Shifting daylength regimes associated with range shifts alter aphid‐parasitoid community dynamics

With climate change leading to poleward range expansion of species, populations are exposed to new daylength regimes along latitudinal gradients. Daylength is a major factor affecting insect life cycles and activity patterns, so a range shift leading to new daylength regimes is likely to affect population dynamics and species interactions; however, the impact of daylength in isolation on ecological communities has not been studied so far. Here, we tested for the direct and indirect effects of two different daylengths on the dynamics of experimental multitrophic insect communities. We compared the community dynamics under “southern” summer conditions of 14.5‐hr daylight to “northern” summer conditions of 22‐hr daylight. We show that food web dynamics indeed respond to daylength with one aphid species (Acyrthosiphon pisum) reaching much lower population sizes at the northern daylength regime compared to under southern conditions. In contrast, in the same communities, another aphid species (Megoura viciae) reached higher population densities under northern conditions. This effect at the aphid level was driven by an indirect effect of daylength causing a change in competitive interaction strengths, with the different aphid species being more competitive at different daylength regimes. Additionally, increasing daylength also increased growth rates in M. viciae making it more competitive under summer long days. As such, the shift in daylength affected aphid population sizes by both direct and indirect effects, propagating through species interactions. However, contrary to expectations, parasitoids were not affected by daylength. Our results demonstrate that range expansion of whole communities due to climate change can indeed change interaction strengths between species within ecological communities with consequences for community dynamics. This study provides the first evidence of daylength affecting community dynamics, which could not be predicted from studying single species separately.     

Predator diversity and environmental change modify the strengths of trophic and nontrophic interactions

Understanding the dependence of species interaction strengths on environmental factors and species diversity is crucial to predict community dynamics and persistence in a rapidly changing world. Nontrophic (e.g. predator interference) and trophic components together determine species interaction strengths, but the effects of environmental factors on these two components remain largely unknown. This impedes our ability to fully understand the links between environmental drivers and species interactions. Here, we used a dynamical modelling framework based on measured predator functional responses to investigate the effects of predator diversity, prey density, and temperature on trophic and nontrophic interaction strengths within a freshwater food web. We found that (i) species interaction strengths cannot be predicted from trophic interactions alone, (ii) nontrophic interaction strengths vary strongly among predator assemblages, (iii) temperature has opposite effects on trophic and nontrophic interaction strengths, and (iv) trophic interaction strengths decrease with prey density, whereas the dependence of nontrophic interaction strengths on prey density is concave up. Interestingly, the qualitative impacts of temperature and prey density on the strengths of trophic and nontrophic interactions were independent of predator identity, suggesting a general pattern. Our results indicate that taking multiple environmental factors and the nonlinearity of density‐dependent species interactions into account is an important step towards a better understanding of the effects of environmental variations on complex ecological communities. The functional response approach used in this study opens new avenues for (i) the quantification of the relative importance of the trophic and nontrophic components in species interactions and (ii) a better understanding how environmental factors affect these interactions and the dynamics of ecological communities.     

Individual phenotypic variation reduces interaction strengths in a consumer–resource system

Natural populations often show variation in traits that can affect the strength of interspecific interactions. Interaction strengths in turn influence the fate of pairwise interacting populations and the stability of food webs. Understanding the mechanisms relating individual phenotypic variation to interaction strengths is thus central to assess how trait variation affects population and community dynamics. We incorporated nonheritable variation in attack rates and handling times into a classical consumer–resource model to investigate how variation may alter interaction strengths, population dynamics, species persistence, and invasiveness. We found that individual variation influences species persistence through its effect on interaction strengths. In many scenarios, interaction strengths decrease with variation, which in turn affects species coexistence and stability. Because environmental change alters the direction and strength of selection acting upon phenotypic traits, our results have implications for species coexistence in a context of habitat fragmentation, climate change, and the arrival of exotic species to native ecosystems.     

Using sensitivity analysis to identify keystone species and keystone links in size‐based food webs

Human‐induced alterations in the birth and mortality rates of species and in the strength of interactions within and between species can lead to changes in the structure and resilience of ecological communities. Recent research points to the importance of considering the distribution of body sizes of species when exploring the response of communities to such perturbations. Here, we present a new size‐based approach for assessing the sensitivity and elasticity of community structure (species equilibrium abundances) and resilience (rate of return to equilibrium) to changes in the intrinsic growth rate of species and in the strengths of species interactions. We apply this approach on two natural systems, the pelagic communities of the Baltic Sea and Lake Vättern, to illustrate how it can be used to identify potential keystone species and keystone links. We find that the keystone status of a species is closely linked to its body size. The analysis also suggests that communities are structurally and dynamically more sensitive to changes in the effects of prey on their consumers than in the effects of consumers on their prey. Moreover, we discuss how community sensitivity analysis can be used to study and compare the fragility of communities with different body size distributions by measuring the mean sensitivity or elasticity over all species or all interaction links in a community. We believe that the community sensitivity analysis developed here holds some promise for identifying species and links that are critical for the structural and dynamic robustness of ecological communities.     

Mutualistic networks: moving closer to a predictive theory

Plant–animal mutualistic networks sustain terrestrial biodiversity and human food security. Global environmental changes threaten these networks, underscoring the urgency for developing a predictive theory on how networks respond to perturbations. Here, I synthesise theoretical advances towards predicting network structure, dynamics, interaction strengths and responses to perturbations. I find that mathematical models incorporating biological mechanisms of mutualistic interactions provide better predictions of network dynamics. Those mechanisms include trait matching, adaptive foraging, and the dynamic consumption and production of both resources and services provided by mutualisms. Models incorporating species traits better predict the potential structure of networks (fundamental niche), while theory based on the dynamics of species abundances, rewards, foraging preferences and reproductive services can predict the extremely dynamic realised structures of networks, and may successfully predict network responses to perturbations. From a theoretician's standpoint, model development must more realistically represent empirical data on interaction strengths, population dynamics and how these vary with perturbations from global change. From an empiricist's standpoint, theory needs to make specific predictions that can be tested by observation or experiments. Developing models using short‐term empirical data allows models to make longer term predictions of community dynamics. As more longer term data become available, rigorous tests of model predictions will improve.     

Evolution in plant populations as a driver of ecological changes in arthropod communities

Heritable variation in traits can have wide-ranging impacts on species interactions, but the effects that ongoing evolution has on the temporal ecological dynamics of communities are not well understood. Here, we identify three conditions that, if experimentally satisfied, support the hypothesis that evolution by natural selection can drive ecological changes in communities. These conditions are: (i) a focal population exhibits genetic variation in a trait(s), (ii) there is measurable directional selection on the trait(s), and (iii) the trait(s) under selection affects variation in a community variable(s). When these conditions are met, we expect evolution by natural selection to cause ecological changes in the community. We tested these conditions in a field experiment examining the interactions between a native plant (Oenothera biennis) and its associated arthropod community (more than 90 spp.). Oenothera biennis exhibited genetic variation in several plant traits and there was directional selection on plant biomass, life-history strategy (annual versus biennial reproduction) and herbivore resistance. Genetically based variation in biomass and life-history strategy consistently affected the abundance of common arthropod species, total arthropod abundance and arthropod species richness. Using two modelling approaches, we show that evolution by natural selection in large O. biennis populations is predicted to cause changes in the abundance of individual arthropod species, increases in the total abundance of arthropods and a decline in the number of arthropod species. In small O. biennis populations, genetic drift is predicted to swamp out the effects of selection, making the evolution of plant populations unpredictable. In short, evolution by natural selection can play an important role in affecting the dynamics of communities, but these effects depend on several ecological factors. The framework presented here is general and can be applied to other systems to examine the community-level effects of ongoing evolution.     

Global change alters the stability of food webs

Recent research has generally shown that a small change in the number of species in a food web can have consequences both for community structure and ecosystem processes. However ‘change’ is not limited to just the number of species in a community, but might include an alteration to such properties as precipitation, nutrient cycling and temperature. How such changes might affect species interactions is important, not just through the presence or absence of interactions, but also because the patterning of interaction strengths among species is intimately associated with community stability. Interaction strengths encompass such properties as feeding rates and assimilation efficiencies, and encapsulate functionally important information with regard to ecosystem processes. Interaction strengths represent the pathways and transfer of energy through an ecosystem. We review the best empirical data available detailing the frequency distribution of interaction strengths in communities. We present the underlying (but consistent) pattern of species interactions and discuss the implications of this patterning. We then examine how such a basic pattern might be affected given various scenarios of ‘change’ and discuss the consequences for community stability and ecosystem functioning.     

Rapid contemporary evolution and clonal food web dynamics

Character evolution that affects ecological community interactions often occurs contemporaneously with temporal changes in population size, potentially altering the very nature of those dynamics. Such eco-evolutionary processes may be most readily explored in systems with short generations and simple genetics. Asexual and cyclically parthenogenetic organisms such as microalgae, cladocerans and rotifers, which frequently dominate freshwater plankton communities, meet these requirements. Multiple clonal lines can coexist within each species over extended periods, until either fixation occurs or a sexual phase reshuffles the genetic material. When clones differ in traits affecting interspecific interactions, within-species clonal dynamics can have major effects on the population dynamics. We first consider a simple predator–prey system with two prey genotypes, parametrized with data from a well-studied experimental system, and explore how the extent of differences in defence against predation within the prey population determine dynamic stability versus instability of the system. We then explore how increased potential for evolution affects the community dynamics in a more general community model with multiple predator and multiple prey genotypes. These examples illustrate how microevolutionary ‘details’ that enhance or limit the potential for heritable phenotypic change can have significant effects on contemporaneous community-level dynamics and the persistence and coexistence of species.     

The role of phenotypic plasticity in shaping ecological networks

Plasticity-mediated changes in interaction dynamics and structure may scale up and affect the ecological network in which the plastic species are embedded. Despite their potential relevance for understanding the effects of plasticity on ecological communities, these effects have seldom been analysed. We argue here that, by boosting the magnitude of intra-individual phenotypic variation, plasticity may have three possible direct effects on the interactions that the plastic species maintains with other species in the community: may expand the interaction niche, may cause a shift from one interaction niche to another or may even cause the colonization of a new niche. The combined action of these three factors can scale to the community level and eventually expresses itself as a modification in the topology and functionality of the entire ecological network. We propose that this causal pathway can be more widespread than previously thought and may explain how interaction niches evolve quickly in response to rapid changes in environmental conditions. The implication of this idea is not solely eco-evolutionary but may also help to understand how ecological interactions rewire and evolve in response to global change.     

Increasing community size and connectance can increase stability in competitive communities

The relationship between community complexity and stability has been the subject of an enduring debate in ecology over the last 50 years. Results from early model communities showed that increased complexity is associated with decreased local stability. I demonstrate that increasing both the number of species in a community and the connectance between these species results in an increased probability of local stability in discrete-time competitive communities, when some species would show unstable dynamics in the absence of competition. This is shown analytically for a simple case and across a wider range of community sizes using simulations, where individual species have dynamics that can range from stable point equilibria to periodic or more complex. Increasing the number of competitive links in the community reduces per-capita growth rates through an increase in competitive feedback, stabilising oscillating dynamics. This result was robust to the introduction of a trade-off between competitive ability and intrinsic growth rate and changes in species interaction strengths. This throws new light on the discrepancy between the theoretical view that increased complexity reduces stability and the empirical view that more complex systems are more likely to be stable, giving one explanation for the relative lack of complex dynamics found in natural systems. I examine how these results relate to diversity-biomass stability relationships and show that an analytical solution derived in the region of stable equilibrium dynamics captures many features of the change in biomass fluctuations with community size in communities including species with oscillating dynamics.     

Leveraging nature's backup plans to incorporate interspecific interactions and resilience into restoration

Interspecific interactions are important structuring forces in ecological communities. Interactions can be disturbed when species are lost from a community. When interactions result in fitness gains for at least one participating organism, that organism may experience reduced fitness as a result of interaction disturbance. However, many species exhibit traits that enable individuals to persist and reproduce in spite of such disruptions, resulting in resilience to interaction disturbance. Such traits can result in interaction generalization, phenotypic and behavioral plasticity, and adaptive capacity. We discuss examples of these traits and use case studies to illustrate how restoration practitioners can use a trait‐based approach to examine species of concern, identify traits that are associated with interspecific interactions and are relevant to resilience, and target such traits in restoration. Restoration activities that bolster interaction resilience could include, for example, reintroducing or supporting specific functional groups or managing abiotic conditions to reduce interaction dependence by at‐risk species (e.g. providing structural complexity offering shelter and cover). Resilience may also be an important consideration in species selection for restoration. Establishment of resilient species, able to persist after interaction disturbance, may be essential to restoring to a functioning ecological community. Once such species are present, they could help support more specialized species that lack resilience traits, such as many species of concern. Understanding the conditions under which processes linked to resilience may enable species to persist and communities to reform following interaction disturbance is a key application of community ecology to ecological restoration.     

Contemporary climate change in the Sonoran Desert favors cold‐adapted species

Impacts of long‐term climate shifts on the dynamics of intact communities within species ranges are not well understood. Here, we show that warming and drying of the Southwestern United States over the last 25 years has corresponded to a shift in the species composition of Sonoran Desert winter annuals, paradoxically favoring species that germinate and grow best in cold temperatures. Winter rains have been arriving later in the season, during December rather than October, leading to the unexpected result that plants are germinating under colder temperatures, shifting community composition to favor slow growing, water‐use efficient, cold‐adapted species. Our results demonstrate how detailed ecophysiological knowledge of individual species, combined with long‐term demographic data, can reveal complex and sometimes unexpected shifts in community composition in response to climate change. Further, these results highlight the potentially overwhelming impact of changes in phenology on the response of biota to a changing climate.     

Patterns of species interaction strength in assembled theoretical competition communities

Strength of interactions between species may be an important tool in our effort to understand community structure. Recent theoretical and empirical findings suggest that despite the presence of some strong interactions, weak interactions prevail in communities. Here, we examine how mean interaction strengths change as theoretical competition communities assemble and what the distribution of interaction coefficients is in the communities that are formed during the assembly process. Our results show that the mean competition strengths fall as assembly progresses and that most interactions in the communities formed are weak. Communities that are invulnerable to further invasions are those where interspecific interactions are weaker than the average interaction strength between species in the pool. If these results can be generalized to more than one trophic level, implications for management and conservation of natural communities are substantial.     

Scaling environmental change through the community-level: a trait-based response-and-effect framework for plants

Predicting ecosystem responses to global change is a major challenge in ecology. A critical step in that challenge is to understand how changing environmental conditions influence processes across levels of ecological organization. While direct scaling from individual to ecosystem dynamics can lead to robust and mechanistic predictions, new approaches are needed to appropriately translate questions through the community level. Species invasion, loss, and turnover all necessitate this scaling through community processes, but predicting how such changes may influence ecosystem function is notoriously difficult. We suggest that community‐level dynamics can be incorporated into scaling predictions using a trait‐based response–effect framework that differentiates the community response to environmental change (predicted by response traits) and the effect of that change on ecosystem processes (predicted by effect traits). We develop a response‐and‐effect functional framework, concentrating on how the relationships among species' response, effect, and abundance can lead to general predictions concerning the magnitude and direction of the influence of environmental change on function. We then detail several key research directions needed to better scale the effects of environmental change through the community level. These include (1) effect and response trait characterization, (2) linkages between response‐and‐effect traits, (3) the importance of species interactions on trait expression, and (4) incorporation of feedbacks across multiple temporal scales. Increasing rates of extinction and invasion that are modifying communities worldwide make such a research agenda imperative.     

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.     

Upstream trophic structure modulates downstream community dynamics via resource subsidies

In many natural systems, the physical structure of the landscape dictates the flow of resources. Despite mounting evidence that communities’ dynamics can be indirectly coupled by reciprocal among ecosystem resource flows, our understanding of how directional resource flows might indirectly link biological communities is limited. We here propose that differences in community structure upstream should lead to different downstream dynamics, even in the absence of dispersal of organisms. We report an experimental test of the effect of upstream community structure on downstream community dynamics in a simplified but highly controlled setting, using protist microcosms. We implemented directional flows of resources, without dispersal, from a standard resource pool into upstream communities of contrasting interaction structure and then to further downstream communities of either one or two trophic levels. Our results demonstrate that different types of species interactions in upstream habitats may lead to different population sizes and levels of biomass in these upstream habitats. This, in turn, leads to varying levels of detritus transfer (dead biomass) to the downstream communities, thus influencing their population densities and trophic interactions in predictable ways. Our results suggest that the structure of species interactions in directionally structured ecosystems can be a key mediator of alterations to downstream habitats. Alterations to upstream habitats can thus cascade down to downstream communities, even without dispersal.     

Species identity and negative density dependence lead to convergence in designed plant mixtures of twelve species

Competition effects on community development are difficult to quantify in species-rich plant communities due to the complexity of possible interactions. We used multispecies mixtures to investigate how species identity and competitive interactions influence the development of plant communities. Given the same set of species with differing initial abundance in various communities, we tested whether communities would become more similar (converge) or dissimilar (diverge) over time depending on the relative importance of species identity and competition. Twenty-four experimental communities were established by planting seedlings of twelve wetland species at different relative abundances and absolute densities. The development of the communities was monitored over three years, and yearly changes in biomass were modelled as a linear function of the species biomass at the start of each period. After three years, a clear dominance structure had developed, with four species making up 80% of the aboveground biomass. In all three years, community dynamics was driven by differences in relative growth rates among the species (i.e. an effect of species identity). However, in the second and third years negative density dependence was also important, with changes in the relative abundance of the most abundant species being negatively related to their biomass at the start of the period. Multiple species interactions – though generally weaker than effects of species identity and intraspecific competition – became increasingly important and also contributed to the dominance pattern. It is concluded that species identity and negative density dependence of the dominant species were the most important factors causing the experimental plant communities to converge. We suggest that model systems composed of several species offer a useful method for investigating the influence of functional traits upon community dynamics.     

Plant species’ coexistence relationships may shift according to life history traits and seasons

Although several studies have demonstrated that disturbance contributes to species’ diversity, little emphasis has been placed on the identification of species’ coexistence mechanisms related to life history traits. In this study, we compared species’ richness and components of plant communities around river confluences to explore how disturbance promotes the coexistence of species with different life history traits. Sites upstream and downstream of confluences are ideal for such comparisons because they draw on the same species’ pools and have similar ambient conditions, but differ markedly in the extents of flooding disturbance. We compared sites upstream and downstream of confluences by calculating species’ richness and community similarity indices for several life history traits in both summer and spring. In summer, the combined richness of all the species, of annual- and summer-flowering species, was higher downstream from confluences than upstream, but this was not the case for perennials. Similarity analyses suggested that plant communities are constructed according to a neutral process, whereby interactions between the coexisting species are neutral. However, in spring, species’ richness was similar upstream and downstream of confluences for all life history traits. Similarity analyses suggested that under these circumstances, the communities were constructed through a species-sorting process; i.e., each life history trait had a distinct habitat preference. Thus, the relative strengths of different community assembly processes may change seasonally. We concluded that species groups differing in their responses to disturbance may coexist in a single community. Thus, community structuring following disturbance may involve two processes: a neutral and a species-sorting process. The relative importance of each may vary between species’ life history traits and between seasons, and the interaction may account for current community structures.     

Small mammal controls on the climate‐driven range shift of woody plant species

Climate change is resulting in shifts in species’ ranges as species inhabit new climatically suitable areas. A key factor affecting range‐shifts is the interaction with predators. Small mammals, being primary seed predators and dispersers in forest ecosystems, may play a major role in determining which plant species will successfully expand and the rate at which range‐shifts will occur. Plants dispersing seeds beyond the species’ current range limits will encounter seed predators to which these seeds are novel; however, empirical studies of seed predator–novel seed interactions are lacking. The aims of our study were to: 1) quantify seed selection by small mammals presented with ‘novel’ seeds; 2) quantify the post‐selection fate of ‘novel’ seeds; and 3) identify seed traits that affect seed selection and post‐selection seed fate. We designed a field experiment exposing small mammal communities to novel seeds produced by plants expected to shift their ranges in response to climate change. We matched novel seeds with reference ‘familiar’ seeds and studied key steps defining interactions between small mammals and novel seeds. We found that the probability of selection of a novel seed varied among species and was, at times, higher than the selection probability of familiar seeds. Key traits that affected seed selection and the distance a seed was dispersed for caching were shell hardness and seed mass. We also found that 33% of dispersed seeds were cached in optimal germination sites (e.g. within fallen logs and buried under the leaf litter mat). Through seed emergence trials we found that emergence was higher for larger seeds, suggesting that the role of small mammals may be modulated by emergence rates. Our results suggest that the interaction between small mammals and novel seeds may have cascading effects on climate‐induced plant range shifts and community composition.     

Landscape simplification shapes pathogen prevalence in plant‐pollinator networks

Species interaction networks, which play an important role in determining pathogen transmission and spread in ecological communities, can shift in response to agricultural landscape simplification. However, we know surprisingly little about how landscape simplification‐driven changes in network structure impact epidemiological patterns. Here, we combine mathematical modelling and data from eleven bipartite plant‐pollinator networks observed along a landscape simplification gradient to elucidate how changes in network structure shape disease dynamics. Our empirical data show that landscape simplification reduces pathogen prevalence in bee communities via increased diet breadth of the dominant species. Furthermore, our empirical data and theoretical model indicate that increased connectance reduces the likelihood of a disease outbreak and decreases variance in prevalence among bee species in the community, resulting in a dilution effect. Because infectious diseases are implicated in pollinator declines worldwide, a better understanding of how land use change impacts species interactions is therefore critical for conserving pollinator health.