Does climate determine broad‐scale patterns of species richness? A test of the causal link by natural experiment

Aim Broad‐scale spatial patterns of species richness are very strongly correlated with climatic variables. If there is a causal link, i.e. if climate directly or indirectly determines patterns of richness, then when the climatic variables change, richness should change in the manner that spatial correlations between richness and climate would predict. The present study tests this prediction using seasonal changes in climatic variables and bird richness. Location We used a grid of equal area quadrats (37 000 km²) covering North and Central America as far south as Nicaragua. Methods Summer and winter bird distribution data were drawn from monographs and field guides. Climatic data came from published sources. We also used remotely sensed NDVI (normalized difference vegetation index — a measure of greenness). Results Bird species richness changes temporally (between summer and winter) in a manner that is close to, but statistically distinguishable from, the change one would predict from models relating the spatial variation in richness at a single time to climatic variables. If one further takes into account the seasonal changes in NDVI and within‐season variability of temperature and precipitation, then winter and summer richness follow congruent, statistically indistinguishable patterns. Main conclusions Our results are consistent with the hypothesis that climatic variables (temperature and precipitation) and vegetation cover directly or indirectly influence patterns of bird species richness.     

Plant diversity,biogeography and environment in Iberia: Patterns and possible causal factors

Abstract. We associated patterns of plant diversity with possible causal factors by considering 93 local regions in the Iberian Peninsula and Balearic Islands with respect to biogeography, environmental favourability, and environmental heterogeneity, and their relationship with measured species diversity at four different scales: mean local species richness standardized at a grain of 100 m², total species richness in a community type within a region (regional community richness), mean compositional similarity, and mosaic diversity. Local regions in biogeographic transition zones to the North African and Atlantic floras had higher regional community richness and greater mosaic diversity than did non‐transitional regions, whereas no differences existed in mean local species richness or mean compositional similarity. Mean local species richness was positively related to environmental favourability as measured by actual evapotranspiration, but negatively related to total precipitation and temporal heterogeneity in precipitation. Mean local species richness was greatest in annual grassland and dwarf shrubland communities, and on calcareous bedrock types. Regional community richness was similarly related to actual evapotranspiration and total precipitation, but in addition was positively related to spatial heterogeneity in topography and soil water holding capacity. Mean compositional similarity decreased with increasing spatial heterogeneity and temperature seasonality. Mosaic diversity, a measure of complexity, increased with increasing local and regional richness. We hypothesize that these relationships can be explained by four ecological and evolutionary classes of causal factors: numbers of individuals, intermediate environments, limits to adaptation, and niche variation. These factors operate at various scales and manifest themselves in various ways. For example, at the site level, apparently processes that increase the number of individuals increase mean local species richness, but at the level of the entire region no such effects were found.     

Human impacts on environment–diversity relationships: evidence for biotic homogenization from butterfly species richness patterns

Aim Broad‐scale spatial variation in species richness relates to climate and physical heterogeneity but human activities may be changing these patterns. We test whether climate and heterogeneity predict butterfly species richness regionally and across Canada and whether these relationships change in areas of human activity. Location Canada. Methods We modelled the ranges of 102 butterfly species using genetic algorithms for rule‐set production (GARP). We then measured butterfly species richness and potentially important aspects of human activity and the natural environment. These were included in a series of statistical models to determine which factors are likely to affect butterfly species richness in Canada. We considered patterns across Canada, within predominantly natural areas, human‐dominated areas and particular ecozones. We examined independent observations of butterfly species currently listed under Canada's endangered species legislation to test whether these were consistent with findings from statistical models. Results Growing season temperature is the main determinant of butterfly species richness across Canada, with substantial contributions from habitat heterogeneity (measured using elevation). Only in the driest areas does precipitation emerge as a leading predictor of richness. The slope of relationships between all of these variables and butterfly species richness becomes shallower in human‐dominated areas, but butterfly richness is still highest there. Insecticide applications, habitat loss and road networks reduce butterfly richness in human‐dominated areas, but these effects are relatively small. All of Canada's at‐risk butterfly species are located in these human‐dominated areas. Main conclusions Temperature affects butterfly species richness to a greater extent than habitat heterogeneity at fine spatial scales and is generally far more important than precipitation, supporting both the species richness–energy and habitat heterogeneity hypotheses. Human activities, especially in southern Canada, appear to cause surprisingly consistent trends in biotic homogenization across this region, perhaps through range expansion of common species and loss of range‐restricted species.     

Plant species richness–environment relationships across the Subantarctic–Patagonian transition zone

Aim To evaluate the relative importance of climate, productivity, environmental heterogeneity, biotic associations and habitat use by cattle to account for the species richness of trees, shrubs and herbs across the Subantarctic–Patagonian transition. Location An area of c. 150 × 150 km, within the transition zone between the Subantarctic and Patagonian subregions on the eastern slope of the Andes (c. 39–42° S, 70–72° W). Methods All vascular plants found at each one of 50 (10 × 10 m) sampling plots were counted to estimate the local tree, shrub and herb species richness. Path analysis was used to evaluate the relationship between the richness of the three life‐forms and plant cover, dried litter biomass, mean annual temperature, annual precipitation, daily temperature range, substrate heterogeneity and number of faecal pats. Principal coordinates of neighbour matrices was used to model the spatial autocorrelation of the data. Results Total plant species richness showed a unimodal pattern of spatial variation across the transition. Richness responded positively to indirect effects of precipitation mediated through plant cover, but there was a negative overall effect of precipitation on richness towards the west of the transition, most strongly for trees. An increase in substrate heterogeneity promoted a local increase in herb and shrub richness; the richness of trees increased in sites with steeper slopes. Canopy closure had a direct negative impact on herb richness; it also increased the local accumulation of litter, which negatively affected shrub and herb richness. The impact of habitat use by cattle negatively affected herb richness in areas to the east of the biogeographical transition. Main conclusions We suggest that the importance of indirect climatic effects mediated by vegetation cover can account for species richness patterns across this transition, most strongly for woody species, which supports the productivity hypothesis. The southern temperate forests towards the west may represent a deviation from the predictions of the water–energy dynamics hypothesis. Dissimilar spatial patterns of variation in the richness of woody and herbaceous species, and their different responses to climatic and heterogeneity variables across the transition, suggest that plant life‐form influences the plant species richness–environment relationships.     

Exploring anthropogenic and natural processes shaping fern species richness along elevational gradients

Aim (1) To explore the impact of land use, climate and environmental heterogeneity on fern species richness along a complete elevational gradient, and (2) to evaluate the relative importance of the three groups of variables within different elevational intervals. Location A temperate mountain region (55,507 km²) of Italy on the southern border of the European Alps divided into a regular grid of 1476 cells (grain 35.7 km²). Methods We applied multiple regression (spatial and non‐spatial) to determine the relative influence of the three groups of variables on species richness, including variation partitioning at two scales. We considered the whole gradient (all 1476 cells) to explain the overall elevational pattern of species richness, and we grouped the cells into elevational intervals of 500 m in order to evaluate the explanatory power of the predictors within different zones along the gradient. Results Species richness showed a hump‐shaped pattern with elevation, forming a plateau between 800 and 1500 m. The lowest species richness was found in warm and relatively dry disturbed lowlands. Moving upwards, the greatest species richness was found in forest‐dominated mid‐elevations with high environmental heterogeneity. At high elevations dominated by open natural habitats, where temperature and precipitation were relatively low, species richness declined but less sharply than in the lowlands. Although it was impossible to separate the effects of the three groups of predictors along the whole gradient, the analysis of separate elevational intervals shed light on their relative importance. The decline of species richness within lowlands was mainly related to a combined effect of deforestation and low environmental heterogeneity. In the middle part of the gradient, habitat heterogeneity and topographic roughness were positively associated with species richness. The richness decline within high‐elevation areas was related mostly to climatic constraints. Main conclusions Human impact due to land‐use modifications strongly affects the elevational pattern of species richness. It is therefore increasingly important to adopt a multiple‐hypothesis approach, taking anthropogenic effects explicitly into account when describing ecological processes along elevational gradients.     

Biogeographic anomalies in the species richness of Chilean forests: Incorporating evolution into a climatic – historic scenario

Broad‐scale richness gradients are closely associated with temperature and water availability. However, historical and evolutionary processes have also contributed to shape current diversity patterns. In this paper we focus on the potential influences of Pleistocene glaciation and phylogenetic niche conservatism (the tendency for traits to be maintained during diversification) on the tree diversity gradient in Chile, and we quantify its primary climatic correlates. Tree species richness is greatest at mid latitudes, particularly in the Andes and Coastal ranges, and decreases abruptly to the south and north. Regression tree analysis identified annual precipitation and annual temperature as the primary probable drivers of this gradient. Ice cover during the Last Glacial Maximum was also identified as an ‘important’ variable, but the contemporary and historical predictors are strongly collinear. Geographically weighted regression indicated that the relationships between richness and environmental variables vary regionally: the relationship between tree richness and precipitation is stronger in north‐central Chile, whereas tree richness and temperature are most strongly associated in south‐central Chile. By assigning each species the age of the family to which it belongs and averaging all species in each geographical unit, we also found that species from the oldest families are distributed mainly in mid to high latitudes and species from younger families are distributed mainly at lower latitudes. This pattern is closely associated with annual precipitation. Thus, the ecological component of tree richness follows contemporary climatic gradients of both energy and water, but the aridification of the Atacama Desert was an important driver over evolutionary time. The influence of recent Pleistocene glaciation remains unresolved but it cannot be discounted.     

Vegetation and Soil 15N Natural Abundance in Alpine Grasslands on the Tibetan Plateau: Patterns and Implications

The natural abundance of nitrogen (N) stable isotopes (δ¹⁵N) has the potential to enhance our understanding of the ecosystem N cycle at large spatial scales. However, vegetation and soil δ¹⁵N patterns along climatic and edaphic gradients have not yet been fully understood, particularly for high-altitude ecosystems. Here we determined vegetation and soil δ¹⁵N in alpine grasslands on the Tibetan Plateau by conducting four consecutive regional surveys during 2001–2004, and then examined their relationships with both climatic and edaphic variables. Our results showed that both vegetation and soil N in Tibetan alpine grasslands were more ¹⁵N-enriched than global averages. Vegetation δ¹⁵N did not exhibit any significant trend along the temperature gradient, but decreased significantly with an increase in precipitation amount. In contrast, soil δ¹⁵N did not vary with either mean annual temperature or precipitation. Our results also indicated that soil δ¹⁵N exhibited a slight increase with clay content, but decreased with soil carbon:nitrogen ratio. A general linear model analysis revealed that variations in vegetation δ¹⁵N were dominantly determined by climatic variables, whereas soil δ¹⁵N was related to edaphic variables. These results provide clues for potential climatic and edaphic regulations on ecosystem N cycle in these high-altitude regions.     

Grass (Poaceae) richness patterns across China’s nature reserves

Explaining grass richness patterns over broad geographic scales is a central issue of biogeography and macroecology. In this investigation, we document the richness patterns of grasses across China’s nature reserves, and discuss their possible explanations at national and regional scales using grass richness data coupled with information from climatic and topographical variables. Our results suggest that the water- and energy-related variables and elevation range (a surrogate of topographical heterogeneity) represent the primary explanations for the variation in grass richness across China’s nature reserves. Significant relationships were found between energy and all grasses, water and C₃ grasses, and the combination of water and energy and C₄ grasses at the national scale. Topographical heterogeneity is strongly associated with grass richness across China, whereas climatic constraints to grass diversity vary regionally. In regions of high rainfall, the presence of C₄ grasses is determined by annual potential evapotranspiration and sunshine hours; the climatic constraint to C₃ grass richness in a large and varied region is sunshine hours; whereas in regions of low soil temperature, the temperature determines the grass richness pattern. Our results also suggest that water variables alone do not represent the primary predictor of grass richness at the regional scale.     

Latitudinal diversity gradients in bryophytes and woody plants: Roles of temperature and water availability

It remains unclear whether the latitudinal diversity gradients of micro- and macro-organisms are driven by the same macro-environmental variables. We used the newly completed species catalog and distribution information of bryophytes in China to explore their spatial species richness patterns, and to investigate the underlying roles of energy availability, climatic seasonality, and environmental heterogeneity in shaping these patterns. We then compared these patterns to those found for woody plants. We found that, unlike woody plants, mosses and liverworts showed only weakly negative latitudinal trends in species richness. The spatial patterns of liverwort richness and moss richness were overwhelmingly explained by contemporary environmental variables, although explained variation was lower than that for woody plants. Similar to woody plants, energy and climatic seasonality hypotheses dominate as explanatory variables but show high redundancy in shaping the distribution of bryophytes. Water variables, that is, the annual availability, intra-annual variability and spatial heterogeneity in precipitation, played a predominant role in explaining spatial variation of species richness of bryophytes, especially for liverworts, whereas woody plant richness was affected most by temperature variables. We suggest that further research on spatial patterns of bryophytes should incorporate the knowledge on their ecophysiology and evolution.     

A global comparative analysis of elevational species richness patterns of ferns

Aim Elevational gradients offer an outstanding opportunity to assess factors determining patterns of species richness, but along single transects potential explanatory factors often covary, making it difficult to distinguish between competing hypotheses. Many previous studies on plants have interpreted their results as supporting the mid‐domain effect (MDE) as a major determinant of species richness, even when climatic factors showed similarly high explanatory power. We compared fern species richness along 20 elevational transects to quantify the relative contribution of climate and MDE as drivers of elevational richness patterns. Location Twenty transects world‐wide. Methods Ferns were sampled in 1039 plots of 400–2500 m² each. Mean annual precipitation and temperature, epiphytic bryophyte cover (as a proxy for air humidity) and MDE predictions were included as independent variables. For each transect, we calculated multiple linear models and partitioned the variance to assess the relative contribution of the independent variables, selecting the most parsimonious models based on Akaike weights and multi‐model inference. Results Along most individual gradients, nearly all variance of fern species richness that could be attributed to either space or MDEs was collinear with climatic factors. Yet, the comparison across transects showed that elevational richness patterns are most parsimoniously accounted for by climatic conditions, especially by low water availability at low elevations and in dry regions in general, and by low temperatures at high elevations and in extra‐tropical regions. Main conclusions Fern species richness is most closely related to climatic factors, and while MDE, surface area and metapopulation processes may somewhat modify the patterns, their importance has been overstated in the past. Future research challenges include determining whether the richness–climate relationship reflects: (1) a direct relationship through the physiological tolerance of the plants, (2) an indirect influence of climate on ecosystem productivity, or (3) an evolutionary legacy of longer or faster diversification processes under certain climatic conditions.     

Squamate richness in the Brazilian Cerrado and its environmental–climatic associations

We investigate patterns of species richness of squamates (lizards, snakes, and amphisbaenians) in the Brazilian Cerrado, identifying areas of particularly high richness, and testing predictions of large‐scale richness hypotheses by analysing the relationship between species richness and environmental climatic variables. We used point localities from museum collections to produce maps of the predicted distributions for 237 Cerrado squamate species, using niche‐modelling techniques. We superimposed distributions of all species on a composite map, depicting richness across the ecosystem. Then, we performed a multiple regression analysis using eigenvector‐based spatial filtering (Principal Coordinate of Neighbour Matrices) to assess environmental–climatic variables that are best predictors of species richness. We found that the environmental–climatic and spatial filters multiple regression model explained 78% of the variation in Cerrado squamate richness (r² = 0.78; F = 32.66; P < 0.01). Best predictors of species richness were: annual precipitation, precipitation seasonality, altitude, net primary productivity, and precipitation during the driest quarter. A model selection approach revealed that several mechanisms related to the different diversity hypothesis might work together to explain richness variation in the Cerrado. Areas of higher species richness in Cerrado were located mainly in the south‐west, north, extreme east, and scattered areas in the north‐west portions of the biome. Partitioning of energy among species, habitat differentiation, and tolerance to variable environments may be the primary ecological factors determining variation in squamate richness across the Cerrado. High richness areas in northern Cerrado, predicted by our models, are still poorly sampled, and biological surveys are warranted in that region. The south‐western region of the Cerrado exhibits high species richness and is also undergoing high levels of deforestation. Therefore, maintenance of existing reserves, establishment of ecological corridors among reserves, and creation of new reserves are urgently needed to ensure conservation of species in these areas.     

Energy–water and seasonal variations in climate underlie the spatial distribution patterns of gymnosperm species richness in China

Studying the pattern of species richness is crucial in understanding the diversity and distribution of organisms in the earth. Climate and human influences are the major driving factors that directly influence the large‐scale distributions of plant species, including gymnosperms. Understanding how gymnosperms respond to climate, topography, and human‐induced changes is useful in predicting the impacts of global change. Here, we attempt to evaluate how climatic and human‐induced processes could affect the spatial richness patterns of gymnosperms in China. Initially, we divided a map of the country into grid cells of 50 × 50 km² spatial resolution and plotted the geographical coordinate distribution occurrence of 236 native gymnosperm taxa. The gymnosperm taxa were separated into three response variables: (a) all species, (b) endemic species, and (c) nonendemic species, based on their distribution. The species richness patterns of these response variables to four predictor sets were also evaluated: (a) energy–water, (b) climatic seasonality, (c) habitat heterogeneity, and (d) human influences. We performed generalized linear models (GLMs) and variation partitioning analyses to determine the effect of predictors on spatial richness patterns. The results showed that the distribution pattern of species richness was highest in the southwestern mountainous area and Taiwan in China. We found a significant relationship between the predictor variable set and species richness pattern. Further, our findings provide evidence that climatic seasonality is the most important factor in explaining distinct fractions of variations in the species richness patterns of all studied response variables. Moreover, it was found that energy–water was the best predictor set to determine the richness pattern of all species and endemic species, while habitat heterogeneity has a better influence on nonendemic species. Therefore, we conclude that with the current climate fluctuations as a result of climate change and increasing human activities, gymnosperms might face a high risk of extinction.     

中国蚂蚁丰富度地理分布格局及其与环境因子的关系

物种丰富度分布格局及其形成机制的研究对于生物多样性保护具有重要意义。为了了解中国蚂蚁物种丰富度分布格局,利用中国省级尺度蚂蚁物种分布数据和环境信息,结合GIS和数理统计方法,探讨蚂蚁物种丰富度的地理分布格局与环境因子之间的关系。研究结果表明:(1)蚂蚁丰富度随纬度增加呈逐渐递减趋势,但缺乏显著的经度梯度。丰富度最高的地区主要集中在南方省份,我国北方、西北干旱区和青藏高原北部地区丰富度较低;(2)简单线性回归分析表明,能量、水分和季节性因素中,影响蚂蚁物种丰富度最强的因子分别为最冷月均温(TEMmin)(R2adj=0.532)、年均降水量(PREC)(R2adj=0.376)和年温度变化范围(TEMvar)(R2adj=0.539),而单个生境异质性因子对蚂蚁物种丰富度的影响均不显著;(3)最优模型由年均温(TEM)、海拔变化范围(ELErange)和年温度变化范围(TEMvar)组成,能够解释68.4%的蚂蚁丰富度地理分异。鉴于海拔变化范围更多地反映与温度相关的生境异质性,因此温度是限制中国蚂蚁分布的最重要因素。另外,分析结果还表明,海南、贵州、江西、四川、安徽和山西等6省蚂蚁区系调查最不充分,是未来发现蚂蚁新分布的热点地区。     

Net primary productivity and seasonality of temperature and precipitation are predictors of the species richness of the Damselflies in the Amazon

Several hypotheses have been proposed to explain the mechanisms that generate temporal and spatial species richness patterns. We tested four common hypotheses (water, energy, climatic heterogeneity and net primary productivity) to evaluate which factors best explain patterns of Zygoptera species richness. Of these, we predicted that climatic heterogeneity would be the most important predictor for Zygoptera richness patterns. We sampled communities of adult Zygoptera in 100 small Amazonian streams. Based on generalized linear mixed models (GLMM), we found that net primary productivity and climatic heterogeneity comprised the best model of Zygoptera species richness in Amazonian streams, with an pseudo r² of 39.5%. Results indicate that species richness increases by one species per 1 kg of biomass per square meter in NPP, or with an increase of 2 °C in air temperature variability. Our work corroborates a recent study with other taxa in Brazilian Bioms. This suggests that temporal variation in climate and net primary productivity are important predictors of the macroecological patterns of richness for aquatic organisms in tropical regions.     

What drives elevational patterns of diversity? A test of geometric constraints,climate and species pool effects for pteridophytes on an elevational gradient in Costa Rica

Aim We studied pteridophyte species richness between 100 m and 3400 m along a Neotropical elevational gradient and tested competing hypotheses for patterns of species richness. Location Elevational transects were situated at Volcán Barva in the Braulio Carrillo National Park and La Selva Biological Station (100–2800 m) and Cerro de la Muerte (2700–3400 m), both on the Atlantic slope of Costa Rica, Central America. Method We analysed species richness on 156 plots of 20 × 20 m and measured temperature and humidity at four elevations (40, 650, 1800 and 2800 m). Species richness patterns were regressed against climatic variables (temperature, humidity, precipitation and actual evapotranspiration), regional species pool, area and predicted species number of a geometric null model (the mid‐domain effect, MDE). Results The species richness of the 484 recorded species showed a hump‐shaped pattern with elevation with a richness peak at mid‐elevations (c. 1700 m). The MDE was the single most powerful explanatory variable in linear regression models, but species richness was also associated strongly with climatic variables, especially humidity and temperature. Area and species pool were associated less strongly with observed richness patterns. Main conclusions Geometric models and climatic models exclusive of geometric constraints explained comparable amounts of the elevational variation in species richness. Discrimination between these two factor complexes is not possible based on model fits. While overall fits of geometric models were high, large‐ and small‐ranged species were explained by geometric models to different extents. Species with narrow elevational ranges clustered at both ends of the gradient to a greater extent than predicted by the MDE null models used here. While geometric models explained much of the pattern in species richness, we cannot rule out the role of climatic factors (or vice versa) because the predicted peak in richness from geometric models, the empirical peak in richness and the overlap in favourable environmental conditions all coincide at middle elevations. Mid‐elevations offer highest humidity and moderate temperatures, whereas at high elevations richness is reduced due to low temperatures, and at low elevations by reduced water availability due to high temperatures.     

Why do mountains support so many species of birds?

Although topographic complexity is often associated with high bird diversity at broad geographic scales, little is known about the relative contributions of geomorphologic heterogeneity and altitudinal climatic gradients found in mountains. We analysed the birds in the western mountains of the New World to examine the two‐fold effect of topography on species richness patterns, using two grains at the intercontinental extent and within temperate and tropical latitudes. Birds were also classified as montane or lowland, based on their overall distributions in the hemisphere. We estimated range in temperature within each cell and the standard deviation in elevation (topographic roughness) based on all pixels within each cell. We used path analysis to test for the independent effects of topographic roughness and temperature range on species richness while controlling for the collinearity between topographic variables. At the intercontinental extent, actual evapotranspiration (AET) was the primary driver of species richness patterns of all species taken together and of lowland species considered separately. In contrast, within‐cell temperature gradients strongly influenced the richness of montane species. Regional partitioning of the data also suggested that range in temperature either by itself or acting in combination with AET had the strongest “effect” on montane bird species richness everywhere. Topographic roughness had weaker “effects” on richness variation throughout, although its positive relationship with richness increased slightly in the tropics. We conclude that bird diversity gradients in mountains primarily reflect local climatic gradients. Widespread (lowland) species and narrow‐ranged (montane) species respond similarly to changes in the environment, differing only in that the richness of lowland species correlates better with broad‐scale climatic effects (AET), whereas mesoscale climatic variation accounts for richness patterns of montane species. Thus, latitudinal and altitudinal gradients in species richness can be explained through similar climatic‐based processes, as has long been argued.     

Contrasting response of native and alien plant species richness to environmental energy and human impact along alpine elevation gradients

Aim We tested whether the species–energy and species–human relationships vary between native and both naturalized and casual alien species richness when other environmental variables had been taken into account. Location Trento Province, a region (c. 6200 km²) on the southern border of the European Alps (Italy), subdivided into 156 contiguous (c. 37.5 km²) cells and ranging in elevation from 66 to 3769 m. Methods Data were separated into three subsets, representing richness of natives, naturalized aliens and casual aliens and separately related to temperature, human population and various environmental correlates of plant species diversity. We applied ordinary least squares and simultaneous autoregressive regressions to identify potential contrasting responses of the three plant status subsets and hierarchical partitioning to evaluate the relative importance of the predictor variables. Results Variation in alien plant species richness along the region was almost entirely explained by temperature and human population density. The relationships were positive but strongly curvilinear. Native species richness was less strongly related to either factor but was positively related to the presence of calcareous bedrock. Native species richness had a decelerating positive relationship with temperature (R²= 55%), whereas naturalized and casual aliens had a positive accelerating relationship explaining 86% and 62% of the variation in richness, respectively. Native species richness had a positive decelerating relationship with population density (R²= 42%), whilst both alien subsets had a positive accelerating relationship. Main conclusions Alien species richness was higher in areas with the most rich and diverse assemblages of native species. Areas at high altitudes are not especially prone to alien invasion due to energy constraints, low propagule pressure and disturbance, even considering a potential increased in temperature. Thus, if we consider future environmental change, we should expect a stronger response of aliens than natives in the currently warm, urbanized, low‐altitude areas than in cold, high‐altitude areas where human population density is low.     

Water–energy,land‐cover and heterogeneity drivers of the distribution of plant species richness in a mountain region of the European Alps

Aim To evaluate the relative importance of water–energy, land‐cover, environmental heterogeneity and spatial variables on the regional distribution of Red‐Listed and common vascular plant species richness. Location Trento Province (c. 6200 km²) on the southern border of the European Alps (Italy), subdivided regularly into 228 3′ × 5′ quadrants. Methods Data from a floristic inventory were separated into two subsets, representing Red‐Listed and common (i.e. all except Red‐Listed) plant species richness. Both subsets were separately related to water–energy, land‐cover and environmental heterogeneity variables. We simultaneously applied ordinary least squares regression with variation partitioning and hierarchical partitioning, attempting to identify the most important factors controlling species richness. We combined the analysis of environmental variables with a trend surface analysis and a spatial autocorrelation analysis. Results At the regional scale, plant species richness of both Red‐Listed and common species was primarily related to energy availability and land cover, whereas environmental heterogeneity had a lesser effect. The greatest number of species of both subsets was found in quadrants with the largest energy availability and the greatest degree of urbanization. These findings suggest that the elevation range within our study region imposes an energy‐driven control on the distribution of species richness, which resembles that of the broader latitude gradient. Overall, the two species subsets had similar trends concerning the relative importance of water–energy, land cover and environmental heterogeneity, showing a few differences regarding the selection of some predictors of secondary importance. The incorporation of spatial variables did not improve the explanatory power of the environmental models and the high original spatial autocorrelation in the response variables was reduced drastically by including the selected environmental variables. Main conclusions Water–energy and land cover showed significant pure effects in explaining plant species richness, indicating that climate and land cover should both be included as explanatory variables in modelling species richness in human‐affected landscapes. However, the high degree of shared variation between the two groups made the relative effects difficult to separate. The relatively low range of variation in the environmental heterogeneity variables within our sampling domain might have caused the low importance of this complex factor.     

Modelling floristic species richness on a regional scale: a case study in Switzerland

In this paper a multivariate linear regression model is proposed for predicting and mapping regional species richness in areas below the timberline according to environmental variables. The data used in setting up the model were derived from a floristic inventory. Using a stepwise regression technique, five environmental variables were found to explain 48.9% of the variability in the total number of plant species: namely temperature range, proximity to a big river or lake, threshold of minimum annual precipitation, amount of calcareous rock outcrops and number of soil types. A considerable part of the unexplained variability is thought to have been influenced by variations in the quality of the botanical inventory. These results show the importance of systematic floristic sampling in addition to conventional inventories when using floristic data as a basis in nature conservation. Nevertheless it is still possible to interpret the resulting diversity patterns ecologically. Regional species richness in Switzerland appears to be a function of: (i) environmental heterogeneity; (ii) threshold values of minimum precipitation; and (iii) presence of calcareous rock outcrops. According to similar studies, environmental heterogeneity was the strongest determinant of total species richness. In contrast to some studies, high productivity decreased the number of species. Furthermore, the implications of this work for climate change scenarios are discussed.