Relics of the Europe's warm past: phylogeography of the Aesculapian snake

Understanding how species responded to past climate change can provide information about how they may respond to the current global warming. Here we show how a European reptile species responded to the last natural global warming event at the Pleistocene-Holocene transition that led to the Holocene climatic optimum approximately 5000-8000 years ago. The Aesculapian snake, Zamenis longissimus, is a thermophilous species whose present-day distribution in the southern half of Europe is a remnant of much wider range during the Holocene climatic optimum when populations occurred as far north as Denmark. These northern populations went extinct as the climate cooled, and presently the species is extinct from all central Europe, except few relic populations in locally suitable microhabitats in Germany and the Czech Republic. Our phylogenetic and demographic analyses identified two major clades that expanded from their respective western and eastern refugia after the last glacial maximum (18,000-23,000 years ago) and contributed approximately equally to the present range. Snakes from the relic northern populations carried the Eastern clade, showing that it was primarily the snakes from the eastern, probably Balkan, refugium that occupied the central and northern Europe during the Holocene climatic optimum. Two small, deep-branching clades were identified in near the Black Sea and in Greece. These clades provide evidence for two additional refugia, which did not successfully contribute to the colonization of Europe. If, as our results suggest, some populations responded to the mid-Holocene global warming by shifting their ranges further north than other populations of the same species, knowing what populations were able to expand in different species may provide information about what populations will be important for the species' ability to cope with the current global warming.     

Ecological indicators of Tuber aestivum habitats in temperate European beech forests

Although Tuber aestivum is widely distributed across Europe, little is known about its biology and ecology. We assess the vegetation composition of wild T. aestivum sites and use this information to characterise suitable habitats in Fagus sylvatica, Carpinus betulus and Ostrya carpinifolia (FCO) forests. Plant species composition at 16 naturally colonised T. aestivum sites in southwestern Germany and Switzerland was compared with that of 232 reference sites in Swiss FCO forest communities. The vegetation composition of truffle sites exhibits a high proportion of herb and shrub species that are indicators of a warm and dry climate, alkaline soil, and open tree canopy and thus typically occur in thermophilous FCO communities on calcareous bedrock. We conclude that T. aestivum is associated with a set of key species in addition to known host species. Based on their floristic composition, we identified FCO forest communities with high and low suitability for T. aestivum.     

Growth-climate responses indicate shifts in the competitive ability of European beech and Norway spruce under recent climate warming in East-Central Europe

Long-term changes in climate substantially affect the tree growth and species distribution in Europe. In the presented study, the radial growth of Fagus sylvatica (L.) and Picea abies ((L.) Karst.) has been studied along an altitudinal gradient covering six vegetation formations characteristic for sub-montane, montane and high-montane conditions of the western Carpathians. Tree growth responses to temperature and precipitation changes have been analysed based on the sample of increment cores and standard dendroclimatic methods in two time periods, the reference period 1961–1990 and the recent period 1991–2012. The growth responses of spruce and beech to recent changes in climate were similar up to high-montane zones, where the beech shows significantly larger improvements of radial increments in comparison to spruce. The growth responses were mainly temperature driven. In the sub-montane area, the increased effect of precipitation in the recent period was overridden by the negative effects of warming, and the alleviated temperature limitation had an evidently supportive effect on tree growth in montane and high-montane areas. In the near future, the warming will likely cause decline in radial increments of beech and spruce in sub-montane areas due to expected landscape drying. At the same time, the improved competitive ability of beech in the high-montane zones suggests a shift in the leading edge of beech distribution into higher altitudes in East-Central Europe.     

Phylogeography using mitogenomes: A rare Dipodidae,Sicista betulina,in North‐western Europe

Repeated climatic and vegetation changes during the Pleistocene have shaped biodiversity in Northern Europe including Denmark. The Northern Birch Mouse (Sicista betulina) was one of the first small rodent species to colonize Denmark after the Late Glacial Maximum. This study analyses complete mitochondrial genomes and two nuclear genes of the Northern Birch Mouse to investigate the phylogeographical pattern in North‐western Europe and test whether the species colonized Denmark through several colonization events. The latter was prompt by (i) the present‐day distinct northern and southern Danish distribution and (ii) the subfossil record of Northern Birch Mouse, supporting early Weichselian colonization. Samples from Denmark, Norway, Sweden, Russia, Latvia, Estonia, and Slovakia were included. Mitogenomes were obtained from 54 individuals, all representing unique mitogenomes supporting high genetic variation. Bayesian phylogenetic analysis identified two distinct evolutionary linages in Northern Europe diverging within the Elster glaciation period. The results of the two nuclear genomes showed lower genetic differentiation but supported the same evolutionary history. This suggests an allopatric origin of the clades followed by secondary contact. Individuals from southern Denmark were only found in one clade, while individuals from other areas, including northern Denmark, were represented in both clades. Nevertheless, we found no evidence for repeated colonization''s explaining the observed fragmented distribution of the species today. The results indicated that the mitogenome pattern of the Northern Birch Mouse population in southern Denmark was either (i) due to the population being founded from northern Denmark, (ii) a result of climatic and anthropogenic effects reducing population size increasing genetic drift or (iii) caused by sampling bias.     

Potential source and sink locations for climate‐driven species range shifts in Europe since the Last Glacial Maximum

Aim To identify potential source and sink locations for climate‐driven species range shifts in Europe since the Last Glacial Maximum (LGM). Location Europe. Methods We developed a new approach combining past‐climate simulations with the concept of analogous climate space. Our index gives a continuous measure of the potential of a location to have acted as a source or a sink for species that have shifted their ranges since the LGM. High glacial source potential is indicated by LGM climatic conditions that are widespread now; high post‐glacial sink potential is indicated by current climatic conditions that were widespread at the LGM. The degree of isolation of source and sink areas was calculated as the median distance to areas with analogous climate conditions. Results We identified areas of high glacial source potential in the previously recognized refugial areas in the southern European peninsulas, but also in large areas in central‐western Europe. The most climatically isolated source areas were located in northern Spain, in north‐western Europe and in eastern Turkey. From here species would have had to cover substantial distances to find current climate conditions analogous to LGM conditions of these areas. Areas with high post‐glacial sink potential were mainly located in Fennoscandia and in central and south‐eastern Europe. Some of the most isolated sink areas were located in the Spanish highlands and around the Baltic Sea. Main conclusions Our species‐independent approach successfully identified previously recognized glacial refugial areas with high source potential for species range shifts in southern Europe and in addition highlighted other potential source areas in central Europe. This study offers new insights into how the distribution of past and current climatic conditions may have influenced past species range shifts and current large‐scale biodiversity patterns.     

National and European perspectives on climate change sensitivity of the habitats directive characteristic plant species

The main goal of the Habitats Directive, a key document for European conservation, is to maintain a ‘favourable’ conservation status of selected species and habitats. In the face of near-future climatic change this goal may become difficult to achieve. Here, we evaluate the sensitivity to climate change of 84 plant species that characterise the Danish habitat types included in the Habitats Directive. A fuzzy bioclimatic envelope model, linking European and Northwest African species’ distribution data with climate, was used to predict climatically suitable areas for these species in year 2100 under two-climate change scenarios. Climate sensitivity was evaluated at both Danish and European scales to provide an explicit European perspective on the impacts predicted for Denmark. In all 69–99% of the species were predicted to become negatively affected by climate change at either scale. Application of international Red List criteria showed that 43–55% and 17–69% would become vulnerable in Denmark and Europe, respectively. Northwest African atlas data were used to improve the ecological accuracy of the future predictions. For comparison, using only European data added 0–7% to these numbers. No species were predicted to become extinct in Europe, but 4–7% could be lost from Denmark. Some species were predicted to become positively affected in Denmark, but negatively affected in Europe. In addition to nationally endangered species, this group would be an important focus for a Danish conservation strategy. A geographically differentiated Danish conservation strategy is suggested as the eastern part of Denmark was predicted to be more negatively affected than the western part. No differences in the sensitivity of the Habitats Directive habitats were found. We conclude that the conservation strategy of the Habitats Directive needs to integrate the expected shifts in species’ distributions due to climate change.     

欧洲典型树种展叶始期的时空变化及其对气候变化的响应

近年来,全球变暖对植物春季物候期产生了显著影响.很多研究报道了中国地区木本植物春季物候期变化的时空格局,但在同处于北半球温带地区的欧洲则相关研究较少.为了增进物候变化及其对气候变化响应规律的区域对比,本研究利用欧洲地区展叶始期(1980-2014年)数据和相应的气象数据,研究欧洲七叶树、垂枝桦、欧洲山毛榉和夏栎4种典型...     

Modelling the distribution of Ilex aquifolium at the north-eastem edge of its geographical range

Plant distribution borders are key features to characterise the ecological niche of a species and to monitor effects of climate change. Here we focus on an evergreen small tree, Ilex aquifolium, which reaches its north-eastern range edge in Denmark. Our main objectives are to describe and to model the current distribution of the species, to identify the most important climatic and land use factors which shape this distribution pattern, and to analyse the species' habitat requirements. For this purpose we used data from a national mapping project, complemented by information from forest owners. The distribution and abundance of I. aquifolium in Denmark have markedly changed during the past 40 years. It is now found in almost all districts, although the centres of abundance still coincide with the historical records. Our model shows lower habitat suitability for the species in northern and eastern districts, where winters are more severe and spring precipitation is lower. To a lesser extent, land use influences I. aquifolium occurrence, but it is more common in areas with a high proportion of forests and/or urban sites. The analysis of habitat requirements supports these results, since I. aquifolium occurs mainly as a forest species in deciduous stands, on relatively nutrient-rich moist soils, and under moderately high light conditions. However, some records may be the product of seed dispersal from planted individuals nearby. The results suggest that the range edge of the species has moved at least 100 km east within half a century. Since I.aquifolium is sensitive to winter frost, this change in distribution may be due to increasingly mild winter temperatures.     

Mountain landscapes offer few opportunities for high‐elevation tree species migration

Climate change is anticipated to alter plant species distributions. Regional context, notably the spatial complexity of climatic gradients, may influence species migration potential. While high‐elevation species may benefit from steep climate gradients in mountain regions, their persistence may be threatened by limited suitable habitat as land area decreases with elevation. To untangle these apparently contradictory predictions for mountainous regions, we evaluated the climatic suitability of four coniferous forest tree species of the western United States based on species distribution modeling (SDM) and examined changes in climatically suitable areas under predicted climate change. We used forest structural information relating to tree species dominance, productivity, and demography from an extensive forest inventory system to assess the strength of inferences made with a SDM approach. We found that tree species dominance, productivity, and recruitment were highest where climatic suitability (i.e., probability of species occurrence under certain climate conditions) was high, supporting the use of predicted climatic suitability in examining species risk to climate change. By predicting changes in climatic suitability over the next century, we found that climatic suitability will likely decline, both in areas currently occupied by each tree species and in nearby unoccupied areas to which species might migrate in the future. These trends were most dramatic for high elevation species. Climatic changes predicted over the next century will dramatically reduce climatically suitable areas for high‐elevation tree species while a lower elevation species, Pinus ponderosa, will be well positioned to shift upslope across the region. Reductions in suitable area for high‐elevation species imply that even unlimited migration would be insufficient to offset predicted habitat loss, underscoring the vulnerability of these high‐elevation species to climatic changes.     

Specialists in ancient trees are more affected by climate than generalists

Ancient trees are considered one of the most important habitats for biodiversity in Europe and North America. They support exceptional numbers of specialized species, including a range of rare and endangered wood‐living insects. In this study, we use a dataset of 105 sites spanning a climatic gradient along the oak range of Norway and Sweden to investigate the importance of temperature and precipitation on beetle species richness in ancient, hollow oak trees. We expected that increased summer temperature would positively influence all wood‐living beetle species whereas precipitation would be less important with a negligible or negative impact. Surprisingly, only oak‐specialist beetles with a northern distribution increased in species richness with temperature. Few specialist beetles and no generalist beetles responded to the rise of 4°C in summer as covered by our climatic gradient. The negative effect of precipitation affected more specialist species than did temperature, whereas the generalists remained unaffected. In summary, we suggest that increased summer temperature is likely to benefit a few specialist beetles within this dead wood community, but a larger number of specialists are likely to decline due to increased precipitation. In addition, generalist species will remain unaffected. To minimize adverse impacts of climate change on this important community, long‐term management plans for ancient trees are important.     

The effect of climate on the phenology, acorn crop and radial increment of pedunculate oak (Quercus robur) in the middle Volga region, Tatarstan, Russia

Our data, collected in the extreme east of Europe, show that a significant biological effect of climate change has been experienced even in territories where temperature increase has been the lowest. This study documents the climatic response of pedunculate oak (Quercus robur) growing near its north-eastern limits in Europe. It demonstrates the potential of oak trees in old-growth forest to act as proxy climate indicators. Many factors may influence the temporal stability of the growth-climate, acorn crop-climate and first leafing-climate relationships. Climate data, climatic fluctuations, reproduction, genetics and tree-age may relate to this instability. Our results stress that an increase in climate variability or climatic warming resulting from warmer winters or summers could affect the oak population in eastern Europe in a similar way to that in western Europe. These findings, from remnants of oak forest in the middle Volga region of Russia, allow a further understanding of how species could be affected by future climates.     

Models of climate associations and distributions of amphibians in Italy

Analyzing the relationships between the distribution of animal species and climatic variables is not only important for understanding which factors govern species distribution but also for improving our ability to predict future ecological responses to climate change. In the context of global climate change, amphibians are of particular interest because of their extreme sensitivity to the variation of temperature and precipitation regimes. We analyzed species–climate relationships for 17 amphibian species occurring in Italy using species distribution data at the 10 × 10 km resolution. A machine learning method, Random Forests, was used to model the distribution of amphibians in relation to a set of 18 climatic variables. The results showed that the variables which had the highest importance were those related to precipitation, indicating that precipitation is an important factor in determining amphibian distribution. Future projections showed a complex response of species distributions, emphasizing the potential severity of climate change on the distributions of amphibians in Italy. The species that will decrease the most are those occurring in mountainous and Mediterranean areas. Our results provide some preliminary information that could be useful for amphibian conservation, indicating if future conservation priorities for some species should be enhanced.     

Climatic niche of the Saker Falcon Falco cherrug: predicted new areas to direct population surveys in Central Asia

Accurate species distribution data across remote and extensive geographical areas are difficult to obtain. Here, we use bioclimatic envelope models to determine climatic constraints on the distribution of the migratory Saker Falcon Falco cherrug to identify areas in data-deficient regions that may contain unidentified populations. Sakers live at low densities across large ranges in remote regions, making distribution status difficult to assess. Using presence-background data and eight bioclimatic variables within a species distribution modelling framework, we applied MaxEnt to construct models for both breeding and wintering ranges. Occurrence data were spatially filtered and climatic variables tested for multicollinearity before selecting best fit models using the Akaike information criterion by tuning MaxEnt parameters. Model predictive performance tested using the continuous Boyce index (B) was high for both breeding (B_TEST = 0.921) and wintering models (B_TEST = 0.735), with low omission rates and minimal overfitting. The Saker climatic niche was defined by precipitation in the warmest quarter in the breeding range model, and mean temperature in the wettest quarter in the wintering range model. Our models accurately predicted areas of highest climate suitability and defined the climatic constraints on a wide-ranging rare species, suggesting that climate is a key determinant of Saker distribution across macro-scales. We recommend targeted population surveys for the Saker based on model predictions to areas of highest climatic suitability in key regions with distribution knowledge gaps, in particular the Qinghai-Tibet plateau in western China. Further applications of our models could identify protected areas and reintroduction sites, inform development conflicts, and assess the impact of climate change on distributions.     

中国亚热带地区4种极危冷杉属植物的地理分布特征及其与气候的关系

通过野外实地调查和文献调研, 分析4种极危冷杉属(Abies)植物的地理分布现状和特征, 构建其分布区的气候数据库, 采用国际上常用的研究植被与气候关系的指标和方法, 定量分析冷杉属地理分布与水热条件的关系, 并通过因子分析方法确定影响冷杉属地理分布的限制性气候因子。结果表明: 1)百山祖冷杉(Abies beshanzuensis)、梵净山冷杉(A. fanjingshanensis)、元宝山冷杉(A. yuanbaoshanensis)和资源冷杉(A. beshanzuensis var. ziyuanensis)零星分布于我国亚热带地区, 水平分布范围为25°25′-27°55′ N, 108°42′-119°12′ E , 分布范围狭窄; 垂直分布范围为1363-2390 m, 均分布于山体上部。海拔上限高度随着经度的减小而增加。2)整个分布区的气温较低, 年平均气温为7.4-11.2 ℃; 降水较为充沛, 年降水量为1265.4-1945.7 mm, 其中生长季降水量占74.78%; 湿度较大, 年相对湿度为77%-86%。气候指标范围狭窄, 表明对水热条件要求很高。分布区寒冷指数较低, 限制其垂直向下扩散, 呈现局限分布于山体上部的现状。3) 4种冷杉各分布区的气候条件基本相近, 梵净山冷杉分布区气温相对偏低, 元宝山冷杉分布区湿度相对偏大。4) 3个气候因子对这4种冷杉的地理分布影响较大, 其作用排序为: 低温因子>极端低温因子>湿度因子。在全球气候变暖的大背景下, 这4种冷杉自然分布区的适宜生境将进一步缩小, 直至物种灭绝, 急需开展迁地保护工作。     

Climatic Control on Forests and Tree Species Distribution in the Forest Region of Northeast China

North-east （NE） China covers considerable climatic gradients and all major forests types of NE Asia. in the present study, 10 major forest types across the forest region of NE China were sampled to Investigate forest distribution in relation to climate. Canonical correspondence analysis （CCA） revealed that growing season precipitation and energy availability were primary climatic factors for the overall forest pattern of NE China, accounting for 66% of the explanatory power of CCA. Conversely, annual precipitation and winter coldness had minor effects. Generalized additive models revealed that tree species responded to climatic gradients differently and showed three types of response curve： （i） monotonous decline; （ii） monotonous Increase; and （iii） a unimodai pattern. Furthermore, tree species showed remarkable differences in limiting climatic factors for their distribution. The power of climate in explaining species distribution declined significantly with decreasing species dominance, suggesting that the distribution of dominant species was primarily controlled by climate, whereas that of subordinate species was more affected by competition from other species.     

Ice age legacies in the geographical distribution of tree species richness in Europe

Aim This study uses a high‐resolution simulation of the Last Glacial Maximum (LGM) climate to assess: (1) whether LGM climate still affects the geographical species richness patterns in the European tree flora and (2) the relative importance of modern and LGM climate as controls of tree species richness in Europe. Location The parts of Europe that were unglaciated during the LGM. Methods Atlas data on the distributions of 55 tree species were linked with data on modern and LGM climate and climatic heterogeneity in a geographical information system with a 60‐km grid. Four measures of species richness were computed: total richness, and richness of the 18 most restricted species, 19 species of medium incidence (intermediate species) and 18 most widespread species. We used ordinary least‐squares regression and spatial autoregressive modelling to test and estimate the richness–climate relationships. Results LGM climate constituted the best single set of explanatory variables for richness of restricted species, while modern climate and climatic heterogeneity was best for total and widespread species richness and richness of intermediate species, respectively. The autoregressive model with all climatic predictors was supported for all richness measures using an information‐theoretic approach, albeit only weakly so for total species richness. Among the strongest relationships were increases in total and intermediate richness with climatic heterogeneity and in restricted richness with LGM growing‐degree‐days. Partial regression showed that climatic heterogeneity accounted for the largest unique variation fraction for intermediate richness, while LGM climate was particularly important for restricted richness. Main conclusions LGM climate appears to still affect geographical patterns of tree species richness in Europe, albeit the relative importance of modern and LGM climate depends on range size. Notably, LGM climate is a strong richness control for species with a restricted range, which appear to still be associated with their glacial refugia.     

Towards European climate risk surfaces: the extent and distribution of analogous and non-analogous climates 1931–2100

Aim Climate is an important determinant of species distributions. We assess different aspects of risk arising from future climate change by quantifying changes in the spatial distribution of future climatic conditions compared with the recent past. Location Europe. Methods A 10′ × 10′ resolution gridded data set of five climate variables was used to calculate expected changes to the area, distance and direction of 1931–60 climatic conditions under the HadCM3 climate model for four future climate scenarios based on different rates of greenhouse gas emissions (SRES scenarios). Three levels of tolerance ranges determined the thresholds for which future conditions are considered analogous to 1931–60 (pre‐warming) conditions. Results For many parts of Europe, areas with pre‐warming analogous climate conditions will be smaller and further away in the future than they are now. For any location in Europe, areas with pre‐warming analogous mean annual temperature conditions will, on average, be reduced between 23.7% (B1 scenario) and 49.7% (A1FI scenario) by 2100 when assuming a medium tolerance range. The mean distance to these areas will, on average, increase between 272 km (B1) and 645 km (A1FI). These changes are more pronounced for temperature than for water availability variables and also for narrow tolerance ranges compared to wide tolerance ranges. Using a combined measure of both temperature and precipitation variables, areas with prewarming analogous conditions are predicted to be in a more northeasterly direction in the future, but there are considerable regional differences within Europe. Main conclusions The results suggest that, for some parts of Europe, the loss of area with any suitable climatic conditions represents the greatest risk to biodiversity, but in other regions the distances that species may have to move to reach suitable climatic conditions may be a greater problem. Quantifying the distance and direction in analyses of change of climatically suitable areas can add additional information for climate change risk assessments.     

Integrating climate and landscape models to prioritize areas and conservation strategies for an endangered arboreal primate

Species distributions are influenced by both climate conditions and landscape structure. Here we propose an integrated analysis of climatic and landscape niche-based models for a forest-dependent primate, the endangered black lion tamarin (Leontopithecus chrysopygus). We applied both climate and landscape variables to predict the distribution of this tamarin and used this information to prioritize strategic areas more accurately. We anticipated that this approach would be beneficial for the selection of pertinent conservation strategies for this flagship species. First, we built climate and landscape niche-based models separately, combining seven algorithms, to infer processes acting on the species distribution at different scales. Subsequently, we combined climate and landscape models using the EcoLand Analysis. Our results suggest that historic and current landscape fragmentation and modification had profoundly adverse effects on the distribution of the black lion tamarins. The models indicated just 2096 km² (out of an original distribution of 92,239 km²) of suitable areas for both climate and landscape. Of this suitable area, the species is currently present in less than 40%, which represents less than 1% of its original distribution. Based on the combined map, we determined the western and southeast regions of the species range to be priority areas for its conservation. We identified areas with high climatic and high landscape suitability, which overlap with the remaining forest fragments in both regions, for habitat conservation and population management. We suggest that areas with high climatic but low landscape suitability should be prioritized for habitat management and restoration. Areas with high landscape suitability and low climatic suitability, such as the Paranapiacaba mountain range should be considered in light of projected climate change scenarios. Our case study illustrates that a combined approach of climatic and landscape niche-based modeling can be useful for establishing focused conservation measures that may increase the likelihood of success.     

Standing genomic variation within coding and regulatory regions contributes to the adaptive capacity to climate in a foundation tree species

Global climate is rapidly changing, and the ability for tree species to adapt is dependent on standing genomic variation; however, the distribution and abundance of functional and adaptive variants are poorly understood in natural systems. We test key hypotheses regarding the genetics of adaptive variation in a foundation tree: genomic variation is associated with climate, and genomic variation is more likely to be associated with temperature than precipitation or aridity. To test these hypotheses, we used 9,593 independent, genomic single‐nucleotide polymorphisms (SNPs) from 270 individuals sampled from Corymbia calophylla's entire distribution in south‐western Western Australia, spanning orthogonal temperature and precipitation gradients. Environmental association analyses returned 537 unique SNPs putatively adaptive to climate. We identified SNPs associated with climatic variation (i.e., temperature [458], precipitation [75] and aridity [78]) across the landscape. Of these, 78 SNPs were nonsynonymous (NS), while 26 SNPs were found within gene regulatory regions. The NS and regulatory candidate SNPs associated with temperature explained more deviance (27.35%) than precipitation (5.93%) and aridity (4.77%), suggesting that temperature provides stronger adaptive signals than precipitation. Genes associated with adaptive variants include functions important in stress responses to temperature and precipitation. Patterns of allelic turnover of NS and regulatory SNPs show small patterns of change through climate space with the exception of an aldehyde dehydrogenase gene variant with 80% allelic turnover with temperature. Together, these findings provide evidence for the presence of adaptive variation to climate in a foundation species and provide critical information to guide adaptive management practices.