Search

Following severe environmental change that reduces mean population fitness below replacement, populations must adapt to avoid eventual extinction, a process called evolutionary rescue. Models of evolutionary rescue demonstrate that initial size, genetic variation, and degree of maladaptation influence population fates. However, many models feature populations that grow without negative density dependence or with constant genetic diversity despite precipitous population decline, assumptions likely to be violated in conservation settings. We examined the simultaneous influences of density-dependent growth and erosion of genetic diversity on populations adapting to novel environmental change using stochastic, individual-based simulations. Density dependence decreased the probability of rescue and increased the probability of extinction, especially in large and initially well-adapted populations that previously have been predicted to be at low risk. Increased extinction occurred shortly following environmental change, as populations under density dependence experienced more rapid decline and reached smaller sizes. Populations that experienced evolutionary rescue lost genetic diversity through drift and adaptation, particularly under density dependence.Populations that declined to extinction entered an extinction vortex, where small size increased drift, loss of genetic diversity, and the fixation of maladaptive alleles, hindered adaptation, and kept populations at small densities where they were vulnerable to extinctionviademographic stochasticity.

Species expanding into new habitats as a result of climate change or human introductions will frequently encounter resident competitors. Theoretical models suggest that such interspecific competition can alter the speed of expansion and the shape of expanding range boundaries. However, competitive interactions are rarely considered when forecasting the success or speed of expansion, in part because there has been no direct experimental evidence that competition affects either expansion speed or boundary shape. Here we demonstrate that interspecific competition alters both expansion speed and range boundary shape. Using a two-species experimental system of the flour beetles Tribolium castaneum and Tribolium confusum, we show that interspecific competition dramatically slows expansion across a landscape over multiple generations. Using a parameterized stochastic model of expansion, we find that this slowdown can persist over the long term. We also find that the shape of the moving range boundary changes continuously over many generations of expansion, first steepening and then becoming shallower, due to the competitive effect of the resident and density-dependent dispersal of the invader. This dynamic boundary shape suggests that current forecasting approaches assuming a constant shape could be misleading. More broadly, our results demonstrate that interactions between competing species can play a large role during range expansions and thus should be included in models and studies that monitor, forecast, or manage expansions in natural systems.

Dispersal is a key process in shaping species spatial distributions. Species interactions and variation in dispersal probabilities may jointly influence species spatial dynamics. However, many studies examine dispersal as a neutral process, independent of community context or intraspecific variation in dispersal behaviour. Here, we use controlled, replicated communities of two Tribolium species (T. castaneum and T. confusum) to examine how intraspecific variation in dispersal behaviour and community context influence dispersal dynamics in simple experimental landscapes composed of homogeneous habitat patches. We found considerable individual-level variation in dispersal probability that was unrelated to body size variation. Further, the context of dispersal mattered, as T. castaneum dispersal was reduced in two-species communities, while T. confusum dispersal was unaffected by community composition. Incorporating individual-level variation into a two-species stochastic spatial Ricker model, we provide evidence that individual-level variability in dispersal behaviour results in more variable spatial spread than assuming individuals have the same dispersal probability. Further, interspecific competition resulted in more variable spatial spread. The variability in spatial spread observed in our tightly controlled and replicated experimental system and in our stochastic model simulations points to potential fundamental limitations in forecasting species shifting ranges without considering potential interspecific interactions and demographic variability in dispersal behaviour.

The flour beetles Tribolium castaneum (Herbst) and Tribolium confusum (du Val) are model organisms for studying the effects of intra- and interspecific competition. Both species are known to compete directly through egg cannibalism and indirectly through the density-dependent release of allelopathic chemicals. To better characterize these indirect interactions, recent work has focused on quantifying the per-capita effects of intraspecific chemical competition. However, the effects of interspecific chemical competition in this system have not previously been estimated. We used experimental microcosms to examine how interspecific chemical secretions affected the reproductive activity and development time of laboratory populations of T. castaneum and T. confusum. We created replicated habitats containing flour medium that had been occupied and chemically ‘conditioned’ by one of the two beetle species across a range of densities, then examined how beetles of the other species responded in terms of the number of eggs laid by females and, separately, the development time of offspring. We found that T. castaneum fecundity was reduced when beetles experienced flour conditioned by T. confusum. In contrast, fecundity of T. confusum was largely unaffected by flour conditioned by T. castaneum. Additionally, we found that interspecific conditioning decreased development times for beetles of both species, particularly the development of larvae to pupae. Our results indicate that interspecific chemical competition impacts the life history of Tribolium species and suggests that models incorporating chemical competition may more accurately describe the biology of flour beetle communities.

When managing natural systems, the importance of recognizing the role of uncertainty has been formalized as the precautionary approach. However, it is difficult to determine the role of stochasticity in the success or failure of management because there is almost always no replication; typically, only a single observation exists for a particular site or management strategy. Yet, assessing the role of stochasticity is important for providing a strong foundation for the precautionary approach, and learning from past outcomes is critical for implementing adaptive management of species or ecosystems. In addition, adaptive management relies on being able to implement a variety of strategies in order to learn—an often difficult task in natural systems. Here, we show that there is large, stochastically driven variability in success for management treatments to control an invasive species, particularly for moderate, and more feasible, management strategies. This is exactly where the precautionary approach should be important. Even when combining management strategies, we show that moderate effort in management either fails or is highly variable in its success. This variability allows some management treatments to, on average, meet their target, even when failure is probable. Our study is an important quantitative replicated experimental test of the precautionary approach and can serve as a way to understand the variability in management outcomes in natural systems which have the potential to be more variable than our tightly controlled system.

Theory has shown that the effects of demographic stochasticity on communities may depend on the magnitude of fitness differences between species. In particular, it has been suggested that demographic stochasticity has the potential to significantly alter competitive outcomes when fitness differences are small (nearly neutral), but that it has negligible effects when fitness differences are large (highly non‐neutral). Here we test such theory experimentally and extend it to examine how demographic stochasticity affects exclusion frequency and mean densities of consumers in simple, but non‐neutral, consumer–resource communities. We used experimental microcosms of protists and rotifers feeding on a bacterial resource to test how varying absolute population sizes (a driver of demographic stochasticity) affected the probability of competitive exclusion of the weakest competitor. To explore whether demographic stochasticity could explain our experimental results, and to generalize beyond our experiment, we paired the experiment with a continuous‐time stochastic model of resource competition, which we simulated for 11 different fitness inequalities between competiting consumers. Consistent with theory, in both our experiments and our simulations we found that demographic stochasticity altered competitive outcomes in communities where fitness differences were small. However, we also found that demographic stochasticity alone could affect communities in other ways, even when fitness differences between competitors were large. Specifically, demographic stochasticity altered mean densities of both weak and strong competitors in experimental and simulated communities. These findings highlight how demographic stochasticity can change both competitive outcomes in non‐neutral communities and the processes underlying overall community dynamics.

Increasing rates of habitat fragmentation globally underscore the importance of understanding the full spectrum of fragmentation’s ecological consequences. Fragmentation alters the thermal environment of fragments, which may alter the body size of ectothermic organisms and in turn impact survival and reproduction. To determine whether experimental habitat fragmentation alters body size in the heliothermic, ground-dwelling common garden skink (Lampropholis guichenoti). We use body size data spanning 29 years to experimentally test the prediction that lizards will experience morphological changes in forest fragments but not in non-fragmented controls. Lizards were smaller in forest fragments relative to those in the non-fragmented controls after the fragmentation treatment was applied. For lizards within forest fragments, the greater the exposure to deforested areas, the greater the decline in body size. This pattern was strongest in the first 5 years following fragmentation and weakened or reversed over time as the pine plantation matrix surrounding the fragments matured. Using sampling site-scale temperature data for the most recent 5 years of the experiment, we show that temperature predicts lizard body size. Our findings are consistent with predictions made under the temperature-size rule that ectotherms will be smaller in fragmented landscapes because of temperature increases at newly created edges. Our results raise new concerns about the effects of fragmentation on organisms in remnant patches and offer new research priorities, as more evidence is needed to determine the generality of body size declines in fragmented landscapes. Our results also highlight that body size declines, often attributed to climate change, may be amplified by habitat fragmentation, which has been global in its impact.

Despite growing calls to develop data science students’ ethical awareness and expand human-centered approaches to data science education, introductory courses in the field remain largely technical. A new interdisciplinary data science program aims to merge STEM and humanities perspectives starting at the very beginning of the data science curriculum. Existing literature suggests that humanities integration can make STEM courses more appealing to a wider range of students, including women and students of color, and enhance student learning of essential concepts and foundational reasoning skills, such as those collectively known as data acumen. Cultivating students’ data acumen requires a more inclusive vision of how the knowledge and insights generated through computational methods and statistical analysis relates to other ways of knowing.

Understanding the movement of species' ranges is a classic ecological problem that takes on urgency in this era of global change. Historically treated as a purely ecological process, range expansion is now understood to involve eco-evolutionary feedbacks due to spatial genetic structure that emerges as populations spread. We synthesize empirical and theoretical work on the eco-evolutionary dynamics of range expansion, with emphasis on bridging directional, deterministic processes that favor evolved increases in dispersal and demographic traits with stochastic processes that lead to the random fixation of alleles and traits. We develop a framework for understanding the joint influence of these processes in changing the mean and variance of expansion speed and its underlying traits. Our synthesis of recent laboratory experiments supports the consistent role of evolution in accelerating expansion speed on average, and highlights unexpected diversity in how evolution can influence variability in speed: results not well predicted by current theory. We discuss and evaluate support for three classes of modifiers of eco-evolutionary range dynamics (landscape context, trait genetics, and biotic interactions), identify emerging themes, and suggest new directions for future work in a field that stands to increase in relevance as populations move in response to global change.

Soil nitrogen (N) availability is critical for grassland functioning. However, human activities have increased the supply of biologically limiting nutrients, and changed the density and identity of mammalian herbivores. These anthropogenic changes may alter net soil N mineralization (soil net Nmin), that is, the net balance between N mineralization and immobilization, which could severely impact grassland structure and functioning. Yet, to date, little is known about how fertilization and herbivore removal individually, or jointly, affect soil net Nmin across a wide range of grasslands that vary in soil and climatic properties. Here we collected data from 22 grasslands on five continents, all part of a globally replicated experiment, to assess how fertilization and herbivore removal affected potential (laboratory‐based) and realized (field‐based) soil net Nmin. Herbivore removal in the absence of fertilization did not alter potential and realized soil net Nmin. However, fertilization alone and in combination with herbivore removal consistently increased potential soil net Nmin. Realized soil net Nmin, in contrast, significantly decreased in fertilized plots where herbivores were removed. Treatment effects on potential and realized soil net Nmin were contingent on site‐specific soil and climatic properties. Fertilization effects on potential soil net Nmin were larger at sites with higher mean annual precipitation (MAP) and temperature of the wettest quarter (T.q.wet). Reciprocally, realized soil net Nmin declined most strongly with fertilization and herbivore removal at sites with lower MAP and higher T.q.wet. In summary, our findings show that anthropogenic nutrient enrichment, herbivore exclusion and alterations in future climatic conditions can negatively impact soil net Nmin across global grasslands under realistic field conditions. This is an important context‐dependent knowledge for grassland management worldwide.

Predicting whether individuals will colonize a novel habitat is of fundamental ecological interest and is crucial to both conservation efforts and invasive species management. A consistently supported predictor of colonization success is the number of individuals introduced, also called propagule pressure. Propagule pressure increases with the number of introductions and the number of individuals per introduction (the size of the introduction), but it is unresolved which process is a stronger driver of colonization success. Furthermore their relative importance may depend upon the environment, with multiple introductions potentially enhancing colonization of fluctuating environments. To evaluate the relative importance of the number and size of introductions and its dependence upon environmental variability, we paired demographic simulations with a microcosm experiment. Using Tribolium flour beetles as a model system, we introduced a fixed number of individuals into replicated novel habitats of stable or fluctuating quality, varying the number of introductions through time and size of each introduction. We evaluated establishment probability and the size of extant populations after 7 generations. In the simulations and microcosms, we found that establishment probability increased with more, smaller introductions, but was not affected by biologically realistic fluctuations in environmental quality. Population size was not significantly affected by environmental variability in the simulations, but populations in the microcosms grew larger in a stable environment, especially with more introduction events. In general, the microcosm experiment yielded higher establishment probability and larger populations than the demographic simulations. We suggest that genetic mechanisms likely underlie these differences and thus deserve more attention in efforts to parse propagule pressure. Our results highlight the importance of preventing further introductions of undesirable species to invaded sites, and suggest conservation efforts should focus on increasing the number of introductions or re-introductions of desirable species rather than increasing the size of those introduction events.

The demographic rates (e.g., birth, death, migration) of many organisms have been shown to respond strongly to short- and long-term environmental change, including variation in temperature and precipitation. While ecologists have long accounted for such nonhomogeneous demography in deterministic population models, nonhomogeneous stochastic population models are largely absent from the literature. This is especially the case for models that use exact stochastic methods, such as Gillespie’s stochastic simulation algorithm (SSA), which commonly assumes that demographic rates do not respond to external environmental change (i.e., assumes homogeneous demography). In other words, ecologists are currently accounting for the effects of demographic stochasticity or environmental variability, but not both. In this paper, we describe an extension of Gillespie’s SSA (SSA + ) that allows for nonhomogeneous demography and examine how its predictions differ from a method that is partly naive to environmental change (SSAn) for two fundamental ecological models (exponential and logistic growth). We find important differences in the predicted population sizes of SSA + versus SSAn simulations, particularly when demography responds to fluctuating and irregularly changing environments. Further, we outline a computationally inexpensive approach for estimating when and under what circumstances it can be important to fully account for nonhomogeneous demography for any class of model.

Although forest fragmentation is generally thought to impact tree growth and mortality negatively, recent work suggests some forests are resilient. Experimental forests provide an opportunity to examine the timing and extent of forest tree resilience to disturbance from fragmentation. We used the Wog Wog Habitat Fragmentation Experiment in southeastern Australia to test Eucalyptus growth and survivorship responses to forest fragmentation over a 26 year period. We measured 2418 tree diameters and used spline-regression techniques to examine non-monotonic fragmentation effect over two time periods. Over the first 4 years after fragmentation, individual eucalypt tree growth was greater than in continuous forest for large trees and mortality rates were higher only within 10 m of edges. Over the following 22 years only the effects on tree growth remained and on average all fragments rebounded so that their biomass and mortality rates were equivalent to continuous forest. Importantly non-monotonic patterns were observed in growth and mortality with respect to area and distance from edge in both study periods, demonstrating that fragmentation impacts on trees can be strong in localized areas (greatest in 3 ha fragments and 0–30 m edges) and over short time periods. Dry-sclerophyll eucalypt forests join the set of forest types that display resilient growth dynamics post fragmentation. Moreover, persistent non-monotonic impacts on tree growth with respect to tree size, fragment area, and fragment distance from edge, highlighting landscape fragmentation as a driver of habitat heterogeneity within remnant forest fragments.

Understanding pathogen transmission is crucial for predicting and managing disease. Nonetheless, experimental comparisons of alternative functional forms of transmission remain rare, and those experiments that are conducted are often not designed to test the full range of possible forms.To differentiate among 10 candidate transmission functions, we used a novel experimental design in which we independently varied four factors—duration of exposure, numbers of parasites, numbers of hosts and parasite density—in laboratory infection experiments.We used interactions between amphibian hosts and trematode parasites as a model system and all candidate models incorporated parasite depletion. An additional manipulation involving anaesthesia addressed the effects of host behaviour on transmission form.Across all experiments, nonlinear transmission forms involving either a power law or a negative binomial function were the best‐fitting models and consistently outperformed the linear density‐dependent and density‐independent functions. By testing previously published data for two other host–macroparasite systems, we also found support for the same nonlinear transmission forms.Although manipulations of parasite density are common in transmission studies, the comprehensive set of variables tested in our experiments revealed that variation in density alone was least likely to differentiate among competing transmission functions. Across host–pathogen systems, nonlinear functions may often more accurately represent transmission dynamics and thus provide more realistic predictions for infection., The authors tested 10 candidate functions representing the transmission of trematode parasites to amphibian hosts using laboratory experiments varying the duration of exposure, numbers of parasites, numbers of hosts and parasite density. Nonlinear functions outperformed classical frequency‐ and density‐dependent transmission functions and may provide more realistic predictions for infection.

Habitat fragmentation can alter the trophic structure of communities and environmental conditions, thus driving changes in biodiversity and ecosystem functions. Quantifying niches of generalist predators can reveal how fragmentation alters ecosystems. In a habitat fragmentation experiment, we used stable isotopes of a generalist predator skink to test predictions from spatial theory on trophic structure and to quantify abiotic changes associated with fragmentation among continuous forest, fragments, and matrix habitats. We predicted that in fragments and the matrix, isotopic niches would shift due to decreases in skink trophic positions (δ15N) from reductions in trophic structure of arthropod food webs and abiotic changes over time (δ13C) relative to continuous forest. Contrary to theoretical predictions, we did not find evidence of reductions in trophic structure with fragmentation. In fact, skink δ15N values were higher in the matrix and fragments than continuous forest, likely due to changes in distributions of a detritivorous prey species. In addition, δ13C values in the matrix decreased over years since fragmentation due to abiotic changes associated with matrix tree maturation. We show how isotopic niches are influenced by fragmentation via shifts in biotic and abiotic processes. The potential for either or both spatial and abiotic effects of fragmentation present a challenge for theory to better predict ecological changes in fragmented landscapes.

Habitat loss and fragmentation are major threats to biodiversity and ecosystem processes. Our current understanding of the impacts of habitat loss and fragmentation is based largely on studies that focus on either short-term or long-term responses. Short-term responses are often used to predict long-term responses and make management decisions. The lack of studies comparing short- and long-term responses to fragmentation means we do not adequately understand when and how well short-term responses can be extrapolated to predict long-term responses, and when or why they cannot. To address this gap, we used data from one of the world's longest-running fragmentation experiments, The Wog Wog Habitat Fragmentation Experiment. Using data for carabid beetles, we found that responses in the long term (more than 22 yr post-fragmentation ≈22 generations) often contrasted markedly with those in the short term (5 yr post-fragmentation). The total abundance of all carabids, species richness and the occurrence of six species declined in the short term in the fragments but increased over the long term. The occurrence of three species declined initially and continued to decline, whilst another species was positively affected initially but decreased in the long term. Species' responses to the matrix that surrounds the fragments strongly predicted both the direction (increase/decline in occurrence) and magnitude of their responses to fragmentation. Additionally, species' responses to the matrix were somewhat predicted by their preferences for different types of native habitat (open vs. shaded). Our study highlights the degree of the matrix's influence in fragmented landscapes, and how this influence can change over time. We urge caution in using short-term responses to forecast long-term responses in cases where the matrix (1) impacts species' responses to fragmentation (by isolating them, creating new habitat or altering fragment habitat) and (2) is likely to change through time.

Colonisation is a fundamental ecological and evolutionary process that drives the distribution and abundance of organisms. The initial ability of colonists to establish is determined largely by the number of founders and their genetic background. We explore the importance of these demographic and genetic properties for longer term persistence and adaptation of populations colonising a novel habitat using experimental populations of Tribolium castaneum. We introduced individuals from three genetic backgrounds (inbred - outbred) into a novel environment at three founding sizes (2-32), and tracked populations for seven generations. Inbreeding had negative effects, whereas outbreeding generally had positive effects on establishment, population growth and long-term persistence. Severe bottlenecks due to small founding sizes reduced genetic variation and fitness but did not prevent adaptation if the founders originated from genetically diverse populations. Thus, we find important and largely independent roles for both demographic and genetic processes in driving colonisation success.

Summary 1.Checkerboard distributions ? mutually exclusive species co-occurrences ? are a common observation in community ecology and biogeography. While the underlying causes of checkerboard distributions have remained elusive, a longstanding argument is that they are representative of strong competitive interactions and/or dispersal limitation. 2.We explore this using a stochastic two-patch metacommunity model combined with an experimental two-patch system of competing Tribolium species, quantifying checkerboard distributions using the abundance-based index Ast. 3.We find that maintenance of checkerboard distributions is possible in a limited parameter space consisting of low dispersal rates, low population growth rates, and high interspecific competition. Checkerboards were not maintained in experimental metacommunities. 4.Our model, parameterized using independent data, echoed this finding, providing a clear link between model and experiment, and suggested that only small regions of parameter space would allow for checkerboard distributions between patches with equally hospitable environments. These findings may provide insight into when interspecific competition and dispersal limitation would promote checkerboard distributions. This article is protected by copyright. All rights reserved.

1. Ants are a ubiquitous and crucial component of Australian Eucalyptus forests, but responses to long-term habitat fragmentation remain poorly understood. Two ant species were followed across a 21-year history of pine plantation establishment and maturation in a southeast Australian Eucalyptus forest. 2. At Wog Wog in southeastern Australia, a Native Eucalyptus forest was clear-cut to make way for plantation establishment and 12 remnant patches of forest were left intact and subsequently surrounded by a pine matrix. Pitfall traps were placed in the continuous native forest, remnant Eucalyptus patches, and the pine matrix between fragments, and were stratified based on proximity to remnant patch edges and habitat type. Two ant species are focused on that represent the only remaining data for the early years of the experiment. 3. While Leptomyrmex erythrocephalus (Fabricius), the rarer of the two species, was not affected by fragmentation in the short term, 21 years after fragmentation, it was less likely to occur in both the mature pine matrix and fragments than in continuous forest controls. Aphaenogaster longiceps (Smith F.) was equally likely to occur in the fragments, continuous forest, and pine matrix early in the experiment but by year 21 post-fragmentation was less likely to occur in the pine matrix than fragments or controls. 4. Importantly, we only detected negative impacts of fragmentation on ant occurrence as the pine plantation matrix matured and isolated ant populations on fragments. 5. It is concluded that changes in matrix suitability and specific habitat characteristics influence ant persistence in Eucalyptus fragments.

Edge effects due to habitat loss and fragmentation have pervasive impacts on many natural ecosystems worldwide. We aimed to explore whether, in tandem with the resource-based model of edge effects, species feeding-guild and flight-capacity can help explain species responses to an edge. We used a two-sided edge gradient that extended from 1000 m into native Eucalyptus forest to 316 m into an exotic pine plantation. We used generalised additive models to examine the continuous responses of beetle species, feeding-guild species richness and flight-capable group species richness to the edge gradient and environmental covariates. Phytophagous species richness was directly related to variation in vegetation along the edge gradient. There were more flight-capable species in Eucalyptus forest and more flightless species in exotic pine plantation. Many individual species exhibited multiple-peaked edge-profiles. The resource based model for edge effects can be used in tandem with traits such as feeding-guild and flight-capacity to understand drivers of large scale edge responses. Some trait-groups can show generalisable responses that can be linked with drivers such as vegetation richness and habitat structure. Many trait-group responses, however, are less generalisable and not explained by easily measured habitat variables. Difficulties in linking traits with resources along the edge could be due to unmeasured variation and indirect effects. Some species’ responses reached the limits of the edge gradient demonstrating the need to examine edge effects at large scales, such as kilometres.

International audience; Colonization and expansion into novel landscapes determine the distribution and abundance of species in our rapidly changing ecosystems worldwide. Colonization events are crucibles for rapid evolution, but it is not known whether evolutionary changes arise mainly after successful colonization has occurred, or if evolution plays an immediate role, governing the growth and expansion speed of colonizing populations. There is evidence that spatial evolutionary processes can speed range expansion within a few generations because dispersal tendencies may evolve upwards at range edges. Additionally, rapid adaptation to a novel environment can increase population growth rates, which also promotes spread. However, the role of adaptive evolution and the relative contributions of spatial evolution and adaptation to expansion are unclear. Using a model system, red flour beetles (Tribolium castaneum), we either allowed or constrained evolution of populations colonizing a novel environment and measured population growth and spread. At the end of the experiment we assessed the fitness and dispersal tendency of individuals originating either from the core or edge of evolving populations or from nonevolving populations in a common garden. Within six generations, evolving populations grew three times larger and spread 46% faster than populations in which evolution was constrained. Increased size and expansion speed were strongly driven by adaptation, whereas spatial evolutionary processes acting on edge subpopulations contributed less. This experimental evidence demonstrates that rapid evolution drives both population growth and expansion speed and is thus crucial to consider for managing biological invasions and successfully introducing or reintroducing species for management and conservation.

The β-null deviation measure, developed as a null model for β-diversity, is increasingly used in empirical studies to detect the underlying structuring mechanisms in communities (e.g. niche versus neutral and stochastic versus deterministic). Despite growing use, the ecological interpretation of the presence/absence and abundance-based versions of the β-null diversity measure have not been tested against communities with known assembly mechanisms, and thus have not been validated as an appropriate tool for inferring assembly mechanisms. Using a mechanistic model with known assembly mechanisms, we simulated replicate metacommunities and examined β-null deviation values 1) across a gradient of niche (species-sorting) to neutrally structured metacommunities, 2) through time, and 3) we compared the effect of changes in assembly mechanism on the performance of the β-null deviation measures. The impact of stochasticity on assembly outcomes was also considered. β-null deviation measures proved to be interpretable as a measure of niche or neutral assembly. However, the presence/absence version of the β-null deviation measure could not differentiate between niche and neutral metacommunities if demographic stochasticity were present. The abundance-based β-null deviation measure was successful in distinguishing between niche and neutral metacommunities and was robust to the presence of stochasticity, changes through time, and changing assembly mechanisms. However, we suggest that it is not robust to changing abundance evenness distributions or sampling of communities, and so its interpretation still requires some care. We encourage the testing of the assumptions behind null models for ecology and care in their application.

Setting aside high-quality large areas of habitat to protect threatened populations is becoming increasingly difficult as humans fragment and degrade the environment. Biologists and managers therefore must determine the best way to shepherd small populations through the dual challenges of reductions in both the number of individuals and genetic variability. By bringing in additional individuals, threatened populations can be increased in size (demographic rescue) or provided with variation to facilitate adaptation and reduce inbreeding (genetic rescue). The relative strengths of demographic and genetic rescue for reducing extinction and increasing growth of threatened populations are untested, and which type of rescue is effective may vary with population size. Using the flour beetle (Tribolium castaneum) in a microcosm experiment, we disentangled the genetic and demographic components of rescue, and compared them with adaptation from standing genetic variation (evolutionary rescue in the strictest sense) using 244 experimental populations founded at either a smaller (50 individuals) or larger (150 individuals) size. Both types of rescue reduced extinction, and those effects were additive. Over the course of six generations, genetic rescue increased population sizes and intrinsic fitness substantially. Both large and small populations showed evidence of being able to adapt from standing genetic variation. Our results support the practice of genetic rescue in facilitating adaptation and reducing inbreeding depression, and suggest that demographic rescue alone may suffice in larger populations even if only moderately inbred individuals are available for addition.

Humans dominate many important Earth system processes including the nitrogen (N) cycle. Atmospheric N deposition affects fundamental processes such as carbon cycling, climate regulation, and biodiversity, and could result in changes to fundamental Earth system processes such as primary production. Both modelling and experimentation have suggested a role for anthropogenically altered N deposition in increasing productivity, nevertheless, current understanding of the relative strength of N deposition with respect to other controls on production such as edaphic conditions and climate is limited. Here we use an international multiscale data set to show that atmospheric N deposition is positively correlated to aboveground net primary production (ANPP) observed at the 1-m 2 level across a wide range of herbaceous ecosystems. N deposition was a better predictor than climatic drivers and local soil conditions, explaining 16% of observed variation in ANPP globally with an increase of 1 kg Nha � 1 � yr � 1 increasing ANPP by 3%. Soil pH explained 8% of observed variation in ANPP while climatic drivers showed no significant relationship. Our results illustrate that the incorporation of global N deposition patterns in Earth system models are likely to substantially improve estimates of primary production in herbaceous systems. In herbaceous systems across the world, humans appear to be partially driving local ANPP through impacts on the N cycle.

Range expansions are central to two ecological issues reshaping patterns of global biodiversity: biological invasions and climate change. Traditional theory considers range expansion as the outcome of the demographic processes of birth, death and dispersal, while ignoring the evolutionary implications of such processes. Recent research suggests evolution could also play a critical role in determining expansion speed but controlled experiments are lacking. Here we use flour beetles (Tribolium castaneum) to show experimentally that mean expansion speed and stochastic variation in speed are both increased by rapid evolution of traits at the expansion edge. We find that higher dispersal ability and lower intrinsic growth rates evolve at the expansion edge compared with spatially nonevolving controls. Furthermore, evolution of these traits is variable, leading to enhanced variance in speed among replicate population expansions. Our results demonstrate that evolutionary processes must be considered alongside demographic ones to better understand and predict range expansions., Spatial structure provides unique opportunities for evolution during range expansions. Here, the authors show experimentally using the red flour beetle, Tribolium castaneum, that dispersal and growth can evolve through spatial processes, increasing expansion speed and its variance.

Anthropogenic pressures on the global biome are causing widespread species declines and extinctions. Assessing the extinction risk faced by individual species is a critical first step in combating this trend. However, we lack high quality demographic data to do so for the vast majority of plant and animal species. We present an efficient modeling approach to estimate extinction risk based on a statistical framework from the mark-recapture literature. We assessed the model's performance using a combination of simulated data, results from a protist microcosm experiment, and data from a long-term, large-scale habitat fragmentation experiment in southeastern Australia. Simulation experiments showed the model is robust to missing data as well as biological processes not included explicitly in the model's assumptions. Fitting the model to data from the protist experiment yielded accurate predictions of the regional extinction dynamics observed in the system, even with a relatively low level of replication. Finally, the model was able to accurately predict the observed dynamics in the habitat fragmentation experiment. The model provides a robust and accurate method to evaluate a species' extinction risk. Since it only requires presence/absence data, applies to a wide range of survey designs, and allows for observational uncertainty and missing data, it can be applied to many datasets that existing models cannot accommodate. For these reasons, the model should be useful in conservation settings.

International audience; The risk of extinction faced by small isolated populations in changing environments can be reduced by rapid adaptation and subsequent growth to larger, less vulnerable sizes. Whether this process, called evolutionary rescue, is able to reduce extinction risk and sustain population growth over multiple generations is largely unknown. To understand the consequences of adaptive evolution as well as maladaptive processes in small isolated populations, we subjected experimental Tribolium castaneum populations founded with 10 or 40 individuals to novel environments, one more favorable, and one resource poor, and either allowed evolution, or constrained it by replacing individuals one-for-one each generation with those from a large population maintained in the natal environment. Replacement individuals spent one generation in the target novel environment before use to standardize effects due to the parental environment. After eight generations we mixed a subset of surviving populations to facilitate admixture, allowing us to estimate drift load by comparing performance of mixed to unmixed groups. Evolving populations had reduced extinction rates, and increased population sizes in the first four to five generations compared to populations where evolution was constrained. Performance of evolving populations subsequently declined. Admixture restored their performance, indicating high drift load that may have overwhelmed the beneficial effects of adaptation in evolving populations. Our results indicate that evolution may quickly reduce extinction risk and increase population sizes, but suggest that relying solely on adaptation from standing genetic variation may not provide long-term benefits to small isolated populations of diploid sexual species, and that active management facilitating gene flow may be necessary for longer term persistence.

Two formats of plant biodiversity labs were evaluated: a learning cycle format and an expository format. Each had a prelab, a hands-on lab, and a write-to-learn postlab. Bloom's lower- and higher-order cognition and attitudes were assessed. Results showed that the two styles had different costs and benefits. Evidence indicates that a blended style may be best., We compared learning cycle and expository formats for teaching about plant biodiversity in an inquiry-oriented university biology lab class (n = 465). Both formats had preparatory lab activities, a hands-on lab, and a postlab with reflection and argumentation. Learning was assessed with a lab report, a practical quiz in lab, and a multiple-choice exam in the concurrent lecture. Attitudes toward biology and treatments were also assessed. We used linear mixed-effect models to determine impacts of lab style on lower-order cognition (LO) and higher-order cognition (HO) based on Bloom's taxonomy. Relative to the expository treatment, the learning cycle treatment had a positive effect on HO and a negative effect on LO included in lab reports; a positive effect on transfer of LO from the lab report to the quiz; negative impacts on LO quiz performance and on attitudes toward the lab; and a higher degree of perceived difficulty. The learning cycle treatment had no influence on transfer of HO from lab report to quiz or exam; quiz performance on HO questions; exam performance on LO and HO questions; and attitudes toward biology as a science. The importance of LO as a foundation for HO relative to these lab styles is addressed.

Aim Evidence linking the accumulation of exotic species to the suppression of native diversity is equivocal, often relying on data from studies that have used different methods. Plot-level studies often attribute inverse relationships between native and exotic diversity to competition, but regional abiotic filters, including anthropogenic influences, can produce similar patterns. We seek to test these alternatives using identical scale-dependent sampling protocols in multiple grasslands on two continents. Location Thirty-two grassland sites in North America and Australia. Methods We use multiscale observational data, collected identically in grain and extent at each site, to test the association of local and regional factors with the plot-level richness and abundance of native and exotic plants. Sites captured environmental and anthropogenic gradients including land-use intensity, human population density, light and soil resources, climate and elevation. Site selection occurred independently of exotic diversity, meaning that the numbers of exotic species varied randomly thereby reducing potential biases if only highly invaded sites were chosen. Results Regional factors associated directly or indirectly with human activity had the strongest associations with plot-level diversity. These regional drivers had divergent effects: urban-based economic activity was associated with high exotic : native diversity ratios; climate- and landscape-based indicators of lower human population density were associated with low exotic : native ratios. Negative correlations between plot-level native and exotic diversity, a potential signature of competitive interactions, were not prevalent; this result did not change along gradients of productivity or heterogeneity. Main conclusion We show that plot-level diversity of native and exotic plants are more consistently associated with regional-scale factors relating to urbanization and climate suitability than measures indicative of competition. These findings clarify the long-standing difficulty in resolving drivers of exotic diversity using single-factor mechanisms, suggesting that multiple interacting anthropogenic-based processes best explain the accumulation of exotic diversity in modern landscapes.

Despite the ubiquity of bilateral symmetry among animals, a long-standing mystery centers on why parasites that infect paired organs often do so non-randomly. Examples from diverse host and parasite taxa continue to accumulate, yet little is known about their causes or implications for host – parasite fi tness. We combined fi eld surveys, experimental infections, and parasite choice assays to evaluate both competing explanations for – and consequences of – asymmetric infections of amphibian kidneys by echinostome trematodes, which are widespread and potentially pathogenic infections of larval amphibians. Samples from 6001 hosts representing 26 species indicated that echinostome infections exhibit a consistent, right-kidney bias, with ∼ 62% of parasites in the right kidney. Th is pattern could not be explained by variation in kidney size or total infection. Experimental infections of three anuran species reproduced this pattern, with 64% of infections in the right kidney, and indicated it was not the result of diff erential host or parasite mortality. Based on sequential infection experiments and parasite choice assays, we further showed that earlier infections did not aff ect the distribution of subsequently colonizing parasites and that echinostome cercariae followed host-derived cues rather than exhibiting congenital ‘ sidedness ’ . We advance the hypothesis that variation in the position of the right kidney along the anterior – posterior axis controls cue strength in the right nephric duct and thus determines parasite encystment. Correspondingly, anatomical measurements from a subset of larval amphibian hosts revealed that the relative position of the right kidney explained 83% of the variation in infection bias, with no additional contributions associated with kidney volume or host size. We also show that the degree of right-kidney bias associated positively with host growth in experiments. Morphological asymmetries could therefore function as a unique form of tolerance to mitigate the consequences of infection, despite the oft-cited costs of asymmetry for mate selection and enemy vulnerability.

The establishment of commercial tree plantations is a common cause of habitat fragmentation globally, yet the influence of this land use on plant species density in fragmented native forests requires further understanding. Theory predicts species density will be highest in large areas of habitat, and should decline as area is reduced, but whether these effects are scale-dependent is largely unknown. We assessed plant species density (total, common and uncommon species) in experimentally fragmented eucalypt forest remnants (0.25, 0.88, 3.06 ha) surrounded by a pine plantation, at three spatial scales using nested quadrats. Specifically, we consider how plant species density varies across three different sized fragments, and whether the response of species density is magnified in common or uncommon species. Species density in small fragments was higher than continuous forest for all species groupings at the smallest spatial scale (1 m2), and for total and common species at the next smallest spatial scale (16 m2). No species groupings responded to reductions in habitat area at the largest spatial scale (144 m2). We suggest that pine plantations may cause higher species density in small fragments via two mechanisms, either by allowing species to infill unoccupied areas within small fragments, or by buffering small fragments from species losses during a severe and prolonged drought. In both cases we suspect reduced moisture stress (e.g. increased soil moisture, higher shading and reduced temperature) in small fragments has led to the observed changes in species density.

Although recent work has shown that both deterministic and stochastic processes are important in structuring microbial communities, the factors that affect the relative contributions of niche and neutral processes are poorly understood. The macrobiological literature indicates that ecological disturbances can influence assembly processes. Thus, we sampled bacterial communities at 4 and 16 weeks following a wildfire and used null deviation analysis to examine the role that time since disturbance has in community assembly. Fire dramatically altered bacterial community structure and diversity as well as soil chemistry for both time-points. Community structure shifted between 4 and 16 weeks for both burned and unburned communities. Community assembly in burned sites 4 weeks after fire was significantly more stochastic than in unburned sites. After 16 weeks, however, burned communities were significantly less stochastic than unburned communities. Thus, we propose a three-phase model featuring shifts in the relative importance of niche and neutral processes as a function of time since disturbance. Because neutral processes are characterized by a decoupling between environmental parameters and community structure, we hypothesize that a better understanding of community assembly may be important in determining where and when detailed studies of community composition are valuable for predicting ecosystem function.

Plant growth can be limited by resource acquisition and defence against consumers, leading to contrasting trade-off possibilities. The competition-defence hypothesis posits a trade-off between competitive ability and defence against enemies (e.g. herbivores and pathogens). The growth-defence hypothesis suggests that strong competitors for nutrients are also defended against enemies, at a cost to growth rate. We tested these hypotheses using observations of 706 plant populations of over 500 species before and following identical fertilisation and fencing treatments at 39 grassland sites worldwide. Strong positive covariance in species responses to both treatments provided support for a growth-defence trade-off: populations that increased with the removal of nutrient limitation (poor competitors) also increased following removal of consumers. This result held globally across 4 years within plant life-history groups and within the majority of individual sites. Thus, a growth-defence trade-off appears to be the norm, and mechanisms maintaining grassland biodiversity may operate within this constraint.

Pan et al . claim that our results actually support a strong linear positive relationship between productivity and richness, whereas Fridley et al . contend that the data support a strong humped relationship. These responses illustrate how preoccupation with bivariate patterns distracts from a deeper understanding of the multivariate mechanisms that control these important ecosystem properties.

Four metacommunity paradigms-usually called neutral, species sorting, mass effects, and patch dynamics, respectively-are widely used for empirical and theoretical studies of spatial community dynamics. The paradigm framework highlights key ecological mechanisms operating in metacommunities, such as dispersal limitation, competition-colonization tradeoffs, or species equivalencies. However, differences in coexistence mechanisms between the paradigms and in situations with combined influences of multiple paradigms are not well understood. Here, we create a common model for competitive metacommunities, with unique parameterizations for each metacommunity paradigm and for scenarios with multiple paradigms operating simultaneously. We derive analytical expressions for the strength of Chesson's spatial coexistence mechanisms and quantify these for each paradigm via simulation. For our model, fitness-density covariance, a concentration effect measuring the importance of intraspecific aggregation of individuals, is the dominant coexistence mechanism in all three niche-based metacommunity paradigms. Increased dispersal between patches erodes intraspecific aggregation, leading to lower coexistence strength in the mass effects paradigm compared to species sorting. Our analysis demonstrates the potential importance of aggregation of individuals (fitness-density covariance) over co-variation in abiotic environments and competition between species (the storage effect), as fitness-density covariance can be stronger than the storage effect and is the sole stabilizing mechanism in the patch dynamics paradigm. As expected, stable coexistence does not occur in the neutral paradigm, which requires species to be equal and emphasizes the role of stochasticity. We show that stochasticity also plays an important role in niche-structured metacommunities by altering coexistence strength. We conclude that Chesson's spatial coexistence mechanisms provide a flexible framework for comparing metacommunities of varying complexity.

Many ecosystems worldwide are dominated by introduced plant species, leading to loss of biodiversity and ecosystem function. A common but rarely tested assumption is that these plants are more abundant in introduced vs. native communities, because ecological or evolutionary-based shifts in populations underlie invasion success. Here, data for 26 herbaceous species at 39 sites, within eight countries, revealed that species abundances were similar at native (home) and introduced (away) sites – grass species were generally abundant home and away, while forbs were low in abundance, but more abundant at home. Sites with six or more of these species had similar community abundance hierarchies, suggesting that suites of introduced species are assembling similarly on different continents. Overall, we found that substantial changes to populations are not necessarily a pre-condition for invasion success and that increases in species abundance are unusual. Instead, abundance at home predicts abundance away, a potentially useful additional criterion for biosecurity programmes.

Terrestrial ecosystems of sub-Antarctic islands are particularly sensitive to global and local human impacts, including climate change and species invasion. Invertebrates form a central component of these ecosystems. We conducted a stratified survey of 60 sites on sub-Antarctic Heard Island and used Poisson regression models to describe the spatial distribution and abundance of five of the ten free-living species captured. Acari and Collembola were not considered. Five species were not caught in traps in sufficient numbers to model. The distributions of species were described by altitude, vegetation type and aspect. The resulting distribution models can be used to both monitor and predict the effects of climate change and species invasion on this unique and valuable ecosystem.

The expansion of the artificial water-point network and livestock grazing in arid and semi-arid Australia has significantly increased access to water by water limited herbivores and thus has potential to seriously negatively affect the unique endemic flora and fauna. We examined the effects of the expansion of the water-point network on the arid and semi-arid zone bird community, using data from the Atlas of Australia bird surveys of 1977–1981 and 1996–2001. We examined whether traits of species could be used to uncover the critical ecological processes altered by this land use change. We detected large scale declines in individual species. Species reliant on water for nesting and feeding declined more than other groups in the presence of high water-point density, likely through direct effects of livestock degrading habitat of both natural and artificial water points. The arid zone has no natural hoofed animals and livestock have significant impacts. Species that forage and nest on the ground also showed large declines, likely because of trampling and removal of vegetation by livestock and potentially through the indirect effect of water limited predators, dingos and foxes, expanding their ranges in response to the expanding network of water points in the landscape. This result was also apparent at large spatial scales so that these local-scale responses to water points translate into continental-scale population declines. Using traits of species to understand declines of bird species helped us to uncover which critical changes associated with the expansion of the water-point network most affected the bird community and therefore which species are most at risk.

Limits to Invasion Predictions Using mathematical models linked to experiments on laboratory insect populations, Melbourne and Hastings (p. 1536 ) show that current models seriously underestimate variability in spatial spread. Genetic founder effects resulting from small population sizes are probably responsible for the surprising increase in variability and for increased uncertainty of ecological forecasts. Thus, even without exogenous sources of variability such as weather, fundamental uncertainties arise biologically from inherent differences among individuals and small population sizes.

Disturbance is an important factor influencing diversity patterns. Ecological theory predicts that diversity peaks at intermediate levels of disturbance, but this pattern is not present in a majority of empirical tests and can be influenced by the level of ecosystem productivity. We experimentally tested the effects of disturbance on diversity and show that species' autecological traits and community relations predicted species loss. We found that - alone or in concert - increasing disturbance intensity or frequency, or decreasing productivity, reduced diversity. Our species did not exhibit a clear competition-colonization trade-off, and intrinsic growth rate was a more important predictor of response to disturbance and productivity than measures of competitive ability. Furthermore, competitive ability was more important in predicting responses when, in addition to killing individuals, disturbance returned nutrients to the ecosystem. Our results demonstrate that species' traits can help resolve conflicting patterns in the response of diversity to disturbance and productivity.