Search

Biodiversity can stabilize ecological communities through biological insurance, but climate and other environmental changes may disrupt this process via simultaneous ecosystem destabilization and biodiversity loss. While changes to diversity-stability relationships (DSRs) and the underlying mechanisms have been extensively explored in terrestrial plant communities, this topic remains largely unexplored in benthic marine ecosystems that comprise diverse assemblages of producers and consumers. By analyzing two decades of kelp forest biodiversity survey data, we discovered changes in diversity, stability, and their relationships at multiple scales (biological organizational levels, spatial scales, and functional groups) that were linked with the most severe marine heatwave ever documented in the North Pacific Ocean. Moreover, changes in the strength of DSRs during/after the heatwave were more apparent among functional groups than both biological organizational levels (population vs. ecosystem levels) and spatial scales (local vs. broad scales). Specifically, the strength of DSRs decreased for fishes, increased for mobile invertebrates and understory algae, and were unchanged for sessile invertebrates during/after the heatwave. Our findings suggest that biodiversity plays a key role in stabilizing marine ecosystems, but the resilience of DSRs to adverse climate impacts primarily depends on the functional identities of ecological communities., (© 2024 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Spatial synchrony is the tendency for population fluctuations to be correlated among different locations. This phenomenon is a ubiquitous feature of population dynamics and is important for ecosystem stability, but several aspects of synchrony remain unresolved. In particular, the extent to which any particular mechanism, such as dispersal, contributes to observed synchrony in natural populations has been difficult to determine. To address this gap, we leveraged recent methodological improvements to determine how dispersal structures synchrony in giant kelp (Macrocystis pyrifera), a global marine foundation species that has served as a useful system for understanding synchrony. We quantified population synchrony and fecundity with satellite imagery across 11 years and 880 km of coastline in southern California, USA, and estimated propagule dispersal probabilities using a high-resolution ocean circulation model. Using matrix regression models that control for the influence of geographic distance, resources (seawater nitrate), and disturbance (destructive waves), we discovered that dispersal was an important driver of synchrony. Our findings were robust to assumptions about propagule mortality during dispersal and consistent between two metrics of dispersal: (1) the individual probability of dispersal and (2) estimates of demographic connectivity that incorporate fecundity (the number of propagules dispersing). We also found that dispersal and environmental conditions resulted in geographic clusters with distinct patterns of synchrony. This study is among the few to statistically associate synchrony with dispersal in a natural population and the first to do so in a marine organism. The synchronizing effects of dispersal and environmental conditions on foundation species, such as giant kelp, likely have cascading effects on the spatial stability of biodiversity and ecosystem function., (© 2024 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Cross-ecosystem subsidies are critical to ecosystem structure and function, especially in recipient ecosystems where they are the primary source of organic matter to the food web. Subsidies are indicative of processes connecting ecosystems and can couple ecological dynamics across system boundaries. However, the degree to which such flows can induce cross-ecosystem cascades of spatial synchrony, the tendency for system fluctuations to be correlated across locations, is not well understood. Synchrony has destabilizing effects on ecosystems, adding to the importance of understanding spatiotemporal patterns of synchrony transmission. In order to understand whether and how spatial synchrony cascades across the marine-terrestrial boundary via resource subsidies, we studied the relationship between giant kelp forests on rocky nearshore reefs and sandy beach ecosystems that receive resource subsidies in the form of kelp wrack (detritus). We found that synchrony cascades from rocky reefs to sandy beaches, with spatiotemporal patterns mediated by fluctuations in live kelp biomass, wave action, and beach width. Moreover, wrack deposition synchronized local abundances of shorebirds that move among beaches seeking to forage on wrack-associated invertebrates, demonstrating that synchrony due to subsidies propagates across trophic levels in the recipient ecosystem. Synchronizing resource subsidies likely play an underappreciated role in the spatiotemporal structure, functioning, and stability of ecosystems., Competing Interests: Competing interests statement:The authors declare no competing interest.

Species coexistence attracts wide interest in ecology. Modern coexistence theory (MCT) identifies coexistence mechanisms, one of which, storage effects, hinges on relationships between fluctuations in environmental and competitive pressures. However, such relationships are typically measured using covariance, which does not account for the possibility that environment and competition may be more related to each other when they are strong than when weak, or vice versa. Recent work showed that such 'asymmetric tail associations' (ATAs) are common between ecological variables, and are important for extinction risk, ecosystem stability, and other phenomena. We extend MCT, decomposing storage effects to show the influence of ATAs. Analysis of a simple model and an empirical example using diatoms illustrate that ATA influences can be comparable in magnitude to other mechanisms of coexistence and that ATAs can make the difference between species coexistence and competitive exclusion. ATA influences may be an important new mechanism of coexistence., (© 2023 John Wiley & Sons Ltd.)

AbstractPopulation spatial synchrony-the tendency for temporal population fluctuations to be correlated across locations-is common and important to metapopulation stability and persistence. One common cause of spatial synchrony, termed the Moran effect, occurs when populations respond to environmental fluctuations, such as weather, that are correlated over space. Although the degree of spatial synchrony in environmental fluctuations can differ between seasons and different population processes occur in different seasons, the impact on population spatial synchrony is uncertain because prior work has largely assumed that the spatial synchrony of environmental fluctuations and their effect on populations are consistent over annual sampling intervals. We used theoretical models to examine how seasonality in population processes and the spatial synchrony of environmental drivers affect population spatial synchrony. We found that population spatial synchrony can depend not only on the spatial synchrony of environmental drivers but also on the degree to which environmental fluctuations are correlated across seasons, locally, and across space. Moreover, measurements of synchrony from "snapshot" population censuses may not accurately reflect synchrony during other parts of the year. Together, these results show that neglecting seasonality in environmental conditions and population processes is consequential for understanding population spatial synchrony and its driving mechanisms.

Understanding the relationship between biodiversity and ecosystem stability is a central goal of ecologists. Recent studies have concluded that biodiversity increases community temporal stability by increasing the asynchrony between the dynamics of different species. Theoretically, this enhancement can occur through either increased between-species compensatory dynamics, a fundamentally biological mechanism; or through an averaging effect, primarily a statistical mechanism. Yet it remains unclear which mechanism is dominant in explaining the diversity-stability relationship. We address this issue by mathematically decomposing asynchrony into components separately quantifying the compensatory and statistical-averaging effects. We applied the new decomposition approach to plant survey and experimental data from North American grasslands. We show that statistical averaging, rather than compensatory dynamics, was the principal mediator of biodiversity effects on community stability. Our simple decomposition approach helps integrate concepts of stability, asynchrony, statistical averaging, and compensatory dynamics, and suggests that statistical averaging, rather than compensatory dynamics, is the primary means by which biodiversity confers ecological stability., (© 2022. The Author(s).)

Synchronous dynamics (fluctuations that occur in unison) are universal phenomena with widespread implications for ecological stability. Synchronous dynamics can amplify the destabilizing effect of environmental variability on ecosystem functions such as productivity, whereas the inverse, compensatory dynamics, can stabilize function. Here we combine simulation and empirical analyses to elucidate mechanisms that underlie patterns of synchronous versus compensatory dynamics. In both simulated and empirical communities, we show that synchronous and compensatory dynamics are not mutually exclusive but instead can vary by timescale. Our simulations identify multiple mechanisms that can generate timescale-specific patterns, including different environmental drivers, diverse life histories, dispersal, and non-stationary dynamics. We find that traditional metrics for quantifying synchronous dynamics are often biased toward long-term drivers and may miss the importance of short-term drivers. Our findings indicate key mechanisms to consider when assessing synchronous versus compensatory dynamics and our approach provides a pathway for disentangling these dynamics in natural systems., (© 2022 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

AbstractFunctionally variable symbionts commonly co-occur including within the roots of individual plants, in spite of arguments from simple models of the stability of mutualism that predict competitive exclusion among symbionts. We explore this paradox by evaluating the dynamics generated by symbiont competition for plant resources and the plant's preferential allocation to the most beneficial symbiont using a system of differential equations representing the densities of mutualistic and nonmutualistic symbionts and the level of preferentially allocated and nonpreferentially allocated resources for which the symbionts compete. We find that host preferential allocation and costs of mutualism generate resource specialization that makes the coexistence of beneficial and nonbeneficial symbionts possible. Furthermore, coexistence becomes likely because of negative physiological feedbacks in host preferential allocation. We find that biologically realistic models of plant physiology and symbiont competition predict that the coexistence of beneficial and nonbeneficial symbionts should be common in root symbioses and that the density and relative abundance of mutualists should increase in proportion to the needs of the host.

Synchrony is broadly important to population and community dynamics due to its ubiquity and implications for extinction dynamics, system stability, and species diversity. Investigations of synchrony in community ecology have tended to focus on covariance in the abundances of multiple species in a single location. Yet, the importance of regional environmental variation and spatial processes in community dynamics suggests that community properties, such as species richness, could fluctuate synchronously across patches in a metacommunity, in an analog of population spatial synchrony. Here, we test the prevalence of this phenomenon and the conditions under which it may occur using theoretical simulations and empirical data from 20 marine and terrestrial metacommunities. Additionally, given the importance of biodiversity for stability of ecosystem function, we posit that spatial synchrony in species richness is strongly related to stability. Our findings show that metacommunities often exhibit spatial synchrony in species richness. We also found that richness synchrony can be driven by environmental stochasticity and dispersal, two mechanisms of population spatial synchrony. Richness synchrony also depended on community structure, including species evenness and beta diversity. Strikingly, ecosystem stability was more strongly related to richness synchrony than to species richness itself, likely because richness synchrony integrates information about community processes and environmental forcing. Our study highlights a new approach for studying spatiotemporal community dynamics and emphasizes the spatial dimensions of community dynamics and stability., (© 2021 The Authors. Ecology published by Wiley Periodicals LLC on behalf of Ecological Society of America.)

Synchrony among population fluctuations of multiple coexisting species has a major impact on community stability, i.e. on the relative temporal constancy of aggregate properties such as total community biomass. However, synchrony and its impacts are usually measured using covariance methods, which do not account for whether species abundances may be more correlated when species are relatively common than when they are scarce, or vice versa. Recent work showed that species commonly exhibit such 'asymmetric tail associations'. We here consider the influence of asymmetric tail associations on community stability. We develop a 'skewness ratio' which quantifies how much species relationships and tail associations modify stability. The skewness ratio complements the classic variance ratio and related metrics. Using multi-decadal grassland datasets, we show that accounting for tail associations gives new viewpoints on synchrony and stability; e.g. species associations can alter community stability differentially for community crashes or explosions to high values, a fact not previously detectable. Species associations can mitigate explosions of community abundance to high values, increasing one aspect of stability, while simultaneously exacerbating crashes to low values, decreasing another aspect of stability; or vice versa. Our work initiates a new, more flexible paradigm for exploring species relationships and community stability. This article is part of the theme issue 'Synchrony and rhythm interaction: from the brain to behavioural ecology'.

Periodical cicadas exhibit an extraordinary capacity for self-organizing spatially synchronous breeding behavior. The regular emergence of periodical cicada broods across the United States is a phenomenon of longstanding public and scientific interest, as the cicadas of each brood emerge in huge numbers and briefly dominate their ecosystem. During the emergence, the 17-year periodical cicada species Magicicada cassini is found to form synchronized choruses, and we investigated their chorusing behavior from the standpoint of spatial synchrony.Cicada choruses were observed to form in trees, calling regularly every five seconds. In order to determine the limits of this self-organizing behavior, we set out to quantify the spatial synchronization between cicada call choruses in different trees, and how and why this varies in space and time.We performed 20 simultaneous recordings in Clinton State Park, Kansas, in June 2015 (Brood IV), with a team of citizen-science volunteers using consumer equipment (smartphones). We use a wavelet approach to show in detail how spatially synchronous, self-organized chorusing varies across the forest.We show how conditions that increase the strength of audio interactions between cicadas also increase the spatial synchrony of their chorusing. Higher forest canopy light levels increase cicada activity, corresponding to faster and higher-amplitude chorus cycling and to greater synchrony of cycles across space. We implemented a relaxation-oscillator-ensemble model of interacting cicadas, finding that a tendency to call more often, driven by light levels, results in all these effects.Results demonstrate how the capacity to self-organize in ecology depends sensitively on environmental conditions. Spatially correlated modulation of cycling rate by an external driver can also promote self-organization of phase synchrony., Competing Interests: The authors declare no conflict of interest., (© 2020 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Rift Valley fever virus (RVFV) is a mosquito-borne zoonotic arbovirus with important livestock and human health, and economic consequences across Africa and the Arabian Peninsula. Climate and vegetation monitoring guide RVFV forecasting models and early warning systems; however, these approaches make monthly predictions and a need exists to predict primary vector abundances at finer temporal scales. In Kenya, an important primary RVFV vector is the mosquito Aedes mcintoshi. We used a zero-inflated negative binomial regression and multimodel averaging approach with georeferenced Ae. mcintoshi mosquito counts and remotely sensed climate and topographic variables to predict where and when abundances would be high in Kenya and western Somalia. The data supported a positive effect on abundance of minimum wetness index values within 500 m of a sampling site, cumulative precipitation values 0 to 14 days prior to sampling, and elevated land surface temperature values ~3 weeks prior to sampling. The probability of structural zero counts of mosquitoes increased as percentage clay in the soil decreased. Weekly retrospective predictions for unsampled locations across the study area between 1 September and 25 January from 2002 to 2016 predicted high abundances prior to RVFV outbreaks in multiple foci during the 2006-2007 epizootic, except for two districts in Kenya. Additionally, model predictions supported the possibility of high Ae. mcintoshi abundances in Somalia, independent of Kenya. Model-predicted abundances were low during the 2015-2016 period when documented outbreaks did not occur, although several surveillance systems issued warnings. Model predictions prior to the 2018 RVFV outbreak indicated elevated abundances in Wajir County, Kenya, along the border with Somalia, but RVFV activity occurred west of the focus of predicted high Ae. mcintoshi abundances., Competing Interests: The authors have declared that no competing interests exist.

Large-scale spatial synchrony is ubiquitous in ecology. We examined 56 years of data representing chlorophyll density in 26 areas in British seas monitored by the Continuous Plankton Recorder survey. We used wavelet methods to disaggregate synchronous fluctuations by timescale and determine that drivers of synchrony include both biotic and abiotic variables. We tested these drivers for statistical significance by comparison with spatially synchronous surrogate data. Identification of causes of synchrony is distinct from, and goes beyond, determining drivers of local population dynamics. We generated timescale-specific models, accounting for 61% of long-timescale (> 4yrs) synchrony in a chlorophyll density index, but only 3% of observed short-timescale (< 4yrs) synchrony. Thus synchrony and its causes are timescale-specific. The dominant source of long-timescale chlorophyll synchrony was closely related to sea surface temperature, through a climatic Moran effect, though likely via complex oceanographic mechanisms. The top-down action of Calanus finmarchicus predation enhances this environmental synchronising mechanism and interacts with it non-additively to produce more long-timescale synchrony than top-down and climatic drivers would produce independently. Our principal result is therefore a demonstration of interaction effects between Moran drivers of synchrony, a new mechanism for synchrony that may influence many ecosystems at large spatial scales., Competing Interests: The authors have declared that no competing interests exist.

Taylor's law (TL), a commonly observed and applied pattern in ecology, describes variances of population densities as related to mean densities via log(variance) = log(a) + b*log(mean). Variations among datasets in the slope, b, have been associated with multiple factors of central importance in ecology, including strength of competitive interactions and demographic rates. But these associations are not transparent, and the relative importance of these and other factors for TL slope variation is poorly studied. TL is thus a ubiquitously used indicator in ecology, the understanding of which is still opaque. The goal of this study was to provide tools to help fill this gap in understanding by providing proximate determinants of TL slopes, statistical quantities that are correlated to TL slopes but are simpler than the slope itself and are more readily linked to ecological factors. Using numeric simulations and 82 multi-decadal population datasets, we here propose, test and apply two proximate statistical determinants of TL slopes which we argue can become key tools for understanding the nature and ecological causes of TL slope variation. We find that measures based on population skewness, coefficient of variation and synchrony are effective proximate determinants. We demonstrate their potential for application by using them to help explain covariation in slopes of spatial and temporal TL (two common types of TL). This study provides tools for understanding TL, and demonstrates their usefulness., (© 2018 The Authors. Journal of Animal Ecology © 2018 British Ecological Society.)

Spatial synchrony, defined as correlated temporal fluctuations among populations, is a fundamental feature of population dynamics, but many aspects of synchrony remain poorly understood. Few studies have examined detailed geographical patterns of synchrony; instead most focus on how synchrony declines with increasing linear distance between locations, making the simplifying assumption that distance decay is isotropic. By synthesising and extending prior work, we show how geography of synchrony, a term which we use to refer to detailed spatial variation in patterns of synchrony, can be leveraged to understand ecological processes including identification of drivers of synchrony, a long-standing challenge. We focus on three main objectives: (1) showing conceptually and theoretically four mechanisms that can generate geographies of synchrony; (2) documenting complex and pronounced geographies of synchrony in two important study systems; and (3) demonstrating a variety of methods capable of revealing the geography of synchrony and, through it, underlying organism ecology. For example, we introduce a new type of network, the synchrony network, the structure of which provides ecological insight. By documenting the importance of geographies of synchrony, advancing conceptual frameworks, and demonstrating powerful methods, we aim to help elevate the geography of synchrony into a mainstream area of study and application., (© 2017 John Wiley & Sons Ltd/CNRS.)

Taylor's law (TL) is a widely observed empirical pattern that relates the variances to the means of groups of nonnegative measurements via an approximate power law: variance g ≈ a [Formula: see text] mean g b , where g indexes the group of measurements. When each group of measurements is distributed in space, the exponent b of this power law is conjectured to reflect aggregation in the spatial distribution. TL has had practical application in many areas since its initial demonstrations for the population density of spatially distributed species in population ecology. Another widely observed aspect of populations is spatial synchrony, which is the tendency for time series of population densities measured in different locations to be correlated through time. Recent studies showed that patterns of population synchrony are changing, possibly as a consequence of climate change. We use mathematical, numerical, and empirical approaches to show that synchrony affects the validity and parameters of TL. Greater synchrony typically decreases the exponent b of TL. Synchrony influenced TL in essentially all of our analytic, numerical, randomization-based, and empirical examples. Given the near ubiquity of synchrony in nature, it seems likely that synchrony influences the exponent of TL widely in ecologically and economically important systems., Competing Interests: The authors declare no conflict of interest.

Multidimensional data are integral to many community-ecological studies and come in various forms, such as stable isotopes, compound specific analyses (e.g., amino acids and fatty acids), and both biodiversity and life history traits. Scientists employing such data often lack standardized metrics to evaluate communities in niche space where more than 2 dimensions are involved. To alleviate this problem, we developed a graphing and analytical approach for use with more than two variables, based on previously established stable isotope bi-plot metrics. We introduce here our community metrics as R scripts. By extending the original metrics to multiple dimensions, we created n-dimensional plots and metrics to characterize any set of quantitative measurements of a community. We demonstrate the utility of these metrics using stable isotope data; however, the approaches are applicable to many types of data. The resulting metrics provide more and better information compared to traditional analytic frameworks. The approach can be applied in many branches of community ecology, and it offers accessible metrics to quantitatively analyze the structure of communities across ecosystems and through time., Competing Interests: The authors declare no competing financial interests.

During the 1980s, the North Sea plankton community underwent a well-documented ecosystem regime shift, including both spatial changes (northward species range shifts) and temporal changes (increases in the total abundances of warmer water species). This regime shift has been attributed to climate change. Plankton provide a link between climate and higher trophic-level organisms, which can forage on large spatial and temporal scales. It is therefore important to understand not only whether climate change affects purely spatial or temporal aspects of plankton dynamics, but also whether it affects spatiotemporal aspects such as metapopulation synchrony. If plankton synchrony is altered, higher trophic-level feeding patterns may be modified. A second motivation for investigating changes in synchrony is that the possibility of such alterations has been examined for few organisms, in spite of the fact that synchrony is ubiquitous and of major importance in ecology. This study uses correlation coefficients and spectral analysis to investigate whether synchrony changed between the periods 1959-1980 and 1989-2010. Twenty-three plankton taxa, sea surface temperature (SST), and wind speed were examined. Results revealed that synchrony in SST and plankton was altered. Changes were idiosyncratic, and were not explained by changes in abundance. Changes in the synchrony of Calanus helgolandicus and Para-pseudocalanus spp appeared to be driven by changes in SST synchrony. This study is one of few to document alterations of synchrony and climate-change impacts on synchrony. We discuss why climate-change impacts on synchrony may well be more common and consequential than previously recognized., (© 2016 The Authors. Global Change Biology Published by John Wiley & Sons Ltd.)

Phytoplankton are key components of aquatic ecosystems, fixing CO2 from the atmosphere through photosynthesis and supporting secondary production, yet relatively little is known about how future global warming might alter their biodiversity and associated ecosystem functioning. Here, we explore how the structure, function, and biodiversity of a planktonic metacommunity was altered after five years of experimental warming. Our outdoor mesocosm experiment was open to natural dispersal from the regional species pool, allowing us to explore the effects of experimental warming in the context of metacommunity dynamics. Warming of 4°C led to a 67% increase in the species richness of the phytoplankton, more evenly-distributed abundance, and higher rates of gross primary productivity. Warming elevated productivity indirectly, by increasing the biodiversity and biomass of the local phytoplankton communities. Warming also systematically shifted the taxonomic and functional trait composition of the phytoplankton, favoring large, colonial, inedible phytoplankton taxa, suggesting stronger top-down control, mediated by zooplankton grazing played an important role. Overall, our findings suggest that temperature can modulate species coexistence, and through such mechanisms, global warming could, in some cases, increase the species richness and productivity of phytoplankton communities.

Antibiotics leak constantly into environments due to widespread use in agriculture and human therapy. Although sublethal concentrations are well known to select for antibiotic-resistant bacteria, little is known about how bacterial evolution cascades through food webs, having indirect effect on species not directly affected by antibiotics (e.g. via population dynamics or pleiotropic effects). Here, we used an experimental evolution approach to test how temporal patterns of antibiotic stress, as well as migration within metapopulations, affect the evolution and ecology of microcosms containing one prey bacterium, one phage and two protist predators. We found that environmental variability, autocorrelation and migration had only subtle effects for population and evolutionary dynamics. However, unexpectedly, bacteria evolved greatest fitness increases to both antibiotics and enemies when the sublethal levels of antibiotics were highest, indicating positive pleiotropy. Crucially, bacterial adaptation cascaded through the food web leading to reduced predator-to-prey abundance ratio, lowered predator community diversity and increased instability of populations. Our results show that the presence of natural enemies can modify and even reverse the effects of antibiotics on bacteria, and that antibiotic selection can change the ecological properties of multitrophic microbial communities by having indirect effects on species not directly affected by antibiotics., (© 2015 The Author(s) Published by the Royal Society. All rights reserved.)

Distributions of species body sizes within a taxonomic group, for example, mammals, are widely studied and important because they help illuminate the evolutionary processes that produced these distributions. Distributions of the sizes of species within an assemblage delineated by geography instead of taxonomy (all the species in a region regardless of clade) are much less studied but are equally important and will illuminate a different set of ecological and evolutionary processes. We develop and test a mechanistic model of how diversity varies with body mass in marine ecosystems. The model predicts the form of the 'diversity spectrum', which quantifies the distribution of species' asymptotic body masses, is a species analogue of the classic size spectrum of individuals, and which we have found to be a new and widely applicable description of diversity patterns. The marine diversity spectrum is predicted to be approximately linear across an asymptotic mass range spanning seven orders of magnitude. Slope -0.5 is predicted for the global marine diversity spectrum for all combined pelagic zones of continental shelf seas, and slopes for large regions are predicted to lie between -0.5 and -0.1. Slopes of -0.5 and -0.1 represent markedly different communities: a slope of -0.5 depicts a 10-fold reduction in diversity for every 100-fold increase in asymptotic mass; a slope of -0.1 depicts a 1.6-fold reduction. Steeper slopes are predicted for larger or colder regions, meaning fewer large species per small species for such regions. Predictions were largely validated by a global empirical analysis. Results explain for the first time a new and widespread phenomenon of biodiversity. Results have implications for estimating numbers of species of small asymptotic mass, where taxonomic inventories are far from complete. Results show that the relationship between diversity and body mass can be explained from the dependence of predation behaviour, dispersal, and life history on body mass, and a neutral assumption about speciation and extinction., (© 2014 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Temperature is a key driver of ecological processes and patterns. The ramifications of temperature for ecological communities include not only its direct effects on the physiology of individuals, but also how these effects play out in the context of other processes such as competition. Apparently idiosyncratic or difficult to predict effects of temperature on competitive outcomes are well represented in the literature. General theoretical understanding of how physiological influences of temperature filter through community dynamics to determine outcomes is limited. We present a theoretical framework for predicting the effects of temperature on competition among species, based on understanding the effects of temperature on the physiological and population parameters of the species. The approach helps unify formal resource competition theory with metabolic and physiological ecology. Phytoplankton and many other ectotherms are smaller at higher temperatures. This has been observed experimentally, across geographical gradients, and as change accompanying climate warming, but it has not been explained in terms of competition. As a case study, we apply our theoretical framework to competition for nutrients among differently sized phytoplankton. Based on this analysis, we hypothesize that the prevalence of smaller phytoplankton at higher temperatures is at least partly due to an accentuated competitive advantage of smaller cells at higher temperatures with respect to nutrient uptake and growth. We examine the scope for extending the approach to understand resource competition, generally, among ectotherms of different sizes., (© 2013 The Authors. Journal of Animal Ecology © 2013 British Ecological Society.)

Climate warming has been linked to an apparent general decrease in body sizes of ectotherms, both across and within taxa, especially in aquatic systems. Smaller body size in warmer geographical regions has also been widely observed. Since body size is a fundamental determinant of many biological attributes, climate-warming-related changes in size could ripple across multiple levels of ecological organization. Some recent studies have questioned the ubiquity of temperature-size rules, however, and certain widespread and abundant taxa, such as diatoms, may be important exceptions. We tested the hypothesis that diatoms are smaller at warmer temperatures using a system of geothermally heated streams. There was no consistent relationship between size and temperature at either the population or community level. These field data provide important counterexamples to both James' and Bergmann's temperature-size rules, respectively, undermining the widely held assumption that warming favours the small. This study provides compelling new evidence that diatoms are an important exception to temperature-size rules for three reasons: (i) we use many more species than prior work; (ii) we examine both community and species levels of organization simultaneously; (iii) we work in a natural system with a wide temperature gradient but minimal variation in other factors, to achieve robust tests of hypotheses without relying on laboratory setups, which have limited realism. In addition, we show that interspecific effects were a bigger contributor to whole-community size differences, and are probably more ecologically important than more commonly studied intraspecific effects. These findings highlight the need for multispecies approaches in future studies of climate warming and body size., (© 2013 John Wiley & Sons Ltd.)

Landscape ecology plays a vital role in understanding the impacts of land-use change on biodiversity, but it is not a predictive discipline, lacking theoretical models that quantitatively predict biodiversity patterns from first principles. Here, we draw heavily on ideas from phylogenetics to fill this gap, basing our approach on the insight that habitat fragments have a shared history. We develop a landscape 'terrageny', which represents the historical spatial separation of habitat fragments in the same way that a phylogeny represents evolutionary divergence among species. Combining a random sampling model with a terrageny generates numerical predictions about the expected proportion of species shared between any two fragments, the locations of locally endemic species, and the number of species that have been driven locally extinct. The model predicts that community similarity declines with terragenetic distance, and that local endemics are more likely to be found in terragenetically distinctive fragments than in large fragments. We derive equations to quantify the variance around predictions, and show that ignoring the spatial structure of fragmented landscapes leads to over-estimates of local extinction rates at the landscape scale. We argue that ignoring the shared history of habitat fragments limits our ability to understand biodiversity changes in human-modified landscapes., (© 2013 The Authors. Ecology Letters published by John Wiley & Sons Ltd and CNRS.)

A major goal of ecology is to discover how dynamics and structure of multi-trophic ecological communities are related. This is difficult, because whole-community data are limited and typically comprise only a snapshot of a community instead of a time series of dynamics, and mathematical models of complex system dynamics have a large number of unmeasured parameters and therefore have been only tenuously related to real systems. These are related problems, because long time-series, if they were commonly available, would enable inference of parameters. The resulting 'plague of parameters' means most studies of multi-species population dynamics have been very theoretical. Dynamical models parametrized using physiological allometries may offer a partial cure for the plague of parameters, and these models are increasingly used in theoretical studies. However, physiological allometries cannot determine all parameters, and the models have also rarely been directly tested against data. We confronted a model of community dynamics with data from a lake community. Many important empirical patterns were reproducible as outcomes of dynamics, and were not reproducible when parameters did not follow physiological allometries. Results validate the usefulness, when parameters follow physiological allometries, of classic differential-equation models for understanding whole-community dynamics and the structure-dynamics relationship.

1. The power-law dependence of metabolic rate on body mass has major implications at every level of ecological organization. However, the overwhelming majority of studies examining this relationship have used basal or resting metabolic rates, and/or have used data consisting of species-averaged masses and metabolic rates. Field metabolic rates are more ecologically relevant and are probably more directly subject to natural selection than basal rates. Individual rates might be more important than species-average rates in determining the outcome of ecological interactions, and hence selection. 2. We here provide the first comprehensive database of published field metabolic rates and body masses of individual birds and mammals, containing measurements of 1498 animals of 133 species in 28 orders. We used linear mixed-effects models to answer questions about the body mass scaling of metabolic rate and its taxonomic universality/heterogeneity that have become classic areas of controversy. Our statistical approach allows mean scaling exponents and taxonomic heterogeneity in scaling to be analysed in a unified way while simultaneously accounting for nonindependence in the data due to shared evolutionary history of related species. 3. The mean power-law scaling exponents of metabolic rate vs. body mass relationships were 0.71 [95% confidence intervals (CI) 0.625-0.795] for birds and 0.64 (95% CI 0.564-0.716) for mammals. However, these central tendencies obscured meaningful taxonomic heterogeneity in scaling exponents. The primary taxonomic level at which heterogeneity occurred was the order level. Substantial heterogeneity also occurred at the species level, a fact that cannot be revealed by species-averaged data sets used in prior work. Variability in scaling exponents at both order and species levels was comparable to or exceeded the differences 3/4-2/3 = 1/12 and 0.71-0.64. 4. Results are interpreted in the light of a variety of existing theories. In particular, results are consistent with the heat dissipation theory of Speakman & Król (2010) and provided some support for the metabolic levels boundary hypothesis of Glazier (2010). 5. Our analysis provides the first comprehensive empirical analysis of the scaling relationship between field metabolic rate and body mass in individual birds and mammals. Our data set is a valuable contribution to those interested in theories of the allometry of metabolic rates., (© 2013 The Authors. Journal of Animal Ecology © 2013 British Ecological Society.)

Population dynamics are affected by changes in both the mean and standard deviation of climate, e.g., changes in average temperature are likely to affect populations, but so are changes in the strength of year-to-year temperature variability. The impacts of increases in average temperature are extensively researched, while the impacts of changes in climate variability are less studied. Is the greater attention given to changes in mean environment justified? To help answer this question we developed a simple population model, explicitly linked to an environmental process. We used the model to compare the sensitivities of a population's long-term stochastic growth rate, a measure of fitness, to changes in the mean and standard deviation of the environment. Results are interpreted in light of a comparative analysis of the relative magnitudes of change in means and standard deviations of biologically relevant climate variables in the United States. Results show that changes in the variability of the environment can be more important for many populations. Changes in mean conditions are likely to have a greater impact than changes in variability on populations far from their ideal environment, for example, populations near species range boundaries and potentially of conservation concern. Populations near range centres and close to their ideal environment are more likely to be affected by changes in variability. Among pest and insect disease vectors, as well as species of commercial value, populations likely to be of greatest economic and public health significance are those near species range centers, living in a near-ideal environment for the species. Observed changes in the variability of climate variables may benefit these populations.

Halley et al. purport to show a power-law relationship between fragment size and relaxation rates. We use a much more extensive data set to show that area dependence of relaxation rates exists only for very small fragment sizes (<60 hectares), which has limited relevance for our analyses conducted using 250,000-hectare grid squares. We also show that the example of Halley et al. is based on an unrealistic fragmentation model with an infinite number of fragments that have average size of zero hectares. A more realistic formulation of the model shows that relaxation is much less dependent on fragmentation than Halley et al. present.

Experimental data from intergenerational field manipulations of entire food webs are scarce, yet such approaches are essential for gauging impacts of environmental change in natural systems. We imposed 2 years of intermittent drought on stream channels in a replicated field trial, to measure food web responses to simulated climate change. Drought triggered widespread losses of species and links, with larger taxa and those that were rare for their size, many of which were predatory, being especially vulnerable. Many network properties, including size-scaling relationships within food chains, changed in response to drought. Other properties, such as connectance, were unaffected. These findings highlight the need for detailed experimental data from different organizational levels, from pairwise links to the entire food web. The loss of not only large species, but also those that were rare for their size, provides a newly refined way to gauge likely impacts that may be applied more generally to other systems and/or impacts.

Predicting when future species extinctions will occur is necessary for directing conservation investments but has proved difficult. We developed a new method for predicting extinctions over time, accounting for the timing and magnitude of habitat loss. We applied this to the Brazilian Amazon, predicting that local extinctions of forest-dependent vertebrate species have thus far been minimal (1% of species by 2008), with more than 80% of extinctions expected to be incurred from historical habitat loss still to come. Realistic deforestation scenarios suggest that local regions will lose an average of nine vertebrate species and have a further 16 committed to extinction by 2050. There is a window of opportunity to dilute the legacy of historical deforestation by concentrating conservation efforts in areas with greatest debt.

1. The effect of selective exploitation of certain age, stage or sex classes (e.g., trophy hunting) on population dynamics is relatively well studied in fisheries and sexually dimorphic mammals. 2. Harvesting of terrestrial species with no morphological differences visible between the different age and sex classes (monomorphic species) is usually assumed to be nonselective because monomorphicity makes intentionally selective harvesting pointless and impractical. But harvesting of the red grouse (Lagopus lagopus scoticus), a monomorphic species, was recently shown to be unintentionally selective. This study uses a sex- and age-specific model to explore the previously unresearched effects of unintentional harvesting selectivity. 3. We examine the effects of selectivity on red grouse dynamics by considering models with and without selectivity. Our models include territoriality and parasitism, two mechanisms known to be important for grouse dynamics. 4. We show that the unintentional selectivity of harvesting that occurs in red grouse decreases population yield compared with unselective harvesting at high harvest rates. Selectivity also dramatically increases extinction risk at high harvest rates. 5. Selective harvesting strengthens the 3- to 13-year red grouse population cycle, suggesting that the selectivity of harvesting is a previously unappreciated factor contributing to the cycle. 6. The additional extinction risk introduced by harvesting selectivity provides a quantitative justification for typically implemented 20-40% harvest rates, which are below the maximum sustainable yield that could be taken, given the observed population growth rates of red grouse. 7. This study shows the possible broad importance of investigating in future research whether unintentionally selective harvesting occurs on other species., (© 2011 The Authors. Journal of Animal Ecology © 2011 British Ecological Society.)

1. The spectral colour of population dynamics and its causes have attracted much interest. The spectral colour of a time series can be determined from its power spectrum, which shows what proportion of the total variance in the time series occurs at each frequency. A time series with a red spectrum (a negative spectral exponent) is dominated by low-frequency oscillations, and a time series with a blue spectrum (a positive spectral exponent) is dominated by high-frequency oscillations. 2. Both climate variables and population time series are characterised by red spectra, suggesting that a population's environment might be partly responsible for its spectral colour. Laboratory experiments and models have been used to investigate this potential link. However, no study using field data has directly tested whether populations in redder environments are redder. 3. This study uses the Global Population Dynamics Database together with climate data to test for this effect. We found that the spectral exponent of mean summer temperatures correlates positively and significantly with population spectral exponent. 4. We also found that over the last century, temperature climate variables on most continents have become bluer. 5. Although population time series are not long or abundant enough to judge directly whether their spectral colours are changing, our two results taken together suggest that population spectral colour may be affected by the changing spectral colour of climate variables. Population spectral colour has been linked to extinction; we discuss the potential implications of our results for extinction probability., (© 2011 The Authors. Journal of Animal Ecology © 2011 British Ecological Society.)

Predator-prey body size relationships influence food chain length, trophic structure, transfer efficiency, interaction strength, and the bioaccumulation of contaminants. Improved quantification of these relationships and their response to the environment is needed to parameterize food web models and describe food web structure and function. A compiled data set comprising 29582 records of individual prey eaten at 21 locations by individual predators that spanned 10 orders of magnitude in mass and lived in marine environments ranging from the poles to the tropics was used to investigate the influence of predator size and environment on predator and prey size relationships. Linear mixed effects models demonstrated that predator-prey mass ratios (PPMR) increased with predator mass. The amount of the increase varied among locations and predator species and individuals but was not significantly influenced by temperature, latitude, depth, or primary production. Increases in PPMR with predator mass implied nonlinear relationships between log body mass and trophic level and reductions in transfer efficiency with increasing body size. The results suggest that very general rules determine dominant trends in PPMR in diverse marine ecosystems, leading to the ubiquity of size-based trophic structuring and the consistency of observed relationships between the relative abundance of individuals and their body size.

Many studies have aimed to understand food webs by investigating components such as trophic links (one consumer taxon eats one resource taxon), tritrophic interactions (one consumer eats an intermediate taxon, which eats a resource), or longer chains of links. We show here that none of these components (links, tritrophic interactions, and longer chains), individually or as an ensemble, accounts fully for the properties of the next higher level of organization. As a cell is more than its molecules, as an organ is more than its cells, and as an organism is more than its organs, in a food web, new structure emerges at every organizational level up to and including the whole web. We demonstrate the emergence of properties at progressively higher levels of structure by using all of the directly observed, appropriately organized, publicly available food web datasets with relatively complete trophic link data and with average body mass and population density data for each taxon. There are only three such webs, those of Tuesday Lake, Michigan, in 1984 and 1986, and Ythan Estuary, Scotland. We make the data freely available online with this report. Differences in web patterns between Tuesday Lake and Ythan Estuary, and similarities of Tuesday Lake in 1984 and 1986 despite 50% turnover of species, suggest that the patterns we describe respond to major differences between ecosystem types.

Individuals tend to interact more strongly with nearby individuals or within particular social groups. Recent theoretical advances have demonstrated that these within-population relationships can have fundamental implications for ecological and evolutionary dynamics. In particular, contact networks are crucial to the spread and evolution of disease. However, the theory remains largely untested experimentally. Here, we manipulate habitat viscosity and thereby the frequency of local interactions in an insect-pathogen model system in which the virus had previously been shown to have little effect on host population dynamics. At high viscosity, the pathogen caused the collapse of dominant and otherwise stable host generation cycles. Modeling shows that this collapse can be explained by an increase in the frequency of intracohort interactions relative to intercohort interactions, leading to more disease transmission. Our work emphasizes that spatial structure can subtly mediate intraspecific competition and the effects of natural enemies. A decrease in dispersal in a population may actually (sometimes rather counterintuitively) intensify the effects of parasites. Broadly, because anthropological and environmental change often cause changes in population mixing, our work highlights the potential for dramatic changes in the effects of parasites on host populations.

Predicting species population density-body mass scaling in community food webs (henceforth webs) is important for conservation and to understand community structure. Very different types of scaling have been studied, based on either individuals or species. The individual size distribution (ISD) describes the distribution of individual-organism body masses regardless of taxonomy, and contains the same information as the abundance spectrum. Focusing instead on species, the local size-density relationship (LSDR) plots population densities vs. mean body masses of species. The distribution of species mean body masses (the species-mean-size distribution, SMSD) is also important but previously little studied in webs. We here combine and formalize theory of several authors to predict: how these three descriptions are related; the forms of the LSDR and ISD; and variation in scaling among webs. We describe empirically the SMSDs of two pelagic, one estuarine, and 146 soil webs by power laws and generalizations. We test theory and find it broadly validated.

International migration will play an increasing role in the demographic future of most nations if fertility continues to decline globally. We developed an algorithm to project future numbers of international migrants from any country or region to any other. The proposed generalized linear model (GLM) used geographic and demographic independent variables only (the population and area of origins and destinations of migrants, the distance between origin and destination, the calendar year, and indicator variables to quantify nonrandom characteristics of individual countries). The dependent variable, yearly numbers of migrants, was quantified by 43653 reports from 11 countries of migration from 228 origins and to 195 destinations during 1960-2004. The final GLM based on all data was selected by the Bayesian information criterion. The number of migrants per year from origin to destination was proportional to (population of origin)(0.86)(area of origin)(-0.21)(population of destination)(0.36)(distance)(-0.97), multiplied by functions of year and country-specific indicator variables. The number of emigrants from an origin depended on both its population and its population density. For a variable initial year and a fixed terminal year 2004, the parameter estimates appeared stable. Multiple R(2), the fraction of variation in log numbers of migrants accounted for by the starting model, improved gradually with recentness of the data: R(2) = 0.57 for data from 1960 to 2004, R(2) = 0.59 for 1985-2004, R(2) = 0.61 for 1995-2004, and R(2) = 0.64 for 2000-2004. The migration estimates generated by the model may be embedded in deterministic or stochastic population projections.

Populations fluctuate because of their internal dynamics, which can be nonlinear and stochastic, and in response to environmental variation. Theory predicts how the colour of environmental stochasticity affects population means, variances and correlations with the environment over time. The theory has not been tested for cycling populations, commonly observed in field systems. We applied noise of different colours to cycling laboratory beetle populations, holding other statistical properties of the noise fixed. Theory was largely validated, but failed to predict observations in sufficient detail. The main period of population cycling was shifted up to 33% by the colour of environmental stochasticity. Noise colour affected population means, variances and dominant periodicities differently for populations that cycled in different ways without noise. Our results show that changes in the colour of climatic variability, partly caused by humans, may affect the main periodicity of cycling populations, possibly impacting industry, pest management and conservation.

Stochasticity alters the nonlinear dynamics of inherently cycling populations. The power spectrum can describe and explain the impacts of stochasticity. We fitted models to short observed time series of flour beetle populations in the frequency domain, then used a well fitting stochastic mechanistic model to generate detailed predictions of population spectra. Some predicted spectral peaks represent periodic phenomena induced or modified by stochasticity and were experimentally confirmed. For one experimental treatment, linearization theory explained that these peaks represent overcompensatory decay of deviations from deterministic oscillation. In another treatment, stochasticity caused frequent directional phase shifting around a cyclic attractor. This directional phase shifting was not explained by linearization theory and modified the periodicity of the system. If field systems exhibit directional phase shifting, then changing the intensity of demographic or environmental noise while holding constant the structure of the noise can change the main frequency of population fluctuations.

It has been suggested that differences in body size between consumer and resource species may have important implications for interaction strengths, population dynamics, and eventually food web structure, function, and evolution. Still, the general distribution of consumer-'resource body-size ratios in real ecosystems, and whether they vary systematically among habitats or broad taxonomic groups, is poorly understood. Using a unique global database on consumer and resource body sizes, we show that the mean body-size ratios of aquatic herbivorous and detritivorous consumers are several orders of magnitude larger than those of carnivorous predators. Carnivorous predator-prey body-size ratios vary across different habitats and predator and prey types (invertebrates, ectotherm, and endotherm vertebrates). Predator-prey body-size ratios are on average significantly higher (1) in freshwater habitats than in marine or terrestrial habitats, (2) for vertebrate than for invertebrate predators, and (3) for invertebrate than for ectotherm vertebrate prey. If recent studies that relate body-size ratios to interaction strengths are general, our results suggest that mean consumer-resource interaction strengths may vary systematically across different habitat categories and consumer types.