Forecasting fine‐scale changes in the food‐web structure of coastal marine communities under climate change

Climate change is inducing deep modifications in local communities worldwide as a consequence of individualistic species range shifts. Understanding how complex interaction networks will be reorganized under climate change represents a major challenge in the fields of ecology and biogeography. However, forecasting the potential effects of climate change on local communities, and more particularly on food‐web structure, requires the consideration of highly structuring processes, such as trophic interactions. A major breakthrough is therefore expected by combining predictive models integrating habitat selection processes, the physiological limits of marine species and their trophic interactions. In this study, we forecasted the potential impacts of climate change on the local food‐web structure of the highly threatened Gulf of Gabes ecosystem located in the south of the Mediterranean Sea. We coupled the climatic envelope and habitat models to an allometric niche food web model, hence taking into account the different processes acting at regional (climate) and local scales (habitat selection and trophic interactions). Our projections under the A2 climate change scenario showed that future food webs would be composed of smaller species with fewer links, resulting in a decrease of connectance, generality, vulnerability and mean trophic level of communities and an increase of the average path length, which may have large consequences on ecosystem functioning. The unified framework presented here, by connecting food‐web ecology, biogeography and seascape ecology, allows the exploration of spatial aspects of interspecific interactions under climate change and improves our current understanding of climate change impacts on local marine food webs.     

Habitat availability and gene flow influence diverging local population trajectories under scenarios of climate change: a place‐based approach

Ecological niche theory holds that species distributions are shaped by a large and complex suite of interacting factors. Species distribution models (SDMs) are increasingly used to describe species’ niches and predict the effects of future environmental change, including climate change. Currently, SDMs often fail to capture the complexity of species’ niches, resulting in predictions that are generally limited to climate‐occupancy interactions. Here, we explore the potential impact of climate change on the American pika using a replicated place‐based approach that incorporates climate, gene flow, habitat configuration, and microhabitat complexity into SDMs. Using contemporary presence–absence data from occupancy surveys, genetic data to infer connectivity between habitat patches, and 21 environmental niche variables, we built separate SDMs for pika populations inhabiting eight US National Park Service units representing the habitat and climatic breadth of the species across the western United States. We then predicted occurrence probability under current (1981–2010) and three future time periods (out to 2100). Occurrence probabilities and the relative importance of predictor variables varied widely among study areas, revealing important local‐scale differences in the realized niche of the American pika. This variation resulted in diverse and – in some cases – highly divergent future potential occupancy patterns for pikas, ranging from complete extirpation in some study areas to stable occupancy patterns in others. Habitat composition and connectivity, which are rarely incorporated in SDM projections, were influential in predicting pika occupancy in all study areas and frequently outranked climate variables. Our findings illustrate the importance of a place‐based approach to species distribution modeling that includes fine‐scale factors when assessing current and future climate impacts on species’ distributions, especially when predictions are intended to manage and conserve species of concern within individual protected areas.     

Evidence for shared broad‐scale climatic niches of diploid and polyploid plants

Whole‐genome duplication (polyploidy) occurs frequently and repeatedly within species, often forming new lineages that contribute to biodiversity, particularly in plants. Establishment and persistence of new polyploids may be thwarted by competition with surrounding diploids; however, climatic niche shifts, where polyploids occupy different niches than diploid progenitors, may help polyploids overcome this challenge. We tested for climatic niche shifts between cytotypes using a new ordination approach and an unprecedentedly large data set containing young, conspecific diploids and polyploids. Despite expectations of frequent niche shifts, we show evidence for alternative patterns, such as niche conservatism and contraction, rather than a prevalent pattern of niche shifts. In addition, we explore how interpreting climatic niches plotted on environmental niche (principal component) axes can generate hypotheses about processes underlying niche dynamics. Dispersal capabilities or other life‐history traits, rather than shifts to new climatic niches, could better explain polyploid persistence in the long term.     

Identifying climatic niche shifts using coarse‐grained occurrence data: a test with non‐native freshwater fish

Aim We tested whether coarse‐grained occurrence data can be used to detect climatic niche shifts between native and non‐native ranges for a set of widely introduced freshwater fishes. Location World‐wide. Methods We used a global database of freshwater fish occurrences at the river basin scale to identify native and non‐native ranges for 18 of the most widely introduced fish species. We also examined climatic conditions within each river basin using fine‐grained climate data. We combined this information to test whether climatic niche shifts have occurred between native and non‐native ranges. We defined climatic niche shifts as instances where the ranges of a climatic variable within native and non‐native basins exhibit zero overlap. Results We detected at least one climatic niche shift for each of the 18 studied species. However, we did not detect common patterns in the thermal preference or biogeographic origin of the non‐native fish, hence suggesting a species‐specific response. Main conclusions Coarse‐grained occurrence data can be used to detect climatic niche shifts. They also enable the identification of the species experiencing niche shifts, although the mechanisms responsible for these shifts (e.g. local adaptation, dispersal limitation or physiological constraints) have yet to be determined. Furthermore, the coarse‐grained approach, which highlights regions where climatic niche shifts have occurred, can be used to select specific river basins for more detailed, fine‐grained studies.     

Spatiotemporally explicit demographic modelling supports a joint effect of historical barriers to dispersal and contemporary landscape composition on structuring genomic variation in a red‐listed grasshopper

Inferring the processes underlying spatial patterns of genomic variation is fundamental to understand how organisms interact with landscape heterogeneity and to identify the factors determining species distributional shifts. Here, we use genomic data (restriction site‐associated DNA sequencing) to test biologically informed models representing historical and contemporary demographic scenarios of population connectivity for the Iberian cross‐backed grasshopper Dociostaurus hispanicus, a species with a narrow distribution that currently forms highly fragmented populations. All models incorporated biological aspects of the focal taxon that could hypothetically impact its geographical patterns of genomic variation, including (a) spatial configuration of impassable barriers to dispersal defined by topographic landscapes not occupied by the species; (b) distributional shifts resulting from the interaction between the species bioclimatic envelope and Pleistocene glacial cycles; and (c) contemporary distribution of suitable habitats after extensive land clearing for agriculture. Spatiotemporally explicit simulations under different scenarios considering these aspects and statistical evaluation of competing models within an Approximate Bayesian Computation framework supported spatial configuration of topographic barriers to dispersal and human‐driven habitat fragmentation as the main factors explaining the geographical distribution of genomic variation in the species, with no apparent impact of hypothetical distributional shifts linked to Pleistocene climatic oscillations. Collectively, this study supports that both historical (i.e., topographic barriers) and contemporary (i.e., anthropogenic habitat fragmentation) aspects of landscape composition have shaped major axes of genomic variation in the studied species and emphasizes the potential of model‐based approaches to gain insights into the temporal scale at which different processes impact the demography of natural populations.     

Glacial refugia,recolonization patterns and diversification forces in Alpine‐endemic Megabunus harvestmen

The Pleistocene climatic fluctuations had a huge impact on all life forms, and various hypotheses regarding the survival of organisms during glacial periods have been postulated. In the European Alps, evidence has been found in support of refugia outside the ice shield (massifs de refuge) acting as sources for postglacial recolonization of inner‐Alpine areas. In contrast, evidence for survival on nunataks, ice‐free areas above the glacier, remains scarce. Here, we combine multivariate genetic analyses with ecological niche models (ENMs) through multiple timescales to elucidate the history of Alpine Megabunus harvestmen throughout the ice ages, a genus that comprises eight high‐altitude endemics. ENMs suggest two types of refugia throughout the last glacial maximum, inner‐Alpine survival on nunataks for four species and peripheral refugia for further four species. In some geographic regions, the patterns of genetic variation are consistent with long‐distance dispersal out of massifs de refuge, repeatedly coupled with geographic parthenogenesis. In other regions, long‐term persistence in nunataks may dominate the patterns of genetic divergence. Overall, our results suggest that glacial cycles contributed to allopatric diversification in Alpine Megabunus, both within and at the margins of the ice shield. These findings exemplify the power of ENM projections coupled with genetic analyses to identify hypotheses about the position and the number of glacial refugia and thus to evaluate the role of Pleistocene glaciations in driving species‐specific responses of recolonization or persistence that may have contributed to observed patterns of biodiversity.     

Spatio‐temporal variation of biotic factors underpins contemporary range dynamics of congeners

Species’ ranges are complex often exhibiting multidirectional shifts over space and time. Despite the strong fingerprint of recent historical climate change on species’ distributions, biotic factors such as loss of vegetative habitat and the presence of potential competitors constitute important yet often overlooked drivers of range dynamics. Furthermore, short‐term changes in environmental conditions can influence the underlying processes of local extinction and local colonization that drive range shifts, yet are rarely considered at broad scales. We used dynamic state‐space occupancy models to test multiple hypotheses of the relative importance of major drivers of range shifts of Golden‐winged Warblers (Vermivora chrysoptera) and Blue‐winged Warblers (V. cyanoptera) between 1983 and 2012 across North America: warming temperatures; habitat changes; and occurrence of congeneric species, used here as proxy for biotic interactions. Dynamic occupancies for both species were most influenced by spatial relative to temporal variation in temperature and habitat. However, temporal variation in temperature anomalies and biotic interactions remained important. The two biotic factors considered, habitat change and biotic interactions, had the largest relative effect on estimated extinction rates followed by abiotic temperature anomalies. For the Golden‐winged Warbler, the predicted presence of the Blue‐winged Warbler, a hypothesized competitor, most influenced extinction probabilities, contributing to evidence supporting its role in site‐level species replacement. Given the overall importance of biotic factors on range‐wide dynamic occupancies, their consideration alongside abiotic factors should not be overlooked. Our results suggest that warming compounds the negative effect of habitat loss emphasizing species’ need for habitat to adapt to a changing climate. Notably, even closely related species exhibited individual responses to abiotic and biotic factors considered.     

Correlation between genetic diversity and environmental suitability: taking uncertainty from ecological niche models into account

The hindcast of shifts in the geographical ranges of species as estimated by ecological niche modelling (ENM) has been coupled with phylogeographical patterns, allowing the inference of past processes that drove population differentiation and genetic variability. However, more recently, some studies have suggested that maps of environmental suitability estimated by ENM may be correlated to species' abundance, raising the possibility of using environmental suitability to infer processes related to population demographic dynamics and genetic variability. In both cases, one of the main problems is that there is a wide variation in ENM development methods and climatic models. In this study, we analyse the relationship between heterozygosity (He) and environmental suitability from multiple ENMs for 25 population estimates for Dipteryx alata, a widely distributed, endemic tree species of the Cerrado region of central Brazil. We propose a new approach for generating a statistical distribution of correlations under randomly generated ENM. The confidence intervals from these distributions indicate how model selection with different properties affects the ability to detect a correlation of interest (e.g. the correlation between He and suitability). Additionally, our approach allows us to explore which particular ensemble of ENMs produces the better result for finding an association between environmental suitability and He. Caution is necessary when choosing a method or a climatic data set for modelling geographical distributions, but the new approach proposed here provides a conservative way to evaluate the ability of ensembles to detect patterns of interest.     

Climatic control of dispersal–ecological specialization trade‐offs: a metacommunity process at the heart of the latitudinal diversity gradient?

We outline the potentially important role of dispersal in linking diversity patterns at different spatial and temporal scales, and the resulting potential to link hypotheses explaining macroscale patterns of diversity. We do this by proposing a possible mechanism linking climate to diversity patterns: we argue that climate, via effects of continuity of habitat availability in space and time, mediates a dispersal–ecological specialization trade‐off at the metacommunity level that leads to latitudinal trends in dispersal ability, ecological specialization, range sizes, speciation and species richness, ultimately driving the latitudinal diversity gradient. This trade‐off constitutes a possible mechanism for the strong macroscale correlation between climate and species richness that is consistent with recent ideas about niche conservatism and gradient lengths, as well as other leading hypotheses. We present an overview of predictions derived from our ideas. Of these, some have already been tested and supported and others are still open to debate or need testing. Together they provide a unique set of predictions that allows falsification.     

Reconstructing changes in the genotype,phenotype, and climatic niche of an introduced species

An introduced species must contend with enormous environmental variation in its introduced range. In this study, we use niche models and ordination analyses to reconstruct changes in genotype, phenotype, and climatic niche of Johnsongrass Sorghum halepense, which is regarded as one of the world's most threatening invasive plants. In the United States, Johnsongrass has rapidly evolved within‐ and among‐population genetic diversity; our results show that genetic differentiation in expanding Johnsongrass populations has resulted in phenotypic variation that is consistent with habitat and climatic variation encountered during its expansion. Moreover, Johnsongrass expanded from agricultural to non‐agricultural habitat, and now, despite occupying overlapping ranges, extant agricultural and non‐agricultural populations are genetically and phenotypically distinct and manifest different plastic responses when encountering environmental variation. Non‐agricultural accessions are broadly distributed in climatic and geographic space and their fitness traits demonstrate plastic responses to common garden conditions that are consistent with local specialization. In contrast, agricultural accessions demonstrate ‘general purpose’ plastic responses and have more restricted climatic niches and geographic distributions. They also grow much larger than non‐agricultural accessions. If these differences are adaptive, our results suggest that adaptation to local habitat variation plays a crucial role in the ecology of this invader. Further, its success relates to its ability to succeed on dual fronts, by responding simultaneously to habitat and climate variability and by capitalizing on differential responses to these factors during its range expansion.     

The shaping of genetic variation in edge‐of‐range populations under past and future climate change

With rates of climate change exceeding the rate at which many species are able to shift their range or adapt, it is important to understand how future changes are likely to affect biodiversity at all levels of organisation. Understanding past responses and extent of niche conservatism in climatic tolerance can help predict future consequences. We use an integrated approach to determine the genetic consequences of past and future climate changes on a bat species, Plecotus austriacus. Glacial refugia predicted by palaeo‐modelling match those identified from analyses of extant genetic diversity and model‐based inference of demographic history. Former refugial populations currently contain disproportionately high genetic diversity, but niche conservatism, shifts in suitable areas and barriers to migration mean that these hotspots of genetic diversity are under threat from future climate change. Evidence of population decline despite recent northward migration highlights the need to conserve leading‐edge populations for spearheading future range shifts.     

Tree hydraulic traits are coordinated and strongly linked to climate‐of‐origin across a rainfall gradient

Plant hydraulic traits capture the impacts of drought stress on plant function, yet vegetation models lack sufficient information regarding trait coordination and variation with climate‐of‐origin across species. Here, we investigated key hydraulic and carbon economy traits of 12 woody species in Australia from a broad climatic gradient, with the aim of identifying the coordination among these traits and the role of climate in shaping cross‐species trait variation. The influence of environmental variation was minimized by a common garden approach, allowing us to factor out the influence of environment on phenotypic variation across species. We found that hydraulic traits (leaf turgor loss point, stomatal sensitivity to drought [P_gs], xylem vulnerability to cavitation [P_x], and branch capacitance [C_branch]) were highly coordinated across species and strongly related to rainfall and aridity in the species native distributional range. In addition, trade‐offs between drought tolerance and plant growth rate were observed across species. Collectively, these results provide critical insight into the coordination among hydraulic traits in modulating drought adaptation and will significantly advance our ability to predict drought vulnerability in these dominant trees species.     

Comparing macroecological patterns across continents: evolution of climatic niche breadth in varanid lizards

Macroecological analyses often test hypotheses at the global scale, or among more closely related species in a single region (e.g. continent). Here, we test several hypotheses about climatic niche widths among relatively closely related species that occur across multiple continents, and compare patterns within and across continents to see if they differ. We focus on the lizard genus Varanus (monitor lizards), which occurs in diverse environments in Africa, Asia, and Australia. We address three main questions. 1) How do climatic niche breadths of species on a given niche axis change based on the position of species along that niche axis? (E.g. are species that occur in more extreme environments more narrowly specialized for those conditions?) 2) Are there trade‐offs in niche breadths on temperature and precipitation axes among species, or are niche widths on different axes positively related? 3) Is variation in niche breadths among species explained primarily by within‐locality seasonal variation, or by differences in climatic conditions among localities across the species range? We generate a new time‐calibrated phylogeny for Varanus and test these hypotheses within and between continents using climatic data and phylogenetic methods. Our results show that patterns on each continent often parallel each other and global patterns. However, in many other cases, the strength of relationships can change dramatically among closely related species on different continents. Overall, we found that: 1) species in warmer environments have narrower temperature niche breadths, but there is no relationship between precipitation niche breadth and niche position; 2) temperature and precipitation niche breadths tend to be positively related among species, rather than showing trade‐offs; and 3) within‐locality seasonal variation explains most variation in climatic niche breadths. Some of these results are concordant with previous studies (in amphibians and North American lizards), and might represent general macroecological patterns.     

Ecophysiological variation across a forest‐ecotone gradient produces divergent climate change vulnerability within species

Climate change related risks and impacts on ectotherms will be mediated by habitats and their influence on local thermal environments. While many studies have documented morphological and genetic aspects of niche divergence across habitats, few have examined thermal performance across such gradients and directly linked this variation to contemporary climate change impacts. In this study, we quantified variation in thermal performance across a gradient from forest to gallery forest‐savanna mosaic in Cameroon for a skink species (Trachylepis affinis) known to be diverging genetically and morphologically across that habitat gradient. Based on these results, we then applied a mechanistic modelling approach (NicheMapR) to project changes in potential activity, as constrained by thermal performance, in response to climate change. As a complimentary approach, we also compared mechanistic projections with climate‐driven changes in habitat suitability based on species distribution models of forest and ecotone skinks. We found that ecotone skinks may benefit from warming and experience increased activity while forest skinks will likely face a drastic decrease in thermal suitability across the forest zone. Species distribution models projected that thermal suitability for forest skinks in coastal forests would decline but in other parts of the forest zone skinks are projected to experience increased thermal suitability. The results here highlight the utility of mechanistic approaches in revealing and understanding patterns of climate change vulnerability which may not be detected with species distribution models alone. This study also emphasizes the importance of intra‐specific physiological variation, and habitat‐specific thermal performance relationships in particular, in determining warming responses.     

Spatial genetic variation and habitat association of <Emphasis Type="Italic">Rhinichthys cataractae</Emphasis>, the longnose dace,in the Driftless Area of the upper Mississippi River basin

The Driftless Area of the upper Mississippi River drainage is a unique geographic region because of its complex geological history and the influence of recent, intensive human activities. The longnose dace, Rhinichthys cataractae, is a relatively common, small freshwater fish that is distributed in swift, cool streams within the region. The aim of this study was to determine the spatial genetic differentiation of the longnose dace and define the broad scale environmental variables that shape the distribution of the species in the southwestern portion of the Driftless Area. Genotypic data from seven microsatellite loci were analyzed for 276 individuals from 15 localities representing major drainages within the region in northeast Iowa. Broad scale environmental variables including hydrologic, soil, and climatic factors were evaluated to construct an ecological niche model (ENM) to predict the suitability of habitat for the species within the region. Results of the genetic analyses revealed two distinct, but somewhat admixed genetic clusters of longnose dace in Iowa. The genetic differentiation between localities and between drainages was low to moderate with some evidence of isolation by distance. Most of the variation was observed by differences between individuals within local populations. The ENM generated largely reflected the known distribution of the species in Iowa with a decreasing probability of suitable habitat from northern to southern drainages. Geologic factors played a key role in the model. The distribution and population structure of the longnose dace in the northeast Iowa revealed that isolation by distance, historical processes and the underlying geology are primarily responsible for the observed spatial distribution of genetic variation.     

Acido‐ and neutrophilic temperate forest plants display distinct shifts in ecological pH niche across north‐western Europe

Ecological niches of organisms vary across geographical space, but niche shift patterns between regions and the underlying mechanisms remain largely unexplored. We studied shifts in the pH niche of 42 temperate forest plant species across a latitudinal gradient from northern France to boreo‐nemoral Sweden. We asked 1) whether species restrict their niches with increasing latitude as they reach their northern range margin (environmental constraints); 2) whether species expand their niches with increasing latitude as regional plant species richness decreases (competitive release); and 3) whether species shift their niche position toward more acidic sites with increasing latitude as the relative proportion of acidic soils increases (local adaptation). Based on 1458 vegetation plots and corresponding soil pH values, we modelled species response curves using Huisman–Olff–Fresco models. Four niche measures (width, position, left and right border) were compared among regions by randomization tests. We found that with increasing latitude, neutrophilic species tended to retreat from acidic sites, indicating that these species retreat to more favorable sites when approaching their range margin. Alternatively, these species might benefit from enhanced nitrogen deposition on formerly nutrient‐poor, acidic sites in southern regions or lag behind in post‐glacial recolonization of potential habitats in northern regions. Most acidophilic species extended their niche toward more base‐rich sites with increasing latitude, indicating competitive release from neutrophilic species. Alternatively, acidophilic species might benefit from optimal climatic conditions in the north where some have their core distribution area. Shifts in the niche position suggested that local adaptation is of minor importance. We conclude that shifts in the pH niche of temperate forest plants are the rule, but the directions of the niche shifts and possible explanations vary. Our study demonstrates that differentiating between acidophilic and neutrophilic species is crucial to identify general patterns and underlying mechanisms.     

Niche models tell half the story: spatial context and life‐history traits influence species responses to global change

Aim While niche models are typically used to assess the vulnerability of species to climate change, they have been criticized for their limited assessment of threats other than climate change. We attempt to evaluate this limitation by combining niche models with life‐history models to investigate the relative influence of climate change and a range of fire regimes on the viability of a long‐lived plant population. Specifically, we investigate whether range shift due to climate change is a greater threat to an obligate seeding fire‐prone shrub than altered fire frequency and how these two threatening processes might interact. Location Australian sclerophyll woodland and heathland. Methods The study species is Leucopogon setiger, an obligate seeding fire‐prone shrub. A spatially explicit stochastic matrix model was constructed for this species and linked with a dynamic niche model and fire risk functions representing a suite of average fire return intervals. We compared scenarios with a variety of hypothetical patches, a patch framework based upon current habitat suitability and one with dynamic habitat suitability based on climate change scenarios A1FI and A2. Results Leucopogon setiger was found to be sensitive to fire frequency, with shorter intervals reducing expected minimum abundances (EMAs). Spatial decoupling of fires across the landscape reduced the vulnerability of the species to shortened fire frequencies. Shifting habitat, while reducing EMAs, was less of a threat to the species than frequent fire. Main conclusions Altered fire regime, in particular more frequent fires relative to the historical regime, was predicted to be a strong threat to this species, which may reflect a vulnerability of obligate seeders in general. Range shifts induced by climate change were a secondary threat when habitat reductions were predicted. Incorporating life‐history traits into habitat suitability models by linking species distribution models with population models allowed for the population‐level evaluation of multiple stressors that affect population dynamics and habitat, ultimately providing a greater understanding of the impacts of global change than would be gained by niche models alone. Further investigations of this type could elucidate how particular bioecological factors can affect certain types of species under global change.     

Early stages of divergence: phylogeography, climate modeling, and morphological differentiation in the South American lizard Liolaemus petrophilus (Squamata: Liolaemidae)

This study examines the phylogeographic structure within the Patagonian lizard Liolaemus petrophilus and tests for patterns of between-clade morphological divergence and sexual dimorphism, as well as demographic and niche changes associated with Pleistocene climate changes. We inferred intraspecific relationships, tested hypotheses for historical patterns of population expansion, and incorporated ecological niche modeling (ENM) with standard morphological and geometric morphometric analyses to examine between-clade divergence as indirect evidence for adaptation to different niches. The two inferred haploclades diverged during the early Pleistocene with the Southern clade depicting the genetic signature of a recent population increase associated with expanding niche envelope, whereas the Northern clade shows stable populations in a shrinking niche envelope. The combination of molecular evidence for postisolation demographic change and ENM, suggest that the two haploclades have responded differently to Pleistocene climatic events.