established body of theory may greatly improve the predictability of species persistence in competitive communities under climate change.

Publisher correction: The spatial scales of species coexistence.

An error during production led to a truncation of the final two sentences in the abstract, which should have read 'In so doing, this framework substantially reframes current approaches to spatial community ecology. Quantifying the spatial scales of species coexistence will permit the next important advance in our understanding of the maintenance of diversity in nature, and should improve the contribution of community ecology to biodiversity conservation.' These have been corrected in all versions of the Perspective.

The spatial scales of species coexistence.

Understanding how species diversity is maintained is a foundational problem in ecology and an essential requirement for the discipline to be effective as an applied science. Ecologists' understanding of this problem has rapidly matured, but this has exposed profound uncertainty about the spatial scales required to maintain species diversity. Here we define and develop this frontier by proposing the coexistence-area relationship-a real relationship in nature that can be used to understand the determinants of the scale-dependence of diversity maintenance. The coexistence-area relationship motivates new empirical techniques for addressing important, unresolved problems about the influence of demographic stochasticity, environmental heterogeneity and dispersal on scale-dependent patterns of diversity. In so doing, this framework substantially reframes current approaches to spatial community ecology. Quantifying the spatial scales of species coexistence will permit the next important advance in our understanding of the maintenance of diversity in nature, and should improve the contribution of community ecology to biodiversity conservation.

Temporal coexistence mechanisms contribute to the latitudinal gradient in forest diversity.

The tropical forests of Borneo and Amazonia may each contain more tree species diversity in half a square kilometre than do all the temperate forests of Europe, North America, and Asia combined. Biologists have long been fascinated by this disparity, using it to investigate potential drivers of biodiversity. Latitudinal variation in many of these drivers is expected to create geographic differences in ecological and evolutionary processes, and evidence increasingly shows that tropical ecosystems have higher rates of diversification, clade origination, and clade dispersal. However, there is currently no evidence to link gradients in ecological processes within communities at a local scale directly to the geographic gradient in biodiversity. Here, we show geographic variation in the storage effect, an ecological mechanism that reduces the potential for competitive exclusion more strongly in the tropics than it does in temperate and boreal zones, decreasing the ratio of interspecific-to-intraspecific competition by 0.25% for each degree of latitude that an ecosystem is located closer to the Equator. Additionally, we find evidence that latitudinal variation in climate underpins these differences; longer growing seasons in the tropics reduce constraints on the seasonal timing of reproduction, permitting lower recruitment synchrony between species and thereby enhancing niche partitioning through the storage effect. Our results demonstrate that the strength of the storage effect, and therefore its impact on diversity within communities, varies latitudinally in association with climate. This finding highlights the importance of biotic interactions in shaping geographic diversity patterns, and emphasizes the need to understand the mechanisms underpinning ecological processes in greater detail than has previously been appreciated.

Limited Dispersal Drives Clustering and Reduces Coexistence by the Storage Effect.

Temporal variation can facilitate the coexistence of competitors through the temporal storage effect. However, this theoretical result was derived with the assumption that species have high dispersal rates. Here, I show that limited dispersal diminishes the storage effect in the classical lottery model. Populations become highly clustered during invasion, and population growth rates and extinction probabilities are functions of cluster size. I adopt the term "nucleation" from the physics literature to describe these characteristics. I developed approximations that incorporated nucleation to capture the spatiotemporal dynamics of the simulated model. Using analytical results from these approximations, I show that limited dispersal dampens asynchronous fluctuations in reproduction between species. This makes species appear to be more similar in their growth rate responses to the environment, thus reducing the potential for the storage effect. Theoretical results lead to simple rules relating average dispersal distances to relative reductions in potential coexistence. To demonstrate their use, I perform a preliminary analysis of two plant communities: tropical trees and desert annuals. In both communities, small-seeded species that disperse short distances on average have the strongest reductions in potential coexistence; species with wind- or animal-driven dispersal disperse farther distances, on average, and experience moderate or small reductions.

The rise of novelty in ecosystems.

Rapid and ongoing change creates novelty in ecosystems everywhere, both when comparing contemporary systems to their historical baselines, and predicted future systems to the present. However, the level of novelty varies greatly among places. Here we propose a formal and quantifiable definition of abiotic and biotic novelty in ecosystems, map abiotic novelty globally, and discuss the implications of novelty for the science of ecology and for biodiversity conservation. We define novelty as the degree of dissimilarity of a system, measured in one or more dimensions relative to a reference baseline, usually defined as either the present or a time window in the past. In this conceptualization, novelty varies in degree, it is multidimensional, can be measured, and requires a temporal and spatial reference. This definition moves beyond prior categorical definitions of novel ecosystems, and does not include human agency, self-perpetuation, or irreversibility as criteria. Our global assessment of novelty was based on abiotic factors (temperature, precipitation, and nitrogen deposition) plus human population, and shows that there are already large areas with high novelty today relative to the early 20th century, and that there will even be more such areas by 2050. Interestingly, the places that are most novel are often not the places where absolute changes are largest; highlighting that novelty is inherently different from change. For the ecological sciences, highly novel ecosystems present new opportunities to test ecological theories, but also challenge the predictive ability of ecological models and their validation. For biodiversity conservation, increasing novelty presents some opportunities, but largely challenges. Conservation action is necessary along the entire continuum of novelty, by redoubling efforts to protect areas where novelty is low, identifying conservation opportunities where novelty is high, developing flexible yet strong regulations and policies, and establishing long-term experiments to test management approaches. Meeting the challenge of novelty will require advances in the science of ecology, and new and creative. conservation approaches.

Coexistence in tropical forests through asynchronous variation in annual seed production.

The storage effect is a mechanism that can facilitate the coexistence of competing species through temporal fluctuations in reproductive output. Numerous natural systems have the prerequisites for the storage effect, yet it has rarely been quantitatively assessed. Here, we investigate the possible importance of the storage effect in explaining the coexistence of tree species in the diverse tropical forest on Barro Colorado Island, Panama. This tropical forest has been monitored for more than 20 years, and annual seed production is asynchronous among species, a primary requirement for the storage effect. We constructed a model of forest regeneration that includes species-specific recruitment through seed, sapling, and adult stages, and we parameterized the model using data for 28 species for which information is known about seedling germination and survival. Simulations of the model demonstrated that the storage effect alone can be a strong mechanism allowing long-term persistence of species. We also developed a metric to quantify the strength of the storage effect in a way comparable to classical resource partitioning. Applying this metric to seed production data from 108 species, the storage effect reduces the strength of pairwise interspecific competition to 11-43% of the strength of intraspecific competition, thereby demonstrating strong potential to facilitate coexistence. Finally, for a subset of 51 species whose phylogenetic relationships are known, we compared the strength of the storage effect between pairs of species to their phylogenetic similarity. The strength of the storage effect between closely related species was on average no different from distantly related species, implying that the storage effect can be important in promoting the coexistence of even closely related species.