The relative roles of environment and history as controls of tree species composition and richness in Europe

Aim This study investigates the determinants of European‐scale patterns in tree species composition and richness, addressing the following questions: (1) What is the relative importance of environment and history? History refers to lasting effects of past large‐scale events and time‐dependent cumulative effects of ongoing processes, notably dispersal limited range dynamics. (2) Among the environmental determinants, what is the relative importance of climate, soils, and forest cover? (3) Do the answers to questions 1 and 2 differ between conifers and Fagales, the two major monophyletic groups of European trees? Location The study area comprises most of Europe (34° N–72° N and 11° W–32° E). Methods Atlas data on native distributions of 54 large tree species at 50 × 50 km resolution were linked with climatic, edaphic, and forest cover maps in a geographical information system. Unconstrained (principal components analysis using Hellinger distance transformation and detrended correspondence analysis) and constrained ordinations (redundancy analysis using Hellinger distance transformation and canonical correspondence analysis) and multiple linear regressions were used to investigate the determinants of species composition and species richness, respectively. History is expected to leave its mark as broad spatial patterns and was represented by the nine spatial terms of a cubic trend surface polynomial. Results The main floristic pattern identified by all ordinations was a latitude‐temperature gradient, while the lower axes corresponded mostly to spatial variables. Partitioning the floristic variation using constrained ordinations showed the mixed spatial‐environmental and pure spatial fractions to be much greater than the pure environmental fraction. Biplots, forward variable selection, and partial analyses all suggested climatic variables as more important floristic determinants than forest cover or soil variables. Tree species richness peaked in the mountainous regions of East‐Central and Southern Europe, except the Far West. Variation partitioning of species richness found the mixed spatial‐environmental and pure spatial fractions to be much greater than the pure environmental fraction for all species combined and Fagales, but not for conifers. The scaled regression coefficients indicated climate as a stronger determinant of richness than soils or forest cover. While the dominant patterns were similar for conifers and Fagales, conifers exhibited less predictable patterns overall, a smaller pure spatial variation fraction relative to pure environmental fraction, and a greater relative importance of climate; all differences being more pronounced for species richness than for species composition. Main conclusions The analyses suggest that history is at least as important as current environment in controlling species composition and richness of European trees, with the exception of conifer species richness. Strong support for interpreting the spatial patterns as outcomes of historical processes, notably dispersal limitation, came from the observation that many European tree species naturalize extensively outside their native ranges. Furthermore, it was confirmed that climate predominates among environmental determinants of distribution and diversity patterns at large spatial scales. Finally, the particular patterns exhibited by conifers probably reflect greater environmental specialization and greater human impact. These findings warn against expecting the European tree flora to be able track fast future climate changes on its own.     

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.     

Global species–energy relationship in forest plots: role of abundance,temperature and species climatic tolerances

Aim To evaluate the strength of evidence for hypotheses explaining the relationship between climate and species richness in forest plots. We focused on the effect of energy availability which has been hypothesized to influence species richness: (1) via the effect of productivity on the total number of individuals (the more individuals hypothesis, MIH); (2) through the effect of temperature on metabolic rate (metabolic theory of biodiversity, MTB); or (3) by imposing climatic limits on species distributions. Location Global. Methods We utilized a unique ‘Gentry‐style’ 370 forest plots data set comprising tree counts and individual stem measurements, covering tropical and temperate forests across all six forested continents. We analysed variation in plot species richness and species richness controlled for the number of individuals by using rarefaction. Ordinary least squares (OLS) regression and spatial regressions were used to explore the relative performance of different sets of environmental variables. Results Species richness patterns do not differ whether we use raw number of species or number of species controlled for number of individuals, indicating that number of individuals is not the proximate driver of species richness. Productivity‐related variables (actual evapotranspiration, net primary productivity, normalized difference vegetation index) perform relatively poorly as correlates of tree species richness. The best predictors of species richness consistently include the minimum temperature and precipitation values together with the annual means of these variables. Main conclusion Across the world's forests there is no evidence to support the MIH, and a very limited evidence for a prominent role of productivity as a driver of species richness patterns. The role of temperature is much more important, although this effect is more complex than originally assumed by the MTB. Variation in forest plot diversity appears to be mostly affected by variation in the minimum climatic values. This is consistent with the ‘climatic tolerance hypothesis’ that climatic extremes have acted as a strong constraint on species distribution and diversity.     

Determinants of diversity in a naturally fragmented landscape: humid montane forest avifaunas of Mesoamerica

We used a published data set summarizing avifaunas of 31 montane patches of humid forest in Mesoamerica lo analyze avian distributions with respect to site characteristics. This forest type was originally widespread in the lowlands, and became restricted to mountains during Pleistocene climatic changes. Hierarchical partitioning. a recently developed regression procedure, was used to examine independent factor effects. Total species richness, richness of Mesoamerican endemic species, richness of narrowly endemic species, and richness of habitat specialists were considered separately, each analyzed at three spatial scales. For total richness and Mesoamerican endemics, regional-level variables, notably latitude. were most influential. Narrow endemics exhibited more complex patterns, driven by foci both in western Mexico and in Costa Rica and western Panama. Historical factors are suggested to have contributed to this latitudinal pattern, such that the isthmuses of Tehuantepec and Panama acted as barriers to range expansion and peninsular effects catalyzed speciation. elevating numbers of endemic species. In contrast to many anthropogenic fragmentation studies, area and other local-scale patch attributes had little influence on avifaunas. This discrepancy may be related lo fundamental differences in spatial and temporal scaling, with patterns uncovered herein more indicative of long-term community processes.     

Geographical patterns of community‐based tree species richness in Chinese mountain forests: the effects of contemporary climate and regional history

The relationship between climate/productivity and historical/regional contingency and their relative influence on geographical patterns of species richness (GPSR) are still unresolved. Based on field data from 1494 plots from forests on 63 mountains across China, we document the GPSR for forest communities. Regression tree and generalized linear models were used to explore the discreteness and gradient of the distribution of tree species richness (α‐diversity), and to estimate the correlations of climate, historical floristic region, and local habitat with species richness. The collinearity between climatic variables and region were further disentangled; and the spatial autocorrelation in the patterns of α‐diversity and the residuals of alternative predictive models were compared. Overall, 75% of variation in plot‐based α‐diversity of trees was accounted for by all variables included, and about 66.5%, 64.5% and 27.9% by climate, region, and local habitat respectively. Importantly, the explanatory power of these variables differed in particular for coniferous, deciduous broadleaved and evergreen broadleaved species. Ambient temperature was more important for α‐diversity of trees than were the other climatic variables across China. Spatial autocorrelation in the pattern of α‐diversity could be accounted for mainly by spatial variation climate. The concordance between tree α‐diversity, historical flora, contemporary climate, and Quaternary climate change mode suggests the climate/productivity and historical/regional contingency both contribute to the GPSR in a complimentary manner. Taken together, our results provide unique evidence to link of the effects of contemporary climate and historical climate change on species richness across scales.     

Epiphytic bryophyte diversity and range distributions along an elevational gradient in Marojejy,Madagascar

Describing spatial variation in species richness and understanding its links to ecological mechanisms are complementary approaches for explaining geographical patterns of richness. The study of elevational gradients holds enormous potential for understanding the factors underlying global diversity. This paper investigates the pattern of species richness and range-size distribution of epiphytic bryophytes along an elevational gradient in Marojejy National Park, northeast Madagascar. The main objectives are to describe bryophyte species composition and endemism in Marojejy National Park, to describe the species richness and distribution patterns of epiphytic bryophytes along an elevational gradient from 250 m to 2050 m and to evaluate the explanatory value of environmental variables for the observed patterns. Bryophyte samples were collected following a nested design with four hierarchical levels: elevational belts, plots, quadrats, and microplots. In total, 254 epiphytic bryophyte species were recorded, comprising 157 liverworts and 97 mosses. Twenty-three of these are endemic to Madagascar. Species richness exhibits a hump-shaped pattern along the elevational gradient, peaking at 1,250 m. Eighty-seven percent of the total recorded species have a range distribution lower than 1,000 m, at which point 36% are restricted to these single elevations. Our results suggest that mean temperature, relative humidity, and vapor pressure deficit play important roles in shaping the richness pattern observed in this study. While the liverwort richness pattern did not correlate to vapor pressure deficit and responded only weakly to relative humidity, the richness pattern shown by mosses correlates well with mean temperature, relative humidity, and vapor pressure deficit.     

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.     

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.     

Species richness of epiphytic bryophytes: drivers across scales on the edge of the Mediterranean

Spatial variation in species richness is one of the most frequently studied topics on macroecology. However, the relative importance of the factors affecting richness across scales and their influence on some groups of small‐sized organisms, such as bryophytes, remain unclear. We evaluate the relative importance of biogeographic region, climate, topography, forest structure and abundance in shaping epiphytic bryophyte richness at both local (forest) and sample (trunk) scale on the boundary between the Atlantic and Mediterranean regions in NW Spain. For that purpose we used simple, multiple and partial regressions, hierarchical partitioning and partial least squares path analyses. Although climatic variables related to water availability during spring and summer were the most important predictors of bryophyte richness, their effects were moderated by winter temperature. Abundance, in contrast, was mostly related to forest structure. Biogeographic region was not significantly related to richness. Interestingly, forest richness was the best predictor of trunk richness. Our results highlight the importance of seasonal distribution of rainfall and temperatures and support that the richness of bryophyte communities is constrained by mesoscale climatic factors, in particular the interplay between water and energy availability. In contrast, abundance seems to be controlled by habitat characteristics. We also detected a strong top‐down structure between both scales of measurement evidencing a scaling down of the climatic effect: richness at the sample scale is controlled mainly by local richness and local richness is in turn controlled by climate, so mesoscale climatic gradients are indirectly limiting richness at the smallest scale.     

Structures determining bryophyte species richness in a managed forest landscape in boreo-nemoral Europe

Forest patches with high biological value are protected as woodland key habitats (WKH), which are identified by the presence of forest structures and indicator species. However, management for conservation needs to consider also managed forests as habitats for species. In this respect, there is a need to set quantitative targets for species and structures at different landscape scales. Due to non-intensive methods of forest management used prior to 1940 in Latvia, it might be expected that large areas of forest have developed structures that can support many species characteristic of natural forests. The aim of the study was to create a model that best described the richness of bryophyte species that are characteristic of natural forests, using forest structures as explanatory factors. The structures and bryophyte communities on living trees and coarse woody debris (CWD) were described in plots along transects blindly placed in areas dominated by State forests under commercial management. Explanatory variables related to tree species composition and tree size explained 54% of the variation in WKH indicator species richness on living trees. The best explanatory factors were maximum diameter of deciduous tree species and CWD. Low richness of total bryophyte and indicator species was found on dead wood, and the amount of variation in bryophyte species richness on CWD explained by explanatory variables was low. The study indicates the importance of deciduous tree substrate in managed forests in maintaining the spatial continuity of epiphytic species diversity. However, the forests in the managed forest landscape did not support high diversity of epixylic species, even in the WKHs, due to low diversity of suitable dead wood substrate.     

Predictors of mammal species richness in KwaZulu-Natal,South Africa

The management of multi-functional landscapes warrants better knowledge of environment-richness associations at varying disturbance levels and habitat gradients. Intensive land-use patterns for agricultural purposes lead to fragmentation of natural habitat resulting in biodiversity loss that can be measured using landscape metrics to assess mammalian richness. Since carnivores and herbivores are likely to show different responses to disturbance, we calculated carnivore, non-carnivore, and total mammal species richness from camera surveys using a first order Jackknife Estimator. Richness was compared along a habitat gradient comprising coastal forest, Acacia thicket, and highland in KwaZulu-Natal, South Africa. We used standardized OLS regression models to identify climatic and disturbance variables, and landscape metrics as predictors of species richness. The estimated total and non-carnivore species richness were highest in coastal forest, while carnivore species richness was highest in highland followed by coastal forest and Acacia thicket. Average monthly maximum temperature was a significant predictor of all richness groups, and precipitation of the wettest month and isothermality determined total and non-carnivore species richness, respectively. These climatic variables possibly limit species distribution because of physiological tolerance of the species. Total mammal richness was determined by mean shape (+) and habitat division (−) while diversity (+) and patch richness (−) explained carnivore species richness. Mean shape index (+) influenced non-carnivore richness. However, habitat division and patch richness negatively influenced total mammal richness. Though habitat patch size and contiguity had a weak positive prediction, these metrics demonstrated the importance of habitat connectivity for maintaining mammal richness. The identification of these climatic and landscape patterns is important to facilitate future landscape management for mammal conservation in forest-mosaics.     

Determinants of species richness, endemism and turnover in European longhorn beetles

This study assessed the diversity patterns of a large family of beetles, Cerambycidae, in Europe and tested the following hypotheses: 1) richness gradients of this hyperdiverse taxon are driven by water and energy variables; 2) endemism is explained by the same factors, but variation between areas also reflects post‐glacial re‐colonization processes; and 3) faunal composition is determined by the same climatic variables and, therefore, beta diversity (species turnover) is related to richness gradients. Species richness, endemism and beta diversity were modelled using inventories of 37 European territories, built from a database containing the distributions of 609 species. Area, spatial position, and nine topographical and climatic variables were used as predictors in regression and constrained analysis of principal coordinates modelling. Species richness was mostly explained by a temperature gradient, which produced a south‐to‐north decreasing richness gradient. Endemism followed the same pattern, but was also determined by longitudinal variation, peaking in the southwestern and southeastern corners of the continent. Faunal turnover was explained by an important purely spatial pattern and a spatially structured environmental gradient. Thus, contrary to other groups, cerambycid richness was mostly explained by environmental energy, but not by water availability. Endemism was concentrated in the Iberian and Greek peninsulas, but not in Italy. Thus, the latter area may have been the major source of post‐glacial re‐colonization for European longhorn beetles or, otherwise, a poor refuge during glaciations. Turnover patterns were independent of the richness gradient, because northern faunas are nested in southern ones. Turnover, in contrast to richness, was driven by both the independent effects of climate and geographic constraints that might reflect dispersal limitation or stochastic colonization events, suggesting that richness gradients are more environmentally deterministic phenomena than turnover patterns.     

Tree diversity patterns in successive vegetation types along an elevation gradient in the Mountains of Eastern Mexico

Tree species richness changes along elevation gradients in response to underlying environmental conditions. Our hypothesis was that richness is associated with climatic variables and decreases with elevation. The objective was to identify trends in species, genus and family richness, diversity and vegetation structure in relation to climate variables along an elevation gradient with successive types of forest in Veracruz, Mexico. Trees were identified and measured in 0.1 ha at 15 sites located from 140 to 4000 m a.s.l. Generalized linear models were used to fit richness, diversity, basal area and density as a function of elevation; the best model was selected using Akaike’s Information Criterion. Multivariate analyses were used to explore climatic variables associated to composition of groups of sites along the gradient. Along the entire elevation gradient, species, genus and family richness decreased unimodally, and diversity decreased monotonically. Richness was positively correlated with temperature but not with precipitation. Basal area increased monotonically and highest basal area was associated with high humidity and certain tree species (Quercus and Abies). Ordinations indicated three groups of sites: lower elevation dry forest associated with temperature seasonality, mid-elevation cloud forest associated with precipitation-related variables, and coniferous forest at the top of the gradient associated with elevation. Our study shows that different plant communities are associated with certain climatic conditions and harbour different tree species, genera and families. The results support the hypothesis that species richness is associated with climate, and decreases with elevation.     

Spatial patterns of plant richness across treeline ecotones in the Pyrenees reveal different locations for richness and tree cover boundaries

Aim  To forecast the responses of alpine flora to the expected upward shift of treeline ecotones due to climatic warming, we investigated species richness patterns of vascular plants at small spatial scales across elevational transects.
Location  Richness patterns were assessed at local scales along the elevational gradient in two undisturbed treeline ecotones and one disturbed treeline ecotone in the Spanish Pyrenees.
Methods  We placed a rectangular plot (0.3–0.4 ha) in each treeline ecotone. We estimated and described the spatial patterns of plant richness using the point method and Moran's I correlograms. We delineated boundaries based on plant richness and tree cover using moving split windows and wavelet analysis. Then, to determine if floristic and tree cover boundaries were spatially related, overlap statistics were used.
Results  Plant richness increased above the forest limit and was negatively related to tree cover in the undisturbed sites. The mean size of richness patches in one of these sites was 10–15 m. Moving split windows and wavelets detected the sharpest changes in plant richness above the forest limit at both undisturbed sites. Most tree cover and plant richness boundaries were not spatially related.
Main conclusions  The upslope decrease of tree cover may explain the increase of plant richness across alpine treeline ecotones. However, the detection of abrupt richness boundaries well above the forest limit indicates the importance of local environmental heterogeneity to explain the patterns of plant richness at smaller scales. We found highly diverse microsites dominated by alpine species above the forest limit, which should be monitored to describe their response to the predicted upward shift of forests.     

Contrasting effects of biotic interactions on richness and distribution of vascular plants,bryophytes and lichens in an arctic–alpine landscape

Biotic interactions may strongly affect the distribution of individual species and the resulting patterns of species richness. However, the impacts can vary depending on the species or taxa examined, suggesting that the influences of interactions on species distributions and diversity are not always straightforward and can be taxon-contingent. The aim of this study was therefore to examine how the importance of biotic interactions varies within a community. We incorporated three biotic predictors (cover of the dominant vascular species) into two correlative species richness modelling frameworks to predict spatial variation in the number of vascular plants, bryophytes and lichens in arctic–alpine Fennoscandia, in N Europe. In addition, predictions based on single-species distribution models were used to determine the nature of the impact (negative vs. positive outcome) of the three dominant species on individual vascular plant, bryophyte and lichen species. Our results suggest that biotic variables can be as important as abiotic variables, but their relative contributions in explaining the richness of sub-dominant species vary among dominant species, species group and the modelling framework implemented. Similarly, the impacts of biotic interactions on individual species varied among the three species groups and dominant species, with the observed patterns partly reflecting species’ biogeographic range. Our study provides additional support for the importance of biotic interactions in modifying arctic–alpine biodiversity patterns and highlights that the impacts of interactions are not constant across taxa or biotic drivers. The influence of biotic interactions, including the taxon contingency and range-based impacts, should therefore be accounted for when developing biodiversity forecasts.     

Species distribution modelling as a macroecological tool: a case study using New World amphibians

Although species distribution modelling (SDM) is widely accepted among the scientific community and is increasingly used in ecology, conservation biology and biogeography, methodological limitations generate potential problems for its application in macroecology. Using amphibian species richness in North and South America, we compare species richness patterns derived from SDM maps and ‘expert’ maps to evaluate if: 1) richness patterns derived from SDM are biased toward climate‐based explanations for diversity when compared to expert maps, since SDM methods are typically based on climatic variables; and 2) SDM is a reliable tool for generating richness maps in hyperrich regions where point occurrence data are limited for many species. We found that although three widely used SDM methods overestimated amphibian species richness in grid cells when compared to expert richness maps in both North and South America due to systematic overestimation of range sizes, diversity gradients were reasonably robust at broad scales. Further, climatic variables statistically explained patterns of richness at similar levels among the different richness sources, although climatic relationships were stronger in the much better known North America than in South America. We conclude that in the face of the high deforestation rates coupled with incomplete data on species distributions, especially in the tropics, SDM represents a useful macroecological tool for investigating broad‐scale richness patterns and the dynamics between species richness and climate.     

Mapping forest vegetation patterns in an Atlantic-Mediterranean transitional area by integration of ordination and geostatistical techniques

Aims Forest vegetation variability may be explained by the complex interplay among several spatial structuring factors, including climate and topography. We modelled the spatial variability of forest vegetation assemblages and significant environmental variables along a complex environmental gradient or coenocline to produce a detailed cartographic database portraying the distribution of forests along it.Methods We combined an analysis of ordination coenoclines with kriging over 772 field data plots from the third Spanish National Forest Inventory in an Atlantic–Mediterranean transitional area (northern Spain).Important findings The best fitted empirical semivariogram revealed a strong spatial structure of forest species composition along the complex environmental gradient considered (the climatic–topographic gradient from north to south). The steady and gradual increase of semivariance with a marked lag distance indicates a gradual turnover of forest assemblages according to the climatic–topographic variations (regional or local). Two changes in the slope of the semivariogram suggest the existence of two different scales of spatial variation. The interpolation map by Kriging of forest vegetation assemblages along the main coenocline shows a clear spatial distribution pattern of trees and shrubs in accordance with the spatial variation of significant environmental variables. We concluded that the multivariate geostatistical approach is a suitable technique for spatial analysis of forest systems employing data from national forest inventories based on a regular network of field plots. The development of an assortment of maps describing changes in vegetation assemblages and variation in environmental variables is expected to be a suitable tool for an integrated forest management and planning.     

Fern species richness along a central Himalayan elevational gradient, Nepal

Aim The study explores fern species richness patterns along a central Himalayan elevational gradient (100–4800 m a.s.l.) and evaluates factors influencing the spatial increase and decrease of fern richness. Location The Himalayas stretch from west to east by 20°, i.e. 75–95° east, and Nepal is located from 80 to 88° east in this range. Methods We used published data of the distribution of ferns and fern allies to interpolate species elevational ranges. Defining species presence between upper and lower elevation limit is the basis for richness estimates. The richness pattern was regressed against the total number of rainy days, and gradients that are linearly related to elevation, such as length of the growing season, potential evapotranspiration (PET, energy), and a moisture index (MI = PET/mean annual rainfall). The regressions were performed by generalized linear models. Results A unimodal relationship between species richness and elevation was observed, with maximum species richness at 2000 m. Fern richness has a unimodal response along the energy gradients, and a linear response with moisture gradients. Main conclusions The study confirms the importance of moisture on fern distributions as the peak coincides spatially with climatic factors that enhance moisture levels; the maximum number of rainy days and the cloud zone. Energy‐related variables probably control species richness directly at higher elevations but at the lower end the effect is more probably related to moisture.