Three-dimensional mid-domain predictions: geometric constraints in North American amphibian, bird, mammal and tree species richness patterns

The "mid-domain effect" (MDE) has received much attention as a candidate explanation for patterns in species richness over large geographic areas. Mid-domain models generate a central peak in richness when species ranges are placed randomly within a bounded geographic area (i.e. the domain). Until now, domain limits have been described mostly in one-dimension, usually latitude or elevation, and only occasionally in two-dimensions. Here we test 1-D, 2-D and, for the first time, 3-D mid-domain models and assess the effects of geometric constraints on species richness in North American amphibian, bird, mammal and tree species. Using spatially lagged simultaneous autoregressive models, empirical richness was predicted quite well by the mid-domain predictions and the spatial autoregressive term (45–92% R²). However, our results show that empirical species richness peaks do deviate from those of the MDE predictions in 3 dimensions. Variation explained (R²) by MDE predictions generally increased with increasing mean range size of the different biotic groups (from amphibian, to tree, mammal and finally bird data), and decreased with increasing dimensions being accounted for in the models. The results suggest geometric constraints alone can explain much of the variation in species richness with elevation, specifically with respect to the larger-range taxa, birds and mammals. Our analysis addresses many of the recent methodological criticisms directed at studies testing the MDE, and our results support the hypothesis that species diversity patterns are influenced by geometric constraints.     

The mid‐domain effect matters: simulation analyses of range‐size distribution data from Mount Kinabalu,Borneo

Aim In simulation exercises, mid‐domain peaks in species richness arise as a result of the random placement of modelled species ranges within simulated geometric constraints. This has been called the mid‐domain effect (MDE). Where close correspondence is found between such simulations and empirical data, it is not possible to reject the hypothesis that empirical species richness patterns result from the MDE rather than being the outcome (wholly or largely) of other factors. To separate the influence of the MDE from other factors we therefore need to evaluate variables other than species richness. The distribution of range sizes gives different predictions between models including the MDE or not. Here, we produce predictions for species richness and distribution of range sizes from one model without the MDE and from two MDE models: a classical MDE model encompassing only species with their entire range within the domain (range‐restricted MDE), and a model encompassing all species with the theoretical midpoint within the domain (midpoint‐restricted MDE). These predictions are compared with observations from the elevational pattern of range‐size distributions and species richness of vascular plants. Location Mount Kinabalu, Borneo. Methods The data set analysed comprises more than 28,000 plant specimens with information on elevation. Species ranges are simulated with various assumptions for the three models, and the species simulated are subsequently subjected to a sampling that simulates the actual collection of species on Mount Kinabalu. The resulting pattern of species richness and species range‐size distributions are compared with the observed pattern. Results The comparison of simulated and observed patterns indicates that an underlying monotonically decreasing trend in species richness with elevation is essential to explain fully the observed pattern of richness and range size. When the underlying trend is accounted for, the MDE model that restricts the distributions of theoretical midpoints performs better than both the classical MDE model and the model that does not incorporate geometric constraints. Main conclusions Of the three models evaluated here, the midpoint‐restricted MDE model is found to be the best for explaining species richness and species range‐size distributions on Mount Kinabalu.     

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.     

Explaining Andean megadiversity: the evolutionary and ecological causes of glassfrog elevational richness patterns

The Tropical Andes are an important global biodiversity hotspot, harbouring extraordinarily high richness and endemism. Although elevational richness and speciation have been studied independently in some Andean groups, the evolutionary and ecological processes that explain elevational richness patterns in the Andes have not been analysed together. Herein, we elucidate the processes underlying Andean richness patterns using glassfrogs (Centrolenidae) as a model system. Glassfrogs show the widespread mid‐elevation diversity peak for both local and regional richness. Remarkably, these patterns are explained by greater time (montane museum) rather than faster speciation at mid‐elevations (montane species pump), despite the recency of the major Andean uplift. We also show for the first time that rates of climatic‐niche evolution and elevational change are related, supporting the hypothesis that climatic‐niche conservatism decelerates species' shifts in elevational distributions and underlies the mid‐elevation richness peak. These results may be relevant to other Andean clades and montane systems globally.     

Neutral community dynamics, the mid-domain effect and spatial patterns in species richness

The mid‐domain effect (MDE) aims to explain spatial patterns in species richness invoking only stochasticity and geometrical constraints. In this paper, we used simulations to show that its main qualitative prediction, a hump‐shaped pattern in species richness, converges to the expectation of a spatially bounded neutral model when communities are linked by short‐distance migration. As these two models can be linked under specific situations, neutral theory may provide a mechanistic population level basis for MDE. This link also allows establishing in which situations MDE patterns are more likely to be found. Also, in this situation, MDE models could be used as a first approximation to understand the role of both stochastic (ecological drift and migration) and deterministic (adaptation to environmental conditions) processes driving the spatial structure of species richness.     

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.     

Effects of contemporary environment and Quaternary climate change on drylands plant diversity differ between growth forms

Previous studies on large‐scale patterns in plant richness and underlying mechanisms have mostly focused on forests and mountains, while drylands covering most of the world's grasslands and deserts are more poorly investigated for lack of data. Here, we aim to 1) evaluate the plant richness patterns in Inner Asian drylands; 2) compare the relative importance of contemporary environment, historical climate, vegetation changes, and mid‐domain effect (MDE); and 3) explore whether the dominant drivers of species richness differ across growth forms (woody vs herbaceous) and range sizes (common vs rare). Distribution data and growth forms of 13 248 seed plants were compiled from literature and species range sizes were estimated. Generalized linear models and hierarchical partitioning were used to evaluate the relative contribution of different factors. We found that habitat heterogeneity strongly affected both woody and herbaceous species. Precipitation, climate change since the mid‐Holocene and climate seasonality dominated herbaceous richness patterns, while climate change since the Last Glacial Maximum dominated woody richness patterns. Rare species richness was strongly correlated with precipitation, habitat heterogeneity and historical climatic changes, while common species richness was strongly correlated with MDE (woody) or climate seasonality (herbaceous). Temperature had little effects on the species richness patterns of all groups. This study represents the first evaluation of the large‐scale patterns of plant species richness in the Inner Asian drylands. Our results suggest that increasing water deficit due to anthropogenic activities combined with future global warming may increase the extinction risk of many grassland species. Rare species (both herbaceous and woody) may face severe challenges in the future due to increased habitat destruction caused by urbanization and resource exploitation. Overall, our findings indicate that the hypotheses on species richness patterns based on woody plants alone can be insufficient to explain the richness patterns of herbaceous species.     

Fish distributions along depth gradients of a sea mountain range conform to the mid‐domain effect

Species richness often peaks in the middle of bounded geographic domains (e.g. latitude, altitude or depth). Hump‐shaped richness distributions may be due to deterministic processes, such as adaptations to environmental variation. Alternatively, such distributions might also be due to stochastic process. The mid‐domain effect (MDE) posits that hump‐shaped richness distributions arise when species ranges are randomly arranged within the limits of the domain. We tested whether the MDE could account for the richness of bottom‐associated (demersal) fishes between 200 and 800 m on the Chatham Rise, New Zealand. We quantified the depth distributions of 59 fish species from 1891 research trawl catches made between 1991 and 2007. Results showed a broad plateau of high species richness near the centre of the domain (between 300 and 700 m), which was consistent with expectations of the MDE. Further, empirical species richness was better explained statistically by predictions of the MDE than models incorporating additional abiotic predictor variables. Our results deviated from previous studies that identified a greater richness of fishes in warmer, shallower depths with higher primary production. However, our study was conducted entirely below the euphotic zone, at depths where gradients are relatively weak, suggesting that support for the mid‐domain effect may increase across oceanic domains characterised by weak environmental gradients.     

Elevational gradients of reptile richness in the southern Western Ghats of India: Evaluating spatial and bioclimatic drivers

Exploring elevational patterns in species richness and their underlying mechanisms is a major goal in biogeography and community ecology. Reptiles can be powerful model organisms to examine biogeographical patterns. In this study, we examine the elevational patterns of reptile species richness and test a series of hypotheses that may explain them. We sampled reptile communities along a tropical elevational gradient (100–1,500 m a.s.l.) in the Western Ghats of India using time‐constrained visual encounter surveys at each 100‐m elevation zone for 3 years. First, we investigated species richness patterns across elevation and the support of mid‐domain effect and Rapoport's rule. Second, we tested whether a series of bioclimatic (temperature and tree density) and spatial (mid‐domain effect and area) hypotheses explained species richness. We used linear regression and AICc to compare competing models for all reptiles, and each of the subgroups: snakes, lizards, and Western Ghats’ endemics. Overall reptile richness and lizard richness both displayed linear declines with elevation, which was best explained by temperature. Snake richness and endemic species richness did not systematically vary across elevation, and none of the potential hypotheses explained variation in them. This is the first standardized sampling of reptiles along an elevational gradient in the Western Ghats, and our results agree with the global view that temperature is the primary driver of ectotherm species richness. By establishing strong reptile diversity–temperature associations across elevation, our study also has implications for the impact of future climate change on range‐restricted species in the Western Ghats.     

Do Rapoport's rule,the mid‐domain effect or the source–sink hypotheses predict bathymetric patterns of polychaete richness on the Pacific coast of South America?

Aim We evaluated the bathymetric gradient of benthic polychaete species richness from the Chilean coast, as well as its possible underlying causes. We tested three possible hypotheses to explain the richness gradient: (1) Rapoport's effect; (2) the mid‐domain effect (MDE); and (c) the source–sink hypothesis. Location South‐eastern Pacific coast of Chile. Methods The bathymetric gradient in richness was evaluated using the reported ranges of bathymetric distribution of 498 polychaete species, from the intertidal to abyssal zone (c. 4700 m). Rapoport's effect was evaluated by examining the relationship between bathymetric mid‐point and bathymetric range extent, and species richness and depth. The MDE was tested using the Monte Carlo simulation program. The source–sink hypothesis was tested through nestedness analysis. Results Species richness shows significant exponential decay across the bathymetric gradient. The pattern is characterized by a high presence of short‐ranged species on the continental shelf area; while only a few species reach abyssal depths, and they tend to show extremely wide bathymetric ranges. Our simulation analyses showed that, in general, the pattern is robust to sampling artefacts. This pattern cannot be reproduced by the MDE, which predicts a parabolic richness gradient. Rather, results agree with the predictions of Rapoport's effect. Additionally, the data set is significantly nested at species, genus and family levels, supporting the source–sink hypothesis. Main conclusions The sharp exponential decay in benthic polychaete richness across the bathymetric gradient supports the general idea that abyssal environments should harbour fewer species than shallower zones. This pattern may be the result of colonization–extinction dynamics, characterized by abyssal assemblages acting as ‘sinks’ maintained mainly by shallower ‘sources’. The source–sink hypothesis provides a conceptual and methodological framework that may shed light on the search for general patterns of diversity across large spatial scales.     

Neotropical savanna ants show a reversed latitudinal gradient of species richness,with climatic drivers reflecting the forest origin of the fauna

Aim

To evaluate the extent to which ant species richness in Neotropical savannas varies with macrogeographic variables, and to identify the potential climatic drivers of such variation.

Location

The Cerrado savanna biome of central Brazil, in a region spanning ca. 20° of latitude and 18°of longitude.

Methods

Standardized sampling of the arboreal and ground‐dwelling faunas was performed in 29 well‐preserved savanna sites using pitfall traps. Species were classified according to their habitat affinities: open‐savanna specialists, forest‐associated species or habitat generalists. We used generalized linear models to evaluate the importance of geographic (latitude, longitude and elevation) and climatic (mean temperature and three metrics of rainfall) variables as predictors of species richness.

Results

The total number of species recorded at each site varied more than twofold (from 59 to 144), and latitude was the best geographic correlate of overall species richness. However, contrary to the expected pattern, more species were found at higher than lower latitudes. This reversed latitudinal pattern of diversity occurred for both the arboreal and ground‐dwelling faunas, and for the habitat generalists and forest specialists. The savanna specialists showed a mid‐latitudinal peak in diversity. Overall, there was a significant positive association between rainfall and species richness, but the strength of this relationship varied with ant habitat affinity.

Main conclusions

The Cerrado ant fauna shows a reverse latitudinal gradient in species diversity, and this can be explained by increasing rainfall during the warmest months of the year (and therefore in plant productivity) with increasing latitude. The sensitivity of Cerrado ant diversity to declining rainfall contrasts with the high resilience to aridity of the Australian savanna ant fauna, and this reflects the contrasting evolutionary histories of these faunas. Our findings highlight the importance of historical processes as drivers of intercontinental contrasts in macroecological patterns.     

Small mammal species richness and turnover along elevational gradient in Yulong Mountain,Yunnan, Southwest China

Understanding the species diversity patterns along elevational gradients is critical for biodiversity conservation in mountainous regions. We examined the elevational patterns of species richness and turnover, and evaluated the effects of spatial and environmental factors on nonvolant small mammals (hereafter “small mammal”) predicted a priori by alternative hypotheses (mid‐domain effect [MDE], species–area relationship [SAR], energy, environmental stability, and habitat complexity]) proposed to explain the variation of diversity. We designed a standardized sampling scheme to trap small mammals at ten elevational bands across the entire elevational gradient on Yulong Mountain, southwest China. A total of 1,808 small mammals representing 23 species were trapped. We observed the hump‐shaped distribution pattern of the overall species richness along elevational gradient. Insectivores, rodents, large‐ranged species, and endemic species richness showed the general hump‐shaped pattern but peaked at different elevations, whereas the small‐ranged species and endemic species favored the decreasing richness pattern. The MDE and the energy hypothesis were supported, whereas little support was found for the SAR, the environmental stability hypothesis, and the habitat complexity. However, the primary driver(s) for richness patterns differed among the partitioning groups, with NDVI (the normalized difference vegetation index) and MDE being the most important variables for the total richness pattern. Species turnover for all small mammal groups increased with elevation, and it supported a decrease in community similarity with elevational distance. Our results emphasized for increased conservation efforts in the higher elevation regions of the Yulong Mountain.     

The role of environment and mid-domain effect on moth species richness along a tropical elevational gradient

Aim The biodiversity of geometrid moths (Lepidoptera) along a complete tropical elevational gradient was studied for the first time. The patterns are described, and the role of geometric constraints and environmental factors is explored. Location The study was carried out along the Barva Transect (10° N, 84° W), a complete elevational gradient ranging from 40 to 2730 m a.s.l. in Braulio Carrillo National Park, Costa Rica, and adjacent areas. Methods Moths were sampled manually in 2003 and 2004 at 12 rain forest sites using light ‘towers’, each with two 15 W ultraviolet fluorescent tubes. We used abundance‐based rarefaction, statistical estimation of true richness (Chao 1), geographically interpolated observed richness and Fisher's alpha as measures of local diversity. Results A total of 13,765 specimens representing 739 species were analysed. All four measures showed a hump‐shaped pattern with maxima between 500 and 2100 m elevation. The two subfamilies showed richness and diversity maxima at either lower (Ennominae) or higher (Larentiinae) elevation than Geometridae as a whole. Among the four environmental factors tested, relative humidity yielded the highest correlation over the transect with the rarefaction‐based richness estimates as well as with estimated true species richness of Geometridae as a whole and of Larentiinae, while rainfall explained the greatest variation of Ennominae richness. The elevational pattern of moth richness was discordant with both temperature and with tree species richness. A combination of all environmental factors in a stepwise multiple regression produced high values of r² in Geometridae. The potential effects of geometric constraints (mid‐domain effect, MDE) were investigated by comparing them with observed, interpolated richness. Overall, models fitted very well for Geometridae as a whole and for Ennominae, but less well for Larentiinae. Small‐ranged species showed stronger deviations from model predictions than large‐ranged species, and differed strikingly between the two subfamilies, suggesting that environmental factors play a more pronounced role for small‐ranged species. We hypothesize that small‐ranged species (at least of the Ennominae) may tend to be host specialists, whereas large‐ranged species tend to be polyphagous. Based on interpolated ranges, mean elevational range for these moths was larger with increasing elevation, in accordance with Rapoport's elevational rule, although sampling effects may have exaggerated this pattern. The underlying mechanism remains unknown because Rapoport's ‘rescue’ hypothesis could not explain the observed pattern. Conclusions The results clearly show that moth diversity shows a hump‐shaped pattern. However, remarkable variation exists with regard to taxon and range size. Both environmental and geometric factors are likely to contribute to the observed patterns.     

Altitudinal patterns of plant species richness on the Baekdudaegan Mountains, South Korea: mid-domain effect, area, climate, and Rapoport’s rule

We studied the altitudinal patterns of plant species richness and examined the effects of geometric constraints, area, and climatic factors on the observed richness patterns along the ridge of the Baekdudaegan Mountains, South Korea. Rapoport’s altitudinal rule was evaluated by examining the relationship between altitudinal range size and midpoint. We also examined the latitudinal effect on species richness. Plant data were collected from 1,100 plots along a 200–1,900 m altitudinal gradient along the ridge of the Baekdudaegan. A total of 802 plant species from 97 families and 342 genera were found. The altitudinal patterns of plant species richness along the ridge of the Baekdudaegan depicted distinctly hump-shaped patterns, although the absolute altitudes of the richness peaks vary somewhat among plant groups. While the mid-domain effect (MDE) was the most powerful explanatory variable in simple regression models, species richness was also associated with climatic factors, especially mean annual precipitation (MAP) and temperature (MAT) in multiple regression models. The relative importance of the MDE and climatic factors were different among plant groups. The MDE was more important for woody plants and for large-ranged species, whereas climatic factors were better predictors for total and herbaceous plants and for small-ranged species. Rapoport’s altitudinal rule and a latitudinal effect on species richness were not supported. Our study suggests that a combined interaction of the MDE and climatic factors influences species richness patterns along the altitudinal gradient of the Baekdudaegan Mountains, South Korea.     

Pteridophyte richness, climate and topography in the Iberian Peninsula: comparing spatial and nonspatial models of richness patterns

Aim To describe the spatial variation in pteridophyte species richness; evaluate the importance of macroclimate, topography and within‐grid cell range variables; assess the influence of spatial autocorrelation on the significance of the variables; and to test the prediction of the mid‐domain effect. Location The Iberian Peninsula. Methods We estimated pteridophyte richness on a grid map with c. 2500 km² cell size, using published geocoded data of the individual species. Environmental data were obtained by superimposing the grid system over isoline maps of precipitation, temperature, and altitude. Mean and range values were calculated for each cell. Pteridophyte richness was related to the environmental variables by means of nonspatial and spatial generalized least squares models. We also used ordinary least squares regression, where a variance partitioning was performed to partial out the spatial component, i.e. latitude and longitude. Coastal and central cells were compared to test the mid‐domain effect. Results Both spatial and nonspatial models showed that pteridophyte richness was best explained by a second‐order polynomial of mean annual precipitation and a quadratic elevation‐range term, although the relative importance of these two variables varied when spatial autocorrelation was accounted for. Precipitation range was weakly significant in a nonspatial multiple model (i.e. ordinary regression), and did not remain significant in spatial models. Richness is significantly higher along the coast than in the centre of the peninsula. Main conclusions Spatial autocorrelation affects the statistical significance of explanatory variables, but this did not change the biological interpretation of precipitation and elevation range as the main predictors of pteridophyte richness. Spatial and nonspatial models gave very similar results, which reinforce the idea that water availability and topographic relief control species richness in relatively high‐energy regions. The prediction of the mid‐domain effect is falsified.     

The environmental control of plant species density on a Himalayan elevation gradient

Aim In eastern Nepal, forests occupy an elevation gradient of 4000 m with bioclimatic zones from near tropical to alpine. Understorey plants and trees were censused to measure species density and identify patterns of ecological change. By sampling in a manner robust against spurious mid‐domain effects, I aim to identify biologically valid controls on species density. Location The study area consists of land below 4250 m elevation between 27.1 and 27.8° N latitude, 86.5 and 88.0° E longitude on the southern slopes of the Himalaya Range in eastern Nepal. Sampling sites are limited to intact, natural forest with relatively little human impact. Methods Team members counted species of understorey plants and trees ≥ 10 cm d.b.h. in 0.04 ha plots throughout the study area. In addition, basal area, leafing phenology and species composition were determined for the trees in each plot. Estimates of regional species density were compiled for successive 250 m elevation bands from 250 to 4250 m elevation. Species density trends were identified and compared with the expectations of O'Brien's [Journal of Biogeography 25 (1998) 379–398] climate‐based water–energy dynamics model. Results Stand basal area, tree leafing phenology and taxonomic composition (angiosperm vs. gymnosperm) show non‐random change with elevation. Understorey plant and tree species density both have a humped, unimodal trend with more species near the bottom of the gradient and fewest at the top. These trends are consistent with expected effects of the climatically active water and energy variables. After curve‐fitting, significant spatial structure in the residuals suggests that tree communities within the 1750–2250 m elevation range do not realize their climatic potential species richness. Main conclusions Neither mid‐domain effects nor biologically valid boundary effects like dispersal limitation explain the plant species density trends observed. Trends do fit a model in which species density is controlled by the same ‘active’ climatic variables that predict species richness on continental scales. Patterns of leafing phenology on the elevation gradient provide further support for the hypothesis of environmental control of species density. The productivity–diversity linkage that exists on continental scales may also apply on the smaller scale of a Himalayan elevation gradient. Human activity and possible competitive exclusion by Castanopsis tribuloides are the two best explanations for the observed decline in tree species density at 1750–2250 m elevation. Burning, lopping for fodder and livestock grazing might account for the decline, but this study does not assess the relative importance of these activities. The elevation richest for understorey plant and tree species (500–1500 m) also has the most severe reduction in forest cover. Local farmers deserve credit for sustaining plant biodiversity in forest enclaves, but further loss of forest at these elevations should be discouraged.     

Could temperature and water availability drive elevational species richness patterns? A global case study for bats

Aim A global meta‐analysis was used to elucidate a mechanistic understanding of elevational species richness patterns of bats by examining both regional and local climatic factors, spatial constraints, sampling and interpolation. Based on these results, I propose the first climatic model for elevational gradients in species richness, and test it using preliminary bat data for two previously unexamined mountains. Location Global data set of bat species richness along elevational gradients from Old and New World mountains spanning 12.5° S to 38° N latitude. Methods Bat elevational studies were found through an extensive literature search. Use was made only of studies sampling  70% of the elevational gradient without significant sampling biases or strong anthropogenic disturbance. Undersampling and interpolation were explicitly examined with three levels of error analyses. The influence of spatial constraints was tested with a Monte Carlo simulation program, Mid‐Domain Null. Preliminary bat species richness data sets for two test mountains were compiled from specimen records from 12 US museum collections. Results Equal support was found for decreasing species richness with elevation and mid‐elevation peaks. Patterns were robust to substantial amounts of error, and did not appear to be a consequence of spatial constraints. Bat elevational richness patterns were related to local climatic gradients. Species richness was highest where both temperature and water availability were high, and declined as temperature and water availability decreased. Mid‐elevational peaks occurred on mountains with dry, arid bases, and decreasing species richness occurred on mountains with wet, warm bases. A preliminary analysis of bat richness patterns on elevational gradients in western Peru (dry base) and the Olympic Mountains, WA (wet base), supported the predictions of the climate model. Main conclusions The relationship between species richness and combined temperature and water availability may be due to both direct (thermoregulatory constraints) and indirect (food resources) factors. Abundance was positively correlated with species richness, suggesting that bat species richness may also be related to productivity. The climatic model may be applicable to other taxonomic groups with similar ecological constraints, for instance certain bird, insect and amphibian clades.     

Diversity of Eastern North American Ant Communities along Environmental Gradients

Studies of species diversity patterns across regional environmental gradients seldom consider the impact of habitat type on within-site (alpha) and between-site (beta) diversity. This study is designed to identify the influence of habitat type across geographic and environmental space, on local patterns of species richness and regional turnover patterns of ant diversity in the northeastern United States. Specifically, I aim to 1) compare local species richness in paired open and forested transects and identify the environmental variables that best correlate with richness; and 2) document patterns of beta diversity throughout the region in both open and forested habitat. I systematically sampled ants at 67 sites from May to August 2010, spanning 10 degrees of latitude, and 1000 meters of elevation. Patterns of alpha and beta diversity across the region and along environmental gradients differed between forested and open habitats. Local species richness was higher in the low elevation and warmest sites and was always higher in open habitat than in forest habitat transects. Richness decreased as temperature decreased or elevation increased. Forested transects show strong patterns of decreasing dissimilarity in species composition between sites along the temperature gradient but open habitat transects did not. Maximum temperature of the warmest month better predicted species richness than either latitude or elevation. I find that using environmental variables as key predictors of richness yields more biologically relevant results, and produces simpler macroecological models than commonly used models which use only latitude and elevation as predictors of richness and diversity patterns. This study contributes to the understanding of mechanisms that structure the communities of important terrestrial arthropods which are likely to be influenced by climatic change.     

Continental and regional ranges of North American mammals: Rapoport's rule in real and null worlds

Aim To assess the relationship between species richness and distribution within regions arranged along a latitudinal gradient we use the North American mammalian fauna as a study case for testing theoretical models. Location North America. Methods We propose a conceptual framework based on a fully stochastic mid‐domain model to explore geographical patterns of range size and species richness that emerge when the size and position of species ranges along a one‐dimensional latitudinal gradient are randomly generated. We also analyse patterns for the mammal fauna of North America by comparing empirical results from a biogeographical data base with predictions based on randomization null models. Results We confirmed the validity of Rapoport's rule for the mammals of North America by documenting gradients in the size of the continental ranges of species. Additionally, we demonstrated gradients of mean regional range size that parallel those of continental range. Our data also demonstrated that mean range size, measured both as a continental or a regional variable, is significantly correlated with the geographical pattern in species richness. All these patterns deviated sharply from null models. Main conclusions Rapoport's statement of an areographic relationship between species distribution and richness is highly relevant in modern discussions about ecological patterns at the geographical scale.     

Global analysis of reptile elevational diversity

Aim Latitudinal‐ and regional‐scale studies of reptile diversity suggest a predominant temperature effect, unlike many other vertebrate richness patterns which tend to be highly correlated with both temperature and water variables. Here I examine montane gradients in reptile species richness with separate analyses of snakes and lizards from mountains around the world to assess a predominant temperature effect and three additional theories of diversity, including a temperature–water effect, the species–area effect and the mid‐domain effect (MDE). Location Twenty‐five elevational gradients of reptile diversity from temperate, tropical and desert mountains in both hemispheres, spanning 10.3° N to 46.1° N. Methods Elevational gradients in reptile diversity are based on data from the literature. Of the 63 data sets found or compiled, only those with a high, unbiased sampling effort were used in analyses. Twelve predictions and three interactions of diversity theory were tested using nonparametric statistics, linear regressions and multiple regression with the Akaike information criterion (AIC). Results Reptile richness and, individually, snake and lizard richness on mountains followed four distinct patterns: decreasing, low‐elevation plateaus, low‐elevation plateaus with mid‐elevation peaks, and mid‐elevation peaks. Elevational reptile richness was most strongly correlated with temperature. The temperature effect was mediated by precipitation; reptile richness was more strongly tied to temperature on wet gradients than on arid gradients. Area was a secondary factor of importance, whereas the MDE was not strongly associated with reptile diversity on mountains. Main conclusions Reptile diversity patterns on mountains did not follow the predicted temperature–water effect, as all diversity patterns were found on both wet and dry mountains. But the influence of precipitation on the temperature effect most likely reflects reptiles' use of radiant heat sources (sunning opportunities) that are more widespread on arid mountains than wet mountains due to lower humidity, sparser vegetation and less cloud cover across low and intermediate elevations.