Predicting persistence in a changing climate: flow direction and limitations to redistribution

Synthesis Impending climate changes beg the question: which populations and species will go extinct and which will persist under future environmental conditions? When tolerance in situ is not possible, then species must undergo range shifts to avoid extinction. This synthesis explores ways in which directional air and water flow could impede such redistribution and the characteristics that might allow species to disperse against the flow. Considering flow patterns in tandem with climate and range projections has the potential to improve predictions of persistence for the earth’s many non‐moving foundation and basal species as well as the communities and food webs that they support. Predicting which populations and species will persist (i.e. avoid extinction and continue to exist) in the face of climate change requires an understanding of mechanisms that allow species to cope with altered environmental conditions. When processes of tolerance, acclimation, and adaptation are insufficient to allow persistence in situ, redistribution is required for population or species persistence. Here, I review evidence that directional flows of water and air have the potential to restrict species’ range boundaries under ambient conditions, the spread of introduced species, and the redistribution of native species under changing climatic conditions. I develop the hypothesis that flow patterns, such as the speed and directionality (i.e. poleward vs equatorward) of asymmetric air and water currents, may need to be considered when assessing the vulnerability of populations and species to climate change. To the degree that directional flows are found to limit redistribution, there may be disproportionate losses of diversity where the dominant flow direction opposes that of shifting climate space. Within this context, I highlight flow conditions and life‐history traits that could help the most passively‐dispersed species redistribute to track changing climate. These predictions merit further examination in order to better anticipate which populations, species, and associated communities are likely to persist under climate change.     

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.     

Equipped to cope with climate change: traits associated with range filling across European taxa

In order to understand the ecological effects of climate change it is essential to forecast suitable areas for species in the future. However, species’ ability to reach potentially suitable areas is also critical for species survival. These ‘range‐shift’ abilities can be studied using life‐history traits related to four range‐shift stages: emigration, movement, establishment, and proliferation. Here, we use the extent to which species’ ranges fill the climatically suitable area available (‘range filling’) as a proxy for the ability of European mammals and birds to shift their ranges under climate change. We detect which traits associate most closely with range filling. Drawing comparisons with a recent analysis for plants, we ask whether the latitudinal position of species’ ranges supports the assertion that post‐glacial range‐shift limitations cause disequilibrium between ranges and climate. We also disentangle which of the three taxonomic groups has greatest range filling. For mammals, generalists and early‐reproducing species have the greatest range filling. For birds, generalist species with high annual fecundity, which live longer than expected based on body size, have the greatest range filling. Although we consider traits related to the four range‐shift stages, only traits related to establishment and proliferation ability significantly influence range filling of mammals and birds. Species with the greatest range filling are those whose range centroid falls in the latitudinal centre of Europe, suggesting that post‐glacial range expansion is a leading cause of disequilibrium with climate, although other explanations are also possible. Range filling of plants is lower than that of mammals or birds, suggesting that plants are more range‐limited by non‐climatic factors. Therefore, plants might be face greater non‐climatic restraints on range shifts than mammals or birds.     

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.     

Modelling range dynamics of terricolous lichens of the genus Peltigera in the Alps under a climate change scenario

Climate change is expected to strongly impact biodiversity in Alpine ecosystems and species distribution modelling is increasingly used to provide anticipatory information to guide conservation. In this study, (1) we quantified the range loss, range gain, range change and range turnover caused by climate change in the genus Peltigera a group of terricolous lichens widespread across the Alps, and then (2) we evaluated the relationships between the predictors of range dynamics and functional traits. Our results indicate moderate range dynamics for species of the genus Peltigera across the Alps under a climate change scenario. This would imply a relative stability and resistance of these lichens to climate change that may reflect the local persistence of the species under sub-optimal conditions. Our results also suggest that range dynamics could be associated with functional traits mainly related to water-use strategies and to a trade-off between dispersal and establishment ability. This finding suggests that functional traits may strongly modulate the lichen response to climate change and that species with similar functional traits are prone to similar selective pressures.     

A process‐based model supports an association between dispersal and the prevalence of species traits in tropical reef fish assemblages

Habitat dynamics interacting with species dispersal abilities could generate gradients in species diversity and prevalence of species traits when the latter are associated with species dispersal potential. Using a process‐based model of diversification constrained by a dispersal parameter, we simulated the interplay between reef habitat dynamics during the past 140 million years and dispersal, shaping lineage diversification history and assemblage composition globally. The emerging patterns from the simulations were compared to current prevalence of species traits related to dispersal for 6315 tropical reef fish species. We found a significant spatial congruence between the prevalence of simulated low dispersal values and areas with a large proportion of species characterized by small adult body size, narrow home range mobility behaviour, pelagic larval duration shorter than 21 days and diurnal activity. Species characterized by such traits were found predominantly in the Indo‐Australian Archipelago and the Caribbean Sea. Furthermore, the frequency distribution of the dispersal parameter was found to match empirical distributions for body size, PLD and home range mobility behaviour. Also, the dispersal parameter in the simulations was associated to diversification rates and resulted in trait frequency matching empirical distributions. Overall, our findings suggest that past habitat dynamics, in conjunction with dispersal processes, influenced diversification in tropical reef fishes, which may explain the present‐day geography of species traits.     

A framework integrating physiology,dispersal and land‐use to project species ranges under climate change

To study the potential effects of climate change on species, one of the most popular approaches are species distribution models (SDMs). However, they usually fail to consider important species‐specific biological traits, such as species’ physiological capacities or dispersal ability. Furthermore, there is consensus that climate change does not influence species distributions in isolation, but together with other anthropogenic impacts such as land‐use change, even though studies investigating the relative impacts of different threats on species and their geographic ranges are still rare. Here we propose a novel integrative approach which produces refined future range projections by combining SDMs based on distribution, climate, and physiological tolerance data with empirical data on dispersal ability as well as current and future land‐use. Range projections based on different combinations of these factors show strong variation in projected range size for our study species Emberiza hortulana. Using climate and physiological data alone, strong range gains are projected. However, when we account for land‐use change and dispersal ability, future range‐gain may even turn into a future range loss. Our study highlights the importance of accounting for biological traits and processes in species distribution models and of considering the additive effects of climate and land‐use change to achieve more reliable range projections. Furthermore, with our approach we present a new tool to assess species’ vulnerability to climate change which can be easily applied to multiple species.     

What shapes the altitudinal range of a high mountain Mediterranean plant? Recruitment probabilities from ovule to seedling stage

Recruitment is a complex process consisting of sequential stages affected by biotic interactions and abiotic factors. Assessment of these sequential stages and corresponding subprocesses may be useful in identifying the most critical stages. Accordingly, to assess the factors that may determine the altitudinal range limits of the high mountain Mediterranean plant Silene ciliata, a set of demographic stages, from flower production to establishment of 2‐yr‐old plants, and their influence on recruitment probability were examined using a step‐by‐step approach. We integrated florivory, pollination and pre‐dispersal seed predation as pre‐dispersal factors, and seedling emergence and survival as post‐dispersal determinants of recruitment. Three populations were monitored at the southernmost margin of the species along its local altitudinal range. Previous studies suggest that seediness is strongly limited by summer drought especially at the lower boundary of the species, a situation that may worsen under current global warming. Our results showed that recruitment was mainly limited by low seed production in the pre‐dispersal stage and low seedling emergence and survival in the post‐dispersal stage, probably due to environmental harshness in summer. By contrast, biotic factors responsible for propagule loss, such as flower and fruit predation, had a minor effect on the probability of plant recruitment. Although the relative importance of transition probabilities was similar among populations along the altitudinal range, comparatively lower flower production significantly reduced the number of recruited plants at the lowest altitude population. This demographic bottleneck, together with increased competition with other species favoured by climate warming, might collapse population growth and limit persistence at the lower altitudinal range of the species, raising its low local altitudinal edge.     

Expanding population edges: theories,traits, and trade‐offs

Recent patterns of global change have highlighted the importance of understanding the dynamics and mechanisms of species range shifts and expansions. Unique demographic features, spatial processes, and selective pressures can result in the accumulation and evolution of distinctive phenotypic traits at the leading edges of expansions. We review the characteristics of expanding range margins and highlight possible mechanisms for the appearance of phenotypic differences between individuals at the leading edge and core of the range. The development of life history traits that increase dispersal or reproductive ability is predicted by theory and supported with extensive empirical evidence. Many examples of rapid phenotypic change are associated with trade‐offs that may influence the persistence of the trait once expansion ends. Accounting for the effects of edge phenotypes and related trade‐offs could be critical for predicting the spread of invasive species and population responses to climate change.     

Traits related to species persistence and dispersal explain changes in plant communities subjected to habitat loss

Aim Habitat fragmentation is a major driver of biodiversity loss but it is insufficiently known how much its effects vary among species with different life‐history traits; especially in plant communities, the understanding of the role of traits related to species persistence and dispersal in determining dynamics of species communities in fragmented landscapes is still limited. The primary aim of this study was to test how plant traits related to persistence and dispersal and their interactions modify plant species vulnerability to decreasing habitat area and increasing isolation. Location Five regions distributed over four countries in Central and Northern Europe. Methods Our dataset was composed of primary data from studies on the distribution of plant communities in 300 grassland fragments in five regions. The regional datasets were consolidated by standardizing nomenclature and species life‐history traits and by recalculating standardized landscape measures from the original geographical data. We assessed the responses of plant species richness to habitat area, connectivity, plant life‐history traits and their interactions using linear mixed models. Results We found that the negative effect of habitat loss on plant species richness was pervasive across different regions, whereas the effect of habitat isolation on species richness was not evident. This area effect was, however, not equal for all the species, and life‐history traits related to both species persistence and dispersal modified plant sensitivity to habitat loss, indicating that both landscape and local processes determined large‐scale dynamics of plant communities. High competitive ability for light, annual life cycle and animal dispersal emerged as traits enabling species to cope with habitat loss. Main conclusions In highly fragmented rural landscapes in NW Europe, mitigating the spatial isolation of remaining grasslands should be accompanied by restoration measures aimed at improving habitat quality for low competitors, abiotically dispersed and perennial, clonal species.     

A comparative analysis of dispersal syndromes in terrestrial and semi‐terrestrial animals

Dispersal, the behaviour ensuring gene flow, tends to covary with a number of morphological, ecological and behavioural traits. While species‐specific dispersal behaviours are the product of each species’ unique evolutionary history, there may be distinct interspecific patterns of covariation between dispersal and other traits (‘dispersal syndromes’) due to their shared evolutionary history or shared environments. Using dispersal, phylogeny and trait data for 15 terrestrial and semi‐terrestrial animal Orders (> 700 species), we tested for the existence and consistency of dispersal syndromes across species. At this taxonomic scale, dispersal increased linearly with body size in omnivores, but decreased above a critical length in herbivores and carnivores. Species life history and ecology significantly influenced patterns of covariation, with higher phylogenetic signal of dispersal in aerial dispersers compared with ground dwellers and stronger evidence for dispersal syndromes in aerial dispersers and ectotherms, compared with ground dwellers and endotherms. Our results highlight the complex role of dispersal in the evolution of species life‐history strategies: good dispersal ability was consistently associated with high fecundity and survival, and in aerial dispersers it was associated with early maturation. We discuss the consequences of these findings for species evolution and range shifts in response to future climate change.     

Individualistic species limitations of climate‐induced range expansions generated by meso‐scale dispersal barriers

Aim Evidence indicates that species are responding to climate change through distributional range shifts that track suitable climatic conditions. We aim to elucidate the role of meso‐scale dispersal barriers in climate‐tracking responses. Location South coast of England (the English Channel). Methods Historical distributional data of four intertidal invertebrate species were logistically regressed against sea surface temperature (SST) to determine a climate envelope. This envelope was used to estimate the expected climate‐tracking response since 1990 along the coast, which was compared with observed range expansions. A hydrodynamic modelling approach was used to identify dispersal barriers and explore disparities between expected and observed climate tracking. Results Range shifts detected by field survey over the past 20 years were less than those predicted by the changes that have occurred in SST. Hydrodynamic model simulations indicated that physical barriers produced by complex tidal currents have variably restricted dispersal of pelagic larvae amongst the four species. Main conclusions We provide the first evidence that meso‐scale hydrodynamic barriers have limited climate‐induced range shifts and demonstrate that life history traits affect the ability of species to overcome such barriers. This suggests that current forecasts may be flawed, both by overestimating range shifts and by underestimating climatic tolerances of species. This has implications for our understanding of climate change impacts on global biodiversity.     

Life-history traits are correlated with geographical distribution patterns of western European forest herb species

Aim To investigate whether six plant life‐history traits that have been related to colonization ability at local scales are also related to the geographical range characteristics of 273 forest plant species. Location Continental western Europe, five countries in particular: France, Luxemburg, Belgium, the Netherlands and Germany. The region is situated between 42° and 55°N and 5°W and 15°E and has a summed total area of 971,404 km². Methods Distribution data were compiled from five national data bases and converted to a 10′ grid. Life‐history traits were taken from existing compilations of autecological information of European species. The spatial arrangement of occupied grid cells was investigated using Ripley's K. Cross‐species correlations and phylogenetically independent contrasts were used to investigate the relationships between plant life‐history traits and three range characteristics: area of occupancy, latitudinal extent and centroid latitude. Results For herbaceous species, seed dispersal mode, seed production and seed bank longevity exhibited significant associations with geographical range characteristics, including area of occupancy. Woody plant species exhibited fewer significant associations, although maximum height was positively associated with range centroid latitude within the study area. Furthermore, the ranges of species with limited dispersal ability were found to be more clustered than the ranges of species with morphological adaptations for long‐distance seed dispersal. Main conclusions For western European forest plant species, life‐history traits that are related to colonization ability at local scales are associated with variation in large‐scale geographical range characteristics. This finding implies that the distributions of some forest plant species in the study area may be limited by seed dispersal and colonization capacity rather than climate or other environmental factors.     

Towards an interactive,process‐based approach to understanding range shifts: developmental and environmental dependencies matter

Many species are undergoing distributional shifts in response to climate change. However, wide variability in range shifting rates has been observed across taxa, and even among closely‐related species. Attempts to link climate‐mediated range shifts to traits has often produced weak or conflicting results. Here we investigate interactive effects of developmental processes and environmental stress on the expression of traits relevant to range shifts. We use an individual‐based modelling approach to assess how different developmental strategies affect range shift rates under a range of environmental conditions. We find that under stressful conditions, such as at the margins of the species’ fundamental niche, investment in prolonged development leads to the greatest rates of range shifting, especially when longer time in development leads to improved fecundity and dispersal‐related traits. However, under benign conditions, and when traits are less developmentally plastic, shorter development times are preferred for rapid range shifts, because higher generational frequency increases the number of individual dispersal events occurring over time. Our results suggest that the ability of a species to range shift depends not only on their dispersal and colonisation characteristics but also how these characteristics interact with developmental strategies. Benefits of any trait always depended on the environmental and developmental sensitivity of life history trait combinations, and the environmental conditions under which the range shift takes place. Without considering environmental and developmental sources of variation in the expression of traits relevant to range shifts, there is little hope of developing a general understanding of intrinsic drivers of range shift potential.     

Pleistocene climatic changes drive diversification across a tropical savanna

Spatial responses of species to past climate change depend on both intrinsic traits (climatic niche breadth, dispersal rates) and the scale of climatic fluctuations across the landscape. New capabilities in generating and analysing population genomic data, along with spatial modelling, have unleashed our capacity to infer how past climate changes have shaped populations, and by extension, complex communities. Combining these approaches, we uncover lineage diversity across four codistributed lizards from the Australian Monsoonal Tropics and explore how varying climatic tolerances interact with regional climate history to generate common vs. disparate responses to late Pleistocene change. We find more divergent spatial structuring and temporal demographic responses in the drier Kimberley region compared to the more mesic and consistently suitable Top End. We hypothesize that, in general, the effects of species’ traits on sensitivity to climate fluctuation will be more evident in climatically marginal regions. If true, this points to the need in climatically marginal areas to craft more species‐(or trait)‐specific strategies for persistence under future climate change.     

The odd one out or a hidden generalist: Hawaiian Melicope (Rutaceae) do not share traits associated with successful island colonization

Oceanic islands are unique in their species composition, which is defined by arrival of colonizers via long distance dispersal followed by establishment of species followed in some cases by adaptive radiation. Evolutionary biologists identified traits facilitating successful colonization of islands as including polyploidy, self‐compatibility, herbaceousness and ability for long‐distance dispersal. Successful establishment and evolutionary diversification of lineages on islands often involves shifts to woodiness and shifts in methods of outcrossing as well as changes in dispersal ability. The genus Melicope colonized numerous archipelagos throughout the Pacific including the Hawaiian Islands, where the lineage comprises currently 54 endemic species and represents the largest radiation of woody plants on the islands. The wide distributional range of the genus illustrates its high dispersibility, most likely due to adaption to bird dispersal. Here we investigate ploidy in the genus using flow cytometry and chromosome counting. We find the genus to be paleopolyploid with 2n = 4x = 36, a ploidy level characterizing the entire subfamily Amyridoideae and dating back to at least the Palaeocene. Therefore Hawaiian Melicope have not undergone recent polyploidization prior to colonization of the islands. Thus Melicope retained colonization success while exhibiting a combination of traits that typically characterize well established island specialists while lacking some traits associated to successful colonizers.     

Seeds tend to disperse further in the tropics

Despite the importance of seed dispersal in a plant's life cycle, global patterns in seed dispersal distance have seldom been studied. This paper presents the first geographically and taxonomically broad quantification of the latitudinal gradient in seed dispersal distance. Although there is substantial variation in the seed dispersal distances of different species at a given latitude, seeds disperse on average more than an order of magnitude further at the equator than towards the poles. This pattern is partially explained by plant life‐history traits that simultaneously associate with seed dispersal distance and latitude, including dispersal mode and plant height. The extended seed shadow of tropical plants could increase the distance between conspecific individuals. This could facilitate species coexistence and contribute to the maintenance of high plant diversity in tropical communities. The latitudinal gradient in dispersal distance also has implications for species’ persistence in the face of habitat fragmentation and climate change.     

Do species’ traits predict recent shifts at expanding range edges?

Although some organisms have moved to higher elevations and latitudes in response to recent climate change, there is little consensus regarding the capacity of different species to track rapid climate change via range shifts. Understanding species' abilities to shift ranges has important implications for assessing extinction risk and predicting future community structure. At an expanding front, colonization rates are determined jointly by rates of reproduction and dispersal. In addition, establishment of viable populations requires that individuals find suitable resources in novel habitats. Thus, species with greater dispersal ability, reproductive rate and ecological generalization should be more likely to expand into new regions under climate change. Here, we assess current evidence for the relationship between leading-edge range shifts and species' traits. We found expected relationships for several datasets, including diet breadth in North American Passeriformes and egg-laying habitat in British Odonata. However, models generally had low explanatory power. Thus, even statistically and biologically meaningful relationships are unlikely to be of predictive utility for conservation and management. Trait-based range shift forecasts face several challenges, including quantifying relevant natural history variation across large numbers of species and coupling these data with extrinsic factors such as habitat fragmentation and availability.     

Historical legacies and ecological determinants of grass naturalizations worldwide

The global distribution of exotic species is the result of abiotic, biotic and dispersal filtering processes that shape the movement and success of species outside their native range. In this study we aim to understand how these filtering processes drive the fluxes of grass species among regions, the factors that influence which species establish outside of their native range, and where they do so. We used national and subnational checklists of native and introduced grass species to determine the extent to which each region was a source or recipient of exotic grass species. We asked how species traits may distinguish those grass species that have naturalized outside their native range from those that have not, and how environmental conditions are related to the distribution of exotic grass species. We found that exotic grass establishment is shaped by an array of factors including characteristics of regions, traits of species and their interactions. Regions with a longer history of human occupation and larger numbers of native grass species were generally the most important sources of exotic species. Global flows of species were mostly driven by a climate match between the native and exotic ranges, but were also highly asymmetric, with regions with recent human arrival being the major hosts of exotic grass species. Tall, annual and C4 grass species exhibited particularly high probabilities of establishment outside their native range. Despite the idiosyncrasy and stochasticity characteristic of exotic species establishment, this biogeographical analysis revealed important generalities across this large plant group. Our results suggest that grass species that have co-occurred with humans for a longer time may be better adapted to living in anthropogenic landscapes, explaining the global asymmetry in species introductions.     

Pioneers of post-agricultural forest successions are adapted for herbivory avoidance but not biotic seed dispersal

Question

Natural reforestation is an important component of climate mitigation and adaptation, but the ecological processes promoting or constraining it are poorly understood. In this study we employ a stand reconstruction approach (which uses ages of extant trees to estimate year of establishment for each individual tree) to test for general trait-based effects on tree species arrival order in post-agricultural forest successions.

Location

Naturally reforesting post-agricultural landscapes throughout New Zealand.

Methods

Ages were obtained for 2434 individuals spanning 30 tree species across a nationwide network of 128 plots in 14 naturally reforesting post-agricultural sites. These ages were used to calculate individual-level arrival times (relative to the oldest individual in each plot). We estimated species-level arrival times by fitting linear mixed-effects (LME) regressions (with species identity as the fixed effect, and plots nested within sites as the random effects) to individual arrival time data. We used back-casting (where arrival time data are used to document individual-level presence in plots through time) to track annual changes in species abundance and community-weighted mean (CWM) trait values. We used standardised major axis (SMA) regressions to examine the effect of traits related to resource use strategy, herbivory avoidance, seed dispersal and disturbance response on species-level arrival times. We used LME regressions to test for changes in CWM trait values with stand age.

Results

The earliest-arriving species had traits associated with herbivory avoidance, were abiotically dispersed and had short predicted dispersal distances. There was no evidence that traits linked to resource use strategy or disturbance response affected species arrival times. Every significant species-level relationship was recovered in community-level LME analyses.

Conclusions

Our findings suggest that mammalian herbivore control and enhancement of biotic (bird) seed dispersal may be key management interventions in realising the full climate mitigation and adaptation potential of natural reforestation in post-agricultural landscapes.