Bringing traits back into the equation: A roadmap to understand species redistribution.

Ecological and evolutionary theories have proposed that species traits should be important in mediating species responses to contemporary climate change; yet, empirical evidence has so far provided mixed evidence for the role of behavioral, life history, or ecological characteristics in facilitating or hindering species range shifts. As such, the utility of trait-based approaches to predict species redistribution under climate change has been called into question. We develop the perspective, supported by evidence, that trait variation, if used carefully can have high potential utility, but that past analyses have in many cases failed to identify an explanatory value for traits by not fully embracing the complexity of species range shifts. First, we discuss the relevant theory linking species traits to range shift processes at the leading (expansion) and trailing (contraction) edges of species distributions and highlight the need to clarify the mechanistic basis of trait-based approaches. Second, we provide a brief overview of range shift-trait studies and identify new opportunities for trait integration that consider range-specific processes and intraspecific variability. Third, we explore the circumstances under which environmental and biotic context dependencies are likely to affect our ability to identify the contribution of species traits to range shift processes. Finally, we propose that revealing the role of traits in shaping species redistribution may likely require accounting for methodological variation arising from the range shift estimation process as well as addressing existing functional, geographical, and phylogenetic biases. We provide a series of considerations for more effectively integrating traits as well as extrinsic and methodological factors into species redistribution research. Together, these analytical approaches promise stronger mechanistic and predictive understanding that can help society mitigate and adapt to the effects of climate change on biodiversity.

Climate and land-use changes interact to drive long-term reorganization of riverine fish communities globally.

As climate change unfolds, changes in population dynamics and species distribution ranges are expected to fundamentally reshuffle communities worldwide. Yet, a comprehensive understanding of the mechanisms and extent of community reorganization remains elusive. This is particularly true in riverine systems, which are simultaneously exposed to changing temperature and streamflow, and where land-use change continues to be a major driver of biodiversity loss. Here, we use the most comprehensive compilation of fish abundance time series to date to provide a global synthesis of climate- and LU-induced effects on riverine biota with respect to changes in species thermal and streamflow affinities. We demonstrate that fish communities are increasingly dominated by thermophilic (warm-water) and limnophilic (slow-water) species. Despite being consistent with trends in water temperature and streamflow observed over recent decades, these community changes appear largely decoupled from each other and show wide spatial variation. We further reveal a synergy among climate- and land use-related drivers, such that community thermophilization is heightened in more human-modified systems. Importantly, communities in which species experience thermal and flow regimes that approach or exceed their tolerance thresholds (high community sensitivity), as well as species-poor communities (low community resilience), also display faster rates of compositional change. This research illustrates that quantifying vulnerability of riverine systems to climate change requires a broadening from a narrower thermal focus to more integrative approaches that account for the spatially varying and multifaceted sensitivity of riverine organisms to the interactive effects of water temperature, hydrology, and other anthropogenic changes.

Scale of population synchrony confirms macroecological estimates of minimum viable range size.

Global ecosystems are facing a deepening biodiversity crisis, necessitating robust approaches to quantifying species extinction risk. The lower limit of the macroecological relationship between species range and body size has long been hypothesized as an estimate of the relationship between the minimum viable range size (MVRS) needed for species persistence and the organismal traits that affect space and resource requirements. Here, we perform the first explicit test of this assumption by confronting the MVRS predicted by the range-body size relationship with an independent estimate based on the scale of synchrony in abundance among spatially separated populations of riverine fish. We provide clear evidence of a positive relationship between the scale of synchrony and species body size, and strong support for the MVRS set by the lower limit of the range-body size macroecological relationship. This MVRS may help prioritize first evaluations for unassessed or data-deficient taxa in global conservation assessments.

The geography of metapopulation synchrony in dendritic river networks.

Dendritic habitats, such as river ecosystems, promote the persistence of species by favouring spatial asynchronous dynamics among branches. Yet, our understanding of how network topology influences metapopulation synchrony in these ecosystems remains limited. Here, we introduce the concept of fluvial synchrogram to formulate and test expectations regarding the geography of metapopulation synchrony across watersheds. By combining theoretical simulations and an extensive fish population time-series dataset across Europe, we provide evidence that fish metapopulations can be buffered against synchronous dynamics as a direct consequence of network connectivity and branching complexity. Synchrony was higher between populations connected by direct water flow and decayed faster with distance over the Euclidean than the watercourse dimension. Likewise, synchrony decayed faster with distance in headwater than mainstem populations of the same basin. As network topology and flow directionality generate fundamental spatial patterns of synchrony in fish metapopulations, empirical synchrograms can aid knowledge advancement and inform conservation strategies in complex habitats.

Evidence for dispersal syndromes in freshwater fishes.

Dispersal is a fundamental process defining the distribution of organisms and has long been a topic of inquiry in ecology and evolution. Emerging research points to an interdependency of dispersal with a diverse suite of traits in terrestrial organisms, however the extent to which such dispersal syndromes exist in freshwater species remains uncertain. Here, we test whether dispersal in freshwater fishes (1) is a fixed property of species, and (2) correlates with life-history, morphological, ecological and behavioural traits, using a global dataset of dispersal distances collected from the literature encompassing 116 riverine species and 196 locations. Our meta-analysis revealed a high degree of repeatability and heritability in the dispersal estimates and strong associations with traits related to life-history strategies, energy allocation to reproduction, ecological specialization and swimming skills. Together, these results demonstrate that similar to terrestrial organisms, the multi-dimensional nature of dispersal syndromes in freshwater species offer opportunities for the development of a unifying paradigm of movement ecology that transcend taxonomic and biogeographical realms. The high explanatory power of the models also suggests that trait-based and phylogenetic approaches hold considerable promises to inform conservation efforts in a rapidly changing world.

Evolutionary and environmental determinants of freshwater fish thermal tolerance and plasticity.

Understanding the extent to which phylogenetic constraints and adaptive evolutionary forces help define the physiological sensitivity of species is critical for anticipating climate-related impacts in aquatic environments. Yet, whether upper thermal tolerance and plasticity are shaped by common evolutionary and environmental mechanisms remains to be tested. Based on a systematic literature review, we investigated this question in 82 freshwater fish species (27 families) representing 829 experiments for which data existed on upper thermal limits and it was possible to estimate plasticity using upper thermal tolerance reaction norms. Our findings indicated that there are strong phylogenetic signals in both thermal tolerances and acclimation capacity, although it is weaker in the latter. We found that upper thermal tolerances are correlated with the temperatures experienced by species across their range, likely because of spatially autocorrelated processes in which closely related species share similar selection pressures and limited dispersal from ancestral environments. No association with species thermal habitat was found for acclimation capacity. Instead, species with the lowest physiological plasticity also displayed the highest thermal tolerances, reflecting to some extent an evolutionary trade-off between these two traits. Although our study demonstrates that macroecological climatic niche features measured from species distributions are likely to provide a good approximation of freshwater fish sensitivity to climate change, disentangling the mechanisms underlying both acute and chronic heat tolerances may help to refine predictions regarding climate change-related range shifts and extinctions.

Illuminating geographical patterns in species' range shifts.

Species' range shifts in response to ongoing climate change have been widely documented, but although complex spatial patterns in species' responses are expected to be common, comprehensive comparisons of species' ranges over time have undergone little investigation. Here, we outline a modeling framework based on historical and current species distribution records for disentangling different drivers (i.e. climatic vs. nonclimatic) and assessing distinct facets (i.e. colonization, extirpation, persistence, and lags) of species' range shifts. We used extensive monitoring data for stream fish assemblages throughout France to assess range shifts for 32 fish species between an initial period (1980-1992) and a contemporary one (2003-2009). Our results provide strong evidence that the responses of individual species varied considerably and exhibited complex mosaics of spatial rearrangements. By dissociating range shifts in climatically suitable and unsuitable habitats, we demonstrated that patterns in climate-driven colonization and extirpation were less marked than those attributed to nonclimatic drivers, although this situation could rapidly shift in the near future. We also found evidence that range shifts could be related to some species' traits and that the traits involved varied depending on the facet of range shift considered. The persistence of populations in climatically unsuitable areas was greater for short-lived species, whereas the extent of the lag behind climate change was greater for long-lived, restricted-range, and low-elevation species. We further demonstrated that nonclimatic extirpations were primarily related to the size of the species' range, whereas climate-driven extirpations were better explained by thermal tolerance. Thus, the proposed framework demonstrated its potential for markedly improving our understanding of the key processes involved in range shifting and also offers a template for informing management decisions. Conservation strategies would greatly benefit from identifying both the geographical patterns and the species' traits associated with complex modifications of species' distributions in response to global changes.

Past and recent anthropogenic pressures drive rapid changes in riverine fish communities.

Understanding how and why local communities change is a pressing task for conservation, especially in freshwater systems. It remains challenging because of the complexity of biodiversity changes, driven by the spatio-temporal heterogeneity of human pressures. Using a compilation of riverine fish community time series (93% between 1993 and 2019) across the Palaearctic, Nearctic and Australasia realms, we assessed how past and recent anthropogenic pressures drive community changes across both space and time. We found evidence of rapid changes in community composition of 30% per decade characterized by important changes in the dominant species, together with a 13% increase in total abundance per decade and a 7% increase in species richness per decade. The spatial heterogeneity in these trends could be traced back to the strength and timing of anthropogenic pressures and was mainly mediated by non-native species introductions. Specifically, we demonstrate that the negative effects of anthropogenic pressures on species richness and total abundance were compensated over time by the establishment of non-native species, a pattern consistent with previously reported biotic homogenization at the global scale. Overall, our study suggests that accounting for the complexity of community changes and its drivers is a crucial step to reach global conservation goals.

Understanding temporal variability across trophic levels and spatial scales in freshwater ecosystems.

A tenet of ecology is that temporal variability in ecological structure and processes tends to decrease with increasing spatial scales (from locales to regions) and levels of biological organization (from populations to communities). However, patterns in temporal variability across trophic levels and the mechanisms that produce them remain poorly understood. Here we analyzed the abundance time series of spatially structured communities (i.e., metacommunities) spanning basal resources to top predators from 355 freshwater sites across three continents. Specifically, we used a hierarchical partitioning method to disentangle the propagation of temporal variability in abundance across spatial scales and trophic levels. We then used structural equation modeling to determine if the strength and direction of relationships between temporal variability, synchrony, biodiversity, and environmental and spatial settings depended on trophic level and spatial scale. We found that temporal variability in abundance decreased from producers to tertiary consumers but did so mainly at the local scale. Species population synchrony within sites increased with trophic level, whereas synchrony among communities decreased. At the local scale, temporal variability in precipitation and species diversity were associated with population variability (linear partial coefficient, ß = 0.23) and population synchrony (ß = -0.39) similarly across trophic levels, respectively. At the regional scale, community synchrony was not related to climatic or spatial predictors, but the strength of relationships between metacommunity variability and community synchrony decreased systematically from top predators (ß = 0.73) to secondary consumers (ß = 0.54), to primary consumers (ß = 0.30) to producers (ß = 0). Our results suggest that mobile predators may often stabilize metacommunities by buffering variability that originates at the base of food webs. This finding illustrates that the trophic structure of metacommunities, which integrates variation in organismal body size and its correlates, should be considered when investigating ecological stability in natural systems. More broadly, our work advances the notion that temporal stability is an emergent property of ecosystems that may be threatened in complex ways by biodiversity loss and habitat fragmentation.

The color of environmental noise in river networks.

Despite its far-reaching implications for conservation and natural resource management, little is known about the color of environmental noise, or the structure of temporal autocorrelation in random environmental variation, in streams and rivers. Here, we analyze the geography, drivers, and timescale-dependence of noise color in streamflow across the U.S. hydrography, using streamflow time series from 7504 gages. We find that daily and annual flows are dominated by red and white spectra respectively, and spatial variation in noise color is explained by a combination of geographic, hydroclimatic, and anthropogenic variables. Noise color at the daily scale is influenced by stream network position, and land use and water management explain around one third of the spatial variation in noise color irrespective of the timescale considered. Our results highlight the peculiarities of environmental variation regimes in riverine systems, and reveal a strong human fingerprint on the stochastic patterns of streamflow variation in river networks.

Climate change and the global redistribution of biodiversity: substantial variation in empirical support for expected range shifts.

BACKGROUND: Among the most widely predicted climate change-related impacts to biodiversity are geographic range shifts, whereby species shift their spatial distribution to track their climate niches. A series of commonly articulated hypotheses have emerged in the scientific literature suggesting species are expected to shift their distributions to higher latitudes, greater elevations, and deeper depths in response to rising temperatures associated with climate change. Yet, many species are not demonstrating range shifts consistent with these expectations. Here, we evaluate the impact of anthropogenic climate change (specifically, changes in temperature and precipitation) on species' ranges, and assess whether expected range shifts are supported by the body of empirical evidence. METHODS: We conducted a Systematic Review, searching online databases and search engines in English. Studies were screened in a two-stage process (title/abstract review, followed by full-text review) to evaluate whether they met a list of eligibility criteria. Data coding, extraction, and study validity assessment was completed by a team of trained reviewers and each entry was validated by at least one secondary reviewer. We used logistic regression models to assess whether the direction of shift supported common range-shift expectations (i.e., shifts to higher latitudes and elevations, and deeper depths). We also estimated the magnitude of shifts for the subset of available range-shift data expressed in distance per time (i.e., km/decade). We accounted for methodological attributes at the study level as potential sources of variation. This allowed us to answer two questions: (1) are most species shifting in the direction we expect (i.e., each observation is assessed as support/fail to support our expectation); and (2) what is the average speed of range shifts? REVIEW FINDINGS: We found that less than half of all range-shift observations (46.60%) documented shifts towards higher latitudes, higher elevations, and greater marine depths, demonstrating significant variation in the empirical evidence for general range shift expectations. For the subset of studies looking at range shift rates, we found that species demonstrated significant average shifts towards higher latitudes (average = 11.8 km/dec) and higher elevations (average = 9 m/dec), although we failed to find significant evidence for shifts to greater marine depths. We found that methodological factors in individual range-shift studies had a significant impact on the reported direction and magnitude of shifts. Finally, we identified important variation across dimensions of range shifts (e.g., greater support for latitude and elevation shifts than depth), parameters (e.g., leading edge shifts faster than trailing edge for latitude), and taxonomic groups (e.g., faster latitudinal shifts for insects than plants). CONCLUSIONS: Despite growing evidence that species are shifting their ranges in response to climate change, substantial variation exists in the extent to which definitively empirical observations confirm these expectations. Even though on average, rates of shift show significant movement to higher elevations and latitudes for many taxa, most species are not shifting in expected directions. Variation across dimensions and parameters of range shifts, as well as differences across taxonomic groups and variation driven by methodological factors, should be considered when assessing overall confidence in range-shift hypotheses. In order for managers to effectively plan for species redistribution, we need to better account for and predict which species will shift and by how much. The dataset produced for this analysis can be used for future research to explore additional hypotheses to better understand species range shifts.

Unifying climate change biology across realms and taxa.

A major challenge in modern biology is to understand extinction risk from climate change across all realms. Recent research has revealed that physiological tolerance, behavioral thermoregulation, and small elevation shifts are dominant coping strategies on land, whereas large-scale latitudinal shifts are more important in the ocean. Freshwater taxa may face the highest global extinction risks. Nevertheless, some species in each realm face similar risks because of shared adaptive, dispersal, or physiological tolerances and abilities. Taking a cross-realm perspective offers unique research opportunities because confounding physical factors in one realm are often disaggregated in another realm. Cross-realm, across taxa, and other forms of climate change biology synthesis are needed to advance our understanding of emergent patterns of risk across all life.

Species better track climate warming in the oceans than on land.

There is mounting evidence of species redistribution as climate warms. Yet, our knowledge of the coupling between species range shifts and isotherm shifts remains limited. Here, we introduce BioShifts-a global geo-database of 30,534 range shifts. Despite a spatial imbalance towards the most developed regions of the Northern Hemisphere and a taxonomic bias towards the most charismatic animals and plants of the planet, data show that marine species are better at tracking isotherm shifts, and move towards the pole six times faster than terrestrial species. More specifically, we find that marine species closely track shifting isotherms in warm and relatively undisturbed waters (for example, the Central Pacific Basin) or in cold waters subject to high human pressures (for example, the North Sea). On land, human activities impede the capacity of terrestrial species to track isotherm shifts in latitude, with some species shifting in the opposite direction to isotherms. Along elevational gradients, species follow the direction of isotherm shifts but at a pace that is much slower than expected, especially in areas with warm climates. Our results suggest that terrestrial species are lagging behind shifting isotherms more than marine species, which is probably related to the interplay between the wider thermal safety margin of terrestrial versus marine species and the more constrained physical environment for dispersal in terrestrial versus marine habitats.

Publisher Correction: Species traits and phylogenetic conservatism of climate-induced range shifts in stream fishes.

This corrects the article DOI: 10.1038/ncomms6053.

Species traits and phylogenetic conservatism of climate-induced range shifts in stream fishes.

Understanding climate-induced range shifts is crucial for biodiversity conservation. However, no general consensus has so far emerged about the mechanisms involved and the role of phylogeny in shaping species responses has been poorly explored. Here, we investigate whether species traits and their underlying phylogenetic constraints explain altitudinal shifts at the trailing and leading edges of stream fish species ranges. We demonstrate that these shifts are related to dissimilar mechanisms: whereas range retractions show some support for phylogenetic clustering due to a high level of conservatism in thermal safety margins, range expansions are underpinned by both evolutionarily conserved and labile traits, notably trophic position and life-history strategy, hence decreasing the strength of phylogenetic signal. Therefore, while climate change brings many difficulties in establishing a general understanding of species vulnerability, these findings emphasize how combining trait-based approaches in light of the species evolutionary history may offer new opportunities in facing conservation challenges.

Combining genetic and demographic data for prioritizing conservation actions: insights from a threatened fish species.

Prioritizing and making efficient conservation plans for threatened populations requires information at both evolutionary and ecological timescales. Nevertheless, few studies integrate multidisciplinary approaches, mainly because of the difficulty for conservationists to assess simultaneously the evolutionary and ecological status of populations. Here, we sought to demonstrate how combining genetic and demographic analyses allows prioritizing and initiating conservation plans. To do so, we combined snapshot microsatellite data and a 30-year-long demographic survey on a threatened freshwater fish species (Parachondrostoma toxostoma) at the river basin scale. Our results revealed low levels of genetic diversity and weak effective population sizes (<63 individuals) in all populations. We further detected severe bottlenecks dating back to the last centuries (200-800 years ago), which may explain the differentiation of certain populations. The demographic survey revealed a general decrease in the spatial distribution and abundance of P. toxostoma over the last three decades. We conclude that demo-genetic approaches are essential for (1) identifying populations for which both evolutionary and ecological extinction risks are high; and (2) proposing conservation plans targeted toward these at risk populations, and accounting for the evolutionary history of populations. We suggest that demo-genetic approaches should be the norm in conservation practices. We combined genetic and demographic data from a threatened freshwater fish species (Parachondrostoma toxostoma) at the river basin scale for conservation purposes. Genetic diversity and effective population sizes are very low, probably due to the strong genetic bottlenecks detected in this study. The species spatial distribution and abundance also decreased during the last decades.

Modeling the chemical and toxic water status of the Scheldt basin (Belgium), using aquatic invertebrate assemblages and an advanced modeling method.

Self-Organizing Maps have been used on monitoring sites in several Scheldt sub-basins to identify the main aquatic invertebrate assemblages and relate them to the physico-chemical and toxic water status. 12 physico-chemical variables and 2 estimates of toxic risk were available for a dataset made up of a total of 489 records. Two of the five defining clusters reflecting a relatively clean environment were composed by very well diversified functional feeding groups and sensitive taxa. The cleanest assemblage was mainly linked to the sites from the Nete sub-basin. The three other clusters were inversely described with a dominance of oligochaetes and deposit feeders as well as a bad water quality. Such an analysis can be used to support ecological status assessment of rivers and thus might be useful for decision-makers in the evaluation of chemical and toxic water status, as required by the EU Water Framework Directive.

Assessment of stream biological responses under multiple-stress conditions.

BACKGROUND, AIM AND SCOPE: Due to the numerous anthropogenic stress factors that affect aquatic ecosystems, a better understanding of the adverse consequences on the biological community of combined pressures is needed to attain the objectives of the European Water Framework Directive. In this study we propose an innovative approach to assess the biological impact of toxicants under field conditions on a large spatial scale. MATERIALS AND METHODS: Artificial Neural Network (ANN) analyses, focusing on impacts at the community level, were carried out to identify the relative importance of environmental and toxic stress factors on the patterns observed in the aquatic invertebrate fauna from the Scheldt basin (Belgium). RESULTS AND DISCUSSION: Our results show that the use of the backpropagation algorithm of the ANN is a promising method to highlight the relationship between environmental pollution and biological responses. This method allows the effects of chemical exposure to be distinguished from the effects caused by other stressors in running waters. Moreover, the use of an overall estimate for toxic pressure in predictive models enables the links between toxicants and community alterations in the field to be clarified. The ANN correctly predicts 74% of samples with an area under the curve of 0.89 and a Cohen's kappa coefficient of 0.64. Organic load, oxygen availability, water temperature and the nitrate concentration appeared important factors in predicting aquatic invertebrate assemblages. On the other hand, toxic pressure did not seem relevant for these assemblages, suggesting that the water quality characteristics were therefore more important than exposure to toxicants in the water phase for the aquatic invertebrate communities in the study area. However, we suggest that the high organic load encountered in the Scheldt basin may lead to an underestimation of the impact of toxicity.