Schrödinger's Range-Shifting Cat: How Skewed Temperature Dependence Impacts Persistence with Climate Change.

AbstractThe majority of species display strongly asymmetric responses to climatic variables, yet most analytic models used to investigate how species will respond to climate change assume symmetric responses, with largely unknown consequences. Applying a known mapping of population dynamical equations onto corresponding well-studied problems from quantum mechanics, we extend analytical results to incorporate this asymmetry. We derive expressions in terms of parameters representing climate velocity, dispersal rate, maximum growth rate, niche width, high-frequency climate variability, and environmental performance curve skew for three key responses: (1) population persistence, (2) lag between range displacement and climate displacement, and (3) location of maximum population sensitivity. We find that asymmetry impacts these climate change responses, but surprisingly, under our model assumptions, the direction (i.e., warm skewed or cool skewed) of performance curve asymmetry does not strongly contribute to either persistence or lags. Conservation measures to support range-shifting populations may have most benefit near their environmental optimum or where the environmental dependence is shallow, irrespective of whether this is the leading or trailing edge. A metapopulation simulation corroborates our results. Our results shed fresh light on how key features of a species' environmental performance curve can impact its response to climate change.

Community composition exceeds area as a predictor of long-term conservation value.

Conserving biodiversity often requires deciding which sites to prioritise for protection. Predicting the impact of habitat loss is a major challenge, however, since impacts can be distant from the perturbation in both space and time. Here we study the long-term impacts of habitat loss in a mechanistic metacommunity model. We find that site area is a poor predictor of long-term, regional-scale extinctions following localised perturbation. Knowledge of the compositional distinctness (average between-site Bray-Curtis dissimilarity) of the removed community can markedly improve the prediction of impacts on regional assemblages, even when biotic responses play out at substantial spatial or temporal distance from the initial perturbation. Fitting the model to two empirical datasets, we show that this conclusions holds in the empirically relevant parameter range. Our results robustly demonstrate that site area alone is not sufficient to gauge conservation priorities; analysis of compositional distinctness permits improved prioritisation at low cost.

Evolution of prudent predation in complex food webs.

Prudent predators catch sufficient prey to sustain their populations but not as much as to undermine their populations' survival. The idea that predators evolve to be prudent has been dismissed in the 1970s, but the arguments invoked then are untenable in the light of modern evolution theory. The evolution of prudent predation has repeatedly been demonstrated in two-species predator-prey metacommunity models. However, the vigorous population fluctuations that these models predict are not widely observed. Here we show that in complex model food webs prudent predation evolves as a result of consumer-mediated ('apparent') competitive exclusion of resources, which disadvantages aggressive consumers and does not generate such fluctuations. We make testable predictions for empirical signatures of this mechanism and its outcomes. Then we discuss how these predictions are borne out across freshwater, marine and terrestrial ecosystems. Demonstrating explanatory power of evolved prudent predation well beyond the question of predator-prey coexistence, the predicted signatures explain unexpected declines of invasive alien species, the shape of stock-recruitment relations of fish, and the clearance rates of pelagic consumers across the latitudinal gradient and 15 orders of magnitude in body mass. Specific research to further test this theory is proposed.

Metacommunity-scale biodiversity regulation and the self-organised emergence of macroecological patterns.

There exist a number of key macroecological patterns whose ubiquity suggests that the spatio-temporal structure of ecological communities is governed by some universal mechanisms. The nature of these mechanisms, however, remains poorly understood. Here, we probe spatio-temporal patterns in species richness and community composition using a simple metacommunity assembly model. Despite making no a priori assumptions regarding biotic spatial structure or the distribution of biomass across species, model metacommunities self-organise to reproduce well-documented patterns including characteristic species abundance distributions, range size distributions and species area relations. Also in agreement with observations, species richness in our model attains an equilibrium despite continuous species turnover. Crucially, it is in the neighbourhood of the equilibrium that we observe the emergence of these key macroecological patterns. Biodiversity equilibria in models occur due to the onset of ecological structural instability, a population-dynamical mechanism. This strongly suggests a causal link between local community processes and macroecological phenomena.

Quantitative criteria for choosing targets and indicators for sustainable use of ecosystems.

Wide-ranging, indicator-based assessments of large, complex ecosystems are playing an increasing role in guiding environmental policy and management. An example is the EU's Marine Strategy Framework Directive, which requires Member States to take measures to reach "good environmental status" (GES) in European marine waters. However, formulation of indicator targets consistent with the Directive's high-level policy goal of sustainable use has proven challenging. We develop a specific, quantitative interpretation of the concepts of GES and sustainable use in terms of indicators and associated targets, by sharply distinguishing between current uses to satisfy current societal needs and preferences, and unknown future uses. We argue that consistent targets to safeguard future uses derive from a requirement that any environmental state indicator should recover within a defined time (e.g. 30 years) to its pressure-free range of variation when all pressures are hypothetically removed. Within these constraints, specific targets for current uses should be set. Routes to implementation of this proposal for indicators of fish-community size structure, population size of selected species, eutrophication, impacts of non-indigenous species, and genetic diversity are discussed. Important policy implications are that (a) indicator target ranges, which may be wider than natural ranges, systematically and rationally derive from our proposal; (b) because relevant state indicators tend to respond slowly, corresponding pressures should also be monitored and assessed;

Contrasting changes in the abundance and diversity of North American bird assemblages from 1971 to 2010.

Although it is generally recognized that global biodiversity is declining, few studies have examined long-term changes in multiple biodiversity dimensions simultaneously. In this study, we quantified and compared temporal changes in the abundance, taxonomic diversity, functional diversity, and phylogenetic diversity of bird assemblages, using roadside monitoring data of the North American Breeding Bird Survey from 1971 to 2010. We calculated 12 abundance and diversity metrics based on 5-year average abundances of 519 species for each of 768 monitoring routes. We did this for all bird species together as well as for four subgroups based on breeding habitat affinity (grassland, woodland, wetland, and shrubland breeders). The majority of the biodiversity metrics increased or remained constant over the study period, whereas the overall abundance of birds showed a pronounced decrease, primarily driven by declines of the most abundant species. These results highlight how stable or even increasing metrics of taxonomic, functional, or phylogenetic diversity may occur in parallel with substantial losses of individuals. We further found that patterns of change differed among the species subgroups, with both abundance and diversity increasing for woodland birds and decreasing for grassland breeders. The contrasting changes between abundance and diversity and among the breeding habitat groups underscore the relevance of a multifaceted approach to measuring biodiversity change. Our findings further stress the importance of monitoring the overall abundance of individuals in addition to metrics of taxonomic, functional, or phylogenetic diversity, thus confirming the importance of population abundance as an essential biodiversity variable.

Constructing random matrices to represent real ecosystems.

Models of complex systems with n components typically have order n(2) parameters because each component can potentially interact with every other. When it is impractical to measure these parameters, one may choose random parameter values and study the emergent statistical properties at the system level. Many influential results in theoretical ecology have been derived from two key assumptions: that species interact with random partners at random intensities and that intraspecific competition is comparable between species. Under these assumptions, community dynamics can be described by a community matrix that is often amenable to mathematical analysis. We combine empirical data with mathematical theory to show that both of these assumptions lead to results that must be interpreted with caution. We examine 21 empirically derived community matrices constructed using three established, independent methods. The empirically derived systems are more stable by orders of magnitude than results from random matrices. This consistent disparity is not explained by existing results on predator-prey interactions. We investigate the key properties of empirical community matrices that distinguish them from random matrices. We show that network topology is less important than the relationship between a species' trophic position within the food web and its interaction strengths. We identify key features of empirical networks that must be preserved if random matrix models are to capture the features of real ecosystems.

Slower but deeper community change: Intrinsic dynamics regulate anthropogenic impacts on species temporal turnover.

Understanding the mechanisms behind biodiversity dynamics is central to assessing and forecasting anthropogenic impacts on ecological communities. However, the manner in which external environmental drivers act in concert with intrinsic ecological processes to influence local temporal turnover is currently largely unexplored. Here, we determine how human impacts affect multiple metrics of bird community turnover to establish the ecological mechanisms behind compositional change. We used US Breeding Bird Survey data to calculate transect-level rates of three measures of temporal species turnover: (1) "short-term" (initial rate of decline of Sørensen similarity), (2) "long-term" (asymptotic Sørensen similarity), and (3) "throughput" (overall species accumulation rate from species-time relationship exponents) over 2692 transects across 27 regional habitat types. We then hierarchically fit linear models to estimate the effect of anthropogenic impact on these turnover metrics, using the Human Modification Index proxy, while accounting for observed species richness, the size of the species pool, and annual environmental variability. We found broadly consistent impacts of increased anthropogenic pressures across diverse habitat types. The Human Modification Index was associated with greater turnover at long timescales, but marginally slower short-term turnover. The species "throughput" (accumulation rate) was not notably influenced. Examining anthropogenic impacts on different aspects of species turnover in combination allows greater ecological insight. Observed human impacts on short-term turnover were the opposite of existing expectations and suggest humans are disrupting the background turnover of these systems, rather than simply driving rapid directed turnover. The increased long-term turnover without concurrent increases in species accumulation implies human impacts lead to shifts in species occurrence frequency rather than simply greater arrival of "new" species. These results highlight the role of intrinsic dynamics and caution against simple interpretations of increased species turnover as reflections of environmental change.

Steeper size spectra with decreasing phytoplankton biomass indicate strong trophic amplification and future fish declines.

Under climate change, model ensembles suggest that declines in phytoplankton biomass amplify into greater reductions at higher trophic levels, with serious implications for fisheries and carbon storage. However, the extent and mechanisms of this trophic amplification vary greatly among models, and validation is problematic. In situ size spectra offer a novel alternative, comparing biomass of small and larger organisms to quantify the net efficiency of energy transfer through natural food webs that are already challenged with multiple climate change stressors. Our global compilation of pelagic size spectrum slopes supports trophic amplification empirically, independently from model simulations. Thus, even a modest (16%) decline in phytoplankton this century would magnify into a 38% decline in supportable biomass of fish within the intensively-fished mid-latitude ocean. We also show that this amplification stems not from thermal controls on consumers, but mainly from temperature or nutrient controls that structure the phytoplankton baseline of the food web. The lack of evidence for direct thermal effects on size structure contrasts with most current thinking, based often on more acute stress experiments or shorter-timescale responses. Our synthesis of size spectra integrates these short-term dynamics, revealing the net efficiency of food webs acclimating and adapting to climatic stressors.

Are there species smaller than 1 mm?

The rapid advance in genetic sequencing technologies has provided an unprecedented amount of data on the biodiversity of meiofauna. It was hoped that these data would allow the identification and counting of species, distinguished as tight clusters of similar genomes. Surprisingly, this appears not to be the case. Here, we begin a theoretical discussion of this phenomenon, drawing on an individual-based ecological model to inform our arguments. The determining factor in the emergence (or not) of distinguishable genetic clusters in the model is the product of population size with mutation rate-a measure of the adaptability of the population as a whole. This result suggests that indeed one should not expect to observe clearly distinguishable species groupings in data gathered from ultrasequencing of meiofauna.

Phylogeny versus body size as determinants of food web structure.

Food webs are the complex networks of trophic interactions that stoke the metabolic fires of life. To understand what structures these interactions in natural communities, ecologists have developed simple models to capture their main architectural features. However, apparently realistic food webs can be generated by models invoking either predator-prey body-size hierarchies or evolutionary constraints as structuring mechanisms. As a result, this approach has not conclusively revealed which factors are the most important. Here we cut to the heart of this debate by directly comparing the influence of phylogeny and body size on food web architecture. Using data from 13 food webs compiled by direct observation, we confirm the importance of both factors. Nevertheless, phylogeny dominates in most networks. Moreover, path analysis reveals that the size-independent direct effect of phylogeny on trophic structure typically outweighs the indirect effect that could be captured by considering body size alone. Furthermore, the phylogenetic signal is asymmetric: closely related species overlap in their set of consumers far more than in their set of resources. This is at odds with several food web models, which take only the view-point of consumers when assigning interactions. The echo of evolutionary history clearly resonates through current food webs, with implications for our theoretical models and conservation priorities.

Spontaneous genetic clustering in populations of competing organisms.

We introduce and analyse an individual-based evolutionary model, in which a population of genetically diverse organisms compete with each other for limited resources. Through theoretical analysis and stochastic simulations, we show that the model exhibits a pattern-forming instability which is highly amplified by the effects of demographic noise, leading to the spontaneous formation of genotypic clusters. This mechanism supports the thesis that stochasticity has a central role in the formation and coherence of species.

No pervasive relationship between species size and local abundance trends.

Although there is some evidence that larger species could be more prone to population declines, the potential role of size traits in determining changes in community composition has been underexplored in global-scale analyses. Here, we combine a large cross-taxon assemblage time series database (BioTIME) with multiple trait databases to show that there is no clear correlation within communities between size traits and changes in abundance over time, suggesting that there is no consistent tendency for larger species to be doing proportionally better or worse than smaller species at local scales.

Universal power-law diet partitioning by marine fish and squid with surprising stability-diversity implications.

A central question in community ecology is how the number of trophic links relates to community species richness. For simple dynamical food-web models, link density (the ratio of links to species) is bounded from above as the number of species increases; but empirical data suggest that it increases without bounds. We found a new empirical upper bound on link density in large marine communities with emphasis on fish and squid, using novel methods that avoid known sources of bias in traditional approaches. Bounds are expressed in terms of the diet-partitioning function (DPF): the average number of resources contributing more than a fraction f to a consumer's diet, as a function of f. All observed DPF follow a functional form closely related to a power law, with power-law exponents independent of species richness at the measurement accuracy. Results imply universal upper bounds on link density across the oceans. However, the inherently scale-free nature of power-law diet partitioning suggests that the DPF itself is a better defined characterization of network structure than link density.

Rigorous conditions for food-web intervality in high-dimensional trophic niche spaces.

Food webs represent trophic (feeding) interactions in ecosystems. Since the late 1970s, it has been recognized that food-webs have a surprisingly close relationship to interval graphs. One interpretation of food-web intervality is that trophic niche space is low-dimensional, meaning that the trophic character of a species can be expressed by a single or at most a few quantitative traits. In a companion paper we demonstrated, by simulating a minimal food-web model, that food webs are also expected to be interval when niche-space is high-dimensional. Here we characterize the fundamental mechanisms underlying this phenomenon by proving a set of rigorous conditions for food-web intervality in high-dimensional niche spaces. Our results apply to a large class of food-web models, including the special case previously studied numerically.

Intrinsic ecological dynamics drive biodiversity turnover in model metacommunities.

Turnover of species composition through time is frequently observed in ecosystems. It is often interpreted as indicating the impact of changes in the environment. Continuous turnover due solely to ecological dynamics-species interactions and dispersal-is also known to be theoretically possible; however the prevalence of such autonomous turnover in natural communities remains unclear. Here we demonstrate that observed patterns of compositional turnover and other important macroecological phenomena can be reproduced in large spatially explicit model ecosystems, without external forcing such as environmental change or the invasion of new species into the model. We find that autonomous turnover is triggered by the onset of ecological structural instability-the mechanism that also limits local biodiversity. These results imply that the potential role of autonomous turnover as a widespread and important natural process is underappreciated, challenging assumptions implicit in many observation and management tools. Quantifying the baseline level of compositional change would greatly improve ecological status assessments.

Soil resource supply influences faunal size-specific distributions in natural food webs.

The large range of body-mass values of soil organisms provides a tool to assess the ecological organization of soil communities. The goal of this paper is to identify graphical and quantitative indicators of soil community composition and ecosystem functioning, and to illustrate their application to real soil food webs. The relationships between log-transformed mass and abundance of soil organisms in 20 Dutch meadows and heathlands were investigated. Using principles of allometry, maximal use can be made of ecological theory to build and explain food webs. The aggregate contribution of small invertebrates such as nematodes to the entire community is high under low soil phosphorus content and causes shifts in the mass-abundance relationships and in the trophic structures. We show for the first time that the average of the trophic link lengths is a reliable predictor for assessing soil fertility responses. Ordered trophic link pairs suggest a self-organizing structure of food webs according to resource availability and can predict environmental shifts in ecologically meaningful ways.

Dome patterns in pelagic size spectra reveal strong trophic cascades.

In ecological communities, especially the pelagic zones of aquatic ecosystems, certain body-size ranges are often over-represented compared to others. Community size spectra, the distributions of community biomass over the logarithmic body-mass axis, tend to exhibit regularly spaced local maxima, called "domes", separated by steep troughs. Contrasting established theory, we explain these dome patterns as manifestations of top-down trophic cascades along aquatic food chains. Compiling high quality size-spectrum data and comparing these with a size-spectrum model introduced in this study, we test this theory and develop a detailed picture of the mechanisms by which bottom-up and top-down effects interact to generate dome patterns. Results imply that strong top-down trophic cascades are common in freshwater communities, much more than hitherto demonstrated, and may arise in nutrient rich marine systems as well. Transferring insights from the general theory of non-linear pattern formation to domes patterns, we provide new interpretations of past lake-manipulation experiments.

Let's Train More Theoretical Ecologists - Here Is Why.

A tangled web of vicious circles, driven by cultural issues, has prevented ecology from growing strong theoretical roots. Now this hinders development of effective conservation policies. To overcome these barriers in view of urgent societal needs, we propose a global network of postgraduate theoretical training programs.