Multiple stressors,nonlinear effects and the implications of climate change impacts on marine coastal ecosystems

Global climate change will undoubtedly be a pressure on coastal marine ecosystems, affecting not only species distributions and physiology but also ecosystem functioning. In the coastal zone, the environmental variables that may drive ecological responses to climate change include temperature, wave energy, upwelling events and freshwater inputs, and all act and interact at a variety of spatial and temporal scales. To date, we have a poor understanding of how climate‐related environmental changes may affect coastal marine ecosystems or which environmental variables are likely to produce priority effects. Here we use time series data (17 years) of coastal benthic macrofauna to investigate responses to a range of climate‐influenced variables including sea‐surface temperature, southern oscillation indices (SOI, Z4), wind‐wave exposure, freshwater inputs and rainfall. We investigate responses from the abundances of individual species to abundances of functional traits and test whether species that are near the edge of their tolerance to another stressor (in this case sedimentation) may exhibit stronger responses. The responses we observed were all nonlinear and some exhibited thresholds. While temperature was most frequently an important predictor, wave exposure and ENSO‐related variables were also frequently important and most ecological variables responded to interactions between environmental variables. There were also indications that species sensitive to another stressor responded more strongly to weaker climate‐related environmental change at the stressed site than the unstressed site. The observed interactions between climate variables, effects on key species or functional traits, and synergistic effects of additional anthropogenic stressors have important implications for understanding and predicting the ecological consequences of climate change to coastal ecosystems.     

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.     

Resource-based habitat definition, niche overlap and conservation of two sympatric glacial relict butterflies

The precise knowledge of ecological resources and conditions required by species threatened by rapidly changing environmental conditions is of prime importance for conservation biology. Transferability of this knowledge between species with similar ecological requirements is often assumed, but rarely tested. This is especially the case for glacial relict populations confined to climate‐habitat traps from where they cannot move to rejoin areas with suitable environmental conditions. Using two glacial relict butterflies as model organisms, we first quantitatively define larval and adult resource‐based habitat use of each species. Secondly, we test the transferability of ecological profiles (both habitat and ecological niche) between these two species that share both the same biotope and the same host plant. Our results show that both species have markedly different ecological requirements relating to differences in life history and behavioural traits (i.e. egg‐laying strategies and mate‐locating behaviour). Although the two species share many ecological features, they use different functional habitats within our study site. The high degree of interspecific niche overlap should indicate a high interspecific competition. However, we argue that their co‐existence can be explained by the non‐limiting abundance of some resources (e.g. host plants), by the partial separation in time of adult flight periods and by the territorial behaviour of one of the species. We discuss the following general messages: (1) functional habitat of a (threatened) species should be defined in a spatial context corresponding to individual station keeping, and (2) quick diagnosis based on similar ecological requirements may be misleading for the design of reliable conservation and restoration strategies. Detailed mechanistic and quantitative ecological understanding of resource‐use and environmental tolerances across an organism's life cycle is essential for effective conservation in changing environments, like for glacial relict species.     

Comparative thermal adaptation in cicadas (Hemiptera: Cicadidae) inhabiting Mediterranean ecosystems

The thermal responses of cicadas inhabiting the Mediterranean ecosystems in Europe, North America, South Africa, and Australia are investigated. A total of 37 species and two subspecies from 17 genera representing eight tribes and three subfamilies of cicadas are investigated. The analysis includes species that are restricted to the Mediterranean ecosystem as well as those which also inhabit additional environments. The data suggest that cicadas adapt to the climate type regardless of particular types of plants within the various Mediterranean communities. Similarly, cicada thermal responses are independent of body size or taxonomic affinities. There is a wider range of body temperatures for the maximum voluntary tolerance temperature than for heat torpor or minimum flight temperatures. This diversity seems to be determined by the subdivision of the habitat used and the behavior of the species. All species possess relatively elevated heat torpor temperatures adapting to the general thermal characteristics of the Mediterranean ecosystem. The data suggest that cicadas adapt to the Mediterranean climate type regardless of the diversity of particular types of plants within the various communities, of body size or of taxonomic position.     

Floristic patterns and plant traits of Mediterranean communities in fragmented habitats

Aim To contrast floristic spatial patterns and the importance of habitat fragmentation in two plant communities (grassland and scrubland) in the context of ecological succession. We ask whether plant assemblages are affected by habitat fragmentation and, if so, at what spatial scale? Does the relative importance of the niche differentiation and dispersal‐limitation mechanisms change throughout secondary succession? Is the dispersal‐limitation mechanism related to plant functional traits? Location A Mediterranean region, the massif of Albera (Spain). Methods Using a SPOT satellite image to describe the landscape, we tested the effect of habitat fragmentation on species composition, determining the spatial scale of the assemblage response. We then assessed the relative importance of dispersal‐related factors (habitat fragmentation and geographical distance) and environmental constraints (climate‐related variables) influencing species similarity. We tested the association between dispersal‐related factors and plant traits (dispersal mode and life form). Results In both community types, plant composition was partially affected by the surrounding vegetation. In scrublands, animal‐dispersed and woody plants were abundant in landscapes dominated by closed forests, whereas wind‐dispersed annual herbs were poorly represented in those landscapes. Scrubby assemblages were more dependent on geographical distance, habitat fragmentation and climate conditions (temperature, rainfall and solar radiation); grasslands were described only by habitat fragmentation and rainfall. Plant traits did not explain variation in spatial structuring of assemblages. Main conclusions Plant establishment in early Mediterranean communities may be driven primarily by migration from neighbouring established communities, whereas the importance of habitat specialization and community drift increases over time. Plant life forms and dispersal modes did not explain the spatial variation of species distribution, but species richness within the community with differing plant traits was affected by habitat patchiness.     

Species’ traits as predictors of range shifts under contemporary climate change: A review and meta‐analysis

A growing body of literature seeks to explain variation in range shifts using species’ ecological and life‐history traits, with expectations that shifts should be greater in species with greater dispersal ability, reproductive potential, and ecological generalization. Despite strong theoretical support for species’ traits as predictors of range shifts, empirical evidence from contemporary range shift studies remains limited in extent and consensus. We conducted the first comprehensive review of species’ traits as predictors of range shifts, collecting results from 51 studies across multiple taxa encompassing over 11,000 species’ responses for 54 assemblages of taxonomically related species occurring together in space. We used studies of assemblages that directly compared geographic distributions sampled in the 20th century prior to climate change with resurveys of distributions after contemporary climate change and then tested whether species traits accounted for heterogeneity in range shifts. We performed a formal meta‐analysis on study‐level effects of body size, fecundity, diet breadth, habitat breadth, and historic range limit as predictors of range shifts for a subset of 21 studies of 26 assemblages with sufficient data. Range shifts were consistent with predictions based on habitat breadth and historic range limit. However, body size, fecundity, and diet breadth showed no significant effect on range shifts across studies, and multiple studies reported significant relationships that contradicted predictions. Current understanding of species’ traits as predictors of range shifts is limited, and standardized study is needed for traits to be valid indicators of vulnerability in assessments of climate change impacts.     

Differential susceptibility of tree species to oviposition by periodical cicadas

Abstract 1. Periodical cicadas occur widely in deciduous forests of eastern North America and represent the largest insect emergences in the world. Their effects on forest community dynamics, via oviposition damage and root feeding, are potentially great but not well understood. If tree species vary in their susceptibility to cicada oviposition and root colonization, then periodical cicadas may affect ecological interactions among tree species and forest composition. 2. We measured oviposition damage on 63 tree species at 52 sites in southern Indiana, U.S.A. for two broods of periodical cicadas. We compared oviposition damage among different sites, habitats, and tree species. We also measured cicada emergence rates in mature and successional forest habitats and from beneath different tree species. 3. Certain tree species received significantly more oviposition damage than others, and the most and least damaged species were consistent between the two broods. For some species, oviposition damage varied between mature and successional sites. Despite the differences in oviposition rates among species, there was no difference in emergence rates from beneath different tree species, suggesting that initial differences in cicada density among tree species are not maintained through the nymphal life cycle. 4. Cicadas oviposited more at successional sites, but emerged more in mature forest stands, suggesting that successive generations of cicadas represent a shifting mosaic between forests of different ages. As tree species consistently vary in their susceptibility to oviposition damage, periodical cicadas could alter local competitive interactions and therefore affect forest dynamics.     

Leaf‐trait plasticity and species vulnerability to climate change in a Mongolian steppe

Climate change is expected to modify plant assemblages in ways that will have major consequences for ecosystem functions. How climate change will affect community composition will depend on how individual species respond, which is likely related to interspecific differences in functional traits. The extraordinary plasticity of some plant traits is typically neglected in assessing how climate change will affect different species. In the Mongolian steppe, we examined whether leaf functional traits under ambient conditions and whether plasticity in these traits under altered climate could explain climate‐induced biomass responses in 12 co‐occurring plant species. We experimentally created three probable climate change scenarios and used a model selection procedure to determine the set of baseline traits or plasticity values that best explained biomass response. Under all climate change scenarios, plasticity for at least one leaf trait correlated with change in species performance, while functional leaf‐trait values in ambient conditions did not. We demonstrate that trait plasticity could play a critical role in vulnerability of species to a rapidly changing environment. Plasticity should be considered when examining how climate change will affect plant performance, species' niche spaces, and ecological processes that depend on plant community composition.     

Movement,impacts and management of plant distributions in response to climate change: insights from invasions

Synthesis Prediction and management of species responses to climate change is an urgent but relatively young research field. Therefore, climate change ecology must by necessity borrow from other fields. Invasion ecology is particularly well‐suited to informing climate change ecology because both invasion ecology and climate change ecology address the trajectories of rapidly changing novel systems. Here we outline the broad range of active research questions in climate change ecology where research from invasion ecology can stimulate advances. We present ideas for how concepts, case‐studies and methodology from invasion ecology can be adapted to improve prediction and management of species responses to climate change. A major challenge in this era of rapid climate change is to predict changes in species distributions and their impacts on ecosystems, and, if necessary, to recommend management strategies for maintenance of biodiversity or ecosystem services. Biological invasions, studied in most biomes of the world, can provide useful analogs for some of the ecological consequences of species distribution shifts in response to climate change. Invasions illustrate the adaptive and interactive responses that can occur when species are confronted with new environmental conditions. Invasion ecology complements climate change research and provides insights into the following questions: 1) how will species distributions respond to climate change? 2) how will species movement affect recipient ecosystems? And 3) should we, and if so how can we, manage species and ecosystems in the face of climate change? Invasion ecology demonstrates that a trait‐based approach can help to predict spread speeds and impacts on ecosystems, and has the potential to predict climate change impacts on species ranges and recipient ecosystems. However, there is a need to analyse traits in the context of life‐history and demography, the stage in the colonisation process (e.g. spread, establishment or impact), the distribution of suitable habitats in the landscape, and the novel abiotic and biotic conditions under which those traits are expressed. As is the case with climate change, invasion ecology is embedded within complex societal goals. Both disciplines converge on similar questions of ‘when to intervene?‘ and ‘what to do?‘ which call for a better understanding of the ecological processes and social values associated with changing ecosystems.     

Adapt or disperse: understanding species persistence in a changing world

The majority of studies on environmental change focus on the response of single species and neglect fundamental biotic interactions, such as mutualism, competition, predation, and parasitism, which complicate patterns of species persistence. Under global warming, disruption of community interactions can arise when species differ in their sensitivity to rising temperature, leading to mismatched phenologies and/or dispersal patterns. To study species persistence under global climate change, it is critical to consider the ecology and evolution of multispecies interactions; however, the sheer number of potential interactions makes a full study of all interactions unfeasible. One mechanistic approach to solving the problem of complicated community context to global change is to (i) define strategy groups of species based on life‐history traits, trophic position, or location in the ecosystem, (ii) identify species involved in key interactions within these groups, and (iii) determine from the interactions of these key species which traits to study in order to understand the response to global warming. We review the importance of multispecies interactions looking at two trait categories: thermal sensitivity of metabolic rate and associated life‐history traits and dispersal traits of species. A survey of published literature shows pronounced and consistent differences among trophic groups in thermal sensitivity of life‐history traits and in dispersal distances. Our approach increases the feasibility of unraveling such a large and diverse set of community interactions, with the ultimate goal of improving our understanding of community responses to global warming.     

Evidence for evolutionary change associated with the recent range expansion of the British butterfly, Aricia agestis, in response to climate change

Poleward range expansions are widespread responses to recent climate change and are crucial for the future persistence of many species. However, evolutionary change in traits such as colonization history and habitat preference may also be necessary to track environmental change across a fragmented landscape. Understanding the likelihood and speed of such adaptive change is important in determining the rate of species extinction with ongoing climate change. We conducted an amplified fragment length polymorphism (AFLP)‐based genome scan across the recently expanded UK range of the Brown Argus butterfly, Aricia agestis, and used outlier‐based (DFDIST and BayeScan) and association‐based (Isolation‐By‐Adaptation) statistical approaches to identify signatures of evolutionary change associated with range expansion and habitat use. We present evidence for (i) limited effects of range expansion on population genetic structure and (ii) strong signatures of selection at approximately 5% AFLP loci associated with both the poleward range expansion of A. agestis and differences in habitat use across long‐established and recently colonized sites. Patterns of allele frequency variation at these candidate loci suggest that adaptation to new habitats at the range margin has involved selection on genetic variation in habitat use found across the long‐established part of the range. Our results suggest that evolutionary change is likely to affect species’ responses to climate change and that genetic variation in ecological traits across species’ distributions should be maximized to facilitate range shifts across a fragmented landscape, particularly in species that show strong associations with particular habitats.     

Moving forward in global‐change ecology: capitalizing on natural variability

Natural resources managers are being asked to follow practices that accommodate for the impact of climate change on the ecosystems they manage, while global‐ecosystems modelers aim to forecast future responses under different climate scenarios. However, the lack of scientific knowledge about short‐term ecosystem responses to climate change has made it difficult to define set conservation practices or to realistically inform ecosystem models. Until recently, the main goal for ecologists was to study the composition and structure of communities and their implications for ecosystem function, but due to the probable magnitude and irreversibility of climate‐change effects (species extinctions and loss of ecosystem function), a shorter term focus on responses of ecosystems to climate change is needed. We highlight several underutilized approaches for studying the ecological consequences of climate change that capitalize on the natural variability of the climate system at different temporal and spatial scales. For example, studying organismal responses to extreme climatic events can inform about the resilience of populations to global warming and contribute to the assessment of local extinctions. Translocation experiments and gene expression are particular useful to quantitate a species' acclimation potential to global warming. And studies along environmental gradients can guide habitat restoration and protection programs by identifying vulnerable species and sites. These approaches identify the processes and mechanisms underlying species acclimation to changing conditions, combine different analytical approaches, and can be used to improve forecasts of the short‐term impacts of climate change and thus inform conservation practices and ecosystem models in a meaningful way.     

Relationships between life-history strategies of European freshwater fish species and their habitat preferences

1. Focusing on the current environmental characteristics, the ‘habitat template’ theory proposes that life‐history strategies summarise how evolution has shaped species to cope with the temporal and spatial variability of their present environment. The hierarchical ‘landscape filters’ concept predicts that the distribution of species reflects their specific traits that allow them to pass through multiple habitat filters. Together, these theories showed the importance of identifying the functional relationships of species to selective habitat forces in order to predict the organisation and response of communities to the environment. 2. We test the relationships between life‐history traits of European freshwater fish species’ and their habitat preferences, to detect the strategies adopted by fish to cope with their current habitat. For this purpose, we use published data on species traits and habitat preferences (microhabitat hydraulics, temperature and oxygen level). We use multivariate analyses to classify fish species’ strategies and test the relationships between strategies and habitat preferences. 3. We identified a continuum of life‐history patterns between two extremes, with opportunistic and periodic species. Our study supports the idea that microhabitat hydraulics plays a more important role as a template for these species ecological strategies than temperature and oxygen level. Indeed, riffle habitats may select opportunistic species whereas weak relationships are found between species strategies and both their temperature and oxygen level preferences. In addition, the ratio between mortality and growth rate (dimensionless trait), reflecting a trade‐off between growth and survival, varied among species according to the use of their hydraulic habitat, with species using deep habitats exhibiting the highest values. 4. These general relationships between hydraulics and traits may be of importance in community ecology to develop predictive models to understand how fish communities change with the hydraulic environment.     

Spatially structured intraspecific trait variation can foster biodiversity in disturbed,heterogeneous environments

Trait variation within populations is an important area of research for empirical and theoretical ecologists. While differences between individuals are doubtlessly ubiquitous, their role for species coexistence is much less clear and highly debated. Both unstructured (random) and structured (linked to space, time or inheritance) intraspecific trait variation (ITV) may modify species interactions with nontrivial consequences for emerging community compositions. In many ecosystems, these compositions are further driven by prevalent disturbance regimes. I therefore explored the effects of unstructured as well as spatially structured ITV under disturbances in a generic ecological model of competing sessile species. Using spatially explicit, individual‐based simulations, I studied how intraspecific variation in life history traits together with interspecific tradeoffs and disturbance regimes shape long‐term community composition. I found that 1) unstructured ITV does not affect species coexistence in the given context, 2) spatially structured ITV may considerably increase coexistence, but 3) spatially clumped disturbances reduce this effect of spatially structured ITV, especially if interspecific tradeoffs involve dispersal distance. The findings suggest that spatially structured ITV with individual trait responses to local habitat conditions differing among species may create or expand humps in disturbance–diversity relationships. Hence, if present, these forms of spatially structured ITV should be included in ecological models and will be important for reliably assessing community responses to environmental heterogeneity and change.     

Community functional trait composition at the continental scale: the effects of non‐ecological processes

Ecological communities and their response to environmental gradients are increasingly being described by measures of trait composition at the community level – the trait‐based approach. Whether ecological or non‐ecological processes influence trait composition between communities has been debated. Understanding the processes that influence trait composition is important for reconstructing paleoenvironmental conditions from fossil deposits and for understanding changes in community functionality through time. Here, we assess the influence of ecological and non‐ecological processes on the distribution of traits within North American mammals. We found that non‐ecological processes including historical contingency, spatial autocorrelation, and evolutionary history do not influence trait composition; however, the variance in trait composition is highly explained by climate gradients. Our results suggest that habitat breadth, terrestriality, diet breadth, and reproductive traits are strong candidates as proxies for measuring functional aspects of environments in the past and present.     

Flexibility in phenology and habitat use act as buffers to long‐term population declines in UK passerines

Ecological responses to environmental change are wide‐ranging, from alterations in the timing of life‐history events to range and population changes. Explaining the variation across species in these responses is essential for identifying vulnerable species and for developing adequate conservation or mitigation strategies. Using population trend data from the UK Breeding Bird Survey, this study examined the association between long‐term population trends (1994–2007) and phenological, life‐history and resource‐use traits of UK passerine species. Phenology, as well as productivity and resource use were significantly associated with long‐term population trends. Average laying date and first clutch laying period were key predictors, with higher population growth rates associated with earlier laying dates and longer laying periods. This suggests that flexibility in the duration of reproductive periods buffers species against environmental changes. Average laying period was particularly important for migrant species. Flexibility in laying dates for these species is constrained by their arrival dates; mean change in arrival date over a twenty‐five year period strongly predicted population trends amongst migrant species. Besides the key role phenological flexibility plays in buffering population declines, we also showed that more productive, generalist species were less likely to have declining populations than species with specialized habitat requirements, particularly those associated with farmland and urban areas and those reliant on highly seasonal food items (i.e. invertebrate eaters). These results underscore the need for a multi‐faceted approach to understanding the mechanisms governing population trends. Additionally, species’ sensitivity to environmental change is likely to depend on interactions between species‐specific phenology, habitat and resource‐use traits.     

Responses of foliage‐living spider assemblage composition and traits to a climatic gradient in Themeda grasslands

For speciose, but poorly known groups, such as terrestrial arthropods, functional traits present a potential avenue to assist in predicting responses to environmental change. Species turnover is common along environmental gradients, but it is unclear how this is reflected in species traits. Community‐level change in arthropod traits, other than body size, has rarely been explored across spatial scales comparable to those examined here. We hypothesized that the composition and morphological traits of spider assemblages would differ across a gradient of climate and habitat structure. We examined foliage‐living spider assemblages associated with Themeda triandra grasslands along a 900 km climatic gradient in south‐eastern Australia. We used sweep‐netting to collect T. triandra‐associated spiders and counted juveniles and identified adults. We also measured morphological traits of adult spiders and noted their hunting mode. Associations with measures of habitat structure were less consistent than relationships with climate. Both juvenile and adult spiders were more abundant in warmer sites, although species richness was not affected by temperature. We found distinct turnover in species composition along the climatic gradient, with hunting spiders, particularly crab spiders (Thomisidae), making up a greater proportion of assemblages in warmer climates. A range of traits of spiders correlated with the climatic gradient. For example, larger spider species and species that were active hunters were more common in warmer climates. Changes in morphological traits across species, rather than within species drove the morphology‐climate relationship. Strong climate‐trait correlations suggest that it may be possible to predict changes in functional traits of assemblages in response to anthropogenic disturbances such as climate change.     

The effect of range changes on the functional turnover,structure and diversity of bird assemblages under future climate scenarios

Animal assemblages fulfill a critical set of ecological functions for ecosystems that may be altered substantially as climate change‐induced distribution changes lead to community disaggregation and reassembly. We combine species and community perspectives to assess the consequences of projected geographic range changes for the diverse functional attributes of avian assemblages worldwide. Assemblage functional structure is projected to change highly unevenly across space. These differences arise from both changes in the number of species and changes in species’ relative local functional redundancy or distinctness. They sometimes result in substantial losses of functional diversity that could have severe consequences for ecosystem health. Range expansions may counter functional losses in high‐latitude regions, but offer little compensation in many tropical and subtropical biomes. Future management of local community function and ecosystem services thus relies on understanding the global dynamics of species distributions and multiscale approaches that include the biogeographic context of species traits.     

Phylogenetic effects on functional traits and life history strategies of Australian freshwater fish

Understanding the biogeographic and phylogenetic basis to interspecific differences in species’ functional traits is a central goal of evolutionary biology and community ecology. We quantify the extent of phylogenetic influence on functional traits and life‐history strategies of Australian freshwater fish to highlight intercontinental differences as a result of Australia's unique biogeographic and evolutionary history. We assembled data on life history, morphological and ecological traits from published sources for 194 Australian freshwater species. Interspecific variation among species could be described by a specialist–generalist gradient of variation in life‐history strategies associated with spawning frequency, fecundity and spawning migration. In general, Australian fish showed an affinity for life‐history strategies that maximise fitness in hydrologically unpredictable environments. We also observed differences in trait lability between and within life history, morphological and ecological traits where in general morphological and ecological traits were more labile. Our results showed that life‐history strategies are relatively evolutionarily labile and species have potentially evolved or colonised in freshwaters frequently and independently allowing them to maximise population performance in a range of environments. In addition, reproductive guild membership showed strong phylogenetic constraint indicating that evolutionary history is an important component influencing the range and distribution of reproductive strategies in extant species assemblages. For Australian freshwater fish, biogeographic and phylogenetic history contribute to broad taxonomic differences in species functional traits, while finer scale ecological processes contribute to interspecific differences in smaller taxonomic units. These results suggest that the lability or phylogenetic relatedness of different functional traits affects their suitability for testing hypothesis surrounding community level responses to environmental change.     

How phenological tracking shapes species and communities in non-stationary environments

Climate change alters the environments of all species. Predicting species responses requires understanding how species track environmental change, and how such tracking shapes communities. Growing empirical evidence suggests that how species track phenologically – how an organism shifts the timing of major biological events in response to the environment – is linked to species performance and community structure. Such research tantalizingly suggests a potential framework to predict the winners and losers of climate change, and the future communities we can expect. But developing this framework requires far greater efforts to ground empirical studies of phenological tracking in relevant ecological theory. Here we review the concept of phenological tracking in empirical studies and through the lens of coexistence theory to show why a community-level perspective is critical to accurate predictions with climate change. While much current theory for tracking ignores the importance of a multi-species context, basic community assembly theory predicts that competition will drive variation in tracking and trade-offs with other traits. We highlight how existing community assembly theory can help understand tracking in stationary and non-stationary systems. But major advances in predicting the species- and community-level consequences of climate change will require advances in theoretical and empirical studies. We outline a path forward built on greater efforts to integrate priority effects into modern coexistence theory, improved empirical estimates of multivariate environmental change, and clearly defined estimates of phenological tracking and its underlying environmental cues.