Predicting microscale shifts in the distribution of the butterfly Plebejus argus at the northern edge of its range

Species are often observed to occur in restricted patches of particularly warm microclimate at their high latitude/altitude geographic range margin. In these areas, global warming is expected to cause small‐scale expansion of the occupied area, but most previous studies of range expansion have used very coarse scale data. Using high resolution microclimate models together with detailed field surveys, we tested whether the butterfly Plebejus argus, occurring on limestone grassland in north Wales, was responding as might be expected due to climate change in the last 30–40 yr. The abundance of adult Plebejus argus at 100 m resolution in 2011 was strongly affected by elevation and near‐ground temperatures in May. A statistical model including microclimate, fitted to 2011 data, was successful (67% correct) at hindcasting the occurrence of Plebejus argus in 1983 when the average May air temperature was 1.4°C cooler. However, the model was less accurate at hindcasting occurrences in 1972 (50% correct). Given the distribution of micro‐sites in this landscape, we predict that further warming of approximately 1°C would make the majority of sites highly microclimatically suitable for this species. There are a growing number of long‐term studies of range change, and investigations into the mechanisms driving them, but still surprisingly few that explicitly make and test predictions with independent data. Our tests are a valuable example of how accurate predictions of distribution change can be, but also of the inevitable uncertainties. Improved understanding of how well models predict will be very important to plan robust climate change adaptation measures.     

The effect of habitat on the range expansion of a native and an introduced bird species

Aim Range expansion across a heterogeneous landscape may depend on the habitat selected and used by the expanding species. If habitat selection influences range expansion then localities colonized by a species should contain a greater proportion of favoured habitat (and less non‐habitat) than other nearby localities not colonized. White‐winged doves (Zenaida asiatica) and Eurasian collared doves (Streptopelia decaocto) are two bird species that provide an excellent opportunity to test this hypothesis, because the geographic ranges of both species have been expanding in North America for more than two decades. Location Continental USA. Methods We used distribution data from the North American Breeding Bird Survey to test whether the landscapes occupied by each species contained a greater proportion of favoured habitat (urban land, grassland/pasture, shrub land and cropland) and a lower proportion of non‐habitat (forest land) than landscapes where doves were not found. We tested each species separately in each of three broad expansion areas, namely East, Central and West. We also compared rates of spatial spread between expansion areas and between the two species. Results As predicted, both species tended to occupy landscapes with greater proportions of urban land, shrub land and cropland but with less forest land compared with landscapes without doves, in all three expansion areas. Contrary to prediction, occupied landscapes tended to have slightly less grassland/pasture than unoccupied landscapes. Rates of spread differed between the two species and among expansion areas. Main conclusions Range expansion and the extent to which a species fills or saturates its range are influenced by the habitat ecology of the expanding species. Species colonize localities based on the availability of suitable habitat. However, the role of habitat in a species’ range expansion does depend somewhat on the greater geographical setting. Over large regional and geographical scales, range expansion (rate of spread and saturation) may proceed unevenly, suggesting that range expansion is a very dynamic and context‐specific process.     

Breeding success at the range margin of a desert species: implications for a climate‐induced elevational shift

If the breeding range of a species is limited by biotic or abiotic environmental factors that depress breeding success at the range margin, then range expansion is expected when those limiting factors are alleviated. Over a three‐year period, we measured breeding success of a desert species, black‐throated sparrow Amphispiza bilineata, along a steep elevation gradient between the Peninsular Mountains and Colorado Desert (San Diego County, California) that is undergoing a warming trend. We compared breeding success at geographically marginal locations (higher‐elevation chaparral sites) to more central locations (lower‐elevation desert scrub sites) only a short distance apart. Breeding success was measured at the nest level, territory level, and population level. At each level measured, breeding success tended to be greater at higher‐elevation chaparral sites at the distribution margin compared to lower‐elevation sites where the bird was more common. Black‐throated sparrows had 100% reproductive failure at lower‐elevation sites during the two driest years of our study (2006–2007), but did relatively well at higher‐elevation sites. Only in a wetter year (2008) was breeding success improved at lower‐elevation sites. Surprisingly, there was no evidence of an upward elevational shift in distribution over a 26‐year period despite a clear warming trend and drier conditions. Greater territory density at lower‐elevation sites with reproductive failure during dry years suggests the possibility of an ecological trap in this system, which could prevent or delay climate‐induced range shifts. A common presumption has been that desert species will undergo relatively mild negative impacts due to a warming climate, but it is possible that some desert species are already at or near their temperature and aridity tolerance limits within their current range and shifts may not always be possible.     

Plants,Birds and Butterflies: Short-Term Responses of Species Communities to Climate Warming Vary by Taxon and with Altitude

As a consequence of climate warming, species usually shift their distribution towards higher latitudes or altitudes. Yet, it is unclear how different taxonomic groups may respond to climate warming over larger altitudinal ranges. Here, we used data from the national biodiversity monitoring program of Switzerland, collected over an altitudinal range of 2500 m. Within the short period of eight years (2003–2010), we found significant shifts in communities of vascular plants, butterflies and birds. At low altitudes, communities of all species groups changed towards warm-dwelling species, corresponding to an average uphill shift of 8 m, 38 m and 42 m in plant, butterfly and bird communities, respectively. However, rates of community changes decreased with altitude in plants and butterflies, while bird communities changed towards warm-dwelling species at all altitudes. We found no decrease in community variation with respect to temperature niches of species, suggesting that climate warming has not led to more homogenous communities. The different community changes depending on altitude could not be explained by different changes of air temperatures, since during the 16 years between 1995 and 2010, summer temperatures in Switzerland rose by about 0.07°C per year at all altitudes. We discuss that land-use changes or increased disturbances may have prevented alpine plant and butterfly communities from changing towards warm-dwelling species. However, the findings are also consistent with the hypothesis that unlike birds, many alpine plant species in a warming climate could find suitable habitats within just a few metres, due to the highly varied surface of alpine landscapes. Our results may thus support the idea that for plants and butterflies and on a short temporal scale, alpine landscapes are safer places than lowlands in a warming world.     

Natural,human and spatial constraints to expanding populations of otters in the Iberian Peninsula

Aim To determine the relationships between otter (Lutra lutra) distribution dynamics and environmental and spatial constraints over a 20‐year period. Location Andalusia, southern Iberian Peninsula. Methods We synthesized otter distribution data from three otter surveys (1985, 1995 and 2005) using subcatchment areas defined by hydrological barriers. Subcatchments were characterized by two ‘natural’ (climatic and orographic variables) and two ‘human’ (land use and population density) gradients. In addition, we calculated two contagion variables (the distance to previously occupied subcatchments and the percentage of occupied subcatchments within a 50 km buffer) for consecutively surveyed subcatchments. Results Between 1985 and 2005 the percentage of subcatchments with otters present increased from 42% to 72%. Otters tended to be rare or absent from human‐dominated areas. Anthropogenic gradients were better predictors of otter distribution than natural ones. Human and natural gradients showed strong covariation, but for any value of the natural gradients otters tended to be present in subcatchments with lower human impacts. Colonization of new subcatchments was found to be strongly related to contagion variables and expansion rates were slower than those estimated in other studies. Newly colonized areas tended to be located in areas with intermediate human influence, while repeated absences occurred mainly in areas where human impact was most severe. Main conclusions Our results suggest that recent otter expansion across Andalusia is a reflection of large‐scale improvement in environmental conditions. Otter populations that survived the period of strong and generalized declines appear to be acting as sources from which neighbouring areas are colonized, probably aided by improved water quality and increases in food availability. However, the further expansion of otters into their full original range is likely to be constrained by human‐impacted landscapes.     

Linking habitat use to range expansion rates in fragmented landscapes: a metapopulation approach

Temperature increases because of climate change are expected to cause expansions at the high latitude margins of species distributions, but, in practice, fragmented landscapes act as barriers to colonization for most species. Understanding how species distributions will shift in response to climate change therefore requires techniques that incorporate the combined effects of climate and landscape‐scale habitat availability on colonization rates. We use a metapopulation model (Incidence Function Model, IFM) to test effects of fine‐scale habitat use on patterns and rates of range expansion by the butterfly Hesperia comma. At its northern range margin in Britain, this species has increased its breadth of microhabitat use because of climate warming, leading to increased colonization rates. We validated the IFM by reconstructing expansions in five habitat networks between 1982 and 2000, before using it to predict metapopulation dynamics over 100 yr, for three scenarios based on observed changes to habitat use. We define the scenarios as “cold‐world” (only hot, south‐facing 150–250° hillsides are deemed warm enough), “warm‐world” in which 100–300° hillsides can be populated, and “hot‐world”, where the background climate is warm enough to enable use of all aspects (as increasingly observed). In the simulations, increased habitat availability in the hot‐world scenario led to faster range expansion rates, and to long‐term differences in distribution size and pattern. Thus, fine‐scale changes in the distribution of suitable microclimates led to landscape‐scale changes in population size and colonization rate, resulting in coarse‐scale changes to the species distribution. Despite use of a wider range of habitats associated with climate change, H. comma is still expected to occupy a small fraction of available habitat in 100 yr. The research shows that metapopulation models represent a potential framework to identify barriers to range expansion, and to predict the effects of environmental change or conservation interventions on species distributions and persistence.     

Climate change at the landscape scale: predicting fine-grained spatial heterogeneity in warming and potential refugia for vegetation

Current predictions of how species will respond to climate change are based on coarse‐grained climate surfaces or idealized scenarios of uniform warming. These predictions may erroneously estimate the risk of extinction because they neglect to consider spatially heterogenous warming at the landscape scale or identify refugia where species can persist despite unfavourable regional climate. To address this issue, we investigated the heterogeneity in warming that has occurred in a 10 km × 10 km area from 1972 to 2007. We developed estimates by combining long‐term daily observations from a limited number of weather stations with a more spatially comprehensive dataset (40 sites) obtained during 2005–2006. We found that the spatial distribution of warming was greater inland, at lower elevations, away from streams, and at sites exposed to the northwest (NW). These differences corresponded with changes in weather patterns, such as an increasing frequency of hot, dry NW winds. As plant species were biased in the topographic and geographic locations they occupied, these differences meant that some species experienced more warming than others, and are at greater risk from climate change. This species bias could not be detected at coarser scales. The uneven seasonal nature of warming (e.g. more warming in winter, minimums increased more than maximums) means that climate change predictions will vary according to which predictors are selected in species distribution models. Models based on a limited set of predictors will produce erroneous predictions when the correct limiting factor is not selected, and this is difficult to avoid when temperature predictors are correlated because they are produced using elevation‐sensitive interpolations. The results reinforce the importance of downscaling coarse‐grained (∼50 km) temperature surfaces, and suggest that the accuracy of this process could be improved by considering regional weather patterns (wind speed, direction, humidity) and topographic exposure to key wind directions.     

Dissecting insect responses to climate warming: overwintering and post‐diapause performance in the southern green stink bug,Nezara viridula,under simulated climate‐change conditions

The effect of simulated climate change on overwintering and post‐diapause reproductive performance is studied in Nezara viridula (L.) (Heteroptera: Pentatomidae) close to the species' northern range limit in Japan. Insects are reared from October to June under quasi‐natural (i.e. ambient outdoor) conditions and in a transparent incubator, in which climate warming is simulated by adding 2.5°C to the ambient temperatures. Despite the earlier assumption that females of N. viridula overwinter in diapause, whereas males do so in quiescence, regular dissections show that the two sexes overwinter in a state of true diapause. During winter, both sexes are dark‐coloured and have undeveloped reproductive organs. Resumption of development does not start until late March. During winter, the effect of simulated warming on the dynamics and timing of physiological processes appears to be limited. However, the warming significantly enhances winter survival (from 27–31% to 47–70%), which is a key factor in range expansion of N. viridula. In spring, the effect of simulated warming is complex. It advances the post‐diapause colour change and transition from dormancy to reproduction. The earlier resumption of development is more pronounced in females: in April, significantly more females are already in a reproductive state under the simulated warming than under quasi‐natural conditions. In males, the tendency is similar, although the difference is not significant. Warming significantly enhances spring survival and percentage of copulating adults, although not the percentage of ovipositing females and fecundity. The results suggest that, under the expected climate‐warming conditions, N. viridula will likely benefit mostly as a result of increased winter and spring survival and advanced post‐diapause reproduction. Further warming is likely to allow more adults to survive the critical cold season and contribute (both numerically and by increasing heterogeneity) to the post‐overwintering population growth, thus promoting the establishment of this species in newly‐colonized areas.     

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.     

Effect of geographical range size on plant functional traits and the relationships between plant,soil and climate in Chinese grasslands

Aim Our aim was to address the potential effect of the geographical range size of species on the relationships between plant traits, soil and climate in Chinese grasslands. Previous analyses tended to examine plant–environment relationships across many species while ignoring that species with different range sizes may respond differently to the environment. Here we hypothesized that leaf traits of narrow‐ranging species would be more strongly correlated with soil and climatic variables than those of wide‐ranging species. Location Chinese grasslands. Methods Data on leaf traits, including nitrogen and phosphorus concentrations, carbon/nitrogen ratio, nitrogen/phosphorus ratio and specific leaf area, as well as species range sizes for 208 species distributed across 178 sites in Chinese grasslands were collected. Soil and climate information for each study site was also gathered. The effects of range size on leaf traits were tested using one‐way ANOVA. Correlations between leaf traits, soil and climate were calculated for all species pooled together and for species partitioned into range size quartiles, from the first (narrowest‐ ranging 25%) to the fourth (widest‐ranging 25%). Results Narrow‐ranging species tended to occur at high altitude with lower temperature but higher soil nutrient concentrations compared with wide‐ranging species. No direct link between leaf traits and species range sizes was detected. However, patterns of leaf–soil nutrient relationships changed significantly across levels of range size. Narrow‐ranging species tended to be more sensitive to variation in soil nutrient availability than wide‐ranging species, resulting in a shift from a positive leaf–soil nutrient relationship for narrow‐ranging plants to no relationship for wide‐ranging plants. Species responses to climatic variables were unrelated to their range sizes. Main conclusions The close relationship between leaf and soil nutrients indicates a specialization of narrow‐ranging species to particular habitats whereas wide‐ranging species may be able to better withstand changes in environment such as soil fertility over a large area.     

The role of climate in constraining the elevational range of the Water Pipit Anthus spinoletta in an alpine environment

The study of determinants of species’ ranges along elevational gradients may shed light on the ecological factors that constrain their distribution and fundamental niche. We analysed the influence of the climate, habitat at different spatial scales and topography on Water Pipit Anthus spinoletta density in mountain landscapes across a wide elevational gradient. Variables associated with spring and annual temperature values were the main determinants of Water Pipit density, especially at the lower distributional limit (700–1200 m asl), where the species avoided warmer areas. At high‐elevation sites (1600–2300 m asl), the main constraint to the species’ distribution was habitat structure and composition, with steep rocky areas being avoided. Highest densities were found in open but locally heterogeneous habitat at intermediate to high elevations, and the habitat variables that played a major role at the landscape scale were medium‐tall shrublands and woodlands, but with contrasting effects depending on elevation. These results suggest that different sets of variables may constrain density, and effects may differ at the upper and lower elevational limits, with climate being more important at lower elevations and local habitat more important at higher elevations. Ongoing global warming is likely to cause an upward shift in range boundaries of alpine species, but local habitat features could constrain the upward expansion, resulting in range contractions accompanying range shift.     

What is the effect of climate change on marine fish biodiversity in an area of low connectivity,the Baltic Sea?

Aim Climate change could result in an increase in species richness because large‐scale biogeography suggests that more species could be gained from equatorial regions than may be lost pole‐ward. However, the colonization of newly available habitat may lag behind the rate dictated by climatic warming if there exists of a lack of connectivity between ‘donor’ and receiving areas. The objective of this study was to compare how regional warming affected the biodiversity of marine fish in areas that differed in their connectivity in the Baltic Sea. Location North‐east Atlantic, Kattegat and Baltic Sea. Methods The total species richness and the mean species richness from scientific surveys were related to changes in temperature and salinity. Changes in the extent of the distribution of individual fish species were related to the latitudinal distribution, salinity tolerance, maximum body size and exploitation status to assess to what extent climate change and fishing impacts could explain changes in species richness in the Baltic. Results Rising temperatures in the well‐connected Kattegat correlated to an increase in the species richness of fish, due to an increase in low‐latitude species. Unexpectedly, species richness in the poorly connected Baltic Sea also increased. However, the increase seems to be related to higher salinity rather than temperature and there was no influx of low‐latitude species. Main conclusions These results do not support the hypothesis that low‐connectivity areas are less likely to see increases in species richness in response to warming. This indicates that the effect of climate change on biodiversity may be more difficult to predict in areas of low connectivity than in well‐connected areas.     

Population trends of European common birds are predicted by characteristics of their climatic niche

Temperate species are projected to experience the greatest temperature increases across a range of modelled climate change scenarios, and climate warming has been linked to geographical range and population changes of individual species at such latitudes. However, beyond the multiple modelling approaches, we lack empirical evidence of contemporary climate change impacts on populations in broad taxonomic groups and at continental scales. Identifying reliable predictors of species resilience or susceptibility to climate warming is of critical importance in assessing potential risks to species, ecosystems and ecosystem services. Here we analysed long‐term trends of 110 common breeding birds across Europe (20 countries), to identify climate niche characteristics, adjusted to other environmental and life history traits, that predict large‐scale population changes accounting for phylogenetic relatedness among species. Beyond the now well‐documented decline of farmland specialists, we found that species with the lowest thermal maxima (as the mean spring and summer temperature of the hottest part of the breeding distribution in Europe) showed the sharpest declines between 1980 and 2005. Thermal maximum predicted the recent trends independently of other potential predictors. This study emphasizes the need to account for both land‐use and climate changes to assess the fate of species. Moreover, we highlight that thermal maximum appears as a reliable and simple predictor of the long‐term trends of such endothermic species facing climate change.     

Short-term variation in species richness across an altitudinal gradient of alpine summits

In response to climate warming, high altitude alpine vegetation may be replaced by typically lower altitude species, as species re-assemble and migrate to new areas. However, empirical evidence showing vegetation change in response to climate warming is largely unavailable for Australian alpine areas. Here, we examine changes in species richness with respect to climate and altitude over a 7?year period at a range of spatial scales in a re-survey of five alpine summits that are part of the Global Observation Research Initiative in Alpine Environments monitoring network. Eighty species were recorded in 2011 across all summits, an increase of 6 species since 2004. Mean species richness increased at the whole-of-summit scale from 45 to 50 species (about 12?%). At this scale, the rate of species richness increase was almost one new species per year, with 15 new species recorded at one summit. Here, shrub and graminoid species showed the largest increases. At the smaller spatial scales, changes in species richness were less pronounced. Turnover at the species and community level was typically moderate at all spatial scales and on all summits. The strength and direction of species richness change (the difference in species richness between the two sample periods, +/?) was not related to altitude nor variation in climate. Future re-surveys of the summits will confirm whether these short-term variations in species richness, particularly increases in shrubs, are indeed signals of longer-term trends and interactions with a changing climate.     

Response of plant species richness and primary productivity in shrublands along a north–south gradient in Europe to seven years of experimental warming and drought: reductions in primary productivity in the heat and drought year of 2003

We used a nonintrusive field experiment carried out at six sites – Wales (UK), Denmark (DK), the Netherlands (NL), Hungary (HU), Sardinia (Italy – IT), and Catalonia (Spain – SP) – along a climatic and latitudinal gradient to examine the response of plant species richness and primary productivity to warming and drought in shrubland ecosystems. The warming treatment raised the plot daily temperature by ca. 1 °C, while the drought treatment led to a reduction in soil moisture at the peak of the growing season that ranged from 26% at the SP site to 82% in the NL site. During the 7 years the experiment lasted (1999–2005), we used the pin‐point method to measure the species composition of plant communities and plant biomass, litterfall, and shoot growth of the dominant plant species at each site. A significantly lower increase in the number of species pin‐pointed per transect was found in the drought plots at the SP site, where the plant community was still in a process of recovering from a forest fire in 1994. No changes in species richness were found at the other sites, which were at a more mature and stable state of succession and, thus less liable to recruitment of new species. The relationship between annual biomass accumulation and temperature of the growing season was positive at the coldest site and negative at the warmest site. The warming treatment tended to increase the aboveground net primary productivity (ANPP) at the northern sites. The relationship between annual biomass accumulation and soil moisture during the growing season was not significant at the wettest sites, but was positive at the driest sites. The drought treatment tended to reduce the ANPP in the NL, HU, IT, and SP sites. The responses to warming were very strongly related to the Gaussen aridity index (stronger responses the lower the aridity), whereas the responses to drought were not. Changes in the annual aboveground biomass accumulation, litterfall, and, thus, the ANPP, mirrored the interannual variation in climate conditions: the most outstanding change was a decrease in biomass accumulation and an increase in litterfall at most sites during the abnormally hot year of 2003. Species richness also tended to decrease in 2003 at all sites except the cold and wet UK site. Species‐specific responses to warming were found in shoot growth: at the SP site, Globularia alypum was not affected, while the other dominant species, Erica multiflora, grew 30% more; at the UK site, Calluna vulgaris tended to grow more in the warming plots, while Empetrum nigrum tended to grow less. Drought treatment decreased plant growth in several studied species, although there were some species such as Pinus halepensis at the SP site or C. vulgaris at the UK site that were not affected. The magnitude of responses to warming and drought thus depended greatly on the differences between sites, years, and species and these multiple plant responses may be expected to have consequences at ecosystem and community level. Decreases in biodiversity and the increase in E. multiflora growth at the SP site as a response to warming challenge the assumption that sensitivity to warming may be less well developed at more southerly latitudes; likewise, the fact that one of the studied shrublands presented negative ANPP as a response to the 2003 heat wave also challenges the hypothesis that future climate warming will lead to an enhancement of plant growth and carbon sequestration in temperate ecosystems. Extreme events may thus change the general trend of increased productivity in response to warming in the colder sites.     

Detecting climate change induced range shifts: Where and how should we be looking?

Abstract: Global climate warming is expected to cause systematic shifts in the distribution of species and consequently increase extinction risk. Conservation managers must be able to detect, measure and accurately predict range shifts in order to mitigate impacts on biodiversity. However, important responses to climate change may go unnoticed or be dismissed if we fail to collect sufficient baseline data and apply the most sensitive analytical tests. Here we use randomizations of a contemporary data set on rainforest birds of north‐eastern Australia to quantify the sensitivity of three measures for assessing range shifts along altitudinal gradients. We find that smaller range shifts are detectable by analysing change in the mean altitude of presence records rather than upper or lower range boundaries. For a moderate survey effort of 96 surveys, measurements of change in the mean altitude of 34 species have the capacity to provide strong inference for a mean altitudinal range shift as small as 40 m across the species assemblage. We also show that range shifts measured at range boundaries can be potentially misleading when differences in sampling effort between contemporary and historical data sets are not taken into account.     

Range extension and conservation status of Cymbula nigra (Gastropoda: Patellidae) in the Tunisian shores

Cymbula nigra is the largest limpet in the Mediterranean. It has been listed since 1993, as an ‘endangered species’. Its historical range in the Mediterranean includes the southern Spanish, the Moroccan and the Algerian coasts. Recent observations have confirmed its proliferation in southern Spain and its progression along the European and North African Mediterranean coasts. Field surveys carried out along the Tunisian coasts during winter 2011 revealed the presence of this species in the Gulf of Tunis and the Cap Bon. The finding of C. nigra represents a range extension possibly enhanced by the current global warming, as it is a ‘warm‐water species’. The absence of records in previous surveys to the same sampling sites (2006–2007), together with the small sizes recorded in 2011, confirms the recent installation of the species. The limpet was recorded in areas spared of pollution, characterized by a predominant influence of the Atlantic current, and also showed a preference for steep cliffs exposed to strong waves. This is the first population study of this endangered gastropod outside of its original range of distribution. The determination of the ecological characteristics of newly colonized habitats might serve as a starting point for the establishment of appropriate recommendations for its conservation.     

Early signs of range disjunction of submountainous plant species: an unexplored consequence of future and contemporary climate changes

Poleward and upward species range shifts are the most commonly anticipated and studied consequences of climate warming. However, these global responses to climate change obscure more complex distribution change patterns. We hypothesize that the spatial arrangement of mountain ranges and, consequently, climatic gradients in Europe, will result in range disjunctions. This hypothesis was investigated for submountainous forest plant species at two temporal and spatial scales: (i) under future climate change (between 1950–2000 and 2061–2080 periods) at the European scale and (ii) under contemporary climate change (between 1914–1987 and 1997–2013 periods) at the French scale. We selected 97 submountainous forest plant species occurring in France, among which distribution data across Europe are available for 25 species. By projecting future distribution changes for the 25 submountainous plant species across Europe, we demonstrated that range disjunction is a likely consequence of future climate change. To assess whether it is already taking place, we used a large forest vegetation‐plot database covering the entire French territory over 100 years (1914–2013) and found an average decrease in frequency (?0.01 ± 0.004) in lowland areas for the 97 submountainous species – corresponding to a loss of 6% of their historical frequency – along with southward and upward range shifts, suggesting early signs of range disjunctions. Climate‐induced range disjunctions should be considered more carefully since they could have dramatic consequences on population genetics and the ability of species to face future climate changes.     

Seed dispersers help plants to escape global warming

Plants are shifting their ranges towards higher elevations in response to global warming, yet such shifts are occurring at a rate slower than is needed to keep pace with a rapidly changing climate. There is, however, an almost complete lack of knowledge on seed dispersal across altitude, a key process to understand what constrains climate‐driven range shifts. Here, we report the first direct empirical evidence on altitudinal seed dispersal mediated by two common frugivorous mammals: the red fox Vulpes vulpes and the pine marten Martes martes. We conducted a three‐year (bait‐marking) experiment in a mountainous region of Spain. We offered experimental fruits containing colour‐coded seed mimics at feeding stations that simulated source trees. The colour codes allowed us to identify the exact origin of seed mimics found later in mammal scats. Nearly half (47%) of the dispersal events occurred towards higher elevations, despite only ca 25% of the study area being above the average altitude of the feeding stations (1344 m). Seeds dispersed uphill gained an average of 106 m (median = 111 m) and a maximum of 288 m, greatly exceeding the estimated requirements to escape warming (35.4 m per decade). Yet, foxes mediated much more uphill seed dispersal than martens (57% and 26% of dispersal events, respectively), which can be explained by between‐disperser differences in home range size and habitat specificity. Dispersers with larger home ranges move farther and potentially disperse more seeds to higher altitudes, while habitat generalism is necessary to transport seeds above vegetation belts delimiting contrasting habitat types. We discuss how both traits (home range size and habitat specificity) can be used to infer altitudinal seed dispersal across disperser species and mountainous landscapes.     

Elevational shifts,biotic homogenization and time lags in vegetation change during 40 years of climate warming

Many species show evidence of climate‐driven distribution shifts towards higher elevations, but given the tremendous variation among species and regions, we lack an understanding of the community‐level consequences of such shifts. Here we test for signatures of climate warming impacts using a repeat survey of semi‐permanent vegetation plots in 1970 and 2012 in a montane protected area in southern Québec, Canada, where daily maximum and minimum temperatures have increased by ∼1.6°C and ∼2.5°C over the same time period. As predicted, the abundance‐weighted mean elevations of species distributions increased significantly over time (9 m/decade). A community temperature index (CTI) was calculated as the abundance‐weighted mean of the median temperature across occurrences within each species geographic range in eastern North America. CTI did not vary significantly over time, although the raw magnitude of change (+ 0.2°C) matched the expectation based on the upward shift in distributions of 9 m/decade. Species composition of high elevation sites converged over time toward that observed at low elevation, although compositional changes at low elevation sites were more modest. As a consequence, the results of a multivariate analysis showed a decline in among‐plot compositional variability (i.e. beta diversity) over time, thus providing some of the first empirical evidence linking climate warming with biotic homogenization. Finally, plot‐scale species richness showed a marked increase of ∼25% on average. Overall, elevational distribution shifts, biodiversity change, and biotic homogenization over the past four decades have been consistent with predictions based on climate warming, although the rate of change has been relatively slow, suggesting substantial time lags in biotic responses to climate change.