Microbe-mediated plant–soil feedback and its roles in a changing world

Plants affect soil conditions, which in turn alter plant growth and interspecific competition, forming plant?Csoil feedback (PSF) systems. PSF is a good example of bidirectional interactions between biomes and the non-living environments, acting as a major driving force of community structure and ecosystem function. Among the major types of PSF mediated by various soil components, there are many holes in our knowledge of the interactions between PSF mediated by plant species-specific litter and PSF mediated by soil microbes. Here I discuss the role of the functional diversity of microbial decomposers in litter-mediated PSF and also propose new hypotheses on the role of microbial diversity in PSF mediated by pathogenic and mutualistic soil microbes. I also review how PSF interacts with human-induced environmental change, i.e., direct drivers of change in the ecosystem (e.g. climate change and the invasion of alien species). Many authors have suggested that the impact of alien plant species on ecosystems is mediated by PSF, which also interacts with other direct drivers, such as climate change. Using a simple model of litter-mediated PSF with microbial decomposers, I confirm that the interactions between PSF and other direct drivers affect the invasion process of alien species. The model also demonstrates that the functional diversity of microbial decomposers accelerates or decelerates the speed of the invasion depending on the environmental change scenarios. Further theoretical and empirical studies are needed to derive general predictions on how exogenous environmental change induced by human activities alters communities and ecosystems through disturbance or modification of endogenous community?Cecosystem interactions, such as the functioning of PSF.     

Continental‐scale determinants of population trends in European amphibians and reptiles

The continuous decline of biodiversity is determined by the complex and joint effects of multiple environmental drivers. Still, a large part of past global change studies reporting and explaining biodiversity trends have focused on a single driver. Therefore, we are often unable to attribute biodiversity changes to different drivers, since a multivariable design is required to disentangle joint effects and interactions. In this work, we used a meta‐regression within a Bayesian framework to analyze 843 time series of population abundance from 17 European amphibian and reptile species over the last 45 years. We investigated the relative effects of climate change, alien species, habitat availability, and habitat change in driving trends of population abundance over time, and evaluated how the importance of these factors differs across species. A large number of populations (54%) declined, but differences between species were strong, with some species showing positive trends. Populations declined more often in areas with a high number of alien species, and in areas where climate change has caused loss of suitability. Habitat features showed small variation over the last 25 years, with an average loss of suitable habitat of 0.1%/year per population. Still, a strong interaction between habitat availability and the richness of alien species indicated that the negative impact of alien species was particularly strong for populations living in landscapes with less suitable habitat. Furthermore, when excluding the two commonest species, habitat loss was the main correlate of negative population trends for the remaining species. By analyzing trends for multiple species across a broad spatial scale, we identify alien species, climate change, and habitat changes as the major drivers of European amphibian and reptile decline.     

Ecosystem feedbacks and cascade processes: understanding their role in the responses of Arctic and alpine ecosystems to environmental change

Global environmental change, related to climate change and the deposition of airborne N‐containing contaminants, has already resulted in shifts in plant community composition among plant functional types in Arctic and temperate alpine regions. In this paper, we review how key ecosystem processes will be altered by these transformations, the complex biological cascades and feedbacks that might result, and some of the potential broader consequences for the earth system. Firstly, we consider how patterns of growth and allocation, and nutrient uptake, will be altered by the shifts in plant dominance. The ways in which these changes may disproportionately affect the consumer communities, and rates of decomposition, are then discussed. We show that the occurrence of a broad spectrum of plant growth forms in these regions (from cryptogams to deciduous and evergreen dwarf shrubs, graminoids and forbs), together with hypothesized low functional redundancy, will mean that shifts in plant dominance result in a complex series of biotic cascades, couplings and feedbacks which are supplemental to the direct responses of ecosystem components to the primary global change drivers. The nature of these complex interactions is highlighted using the example of the climate‐driven increase in shrub cover in low‐Arctic tundra, and the contrasting transformations in plant functional composition in mid‐latitude alpine systems. Finally, the potential effects of the transformations on ecosystem properties and processes that link with the earth system are reviewed. We conclude that the effects of global change on these ecosystems, and potential climate‐change feedbacks, cannot be predicted from simple empirical relationships between processes and driving variables. Rather, the effects of changes in species distributions and dominances on key ecosystem processes and properties must also be considered, based upon best estimates of the trajectories of key transformations, their magnitude and rates of change.     

A general multi-trait-based framework for studying the effects of biodiversity on ecosystem functioning

Environmental change is as multifaceted as are the species and communities that respond to these changes. Current theoretical approaches to modeling ecosystem response to environmental change often deal only with single environmental drivers or single species traits, simple ecological interactions, and/or steady states, leading to concern about how accurately these approaches will capture future responses to environmental change in real biological systems. To begin addressing this issue, we generalize a previous trait-based framework to incorporate aspects of frequency dependence, functional complementarity, and the dynamics of systems composed of species that are defined by multiple traits that are tied to multiple environmental drivers. The framework is particularly well suited for analyzing the role of temporal environmental fluctuations in maintaining trait variability and the resultant effects on community response to environmental change. Using this framework, we construct simple models to investigate two ecological problems. First, we show how complementary resource use can significantly enhance the nutrient uptake of plant communities through two different mechanisms related to increased productivity (over-yielding) and larger trait variability. Over-yielding is a hallmark of complementarity and increases the total biomass of the community and, thus, the total rate at which nutrients are consumed. Trait variability also increases due to the lower levels of competition associated with complementarity, thus speeding up the rate at which more efficient species emerge as conditions change. Second, we study systems in which multiple environmental drivers act on species defined by multiple, correlated traits. We show that correlations in these systems can increase trait variability within the community and again lead to faster responses to environmental change. The methodological advances provided here will apply to almost any function that relates species traits and environmental drivers to growth, and should prove useful for studying the effects of climate change on the dynamics of biota.     

Disentangling how climate change can affect an aquatic food web by combining multiple experimental approaches

Predicting the biological effects of climate change presents major challenges due to the interplay of potential biotic and abiotic mechanisms. Climate change can create unexpected outcomes by altering species interactions, and uncertainty over the ability of species to develop in situ tolerance or track environmental change further hampers meaningful predictions. As multiple climatic variables shift in concert, their potential interactions further complicate ecosystem responses. Despite awareness of these complexities, we still lack controlled experiments that manipulate multiple climatic stressors, species interactions, and prior exposure of species to future climatic conditions. Particularly studies that address how changes in water availability interact with other climatic stressors to affect aquatic ecosystems are still rare. Using aquatic insect communities of Neotropical tank bromeliads, we combined controlled manipulations of drought length and species interactions with a space‐for‐time transplant (lower elevations represent future climate) and a common garden approach. Manipulating drought length and experiment elevation revealed that adverse effects of drought were amplified at the warmer location, highlighting the potential of climatic stressors to synergistically affect communities. Manipulating the presence of omnivorous tipulid larvae showed that negative interactions from tipulids, presumably from predation, arose under drought, and were stronger at the warmer location, stressing the importance of species interactions in mediating community responses to climate change. The common garden treatments revealed that prior community exposure to potential future climatic conditions did not affect the outcome. In this powerful experiment, we demonstrated how complexities arise from the interplay of biotic and abiotic mechanisms of climate change. We stress that single species can steer ecological outcomes, and suggest that focusing on such disproportionately influential species may improve attempts at making meaningful predictions of climate change impacts on food webs.     

Global change and species interactions in terrestrial ecosystems

The main drivers of global environmental change (CO₂ enrichment, nitrogen deposition, climate, biotic invasions and land use) cause extinctions and alter species distributions, and recent evidence shows that they exert pervasive impacts on various antagonistic and mutualistic interactions among species. In this review, we synthesize data from 688 published studies to show that these drivers often alter competitive interactions among plants and animals, exert multitrophic effects on the decomposer food web, increase intensity of pathogen infection, weaken mutualisms involving plants, and enhance herbivory while having variable effects on predation. A recurrent finding is that there is substantial variability among studies in both the magnitude and direction of effects of any given GEC driver on any given type of biotic interaction. Further, we show that higher order effects among multiple drivers acting simultaneously create challenges in predicting future responses to global environmental change, and that extrapolating these complex impacts across entire networks of species interactions yields unanticipated effects on ecosystems. Finally, we conclude that in order to reliably predict the effects of GEC on community and ecosystem processes, the greatest single challenge will be to determine how biotic and abiotic context alters the direction and magnitude of GEC effects on biotic interactions.     

Vegetation change at high elevation: scale dependence and interactive effects on Niwot Ridge

Background: High-elevation mountain systems may be particularly responsive to climate change.

Aims: Here we investigate how changes along elevation gradients in mountain systems can aid in predicting vegetation distributional changes in time, focusing on how changing climatic controls affect meso-scale transitions at the lower and upper boundaries of alpine vegetation (with forest and subnival zones, respectively) as well as micro-scale transitions among plant communities within the alpine belt. We focus on climate-related drivers, particularly in relation to climate change, but also consider how species interactions, dispersal and responses to disturbance may influence plant responses to these abiotic drivers.

Results: Empirical observations and experimental studies indicate that changing climatic controls influence both meso-scale transitions at the upper and lower boundaries of alpine vegetation and micro-scale transitions among plant communities within tundra. Micro-scale heterogeneity appears to buffer response in many cases, while interactions between climate and other changes may often accelerate change.

Conclusions: Interactions with microtopography and larger edaphic gradients have the capacity to both facilitate rapid changes and reinforce stability, and that these interactions will affect the responsiveness of vegetation to climate change at different spatial scales.     

Simulating plant invasion dynamics in mountain ecosystems under global change scenarios

Across the globe, invasive alien species cause severe environmental changes, altering species composition and ecosystem functions. So far, mountain areas have mostly been spared from large‐scale invasions. However, climate change, land‐use abandonment, the development of tourism and the increasing ornamental trade will weaken the barriers to invasions in these systems. Understanding how alien species will react and how native communities will influence their success is thus of prime importance in a management perspective. Here, we used a spatially and temporally explicit simulation model to forecast invasion risks in a protected mountain area in the French Alps under future conditions. We combined scenarios of climate change, land‐use abandonment and tourism‐linked increases in propagule pressure to test if the spread of alien species in the region will increase in the future. We modelled already naturalized alien species and new ornamental plants, accounting for interactions among global change components, and also competition with the native vegetation. Our results show that propagule pressure and climate change will interact to increase overall species richness of both naturalized aliens and new ornamentals, as well as their upper elevational limits and regional range‐sizes. Under climate change, woody aliens are predicted to more than double in range‐size and herbaceous species to occupy up to 20% of the park area. In contrast, land‐use abandonment will open new invasion opportunities for woody aliens, but decrease invasion probability for naturalized and ornamental alien herbs as a consequence of colonization by native trees. This emphasizes the importance of interactions with the native vegetation either for facilitating or potentially for curbing invasions. Overall, our work highlights an additional and previously underestimated threat for the fragile mountain flora of the Alps already facing climate changes, land‐use transformations and overexploitation by tourism.     

Animal behaviour shapes the ecological effects of ocean acidification and warming: moving from individual to community‐level responses

Biological communities are shaped by complex interactions between organisms and their environment as well as interactions with other species. Humans are rapidly changing the marine environment through increasing greenhouse gas emissions, resulting in ocean warming and acidification. The first response by animals to environmental change is predominantly through modification of their behaviour, which in turn affects species interactions and ecological processes. Yet, many climate change studies ignore animal behaviour. Furthermore, our current knowledge of how global change alters animal behaviour is mostly restricted to single species, life phases and stressors, leading to an incomplete view of how coinciding climate stressors can affect the ecological interactions that structure biological communities. Here, we first review studies on the effects of warming and acidification on the behaviour of marine animals. We demonstrate how pervasive the effects of global change are on a wide range of critical behaviours that determine the persistence of species and their success in ecological communities. We then evaluate several approaches to studying the ecological effects of warming and acidification, and identify knowledge gaps that need to be filled, to better understand how global change will affect marine populations and communities through altered animal behaviours. Our review provides a synthesis of the far‐reaching consequences that behavioural changes could have for marine ecosystems in a rapidly changing environment. Without considering the pervasive effects of climate change on animal behaviour we will limit our ability to forecast the impacts of ocean change and provide insights that can aid management strategies.     

Climate change and biological invasions: evidence,expectations, and response options

A changing climate may directly or indirectly influence biological invasions by altering the likelihood of introduction or establishment, as well as modifying the geographic range, environmental impacts, economic costs or management of alien species. A comprehensive assessment of empirical and theoretical evidence identified how each of these processes is likely to be shaped by climate change for alien plants, animals and pathogens in terrestrial, freshwater and marine environments of Great Britain. The strongest contemporary evidence for the potential role of climate change in the establishment of new alien species is for terrestrial arthropods, as a result of their ectothermic physiology, often high dispersal rate and their strong association with trade as well as commensal relationships with human environments. By contrast, there is little empirical support for higher temperatures increasing the rate of alien plant establishment due to the stronger effects of residence time and propagule pressure. The magnitude of any direct climate effect on the number of new alien species will be small relative to human‐assisted introductions driven by socioeconomic factors. Casual alien species (sleepers) whose population persistence is limited by climate are expected to exhibit greater rates of establishment under climate change assuming that propagule pressure remains at least at current levels. Surveillance and management targeting sleeper pests and diseases may be the most cost‐effective option to reduce future impacts under climate change. Most established alien species will increase their distribution range in Great Britain over the next century. However, such range increases are very likely be the result of natural expansion of populations that have yet to reach equilibrium with their environment, rather than a direct consequence of climate change. To assess the potential realised range of alien species will require a spatially explicit approach that not only integrates bioclimatic suitability and population‐level demographic rates but also simulation of landscape‐level processes (e.g. dispersal, land‐use change, host/habitat distribution, non‐climatic edaphic constraints). In terms of invasive alien species that have known economic or biodiversity impacts, the taxa that are likely to be the most responsive are plant pathogens and insect pests of agricultural crops. However, the extent to which climate adaptation strategies lead to new crops, altered rotations, and different farming practices (e.g. irrigation, fertilization) will all shape the potential agricultural impacts of alien species. The greatest uncertainty in the effects of climate change on biological invasions exists with identifying the future character of new species introductions and predicting ecosystem impacts. Two complementary strategies may work under these conditions of high uncertainty: (i) prioritise ecosystems in terms of their perceived vulnerability to climate change and prevent ingress or expansion of alien species therein that may exacerbate problems; (ii) target those ecosystem already threatened by alien species and implement management to prevent the situation deteriorating under climate change.     

Plant-plant interactions and environmental change

Natural systems are being subjected to unprecedented rates of change and unique pressures from a combination of anthropogenic environmental change drivers. Plant-plant interactions are an important part of the mechanisms governing the response of plant species and communities to these drivers. For example, competition plays a central role in mediating the impacts of atmospheric nitrogen deposition, increased atmospheric carbon dioxide concentrations, climate change and invasive nonnative species. Other plant-plant interaction processes are also being recognized as important factors in determining the impacts of environmental change, including facilitation and evolutionary processes associated with plant-plant interactions. However, plant-plant interactions are not the only factors determining the response of species and communities to environmental change drivers - their activity must be placed within the context of the wide range of factors that regulate species, communities and ecosystems. A major research challenge is to understand when plant-plant interactions play a key role in regulating the impact of environmental change drivers, and the type of role that plant-plant interactions play. Although this is a considerable challenge, some areas of current research may provide the starting point to achieving these goals, and should be pursued through large-scale, integrated, multisite experiments.     

Conceptualising the interactive effects of climate change and biological invasions on subarctic freshwater fish

Climate change and species invasions represent key threats to global biodiversity. Subarctic freshwaters are sentinels for understanding both stressors because the effects of climate change are disproportionately strong at high latitudes and invasion of temperate species is prevalent. Here, we summarize the environmental effects of climate change and illustrate the ecological responses of freshwater fishes to these effects, spanning individual, population, community and ecosystem levels. Climate change is modifying hydrological cycles across atmospheric, terrestrial and aquatic components of subarctic ecosystems, causing increases in ambient water temperature and nutrient availability. These changes affect the individual behavior, habitat use, growth and metabolism, alter population spawning and recruitment dynamics, leading to changes in species abundance and distribution, modify food web structure, trophic interactions and energy flow within communities and change the sources, quantity and quality of energy and nutrients in ecosystems. Increases in temperature and its variability in aquatic environments underpin many ecological responses; however, altered hydrological regimes, increasing nutrient inputs and shortened ice cover are also important drivers of climate change effects and likely contribute to context‐dependent responses. Species invasions are a complex aspect of the ecology of climate change because the phenomena of invasion are both an effect and a driver of the ecological consequences of climate change. Using subarctic freshwaters as an example, we illustrate how climate change can alter three distinct aspects of species invasions: (1) the vulnerability of ecosystems to be invaded, (2) the potential for species to spread and invade new habitats, and (3) the subsequent ecological effects of invaders. We identify three fundamental knowledge gaps focused on the need to determine (1) how environmental and landscape characteristics influence the ecological impact of climate change, (2) the separate and combined effects of climate and non‐native invading species and (3) the underlying ecological processes or mechanisms responsible for changes in patterns of biodiversity.     

Biodiversity change under adaptive community dynamics

Compositional change is a ubiquitous response of ecological communities to environmental drivers of global change, but is often regarded as evidence of declining “biotic integrity” relative to historical baselines. Adaptive compositional change, however, is a foundational idea in evolutionary biology, whereby changes in gene frequencies within species boost population-level fitness, allowing populations to persist as the environment changes. Here, we present an analogous idea for ecological communities based on core concepts of fitness and selection. Changes in community composition (i.e., frequencies of genetic differences among species) in response to environmental change should normally increase the average fitnessof community members. We refer to compositional changes that improve the functional match, or “fit,” between organisms' traits and their environment as adaptive community dynamics. Environmental change (e.g., land-use change) commonly reduces the fit between antecedent communities and new environments. Subsequent change in community composition in response to environmental changes, however, should normally increase community-level fit, as the success of at least some constituent species increases. We argue that adaptive community dynamics are likely to improve or maintain ecosystem function (e.g., by maintaining productivity). Adaptive community responses may simultaneously produce some changes that are considered societally desirable (e.g., increased carbon storage) and others that are undesirable (e.g., declines of certain species), just as evolutionary responses within species may be deemed desirable (e.g., evolutionary rescue of an endangered species) or undesirable (e.g., enhanced virulence of an agricultural pest). When assessing possible management interventions, it is important to distinguish between drivers of environmental change (e.g., undesired climate warming) and adaptive community responses, which may generate some desirable outcomes. Efforts to facilitate, accept, or resist ecological change require separate consideration of drivers and responses, and may highlight the need to reconsider preferences for historical baseline communities over communities that are better adapted to the new conditions.     

Interactive effects of habitat modification and species invasion on native species decline

Different components of global environmental change are often studied and managed independently, but mounting evidence points towards complex non-additive interaction effects between drivers of native species decline. Using the example of interactions between land-use change and biotic exchange, we develop an interpretive framework that will enable global change researchers to identify and discriminate between major interaction pathways. We formalise a distinction between numerically mediated versus functionally moderated causal pathways. Despite superficial similarity of their effects, numerical and functional pathways stem from fundamentally different mechanisms of action and have fundamentally different consequences for conservation management. Our framework is a first step toward building a better quantitative understanding of how interactions between drivers might mitigate or exacerbate the net effects of global environmental change on biotic communities in the future.     

Habitat disturbance selects against both small and large species across varying climates

Global extinction drivers, including habitat disturbance and climate change, are thought to affect larger species more than smaller species. However, it is unclear if such drivers interact to affect assemblage body size distributions. We asked how these two key global change drivers differentially affect the interspecific size distributions of ants, one of the most abundant and ubiquitous animal groups on earth. We also asked whether there is evidence of synergistic interactions and whether effects are related to species’ trophic roles. We generated a global dataset on ant body size from 333 local ant assemblages collected by the authors across a broad range of climates and in disturbed and undisturbed habitats. We used head length (range: 0.22–4.55 mm) as a surrogate of body size and classified species to trophic groups. We used generalized linear models to test whether body size distributions changed with climate and disturbance, independent of species richness. Our analysis yielded three key results: 1) climate and disturbance showed independent associations with body size; 2) assemblages included more small species in warmer climates and fewer large species in wet climates; and 3) both the largest and smallest species were absent from disturbed ecosystems, with predators most affected in both cases. Our results indicate that temperature, precipitation and disturbance have differing effects on the body size distributions of local communities, with no evidence of synergistic interactions. Further, both large and small predators may be vulnerable to global change, particularly through habitat disturbance.     

Coevolution and the Effects of Climate Change on Interacting Species

Background

Recent studies suggest that environmental changes may tip the balance between interacting species, leading to the extinction of one or more species. While it is recognized that evolution will play a role in determining how environmental changes directly affect species, the interactions among species force us to consider the coevolutionary responses of species to environmental changes.

Methodology/Principle Findings

We use simple models of competition, predation, and mutualism to organize and synthesize the ways coevolution modifies species interactions when climatic changes favor one species over another. In cases where species have conflicting interests (i.e., selection for increased interspecific interaction strength on one species is detrimental to the other), we show that coevolution reduces the effects of climate change, leading to smaller changes in abundances and reduced chances of extinction. Conversely, when species have nonconflicting interests (i.e., selection for increased interspecific interaction strength on one species benefits the other), coevolution increases the effects of climate change.

Conclusions/Significance

Coevolution sets up feedback loops that either dampen or amplify the effect of environmental change on species abundances depending on whether coevolution has conflicting or nonconflicting effects on species interactions. Thus, gaining a better understanding of the coevolutionary processes between interacting species is critical for understanding how communities respond to a changing climate. We suggest experimental methods to determine which types of coevolution (conflicting or nonconflicting) drive species interactions, which should lead to better understanding of the effects of coevolution on species adaptation. Conducting these experiments across environmental gradients will test our predictions of the effects of environmental change and coevolution on ecological communities.     

Interspecific interactions affect species and community responses to climate shifts

The response of individual species to climate change may alter the composition and dynamics of communities. Here, we show that the impacts of environmental change on communities can depend on the nature of the interspecific interactions: mutualistic communities typically respond differently than commensalistic or parasitic communities. We model and analyse the geographic range shifting of metapopulations of two interacting species – a host and an obligate species. Different types of interspecific interactions are implemented by modifying local extinction rates according to the presence/absence of the other species. We distinguish and compare three fundamentally different community types: mutualism, commensalism and parasitism. We find that community dynamics during geographic range shifting critically depends on the type of interspecific interactions. Parasitic interactions exacerbate the negative effect of environmental change whereas mutualistic interactions only partly compensate it. Commensalistic interactions exhibit an intermediate response. Based on these model outcomes, we predict that parasitic species interactions may be more vulnerable to geographic range shifting than commensalistic or mutualistic ones. However, we observe that when climate stabilises following a period of change, the rate of community recovery is largely independent of the type of interspecific interactions. These results emphasize that communities respond delicately to environmental change, and that local interspecific interactions can affect range shifting communities at large spatial scales.     

Erosion of community diversity and stability by herbivore removal under warming

Climate change has the potential to influence the persistence of ecological communities by altering their stability properties. One of the major drivers of community stability is species diversity, which is itself expected to be altered by climate change in many systems. The extent to which climatic effects on community stability may be buffered by the influence of species interactions on diversity is, however, poorly understood because of a paucity of studies incorporating interactions between abiotic and biotic factors. Here, I report results of a 10-year field experiment, the past 7 years of which have focused on effects of ongoing warming and herbivore removal on diversity and stability within the plant community, where competitive species interactions are mediated by exploitation through herbivory. Across the entire plant community, stability increased with diversity, but both stability and diversity were reduced by herbivore removal, warming and their interaction. Within the most species-rich functional group in the community, forbs, warming reduced species diversity, and both warming and herbivore removal reduced the strength of the relationship between diversity and stability. Species interactions, such as exploitation, may thus buffer communities against destabilizing influences of climate change, and intact populations of large herbivores, in particular, may prove important in maintaining and promoting plant community diversity and stability in a changing climate.     

Identifying historical and future global change drivers that place species recovery at risk

Ecosystem management in the face of global change requires understanding how co-occurring threats affect species and communities. Such an understanding allows for effective management strategies to be identified and implemented. An important component of this is differentiating between factors that are within (e.g. invasive predators) or outside (e.g. drought, large wildfires) of a local manager's control. In the global biodiversity hotspot of south-western Australia, small- and medium-sized mammal species are severely affected by anthropogenic threats and environmental disturbances, including invasive predators, fire, and declining rainfall. However, the relative importance of different drivers has not been quantified. We used data from a long-term monitoring program to fit Bayesian state-space models that estimated spatial and temporal changes in the relative abundance of four threatened mammal species: the woylie (Bettongia penicillata), chuditch (Dasyurus geoffroii), koomal (Trichosurus vulpecula) and quenda (Isoodon fusciventor). We then use Bayesian structural equation modelling to identify the direct and indirect drivers of population changes, and scenario analysis to forecast population responses to future environmental change. We found that habitat loss or conversion and reduced primary productivity (caused by rainfall declines) had greater effects on species' spatial and temporal population change than the range of fire and invasive predator (the red fox Vulpes vulpes) management actions observed in the study area. Scenario analysis revealed that a greater extent of severe fire and further rainfall declines predicted under climate change, operating in concert are likely to further reduce the abundance of these species, but may be mitigated partially by invasive predator control. Considering both historical and future drivers of population change is necessary to identify the factors that risk species recovery. Given that both anthropogenic pressures and environmental disturbances can undermine conservation efforts, managers must consider how the relative benefit of conservation actions will be shaped by ongoing global change.     

Separating direct and indirect effects of global change: a population dynamic modeling approach using readily available field data

Two sources of complexity make predicting plant community response to global change particularly challenging. First, realistic global change scenarios involve multiple drivers of environmental change that can interact with one another to produce non‐additive effects. Second, in addition to these direct effects, global change drivers can indirectly affect plants by modifying species interactions. In order to tackle both of these challenges, we propose a novel population modeling approach, requiring only measurements of abundance and climate over time. To demonstrate the applicability of this approach, we model population dynamics of eight abundant plant species in a multifactorial global change experiment in alpine tundra where we manipulated nitrogen, precipitation, and temperature over 7 years. We test whether indirect and interactive effects are important to population dynamics and whether explicitly incorporating species interactions can change predictions when models are forecast under future climate change scenarios. For three of the eight species, population dynamics were best explained by direct effect models, for one species neither direct nor indirect effects were important, and for the other four species indirect effects mattered. Overall, global change had negative effects on species population growth, although species responded to different global change drivers, and single‐factor effects were slightly more common than interactive direct effects. When the fitted population dynamic models were extrapolated under changing climatic conditions to the end of the century, forecasts of community dynamics and diversity loss were largely similar using direct effect models that do not explicitly incorporate species interactions or best‐fit models; however, inclusion of species interactions was important in refining the predictions for two of the species. The modeling approach proposed here is a powerful way of analyzing readily available datasets which should be added to our toolbox to tease apart complex drivers of global change.