An investigation into the relationship between water beetle (Coleoptera) distribution and land cover in Scotland and northeast England

Aims To determine the relationship between satellite‐derived land cover data and distribution patterns of water beetle species pools. Location A total of 687 British national grid 10 km squares in Scotland and 588 1 km grid squares in 99 10 km grid squares in northeast England. Methods Multivariate classification and constrained ordination analyses were used with water beetle species and land cover data. Results The major variation in both the Scotland and northeast England 10 km species pools was from squares with upland acid water habitats to squares with lowland fens and ponds whilst for the 1 km northeast England data it was from squares dominated by fast‐flowing streams and rivers to those with fens and ponds. The secondary variations in the 10 and 1 km analyses were the primary ones juxtaposed. Constrained ordination of the Scottish 10 km data showed upland land cover types to be most influential in determining species pool distribution whilst lowland covers were more important with both the 10 and 1 km northeast England pools. The three analyses showed that coastal cover had a relatively high influence on species pool distribution but no species pool group was dominated by coastal species. Main conclusions There were strong relationships between water beetle species pool distributions and the satellite‐derived land cover types dominating upland and lowland areas. The major variation in the northeast England species pool data differed at the two scales analysed. Results indicated that there is considerable potential for the synthesis of water beetle distribution and land cover data for use in environmental and conservation monitoring at both the regional and national scales.     

The relationship between British ground beetles (Coleoptera, Carabidae) and land cover

Aims and methods Ground beetle and satellite‐derived land cover data from 1687 United Kingdom 10 km national grid squares were used to assess the relationship between species pool and cover data in Great Britain using fuzzy classification and constrained ordination. Results Ground beetle species pools classified into nine groups which were related to land cover variables using constrained ordination. There was a strong relationship between upland land cover and three ground beetle groups. Deciduous woodland, coastal and tilled land were associated with three other groups. Three further groups did not appear to be strongly associated with any particular cover, but differed in geographical position. Conclusion The distribution of species pools derived from the British national recording scheme at the 10 km scale was strongly related to satellite‐derived land cover data. There appears to be considerable potential for the use of a synthesis of land cover and ground beetle data in the monitoring of environmental change over a large, countrywide, area.     

A Strategic Interpretation of Beetle (Coleoptera) Assemblages, Biotopes, Habitats and Distribution, and the Conservation Implications

The identification of beetle, and other invertebrate, biotopes based on the recording of species assemblages has become a regular occurrence as a result of increased survey work, with the approach expanded into classifications of grid square pooled species lists at the national scale. A reassessment of beetle biotopes and distribution has been attempted by interpreting the classifications in terms of the major environmental factors of productivity and disturbance, identified as being important drivers in work on habitat templates and strategic triangles. For grassland ground beetle biotopes, productivity was generally related to soil quality whilst disturbance was associated with land management or cover. Productivity of exposed riverine sediment ground beetle biotopes was dependent on deposited organic matter and disturbance on the effects of water flow on site structure. With both ground beetle biotopes, the distribution of assemblages was also affected by substrate water, another abiotic driver. Productivity in aquatic beetle biotopes was a function of base-status, generally reflected by pH, whilst disturbance was mainly due to water flow and wave action. However, disturbance in ditches was also the result of site vegetation management whilst another factor affecting assemblage distribution was water permanence, with temporary water having specific assemblages. For large-scale British, grid-based, classifications productivity and disturbance were interpreted from satellite-derived land cover data. At the national scale, the other important factor influencing distribution was temperature. There is a requirement for better quantification of a number of the factors influencing biotope and assemblage definition. Species strategies employed to cope with the various environmental variables were reflected in traits in such factors as morphology and life-cycles. The effects of environmental perturbations including climate change, pollution and land use are discussed in relation to the environmental variables and to the various species strategies. Biotope classifications, using standardised and reproducible survey methods, allied to a better understanding of the underlying environmental pressures, could produce a unified approach for the conservation of beetle and other invertebrate species.     

Macroinvertebrate species and assemblages in the headwater streams of the River Tyne,northern England in relation to land cover and other environmental variables

The macroinvertebrate species and assemblages of headwater streams of the River Tyne catchment in northern England were classified and their relationship with environmental variables based on stream structure, water acidity, distance from source and land cover investigated using constrained ordination and logistic regression. Fuzzy classification of data from 322 stream sites generated five assemblages. Stream structure, quantified as an exposure index, was found to be the most important environmental variable, with water acidity also important. Distance from source and land cover had less influence on species and assemblage distribution. A considerable amount of variation in assemblage distribution was explained using a two-variable logistic regression with stream structure (exposure index) and water acidity (pH) in a template. Site structure and water acidity appeared to be related to drift, geology and topography with little anthropogenic influence. The applicability of the habitat template concept for explaining the distribution of stream macroinvertebrates is discussed.     

Exploring the relationship between land cover and the distribution of water beetle species (Coleoptera) at the regional scale

The advent of remote-sensed satellite land cover data has provided the opportunity to assess the relationship between invertebrate species distributions and individual land cover types. Water beetle species occur in habitats within specific land cover types and the relationship between the distribution of water beetle species and land covers at the regional scale was investigated using records of 154 species from 1018 sites in north-east England. The land covers of tilled land and urban in the lowlands and of shrub heath and heath grassland in the upland areas proved to be most important in explaining the distribution of species. There were both positive and negative associations between some species and other covers such as woodland and the coast. However, a considerable number of species, generally those with a large number of records, showed no strong relationships with any land cover types. The integration of water beetle species recording data and remote-sensed land cover data as a basis for predicting and monitoring both species distribution and environmental change is discussed.     

Investigating the relationships between the distribution of British ground beetle species (Coleoptera, Carabidae) and temperature, precipitation and altitude

Aims We examine the relationships between the distribution of British ground beetle species and climatic and altitude variables with a view to developing models for evaluating the impact of climate change. Location Data from 1684 10‐km squares in Britain were used to model species–climate/altitude relationships. A validation data set was composed of data from 326 British 10‐km squares not used in the model data set. Methods The relationships between incidence and climate and altitude variables for 137 ground beetle species were investigated using logistic regression. The models produced were subjected to a validation exercise using the Kappa statistic with a second data set of 30 species. Distribution patterns for four species were predicted for Britain using the regression equations generated. Results As many as 136 ground beetle species showed significant relationships with one or more of the altitude and climatic variables but the amount of variation explained by the models was generally poor. Models explaining 20% or more of the variation in species incidence were generated for only 10 species. Mean summer temperature and mean annual temperature were the best predictors for eight and six of these 10 species respectively. Few models based on altitude, annual precipitation and mean winter temperature were good predictors of ground beetle species distribution. The results of the validation exercise were mixed, with models for four species showing good or moderate fits whilst the remainder were poor. Main conclusions Whilst there were many significant relationships between British ground beetle species distributions and altitude and climatic variables, these variables did not appear to be good predictors of ground beetle species distribution. The poor model performance appears to be related to the coarse nature of the response and predictor data sets and the absence of key predictors from the models.     

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.     

Ground beetle species (Coleoptera, Carabidae) associations with land cover variables in northern England and southern Scotland

Distribution data concerning 172 ground beetle species derived from 1145 pitfall trap sites in northern England and southern Scotland were used to assess the relationship between species distribution and 12 satellite-derived land cover variables at the regional scale. A number of species were strongly associated with one cover type and negatively with others. The major variation was for preferences for covers in upland or lowland parts of the region. Other distinct preferences for some species were covers such as those at the coast whilst a number of common species showed no strong preference for any cover variable. The synthesis of ground beetle species distribution and satellite-derived cover data is discussed in relation to environmental assessment and change.     

The influence of multi‐scale environmental variables on the distribution of terricolous lichens in a fog desert

Question: How do environmental variables in a hyper‐arid fog desert influence the distribution patterns of terricolous lichens on both macro‐ and micro‐scales? Location: Namib Desert, Namibia. Methods: Sites with varying lichen species cover were sampled for environmental variables on a macro‐scale (elevation, slope degree, aspect, proximity to river channels, and fog deposition) and on a micro‐scale (soil structure and chemistry). Macro‐scale and micro‐scale variables were analysed separately for associations with lichen species cover using constrained ordination (DCCA) and unconstrained ordination (DCA). Explanatory variables that dominated the first two axes of the constrained ordinations were tested against a lichen cover gradient. Results: Elevation and proximity to river channels were the most significant drivers of lichen species cover in the macro‐scale DCCA, but results of the DCA suggest that a considerable percentage of variation in lichen species cover is unexplained by these variables. On a micro‐scale, sediment particle size explained a majority of lichen community variations, followed by soil pH. When both macro and micro‐scale variables were tested along a lichen cover gradient, soil pH was the only variable to show a significant relationship to lichen cover. Conclusion: The findings suggest that landscape variables contribute to variations in lichen species cover, but that stronger links occur between lichen growth and small‐scale variations in soil characteristics, supporting the need for multi‐scale approaches in the management of threatened biological soil crust communities and related ecosystem functions.     

Local and regional factors determining aquatic and semi-aquatic bug (Heteroptera) assemblages in rivers and streams of Greece

Heteroptera species were collected from 48 sites distributed throughout the mainland and island complexes of Greece during 1999–2004. The aims of this study were to investigate Heteroptera distribution and abundance in Greek streams, identify the environmental factors that are linked to variation in their assemblages and to partition the influence of environmental and spatial components, alone and in combination, on Heteroptera community composition. Canonical ordination techniques (CCA) were used to determine the relationship between environmental variables and species abundance, while variation partitioning was performed using partial CCA to understand the importance of different explanatory variables in Heteroptera variation. Heteroptera variation was decomposed into independent and joint effects of local (physicochemical variables, microhabitat composition, stream width and depth), regional (land use/cover) and geographic variables (longitude, latitude, altitude and distance to source). Land use/cover, aquatic and riparian vegetation, stream size and water chemistry were the most important factors structuring Heteroptera assemblages. At regional scale, bug assemblages were mainly divided into those found in forested and agricultural landscapes, following water quality and microhabitat composition at local scale. Local variables accounted for 48% of the total explained variation, regional variables for 20% whereas geographical position appeared to be the least influencing factor (8.5%). The results of partial constraint analyses suggested that local variables play a major role in Heteroptera variation followed by regional variables. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

Predicting patterns of vascular plant species richness with composite variables: a meso-scale study in Finnish Lapland

This paper is an attempt, using statistical modelling techniques, to understand the patterns of vascular plant species richness at the poorly studied meso-scale within a relatively unexplored subarctic zone. Species richness is related to floristic-environmental composite variables, using occurrence data of vascular plants and environmental and spatial predictor variables in 362 1 km² grid squares in the Kevo Nature Reserve. Species richness is modelled in two different way. First, by detecting the major floristic-environmental gradients with the ordination procedure of canonical correspondence analysis, and subsequently relating these ordination axes to species richness by generalized linear modelling. Second, species richness is directly related to the composite environmental factors of explanatory variables, using partial least squares regression. The most important explanatory variables, as suggested by both approaches, are relatively similar, and largely reflect the influence of altitude or altitudinally related variables in the models. The most prominent floristic gradient in the data runs from alpine habitats to river valleys, and this gradient is the main source of variation in species richness. Some local environmental variables are also relatively important predictors; the grid squares rich in vascular plant taxa are mainly located in the lowlands of the reserve and are characterised by rivers and brooks, as well as by abundant cliff walls. The two statistical models account for approximately the same amount of variation in the species richness, with more than half of the variation unexplained. Potential reasons for the relatively modest fit are discussed, and the results are compared to the characteristics of the diversity-environment relationships at both broader- and finer-scales.     

Threat spots and environmental determinants of red-listed plant,butterfly and bird species in boreal agricultural environments

The aims of this study were (1) to examine the geographic distribution of red-listed species of agricultural environments and identify their national threat spots (areas with high diversity of threatened species) in Finland and (2) to determine the main environmental variables related to the richness and occurrence patterns of red-listed species. Atlas data of 21 plant, 17 butterfly and 11 bird species recorded using 10 km grid squares were employed in the study. Generalized additive models (GAMs) were constructed separately for species richness and occurrence of individual species of the three species groups using climate and land cover predictor variables. The predictive accuracy of models, as measured using correlation between the observed and predicted values and AUC statistics, was generally good. Temperature-related variables were the most important determinants of species richness and occurrence of all three taxa. In addition, land cover variables had a strong effect on the distribution of species. Plants and butterflies were positively related to the cover of grasslands and birds to small-scale agricultural mosaic as well as to arable land. Spatial coincidence of threat spots of plants, butterflies and birds was limited, which emphasizes the importance of considering the potentially contrasting environmental requirements of different taxa in conservation planning. Further, it is obvious that the maintenance of various non-crop habitats and heterogeneous agricultural landscapes has an essential role in the preservation of red-listed species of boreal rural environments.     

Assessing the vulnerability of European butterflies to climate change using multiple criteria

Vulnerability of 100 European butterfly species to climate change was assessed using 13 different criteria and data on species distributions, climate, land cover and topography from 1,608 grid squares 30′ × 60′ in size, and species characteristics increasing the susceptibility to climate change. Four bioclimatic model-based criteria were developed for each species by comparing the present-day distribution and climatic suitability of the occupied grid cells with projected distribution and suitability in the future using the HadCM3-A2 climate scenario for 2051–2080. The proportions of disadvantageous land cover types (bare areas, water, snow and ice, artificial surfaces) and cultivated and managed land in the occupied grid squares and their surroundings were measured to indicate the amount of unfavourable land cover and dispersal barriers for butterflies, and topographical heterogeneity to indicate the availability of potential climatic refugia. Vulnerability was also assessed based on species dispersal ability, geographical localization and habitat specialization. Northern European species appeared to be amongst the most vulnerable European butterflies. However, there is much species-to-species variation, and species appear to be threatened due to different combinations of critical characteristics. Inclusion of additional criteria, such as life-history species characteristics, topography and land cover to complement the bioclimatic model-based species vulnerability measures can significantly deepen the assessments of species susceptibility to climate change.     

Stream macroinvertebrate response to catchment urbanisation (Georgia, U.S.A.)

SUMMARY 1. The effects of catchment urbanisation on water quality were examined for 30 streams (stratified into 15, 50 and 100 km² ± 25% catchments) in the Etowah River basin, Georgia, U.S.A. We examined relationships between land cover (implying cover and use) in these catchments (e.g. urban, forest and agriculture) and macroinvertebrate assemblage attributes using several previously published indices to summarise macroinvertebrate response. Based on a priori predictions as to mechanisms of biotic impairment under changing land cover, additional measurements were made to assess geomorphology, hydrology and chemistry in each stream. 2. We found strong relationships between catchment land cover and stream biota. Taxon richness and other biotic indices that reflected good water quality were negatively related to urban land cover and positively related to forest land cover. Urban land cover alone explained 29–38% of the variation in some macroinvertebrate indices. Reduced water quality was detectable at c. >15% urban land cover. 3. Urban land cover correlated with a number of geomorphic variables such as stream bed sediment size (–) and total suspended solids (+) as well as a number of water chemistry variables including nitrogen and phosphorus concentrations (+), specific conductance (+) and turbidity (+). Biotic indices were better predicted by these reach scale variables than single, catchment scale land cover variables. Multiple regression models explained 69% of variation in total taxon richness and 78% of the variation in the Invertebrate Community Index (ICI) using phi variability, specific conductance and depth, and riffle phi, specific conductance and phi variability, respectively. 4. Indirect ordination analysis was used to describe assemblage and functional group changes among sites and corroborate which environmental variables were most important in driving differences in macroinvertebrate assemblages. The first axis in a non‐metric multidimensional scaling ordination was highly related to environmental variables (slope, specific conductance, phi variability; adj. R²=0.83) that were also important in our multiple regression models. 5. Catchment urbanisation resulted in less diverse and more tolerant stream macroinvertebrate assemblages via increased sediment transport, reduced stream bed sediment size and increased solutes. The biotic indices that were most sensitive to environmental variation were taxon richness, EPT richness and the ICI. Our results were largely consistent over the range in basin size we tested.     

Urban plant ecology patterns and processes: a case study of the flora of the City of Plymouth, Devon, U. K.

Aim Using a large database that has been created over the past 5 years with the RECORDER package, presence/absence data for 829 species of vascular plants in the 103 1-km² squares that cover the city of Plymouth (pop. 243,373) have been analysed by two-way indicator species analysis (TWINSPAN ) and canonical correspondence analysis (CCA) to establish the major species assemblages and to examine their spatial distribution across the city in relation to variation in land use. Location The City of Plymouth. Methods Nine groupings of squares emerged and their distribution was mapped across the city. Interpretation of those groups and their variation in relation to the land use ordination axes showed that TWINSPAN groups lying along the first axis of variation correlated floristic variation with the process of urban development and the historical evolution of urban structure. The second axis appeared to be related to particular remnant semi-natural habitats within the city that could be regarded as ‘hot-spots’ for survival of many plant species. Species were categorized into four types on the basis of their recency and mode of arrival in the city and by using the historical flora of Plymouth produced by T. R. Archer-Briggs in 1880. Variations in the sources of species in relation to the TWINSPAN classification groups were then examined. A second ordination was also computed; this time with geological, altitudinal and distance variables. TWINSPAN groups were then superimposed on the ordinations using polygons to assist with biogeographical interpretation. Results and conclusions The results of both the multivariate analyses and distribution of species by source are then discussed in the context of previous research into urban plant ecology, particularly in Central Europe. The problems of inferring process from pattern in this meso-scale study are reviewed and suggestions are made for further research into urban ecology and biogeography in the city.     

Ecosystem classification and inventory maps as surrogates for ground beetle assemblages in boreal forest

Aims We compare performance of ecosystem classification maps and provincial forest inventory data derived from air photography in reflecting ground beetle (Coleoptera: Carabidae) biodiversity patterns that are related to the forest canopy mosaic. Our biodiversity surrogacy model based on remotely sensed tree canopy cover is validated against field-collected ground data.Methods We used a systematic sampling grid of 198 sites, covering 84 km 2 of boreal mixedwood forest in northwestern Alberta, Canada. For every site, we determined tree basal area, characterized the ground beetle assemblage and obtained corresponding provincial forest inventory and ecosystem classification information. We used variation partitioning, ordination and misclassification matrices to compare beetle biodiversity patterns explained by alternative databases and to determine model biases originating from air photo-interpretation.Important findings Ecosystem classification data performed better than canopy cover derived from forest inventory maps in describing ground beetle biodiversity patterns. The biodiversity surrogacy models based on provincial forest inventory maps and field survey generally detected similar patterns but inaccuracies in air photo-interpretation of relative canopy cover led to differences between the two models. Compared to field survey data, air photo-interpretation tended to confuse two Picea species and two Populus species present and homogenize stand mixtures. This generated divergence in models of ecological association used to predict the relationship between ground beetle assemblages and tree canopy cover. Combination of relative canopy cover from provincial inventory with other geo-referenced land variables to produce the ecosystem classification maps improved biodiversity predictive power. The association observed between uncommon surrogates and uncommon ground beetle species emphasizes the benefits of detecting these surrogates as a part of landscape management. In order to complement conservation efforts established in protected areas, accurate, high resolution, wide ranging and spatially explicit knowledge of landscapes under management is primordial in order to apply effective biodiversity conservation strategies at the stand level as required in the extensively harvested portion of the boreal forest. In development of these strategies, an in-depth understanding of vegetation is key.     

A numerical analysis of the mesoscale distribution patterns of vascular plants in the subarctic Kevo Nature Reserve, northern Finland

Different numerical techniques were used to detect and describe the major ecological-biogeographical patterns of vascular plant distributions at the meso-scale level in a subarctic region in Finland. The distribution patterns of 231 native taxa in 362 1 km² grid squares of the Kevo Nature Reserve were analysed by two-way indicator species analysis and detrended correspondence analysis, and were subsequently related to twenty-eight geographical, topographical, geological, and vegetational variables using simple discriminant functions and canonical correspondence analysis with associated Monte Carlo permutation tests.
The floristic variation detected reflects variations in environmental factors operative at the regional and local scales. No major broad-scale coherent geographical patterns were detected; instead, the spatial distribution of the grids with a similar floristic composition shows a scattered distribution. All the numerical techniques reveal a major gradient from alpine areas to lowland sites with rivers and rocky outcrops, and the most important explanatory variables for predicting the main floristic variation are all associated with altitude. The floristic patterns represented by the second ordination gradient mainly correlate with the abundance of mires. Partial ordinations indicate that both the geographical and geological variables explain relatively little of the species distributional patterns.
Although the meso-scale approach reveals much about the plant-environment relationships in the study area, the floristic variation appears to be determined mainly by fine-scale factors. In the most heterogeneous grids, the grid size used fails to detect accurately the ecological patterns of the species present.     

Landscape patterns of species-level association between ground-beetles and overstory trees in boreal forests of western Canada (Coleoptera, Carabidae)

Spatial associations between species of trees and ground-beetles (Coleoptera: Carabidae) involve many indirect ecological processes, likely reflecting the function of numerous forest ecosystem components. Describing and quantifying these associations at the landscape scale is basic to the development of a surrogate-based framework for biodiversity monitoring and conservation. In this study, we used a systematic sampling grid covering 84 km(2) of boreal mixedwood forest to characterize the ground-beetle assemblage associated with each tree species occurring on this landscape. Projecting the distribution of relative basal area of each tree species on the beetle ordination diagram suggests that the carabid community is structured by the same environmental factors that affects the distribution of trees, or perhaps even by trees per se. Interestingly beetle species are associated with tree species of the same rank order of abundance on this landscape, suggesting that conservation of less abundant trees will concomitantly foster conservation of less abundant beetle species. Landscape patterns of association described here are based on characteristics that can be directly linked to provincial forest inventories, providing a basis that is already available for use of tree species as biodiversity surrogates in boreal forest land management.     

Fine‐scale determinants of butterfly species richness and composition in a mountain region

Aim Global patterns of species richness are often considered to depend primarily on climate. We aimed to determine how topography and land cover affect species richness and composition at finer scales. Location Sierra de Guadarrama (central Iberian Peninsula). Methods We sampled the butterfly fauna of 180 locations (89 in 2004, 91 in 2005) at 600–2300 m elevation in a region of 10800 km². We recorded environmental variables at 100‐m resolution using GIS, and derived generalized linear models for species density (number of species per unit area) and expected richness (number of species standardized to number of individuals) based on variables of topoclimate (elevation and insolation) or land cover (vegetation type, geology and hydrology), or both (combined). We evaluated the models against independent data from the alternative study year. We also tested for differences in species composition among sites and years using constrained ordination (canonical correspondence analysis), and used variation partitioning analyses to quantify the independent and combined roles of topoclimate and land cover. Results Topoclimatic, land cover and combined models were significantly related to observed species density and expected richness. Topoclimatic and combined models outperformed models based on land cover variables, showing a humped elevational diversity gradient. Both topoclimate and land cover made significant contributions to models of species composition. Main conclusions Topoclimatic factors may dominate species richness patterns in regions with pronounced elevational gradients, as long as large areas of natural habitat remain. In contrast, both topoclimate and land cover may have important effects on species composition. Biodiversity conservation in mountainous regions therefore requires protection and management of natural habitats over a wide range of topoclimatic conditions, which may assist in facilitating range shifts and alleviating declines in species richness related to climate change.