The importance of marginal population hotspots of cold‐adapted species for research on climate change and conservation

Areas hosting hotspots of low‐latitude marginal populations of cold‐adapted plant species could be key areas for understanding geographical attributes that result in refugia during climatic shifts as well as the conservation of genetic diversity in the face of climate change. Low‐latitude populations of cold‐adapted plants are important because they may harbour the combination of alleles that foster persistence in a warmer climate. Consequently, identification of areas where arctic‐alpine, circumpolar and circumboreal species reach the low‐latitude ends of their distribution will present a unique opportunity to uncover processes that shaped current biogeographical patterns, as well as prepare for future scenarios. Here, we identify 35 main marginal population hotspots (19 and 16 areas in North America and Europe, respectively) of 183 plant taxa. These hotspots represent areas where southern marginal populations of cold‐adapted species co‐occur. The identification of hotspots was based on geographic overlap of southernmost locations of the target species, in a 50 × 50 km grid. With a threshold of two species in a single grid cell or in two contiguous cells, the analysis revealed that hotspots are in most cases located in the southern portion of major mountain chains. However, hotspots also occur in lowland areas at high latitudes (Fennoscandia, Alaska, Hudson Bay) which do not necessarily correspond to known cold‐ or warm‐stage refugia (e.g. Alps). Rockies and Sierra Nevada both in California and Spain, Apennines, and the southern Scandes, maintain their hotspot status even with more stringent cut‐off thresholds (>3 and >5 species per cell group). From a conservation point of view, our analysis reveals that only a small portion of the hotspots are currently included within protected areas. We discuss the importance of marginal population hotspots to future research on climate change and, finally, outline how conservation strategies can capitalize on the knowledge gained from studying climate change effects on cold‐adapted plants.     

From glacial refugia to hydrological microrefugia: Factors and processes driving the persistence of the climate relict tree Zelkova sicula

With only two tiny populations, the climate relict Zelkova sicula (Sicily, Italy) is one of the rarest trees in the world. It also represents the most marginal member of genus Zelkova that was widespread in the broadleaved forests thriving in warm–temperate climates throughout Eurasia until the Last Glacial Age. Occurring at the westernmost range of the genus under typical Mediterranean climate, the micro‐topographic settings have always appeared crucial for the survival of this relict. However, the factors and processes actually involved in its persistence in the current refugia, as well as the response of similar relict trees in arid environments, are poorly understood worldwide. In the aim to elucidate these aspects, in the two sites hosting Z. sicula analyses of topographical attributes were combined with investigations on soil moisture dynamics. Additionally, plants’ growth and spatial distribution patterns were analyzed to detect fine‐scale differences between populations and assess the possible ecological amplitude of the species. Results revealed that convergent topographies are basic determinants of microrefugia in arid environments. Within the investigated sites, underground moisture never decreases below 25%, buffering seasonal rainfall fluctuations. Therefore, hydrological microrefugia play a key role in decoupling from regional climate, supporting the target species in coping with an unsuitable climatic envelope. Additionally, the inter‐population variability of biometric attributes showed that individual growth is site‐dependent and the species retains a relative ecological plasticity, whereas the strongly clumped spatial patterns confirmed the common clonal growth. On one hand, deeply incised landforms have acted as effective hydrologic microrefugia, on the other clonality coupled with triploidy supposedly improved the resistance of Z. sicula to harsh environments, though entailing inability to reproduce sexually. Most likely, sterility and environmental/physical barriers that have existed for millennia have prevented this relict from leaving the last suitable microrefugia, resulting in the two current rear edge populations.     

Losing your edge: climate change and the conservation value of range‐edge populations

Populations occurring at species' range edges can be locally adapted to unique environmental conditions. From a species' perspective, range‐edge environments generally have higher severity and frequency of extreme climatic events relative to the range core. Under future climates, extreme climatic events are predicted to become increasingly important in defining species' distributions. Therefore, range‐edge genotypes that are better adapted to extreme climates relative to core populations may be essential to species' persistence during periods of rapid climate change. We use relatively simple conceptual models to highlight the importance of locally adapted range‐edge populations (leading and trailing edges) for determining the ability of species to persist under future climates. Using trees as an example, we show how locally adapted populations at species' range edges may expand under future climate change and become more common relative to range‐core populations. We also highlight how large‐scale habitat destruction occurring in some geographic areas where many species range edge converge, such as biome boundaries and ecotones (e.g., the arc of deforestation along the rainforest‐cerrado ecotone in the southern Amazonia), can have major implications for global biodiversity. As climate changes, range‐edge populations will play key roles in helping species to maintain or expand their geographic distributions. The loss of these locally adapted range‐edge populations through anthropogenic disturbance is therefore hypothesized to reduce the ability of species to persist in the face of rapid future climate change.     

The future distribution of river fish: The complex interplay of climate and land use changes,species dispersal and movement barriers

The future distribution of river fishes will be jointly affected by climate and land use changes forcing species to move in space. However, little is known whether fish species will be able to keep pace with predicted climate and land use‐driven habitat shifts, in particular in fragmented river networks. In this study, we coupled species distribution models (stepwise boosted regression trees) of 17 fish species with species‐specific models of their dispersal (fish dispersal model FIDIMO) in the European River Elbe catchment. We quantified (i) the extent and direction (up‐ vs. downstream) of predicted habitat shifts under coupled “moderate” and “severe” climate and land use change scenarios for 2050, and (ii) the dispersal abilities of fishes to track predicted habitat shifts while explicitly considering movement barriers (e.g., weirs, dams). Our results revealed median net losses of suitable habitats of 24 and 94 river kilometers per species for the moderate and severe future scenarios, respectively. Predicted habitat gains and losses and the direction of habitat shifts were highly variable among species. Habitat gains were negatively related to fish body size, i.e., suitable habitats were projected to expand for smaller‐bodied fishes and to contract for larger‐bodied fishes. Moreover, habitats of lowland fish species were predicted to shift downstream, whereas those of headwater species showed upstream shifts. The dispersal model indicated that suitable habitats are likely to shift faster than species might disperse. In particular, smaller‐bodied fish (<200 mm) seem most vulnerable and least able to track future environmental change as their habitat shifted most and they are typically weaker dispersers. Furthermore, fishes and particularly larger‐bodied species might substantially be restricted by movement barriers to respond to predicted climate and land use changes, while smaller‐bodied species are rather restricted by their specific dispersal ability.     

Arctic and boreal plant species decline at their southern range limits in the Rocky Mountains

Climate change is predicted to cause a decline in warm‐margin plant populations, but this hypothesis has rarely been tested. Understanding which species and habitats are most likely to be affected is critical for adaptive management and conservation. We monitored the density of 46 populations representing 28 species of arctic‐alpine or boreal plants at the southern margin of their ranges in the Rocky Mountains of Montana, USA, between 1988 and 2014 and analysed population trends and relationships to phylogeny and habitat. Marginal populations declined overall during the past two decades; however, the mean trend for 18 dicot populations was ?5.8% per year, but only ?0.4% per year for the 28 populations of monocots and pteridophytes. Declines in the size of peripheral populations did not differ significantly among tundra, fen and forest habitats. Results of our study support predicted effects of climate change and suggest that vulnerability may depend on phylogeny or associated anatomical/physiological attributes.     

Thermal adaptation in<Emphasis Type="Italic">Drosophila serrata</Emphasis> under conditions linked to its southern border: Unexpected patterns from laboratory selection suggest limited evolutionary potential

To investigate the ability ofDrosophila serrata to adapt to thermal conditions over winter at the species southern border, replicate lines from three source locations were held as discrete generations over three years at either 19‡C (40 generations) or temperatures fluctuating between 7‡C and 18δC (20 generations). Populations in the fluctuating environment were maintained either with an adult 0‡C cold shock or without a shock. These conditions were expected to result in temperature-specific directional selection for increased viability and productivity under both temperature regimes, and reduced development time under the fluctuating-temperature regime. Selection responses of all lines were tested under both temperature regimes after controlling for carry-over effects by rearing lines in these environments for two generations. When tested in the 19‡C environment, lines evolving at 19‡C showed a faster development time and a lower productivity relative to the other lines, while cold shock reduced development time and productivity of all lines. When tested in the fluctuating environment, productivity of the 7–18‡C lines selected with a cold shock was relatively lower than that of lines selected without a shock, but this pattern was not observed in the other populations. Viability and body size as measured by wing length were not altered by selection or cold shock, although there were consistent effects of source population on wing length. These results provide little evidence for temperature-specific adaptation inD. serrata —although the lines had diverged for some traits, these changes were not consistent with a priori predictions. In particular, there was no evidence for life-history changes reflecting adaptation to winter conditions at the southern border. The potential forD. serrata to adapt to winter conditions may therefore be limited.     

Meta‐analyzing the likely cross‐species responses to climate change

Ecological Niche Models (ENMs) have different performances in predicting potential geographic distributions. Here we meta‐analyzed the likely effects of climate change on the potential geographic distribution of 1,205 bird species from the Neotropical region, modeled using eight ENMs and three Atmosphere‐Ocean General Circulation Models (AOGCM). We considered the variability in ENMs performance to estimate a weighted mean difference between potential geographic distributions for baseline and future climates. On average, potential future ranges were projected to be from 25.7% to 44.5% smaller than current potential ranges across species. However, we found that 0.2% to 18.3% of the total variance in range shifts occurred “within species” (i.e., owing to the use of different modeling techniques and climate models) and 81.7% to 99.8% remained between species (i.e., it could be explained by ecological correlates). Using meta‐analytical techniques akin to regression, we also showed that potential range shifts are barely predicted by bird biological traits. We demonstrated that one can combine and reduce species‐specific effects with high uncertainty in ENMs and also explore potential causes of climate change effect on species using meta‐analytical tools. We also highlight that the search for powerful correlates of climate change‐induced range shifts can be a promising line of investigation.     

Tertiary relict trees in a Mediterranean climate: abiotic constraints on the persistence of Prunus lusitanica at the eroding edge of its range

Aim To investigate the ecophysiological traits allowing persistence of a subtropical relict tree (Prunus lusitanica L.) under a dry Mediterranean climate at the eroding edge of its range. Location A glasshouse for the study under controlled conditions and two marginal populations located in riparian forests of central Spain and exposed to summer drought, in contrast to subtropical populations that grow in mountain cloud forests. Methods Two experiments were conducted to assess tolerance to the abiotic conditions found in riparian habitats. In a glasshouse experiment, gas‐exchange and light‐use parameters were measured to evaluate seedling responses to a factorial combination of irradiance (60%, 10%, 2% and 0.5% full sun) and moisture (well watered control and drought stress). In a parallel field experiment, irradiance and soil moisture were measured as predictors of seedling survival at two sites in which half the seedlings were subjected to a threefold increase in summer precipitation by adding water every 10 days. Results  Soil moisture strongly determined survival both in the glasshouse and in the two field sites. In the field, periodic addition of water failed to increase survival. Water‐use efficiency (WUE) increased with drought. Seedlings did not tolerate deep shade (2–0.5%) and their performance and survival were better when exposed to moderate (10%) or high (60%) irradiance. The effect of water stress on seedling performance was stronger at moderate irradiance, moderate at high irradiance and negligible at very low light. Seedling size (height and number of leaves) attained after 1 month of experimental treatments had a positive effect on survival at the end of the summer, hence greater survival was not achieved at the expense of vertical growth. Main conclusions While studies in Macaronesia have shown that P. lusitanica occupies a wide range of moisture conditions along mountain slopes, it behaves as an obligate riparian species in dry peripheral populations. Intolerance to deep shade and tolerance to moderate and high irradiance allow the species to grow in small and moist gaps, or in treeless river sections. Despite the species’ low range filling in marginal, drought‐prone regions, long‐term persistence might have been achieved through shifts to riparian habitats serving as local refuges.     

Predicting the effects of climate change on population connectivity and genetic diversity of an imperiled freshwater mussel,Cumberlandia monodonta (Bivalvia: Margaritiferidae), in riverine systems

In the face of global climate change, organisms may respond to temperature increases by shifting their ranges poleward or to higher altitudes. However, the direction of range shifts in riverine systems is less clear. Because rivers are dendritic networks, there is only one dispersal route from any given location to another. Thus, range shifts are only possible if branches are connected by suitable habitat, and stream‐dwelling organisms can disperse through these branches. We used Cumberlandia monodonta (Bivalvia: Unionoida: Margaritiferidae) as a model species to investigate the effects of climate change on population connectivity because a majority of contemporary populations are panmictic. We combined ecological niche models (ENMs) with population genetic simulations to investigate the effects of climate change on population connectivity and genetic diversity of C. monodonta. The ENMs were constructed using bioclimatic and landscape data to project shifts in suitable habitat under future climate scenarios. We then used forward‐time simulations to project potential changes in genetic diversity and population connectivity based on these range shifts. ENM results under current conditions indicated long stretches of highly suitable habitat in rivers where C. monodonta persists; populations in the upper Mississippi River remain connected by suitable habitat that does not impede gene flow. Future climate scenarios projected northward and headwater‐ward range contraction and drastic declines in habitat suitability for most extant populations throughout the Mississippi River Basin. Simulations indicated that climate change would greatly reduce genetic diversity and connectivity across populations. Results suggest that a single, large population of C. monodonta will become further fragmented into smaller populations, each of which will be isolated and begin to differentiate genetically. Because C. monodonta is a widely distributed species and purely aquatic, our results suggest that persistence and connectivity of stream‐dwelling organisms will be significantly altered in response to future climate change.     

Niche shifting in response to warming climate after the last glacial maximum: inference from genetic data and niche assessments in the chisel‐toothed kangaroo rat (Dipodomys microps)

During Pleistocene glacial‐interglacial cycles, the geographic range is often assumed to have shifted as a species tracks its climatic niche. Alternatively, the geographic range would not necessarily shift if a species can adapt in situ to a changing environment. The potential for a species to persist in place might increase with the diversity of habitat types that a species exploits. We evaluate evidence for either range shift or range stability between the last glacial maximum (LGM) and present time in the chisel‐toothed kangaroo rat (Dipodomys microps), an endemic of the Great Basin and Mojave deserts. We modeled how the species’ range would have changed if the climatic niche of the species remained conserved between the LGM and present time. The climatic models imply that if D. microps inhabited the same climatic niche during the LGM as it does today, the species would have persisted primarily within the warm Mojave Desert and expanded northwards into the cold Great Basin only after the LGM. Contrary to the climatic models, the mitochondrial DNA assessment revealed signals of population persistence within the current distribution of the species throughout at least the latest glacial‐interglacial cycle. We concluded that D. microps did not track its climatic niche during late Pleistocene oscillations, but rather met the challenge of a changing environment by shifting its niche and retaining large portions of its distribution. We speculate that this kind of response to fluctuating climate was possible because of ‘niche drifting’, an alteration of the species’ realized niche due to plasticity in various biological characters. Our study provides an example of an approach to reconstruct species’ responses to past climatic changes that can be used to evaluate whether and to what extent taxa have capacity to shift their niches in response to the changing environment – information becoming increasingly important to predicting biotic responses to future environmental changes.     

Biogeography of the Amazon molly: ecological niche and range limits of an asexual hybrid species

Aim To understand the relative contributions of environmental factors, dispersal limitations and the presence of sperm donors in determining the distribution of the Amazon molly (Poecilia formosa), a sperm‐dependent unisexual fish species of hybrid origin. To explore niche similarities and/or differences between the hybrid and parental species. To evaluate whether large‐scale abiotic factors can explain a successful introduction of both P. formosa and Poecilia latipinna. Location South‐east United States, Mexico and Central America. Methods We used abiotic variables in ecological niche modelling (ENM) to identify regions with suitable conditions for the presence of the Amazon molly and its two parental species (P. latipinna and Poecilia mexicana). We also used a recently developed metric to calculate the degree of niche overlap between the hybrid and its parental species. Results ENM produced highly significant models [all area under the curve (AUC) > 0.99 for the three species]. Annual mean temperature and minimum temperature of the coldest month were the variables that best explained the distribution of the Amazon molly. With the exception of south Florida, few areas beyond the known distribution of the species were predicted to have suitable environmental conditions. The hybrid species niche overlaps partially with the parental species. However, given the available data, it is neither more similar nor more different than expected by chance. Main conclusions Two different processes are acting to limit the distribution of P. formosa. At the northern limit, although a sperm donor species is present further north, suitable environmental conditions are absent from nearby locations. At the southern limit, a sperm donor species is present and areas with good environmental conditions are present at nearby locations, suggesting that dispersal ability is the limiting factor. We found that the hybrid species overlaps in a similar way with both parental species while still having its own niche identity. This result may be explained by the fact that hybrid species inherit characteristics of two ecologically divergent species, which can result in intermediate or even transgressive phenotypes. These results support recent work on the role of hybridization in diversification.     

Geographic variation in genetic and demographic performance: new insights from an old biogeographical paradigm

The ‘centre–periphery hypothesis’ (CPH) is a long‐standing postulate in ecology that states that genetic variation and demographic performance of a species decrease from the centre to the edge of its geographic range. This hypothesis is based on an assumed concordance between geographical peripherality and ecological marginality such that environmental conditions become harsher towards the limits of a species range. In this way, the CPH sets the stage for understanding the causes of distribution limits. To date, no study has examined conjointly the consistency of these postulates. In an extensive literature review we discuss the birth and development of the CPH and provide an assessment of the CPH by reviewing 248 empirical studies in the context of three main themes. First, a decrease in species occurrence towards their range limits was observed in 81% of studies, while only 51% demonstrated reduced abundance of individuals. A decline in genetic variation, increased differentiation among populations and higher rates of inbreeding were demonstrated by roughly one in two studies (47, 45 and 48%, respectively). However, demographic rates, size and population performance less often followed CPH expectations (20–30% of studies). We highlight the impact of important methodological, taxonomic, and biogeographical biases on such validation rates. Second, we found that geographic and ecological marginality gradients are not systematically concordant, which casts doubt on the reliability of a main assumption of the CPH. Finally, we attempt to disentangle the relative contribution of geographical, ecological and historical processes on the spatial distribution of genetic and demographic parameters. While ecological marginality gradients explain variation in species' demographic performance better than geographic gradients, contemporary and historical factors may contribute interactively to spatial patterns of genetic variation. We thereby propose a framework that integrates species' ecological niche characteristics together with current and past range structure to investigate spatial patterns of genetic and demographic variation across species ranges.     

Birds on the move in the face of climate change: High species turnover in northern Europe

Species richness is predicted to increase in the northern latitudes in the warming climate due to ranges of many southern species expanding northwards. We studied changes in the composition of the whole avifauna and in bird species richness in a period of already warming climate in Finland (in northern Europe) covering 1,100 km in south–north gradient across the boreal zone (over 300,000 km²). We compared bird species richness and species‐specific changes (for all 235 bird species that occur in Finland) in range size (number of squares occupied) and range shifts (measured as median of area of occupancy) based on bird atlas studies between 1974–1989 and 2006–2010. In addition, we tested how the habitat preference and migration strategy of species explain species‐specific variation in the change of the range size. The study was carried out in 10 km squares with similar research intensity in both time periods. The species richness did not change significantly between the two time periods. The composition of the bird fauna, however, changed considerably with 37.0% of species showing an increase and 34.9% a decrease in the numbers of occupied squares, that is, about equal number of species gained and lost their range. Altogether 95.7% of all species (225/235) showed changes either in the numbers of occupied squares or they experienced a range shift (or both). The range size of archipelago birds increased and long‐distance migrants declined significantly. Range loss observed in long‐distance migrants is in line with the observed population declines of long‐distance migrants in the whole Europe. The results show that there is an ongoing considerable species turnover due to climate change and due to land use and other direct human influence. High bird species turnover observed in northern Europe may also affect the functional diversity of species communities.     

Exploring tree species colonization potentials using a spatially explicit simulation model: implications for four oaks under climate change

Climate change impacts tree species differentially by exerting unique pressures and altering their suitable habitats. We previously predicted these changes in suitable habitat for current and future climates using a species habitat model (DISTRIB) in the eastern United States. Based on the accuracy of the model, the species assemblages should eventually reflect the new quasi‐equilibrium suitable habitats (~2100) after accounting for the lag in colonization. However, it is an open question if and when these newly suitable habitats will be colonized under current fragmented landscapes and realistic migration rates. To evaluate this, we used a spatially explicit cell‐based model (SHIFT) that estimates colonization potentials under current fragmented habitats and several estimates of historical migration rates at a 1 km resolution. Computation time, which was previously the biggest constraint, was overcome by a novel application of convolution and Fast Fourier Transforms. SHIFT outputs, when intersected with future suitable habitats predicted by DISTRIB, allow assessment of colonization potential under future climates. In this article, we show how our approach can be used to screen multiple tree species for their colonization potentials under climate change. In particular, we use the DISTRIB and SHIFT models in combination to assess if the future dominant forest types in the north will really be dominated by oaks, as modelled via DISTRIB. Even under optimistic scenarios, we conclude that only a small fraction of the suitable habitats of oaks predicted by DISTRIB is likely to be occupied within 100 years, and this will be concentrated in the first 10–20 km from the current boundary. We also show how DISTRIB and SHIFT can be used to evaluate the potential for assisted migration of vulnerable tree species, and discuss the dynamics of colonization at range limits.     

No‐analog climates and shifting realized niches during the late quaternary: implications for 21st‐century predictions by species distribution models

Empirically derived species distributions models (SDMs) are increasingly relied upon to forecast species vulnerabilities to future climate change. However, many of the assumptions of SDMs may be violated when they are used to project species distributions across significant climate change events. In particular, SDM's in theory assume stable fundamental niches, but in practice, they assume stable realized niches. The assumption of a fixed realized niche relative to climate variables remains unlikely for various reasons, particularly if novel future climates open up currently unavailable portions of species’ fundamental niches. To demonstrate this effect, we compare the climate distributions for fossil‐pollen data from 21 to 15 ka bp (relying on paleoclimate simulations) when communities and climates with no modern analog were common across North America to observed modern pollen assemblages. We test how well SDMs are able to project 20th century pollen‐based taxon distributions with models calibrated using data from 21 to 15 ka. We find that taxa which were abundant in areas with no‐analog late glacial climates, such as Fraxinus, Ostrya/Carpinus and Ulmus, substantially shifted their realized niches from the late glacial period to present. SDMs for these taxa had low predictive accuracy when projected to modern climates despite demonstrating high predictive accuracy for late glacial pollen distributions. For other taxa, e.g. Quercus, Picea, Pinus strobus, had relatively stable realized niches and models for these taxa tended to have higher predictive accuracy when projected to present. Our findings reinforce the point that a realized niche at any one time often represents only a subset of the climate conditions in which a taxon can persist. Projections from SDMs into future climate conditions that are based solely on contemporary realized distributions are potentially misleading for assessing the vulnerability of species to future climate change.