New approaches to direct gradient analysis using environmental scalars and statistical curve-fitting procedures

The conceptual framework of direct gradient analysis (DGA) is discussed in relation to the functional, factorial approach to vegetation. Both approaches use abstract simplified environment gradients with which to correlate vegetation response. Environmental scalars based on physical process models of environment and/or known biological growth processes can be incorporated to make analyses less location specific. An example of an environmental scalar (radiation index) for converting aspect and slope measurements to the more biologically relevant radiation input at a site is given. The problem of the shape of species response curves to environmental gradients is examined using a sample of 1 286 plots from eucalypt forest in southern New South Wales. An important conclusion is that skewed or bimodal response curves may be due to unsatisfactory distribution of observations and/or unrecognized environmental factors. The use of Generalized Linear Modelling (GLM) as a method for providing a statistical basis for DGA is presented. Analyses using GLM, and presence/absence data are presented for a range of eucalypt species (Eucalyptus rossii, E. dalrympleana, E. fastigata etc.). Successful prediction of species distributions (realized niches) can be achieved with mean annual temperature, mean annual rainfall, radiation index and geology. Quadratic terms are required in many cases, indicating bell-shaped response curves. The major variability associated with species niches is shown to be related to a limited number (4) of environmental factors. DGA with biologically relevant scalars and appropriate statistical methods is suitable for studying many problems of species' realized niches and plant community composition.     

No adaptation to altitude in the invasive plant Erigeron annuus in the Swiss Alps

The altitudinal distribution of plants is restricted by various environmental factors, with climatic conditions being one of the primary constraints. Here, we investigate what limits the altitudinal range of the introduced species Erigeron annuus in the Swiss Alps. We planted offspring of E. annuus plants originating from different altitudes into two common gardens, one located at an altitude representing the main area of distribution (400 m) and the other close to the current altitudinal limit of E. annuus in Switzerland (1000 m). In both common gardens all established plants survived and grew vigorously during the growing season. However, there was high winter mortality of seedlings at 1000 m. Furthermore, plant phenology was delayed and reproductive output was reduced at 1000 m, although the seeds produced were larger. The general lack of adaptation to altitude and only moderate levels of plasticity suggest that there is little potential for E. annuus to persist beyond its current altitudinal limit in the Swiss Alps. However, climate warming might promote the upward range expansion of E. annuus by reducing winter mortality and by increasing the chance of producing seeds within the growing season.     

Relationships between environmental variables and vegetation across mountain wetland sites,N. Iran

The mountain wetlands studied represent a unique habitat on the southern slopes of the Alborz mountain range, the second largest range in Iran. In comparison with other parts of this range the western section is ecologically and botanically unknown. Floristic and vegetation variation were assessed using diverse environmental variables along a broad altitudinal span (350 m to 3200 m a.s.l.). Using both statistical and ordination analyses floristic variation was assessed on three defined altitudinal belts which were delimited based on Alborz macro-climatic boundaries. The distribution of individual wetland plant species, of phytogeographic elements and of life-forms all differ among altitudinal belts. This result is also shown in both direct and indirect analyses of ordinations. The proportion of geophytes significantly increases with altitude and geophytes are very well represented in the upper altitudinal belt. The number of species of a narrow phytogeographical distribution (e.g. endemics) increases with altitude, soil pH and EC declined with altitude. The first axis of DCA ordination with passively projected environmental variables indicates that, organic matter and concentration of Fe²⁺ are increased toward higher altitude. The second axis of ordination is related to both soil texture and slope inclination. The distribution of species in the CCA species plot is also close to the distribution of those in the DCA ordination. This study indicates that altitude and slope together with other dependent environmental variables (pH, EC, Ca²⁺ and soil texture) are the main ecological factors controlling species distribution across the Western Alborz wetland sites.     

Nematode distributions as spatial null models for macroinvertebrate species richness across environmental gradients: A case from mountain lakes

Nematode species are widely tolerant of environmental conditions and disperse passively. Therefore, the species richness distribution in this group might largely depend on the topological distribution of the habitats and main aerial and aquatic dispersal pathways connecting them. If so, the nematode species richness distributions may serve as null models for evaluating that of other groups more affected by environmental gradients. We investigated this hypothesis in lakes across an altitudinal gradient in the Pyrenees. We compared the altitudinal distribution, environmental tolerance, and species richness, of nematodes with that of three other invertebrate groups collected during the same sampling: oligochaetes, chironomids, and nonchironomid insects. We tested the altitudinal bias in distributions with t‐tests and the significance of narrow‐ranging altitudinal distributions with randomizations. We compared results between groups with Fisher's exact tests. We then explored the influence of environmental factors on species assemblages in all groups with redundancy analysis (RDA), using 28 environmental variables. And, finally, we analyzed species richness patterns across altitude with simple linear and quadratic regressions. Nematode species were rarely biased from random distributions (5% of species) in contrast with other groups (35%, 47%, and 50%, respectively). The altitudinal bias most often shifted toward low altitudes (85% of biased species). Nematodes showed a lower portion of narrow‐ranging species than any other group, and differed significantly from nonchironomid insects (10% and 43%, respectively). Environmental variables barely explained nematode assemblages (RDA adjusted R² = 0.02), in contrast with other groups (0.13, 0.19 and 0.24). Despite these substantial differences in the response to environmental factors, species richness across altitude was unimodal, peaking at mid elevations, in all groups. This similarity indicates that the spatial distribution of lakes across altitude is a primary driver of invertebrate richness. Provided that nematodes are ubiquitous, their distribution offers potential null models to investigate species richness across environmental gradients in other ecosystem types and biogeographic regions.     

辽东栎林不同层植物沿海拔梯度分布的DCCA分析

 为研究北京东灵山辽东栎（Quercus liaotungensis）不同层植物的分布与环境因子之间的关系,对东灵山辽东栎林进行了植物群落学调查。沿 海拔梯度在乔木层、灌木层和草本层各调查119个样方,同时调查了每个样方的环境因子如海拔、坡度和坡位。物种数据采用二元属性数据,对 非数值环境因子进行赋值。对东灵山辽东栎林和各环境因子进行去趋势典范对应分析（Detrended canonical correspondence analysis, DCCA）,结果表明：DCCA 第一轴基本上反映了植物群落所处环境的海拔的梯度变化,第二轴基本上反映了坡位的梯度变化。同时发现辽东栎林 大多数植物分布在低海拔区域。乔木层的物种分布比较均匀。草本层的物种分布重叠率是最大的,说明草本植物的生境相似性很高。低海拔区 域的生物多样性要高于高海拔区域的生物多样性。结合环境因子间的相关关系和DCCA排序结果,发现海拔是影响东灵山辽东栎林物种分布的主 要环境因子。     

The distribution of ground spiders (Araneae, Gnaphosidae) along the altitudinal gradient of Crete, Greece: species richness, activity and altitudinal range

Aim To study the altitudinal variation of ground spiders (Araneae, Gnaphosidae) of Crete, Greece, as far as species composition, species richness, activity and range of distribution are concerned. Location Altitudinal zones (0–2400 m) along the three main mountain massifs of the island of Crete. Methods Thirty‐three sampling sites were located from 0 to 2400 m a.s.l. on Crete, and sampled using pitfall traps. Material from the high‐activity period of Gnaphosidae (mid‐spring to mid‐autumn) was analysed. Sampling sites were divided into five altitudinal zones of 500 m each. Statistical analysis involved univariate statistics (anova ) and multivariate statistics, such as multidimensional scaling (MDS) and cluster analysis (UPGMA) using binomial data of species presence or absence. Results Species richness declines with altitude and follows a hump‐shaped pattern. The activity pattern of the family, as a whole, is not correlated with altitude and is highly species‐specific. In the highest zone, both species richness and activity decline dramatically. The altitudinal range of species distribution increases with altitude. On the Cretan summits live highly tolerant lowland species and isolated residents of the high mountains of Crete. Two different patterns of community structure are recorded. Main conclusions Communities of Gnaphosidae on Crete present two distinct structures following the altitudinal gradient, these being separated by a transitional zone between 1600 and 2000 m. This study supports previous results which show a hump‐shaped decline in species richness of Gnaphosidae along altitudinal gradients, leading to a peak at 400–700 m, where an optimum of environmental factors exists. This makes this zone the meeting point of the often opportunistic lowland species with the older and most permanent residents of the island. Rapoport's rule on the positive correlation of the altitudinal range of species distributions with altitude is also supported. The high activity recorded for the species that persist on the high mountains of Crete is indicative of a tolerant arachnofauna, and is considered to result from relaxation of competitive interactions with other species. This is related to a reduction in species numbers, shortening of the activity period on high mountains and the unique presence of high mountain species that thrive only there. As shown in our study, strategies to cope with altitude are species‐specific. Therefore, there cannot exist one single model to describe how animals react to the change in altitude, even under the same environmental conditions.     

Plant species and growth form richness along altitudinal gradients in the southwest Ethiopian highlands

Questions: Do growth forms and vascular plant richness follow similar patterns along an altitudinal gradient? What are the driving mechanisms that structure richness patterns at the landscape scale? Location: Southwest Ethiopian highlands. Methods: Floristic and environmental data were collected from 74 plots, each covering 400 m². The plots were distributed along altitudinal gradients. Boosted regression trees were used to derive the patterns of richness distribution along altitudinal gradients. Results: Total vascular plant richness did not show any strong response to altitude. Contrasting patterns of richness were observed for several growth forms. Woody, graminoid and climber species richness showed a unimodal structure. However, each of these morphological groups had a peak of richness at different altitudes: graminoid species attained maximum importance at a lower elevations, followed by climbers and finally woody species at higher elevations. Fern species richness increased monotonically towards higher altitudes, but herbaceous richness had a dented structure at mid‐altitudes. Soil sand fraction, silt, slope and organic matter were found to contribute a considerable amount of the predicted variance of richness for total vascular plants and growth forms. Main Conclusions: Hump‐shaped species richness patterns were observed for several growth forms. A mid‐altitudinal richness peak was the result of a combination of climate‐related water–energy dynamics, species–area relationships and local environmental factors, which have direct effects on plant physiological performance. However, altitude represents the composite gradient of several environmental variables that were interrelated. Thus, considering multiple gradients would provide a better picture of richness and the potential mechanisms responsible for the distribution of biodiversity in high‐mountain regions of the tropics.     

The role of functional traits in species distributions revealed through a hierarchical model

Quantifying how functional traits relate to environmental gradients provides insight into mechanisms governing species distributions. Here, we bring together the fields of species distribution modelling and functional trait ecology with hierarchical modelling by explicitly incorporating traits into a multi‐species distribution model. We combined traits from the leaf‐height‐seed strategy scheme (specific leaf area (SLA), plant height and seed mass) with a distribution model for 20 eucalypt taxa in Victoria, Australia. The key insight of this approach is how traits modulate species responses to environmental gradients. The strongest link was between SLA and percent rock cover (species with low SLA had positive responses to rockiness, whereas high SLA species responded negatively to rockiness). We found evidence for complex yet potentially important interactions. For instance, the probability of species occurrence increased with rainfall and solar radiation on average yet the response varied depending on species height and SLA. Tall species were predicted to increase with rainfall and solar radiation across the range of SLA values (tall species with low SLA were especially sensitive to rainfall). Short species responded positively to rainfall and solar radiation only if they had low SLA. This framework readily accounts for interactions between combinations of traits and environmental variables unlike multi‐step approaches. Further application of this concept will contribute to a generalized mechanistic understanding of how traits influence species distributions along environmental gradients, with implications for understanding the response of species to global change.     

中国翼手目地理分布的环境因子影响分析

利用地理信息系统提取了中国翼手目90个物种已知地理分布点的环境因子数据,并利用主成分分析和逐步回归分析研究了这些环境因子对中国翼手目动物地理分布和种数密度分布的影响程度。结果表明,按其影响程度大小,影响中国翼手目动物地理分布的连续型环境因子依次为年平均气温、大于零度日数、一月份平均气温、无霜期日数、年降雨量、年平均相对湿度、七月份平均气温、海拔高度和年日照时数,其中年日照时数和海拔高度为负影响;影响翼手目种数密度分布的决定环境因子是年降雨量,但年平均气温也有明显作用。作者认为,气候因子影响翼手目动物地理分布的机制是限制性因子的作用,主要受冬眠和活动及摄食时间长短的限制,而种数密度分布则决定于环境资源的丰富度,即主要受到由降雨量决定的食物种类多少的影响。另外,对类别环境因子的研究表明：植被类型对翼手目的种数分布没有明显影响;具有多洞穴特点的喀斯特和火山熔岩地貌具有较多的翼手目种类,但红层地貌由于分布于南方和分散分布的特点而具有最高的翼手目种数密度[动物学报51(3)：413—422,2005]。     

Rotifer species richness along an altitudinal gradient in the Alps

Aim Biodiversity patterns along altitudinal gradients are less studied in aquatic than terrestrial systems, even though aquatic sites provide a more homogeneous environment independent of moisture constraints. We studied the altitudinal species richness pattern for planktonic rotifers in freshwater lakes and identified the environmental predictors for which altitude is a proxy. Location Two hundred and eighteen lakes of Trentino–South Tyrol (Italy) in the eastern Alps; lakes covered 98% (range 65–2960 m above sea level) of the altitudinal gradient in the Alps. Methods We performed: (1) linear regression between species richness and altitude to evaluate the general pattern, (2) multiple linear regression between species richness and environmental predictors excluding altitude to identify the most important predictors, and (3) linear regression between the residuals of the best model of step (2) and altitude to investigate any additional explanatory power of altitude. Selection of environmental predictors was based on limnological importance and non‐parametric Spearman correlations. We applied ordinary least squares regression, generalized linear, and generalized least squares modelling to select the most statistically appropriate model. Results Rotifer species richness showed a monotonic decrease with altitude independent of scale effects. Species richness could be explained (R²= 51%) by lake area as a proxy for habitat diversity, reactive silica and total phosphorus as proxies for productivity, water temperature as a proxy for energy, nitrate as a proxy for human influence and north–south and east–west directions as covariates. These predictors completely accounted for the species richness–altitude pattern, and altitude had no additional effect on species richness. Main conclusions The linear decrease of species richness along the altitudinal gradient was related to the interplay of habitat diversity, productivity, heat content and human influence. These factors are the same in terrestrial and aquatic habitats, but the greater environmental stability of aquatic systems seems to favour a linear pattern.     

The behaviour of scribbly gum moth larvae Ogmograptis sp. Meyrick (Lepidoptera: Bucculatricidae) in the Australian Capital Territory

Abstract  Scribbles, the damage caused by mining larvae of the moth Ogmograptis sp. (Bucculatricidae), are a common sight on the trunks of many Eucalyptus species in eastern Australia. They provide a record of larval behaviour that can be easily measured when the trees shed outer bark layers. Measurements were made of scribbles on three species of eucalypt ( Eucalyptus racemosa ssp. rossii , E. pauciflora and E. delegatensis ) in the Australian Capital Territory. Scribble density varied between tree species and study sites but was consistently higher on the southern and eastern aspects of tree trunks. Some characteristics of mining behaviour, such as initial mining direction, were found to be random, but others including mine length and number of direction changes follow distinct patterns but differ between eucalypt species. It is likely that there is more than one species of Ogmograptis Meyrick as currently described ( O. scribula Meyrick, from E. pauciflora ) and each eucalypt species may be host to a different species of Ogmograptis .     

Relationships between vegetation zonation and altitude in a salt-marsh system in northwest Spain

Abstract. Vegetation zonation in salt marshes has traditionally been attributed largely to altitudinal differences, since altitude determines the temporal pattern of tidal flooding and is thus closely related to proximate determinants of the distribution of species and plant communities. We investigated the distribution of vascular plants and plant communities along a series of altitudinal transects in two salt marshes in the northwestern Iberian Peninsula. Our results indicate that altitudinal range varies significantly both among species and among communities, and confirm that salt-marsh vegetation characteristics (species cover and composition) can be predicted on the basis of altitude, particularly at the lower levels of the profile.     

中国地卷属地衣海拔分布分析

海拔是地衣多样性的重要影响因素。了解地衣的海拔分布格局是地衣多样性保护的基础。研究表明中国地卷属地衣的物种丰富度和个体丰富度沿海拔梯度呈单峰曲线,它们倾向于分布在受全球变暖影响较高的高海拔地区[（2,022±995）m],且具有较窄的海拔分布幅（约68%的物种分布于海拔为1,694－2,954m的地带）,因而对其进行保护生物学的研究已十分迫切。地卷属11种地衣具有不同的海拔分布规律,这些分布规律与各自分布的海拔区间有关。基于11物种的海拔分析支持Rapoport法则。全球暖化对4种地衣的潜在威胁尤其强烈,即：大陆地卷Peltigera continentalis、长孢地卷P. dolichospora、克氏地卷P. kristinssonii和小地卷P. venosa。     

Species richness and trait composition of butterfly assemblages change along an altitudinal gradient

Species richness patterns along altitudinal gradients are well-documented ecological phenomena, yet very little data are available on how environmental filtering processes influence the composition and traits of butterfly assemblages at high altitudes. We have studied the diversity patterns of butterfly species at 34 sites along an altitudinal gradient ranging from 600 to 2,000 m a.s.l. in the National Park Berchtesgaden (Germany) and analysed traits of butterfly assemblages associated with dispersal capacity, reproductive strategies and developmental time from lowlands to highlands, including phylogenetic analyses. We found a linear decline in butterfly species richness along the altitudinal gradient, but the phylogenetic relatedness of the butterfly assemblages did not increase with altitude. Compared to butterfly assemblages at lower altitudes, those at higher altitudes were composed of species with larger wings (on average 9 %) which laid an average of 68 % more eggs. In contrast, egg maturation time in butterfly assemblages decreased by about 22 % along the altitudinal gradient. Further, butterfly assemblages at higher altitudes were increasingly dominated by less widespread species. Based on our abundance data, but not on data in the literature, population density increased with altitude, suggesting a reversed density–distribution relationship, with higher population densities of habitat specialists in harsh environments. In conclusion, our data provide evidence for significant shifts in the composition of butterfly assemblages and for the dominance of different traits along the altitudinal gradient. In our study, these changes were mainly driven by environmental factors, whereas phylogenetic filtering played a minor role along the studied altitudinal range.     

Continuum or zonation? Altitudinal gradients in the forest vegetation of Mt. Kilimanjaro

Based on the analysis of 600 vegetation plots using the method of Braun-Blanquet (1964) the composition of the whole vascular forest plant flora with about 1220 species was studied in the forests of Mt. Kilimanjaro. The altitudinal distribution of all strata (trees, shrubs, epiphytes, lianas and herbs) along a transect of 2400 m is discussed with respect to altitudinal zonation and ecological factors. With uni-dimensionally constraint clustering significant discontinuities were revealed that occurred simultaneously in the different strata. Thus even in structurally highly complex, multilayered tropical montane forests distinct community units exist that can be surveyed and classified by the Braun-Blanquet approach. This observed zonation was significantly correlated with altitude, temperature and soil acidity (pH); rainfall was of importance in particular for the zonation of epiphytes. Other key factors were humidity (influenced by stable cloud condensation belts) and minimum temperature (in particular the occurrence of frost at 2700 m altitude upslope). The contrary results of other transect studies in East Africa in respect to continuity of change in floristic composition appear to be caused by different sampling methods and intensities or mixing of data from areas with different climate conditions, whereas species richness did not influence the clarity of floristic discontinuities on Kilimanjaro and other parts of East Africa.     

Explaining grass-nutrient patterns in a savanna rangeland of southern Africa

Aim The search for possible factors influencing the spatial variation of grass quality is an important step towards understanding the distribution of herbivores, as well as a step towards identifying crucial areas for conservation and restoration. A number of studies have shown that grass quality at a regional scale is influenced by climatic variables. At a local scale, site factors and their interaction are considered important. In this study, we aimed at examining environmental correlates of grass quality at a local scale. The study also sought to establish if biotic factors interact significantly with abiotic factors in influencing a variation in grass quality. Location The study area is located in the Kruger National Park of South Africa. The study area stretches from west (22°49′ S and 31°01′ E) to east, (22°44′ S and 31°22′ E) covering an area of about 25 × 6 km in the far northern region of the Kruger National Park. Methods We collected environmental data such as soil texture, percentage grass cover and biomass as well as grass samples for chemical analysis from specific locations in the study area. In addition, a digital elevation model (DEM) with a resolution of 5 m was used to derive elevation, slope and aspect using a geographic information system (GIS), which were related to grass quality. We used correlation analysis and anova to relate environmental variables to grass quality. Multivariate analysis techniques were used to simultaneously analyse and explore the complex interactions between variables. Results and conclusions Our results indicate that there is a significant relationship between grass quality parameters and site‐specific factors such as slope, altitude, percentage grass cover, aspect and soil texture. Relatively, percentage grass cover and soil texture were more critical in explaining a variation in grass quality. Plant characteristics such as species type interact significantly with slope, altitude and geology in influencing nutrient distribution. The results of this study may give a better insight on foliar nutrient distribution patterns at a landscape scale in savanna rangelands. Furthermore, the results of this study may help in the selection of ancillary information, which could be used in conjunction with other data such as remotely sensed data to map grass quality – an important step towards understanding the distribution and feeding patterns of wildlife. However, we acknowledge that this study is based on one seasonal snapshot, therefore some slightly different findings may be obtained during other times of the year. Nevertheless, the study has revealed that under the conditions experienced during the study period, nutrient distribution varies with varying biotic and abiotic factors.     

澜沧县两栖动物多样性及海拔分布格局

海拔梯度综合了温度、降水等气候因子,是影响物种多样性分布格局的重要环境因子,两栖动物多样性及其海拔分布格局一直是生物地理学和生态学研究的重要内容。为了全面掌握云南澜沧县两栖动物多样性现状及海拔分布格局,在调查并掌握澜沧县两栖动物多样性本底数据的基础上,根据地形地貌和生境的不同,在澜沧县99个10km×10km有效网格中按照分层抽样的方法选取45个网格,每个网格设置3-5条样线,于2016-2017年开展野外调查。调查共发现两栖动物1842只,隶属于2目9科30属60种,以树蛙科为主,占两栖动物物种总数的28.3%,其中云南纤树蛙（Gracixalus yunnanensis）为新种,清迈泽陆蛙（Fejervarya qingmaiensis）为中国新纪录种;调查到中国特有种有15种,三有保护动物有25种,受威胁物种有19种,其中极危物种1种,濒危物种4种,易危物种14种;澜沧县两栖动物在海拔600-2400m范围内均有分布,其中,在海拔1800-2000m之间的范围内调查到的两栖动物物种数目和多度均较高,其次是在海拔1600-1800m和1200-1400m范围内;本次调查记录到的两栖动物物种数是历史记录物种数的3倍,属数比历史记录多了1倍,铃蟾科为新记录科;丰富度较高的网格尤其是丰富度在20以上的网格均处于中高海拔,海拔相对较低和较高的网格丰富度相对较低;两栖动物多样性科、属、种的丰富度在海拔梯度上分布格局不太一致,种丰富度的海拔分布格局大体呈双峰型,是由包括平均降水量、平均温度和植被覆盖指数等环境因子的综合作用,各因子与物种丰富度相关性从大到小依次为平均降水量 > 平均海拔=平均温度 > 植被覆盖指数 > 水网密度指数 > 样线距离水域的平均距离 > 植被类型数量。研究摸清了澜沧县两栖动物的多样性状况,分析了两栖动物多样性海拔分布格局,为澜沧县生物多样性保护提供基础数据。     

Endemic flora biodiversity in the south of the Iberian Peninsula: altitudinal distribution,life forms and dispersal modes

The south of the Iberian Peninsula, with an altitudinal range varying from sea level to 3482m and annual average rainfall ranging from 206 to 2223mm, has 516 vascular endemic species or subspecies, that is, an endemicity rate of 13%. This survey deals with parameters such as species richness, originality, life forms and dispersal modes of these plants, in relation to altitudinal and rainfall gradients. Although most of the endemic plants occur between 600 and 1400m a.s.l. and in the range of 600–1000mm annual average rainfall, floristic originality (rate of endemic taxa per area unit) increases exponentially with altitude. The biological spectrum of this endemic flora does not follow the usual patterns observed either in local floras in the south of the Iberian Peninsula or in other regions of the Mediterranean Basin. Chamaephytes (46.08%) and hemicryptophytes (31.37%) are very abundant, whereas therophytes (11.96%) and phanerophytes (0.98%) are comparatively rare. There is a statistically significant correlation between life form and both altitudinal and rainfall gradients. Chamaephytes reach their highest density rates within 1400–2000m a.s.l., but these records tend to decrease as rainfall rates increase. Abundance of hemicryptophytes is directly dependent on rainfall rates and inversely dependent on temperature. The altitudinal distribution pattern of therophytes is opposite to that of hemicryptophytes, but there is no clear correlation as far as rainfall gradient is concerned. Considering both the endemic plants as a whole and each of the life form groups, the relationships between the dispersal modes used and the altitudinal and rainfall gradients are analysed. Up to 44.51% of the endemic plants do not present evident adaptations to promote the dispersion of their diaspores. However, only in the group of therophytes, whose occurrence is positively related to areas of minor altitude, does this difficulty of dispersion play a significant role in the maintenance of stenochory.     

Predicting potential responses to future climate in an alpine ungulate: interspecific interactions exceed climate effects

The altitudinal shifts of many montane populations are lagging behind climate change. Understanding habitual, daily behavioural rhythms, and their climatic and environmental influences, could shed light on the constraints on long‐term upslope range‐shifts. In addition, behavioural rhythms can be affected by interspecific interactions, which can ameliorate or exacerbate climate‐driven effects on ecology. Here, we investigate the relative influences of ambient temperature and an interaction with domestic sheep (Ovis aries) on the altitude use and activity budgets of a mountain ungulate, the Alpine chamois (Rupicapra rupicapra). Chamois moved upslope when it was hotter but this effect was modest compared to that of the presence of sheep, to which they reacted by moving 89–103 m upslope, into an entirely novel altitudinal range. Across the European Alps, a range‐shift of this magnitude corresponds to a 46% decrease in the availability of suitable foraging habitat. This highlights the importance of understanding how factors such as competition and disturbance shape a given species’ realised niche when predicting potential future responses to change. Furthermore, it exposes the potential for manipulations of species interactions to ameliorate the impacts of climate change, in this case by the careful management of livestock. Such manipulations could be particularly appropriate for species where competition or disturbance already strongly restricts their available niche. Our results also reveal the potential role of behavioural flexibility in responses to climate change. Chamois reduced their activity when it was warmer, which could explain their modest altitudinal migrations. Considering this behavioural flexibility, our model predicts a small 15–30 m upslope shift by 2100 in response to climate change, less than 4% of the altitudinal shift that would be predicted using a traditional species distribution model‐type approach (SDM), which assumes that species’ behaviour remains unchanged as climate changes. Behavioural modifications could strongly affect how species respond to a changing climate.