Assessment of the relationships of geographic variation in species richness to climate and landscape variables within and among lineages of North American freshwater fishes

Aim Geographic variation in species richness is a well‐studied phenomenon. However, the unique response of individual lineages to environmental gradients in the context of general patterns of biodiversity across broad spatial scales has received limited attention. The focus of this research is to examine relationships between species richness and climate, topographic heterogeneity and stream channel characteristics within and among families of North American freshwater fishes. Location The United States and Canada. Methods Distribution maps of 828 native species of freshwater fishes were used to generate species richness estimates across the United States and Canada. Variation in species richness was predicted using spatially explicit models incorporating variation in climate, topography and/or stream channel length and stream channel diversity for all 828 species as well as for the seven largest families of freshwater fishes. Results The overall gradient of species richness in North American freshwater fishes is best predicted by a model incorporating variables describing climate and topography. However, the response of species richness to particular climate or landscape variables differed among families, with models possessing the highest predictive ability incorporating data on climate, topography and/or stream channel characteristics within a region. Main conclusions The correlations between species richness and abiotic variables suggest a strong influence of climate and physical habitat on the structuring of regional assemblages of North American freshwater fishes. However, the relationship between these variables and species richness varies among families, suggesting the importance of phylogenetic constraints on the regulation of geographic distributions of species.     

The future representativeness of Madagascar's protected area network in the face of climate change

With many species predicted to respond to a changing climate by shifting their distribution to climatically suitable areas, the effectiveness of static protected areas (PAs) is in question. The Madagascan PA network area has quadrupled over the past 15 years, and, although conservation planning techniques were employed to prioritise suitable areas for protection during this process, climate change impacts were not considered. We make use of species distribution models for 750 Madagascan vertebrate species to assess the potential impacts of climate change on (1) species richness across Madagascar, (2) species gain, loss and turnover in Madagascar's PAs and (3) PA network representativeness. Results indicate that Madagascar is predicted to experience substantial shifts in species richness, with most PAs predicted to experience high rates of species turnover. Provided there are no barriers to species movements, the representativeness of the current PA network will remain high for the species that are predicted to survive changes in climate by 2070, suggesting that little benefit will be gained from establishing new PAs. However, this rests on the assumption of mobility through areas currently characterised by fragmentation and anthropogenic activity, something that will require considerable expansion in conservation efforts in order to achieve.     

Climate and Species Richness Predict the Phylogenetic Structure of African Mammal Communities

We have little knowledge of how climatic variation (and by proxy, habitat variation) influences the phylogenetic structure of tropical communities. Here, we quantified the phylogenetic structure of mammal communities in Africa to investigate how community structure varies with respect to climate and species richness variation across the continent. In addition, we investigated how phylogenetic patterns vary across carnivores, primates, and ungulates. We predicted that climate would differentially affect the structure of communities from different clades due to between-clade biological variation. We examined 203 communities using two metrics, the net relatedness (NRI) and nearest taxon (NTI) indices. We used simultaneous autoregressive models to predict community phylogenetic structure from climate variables and species richness. We found that most individual communities exhibited a phylogenetic structure consistent with a null model, but both climate and species richness significantly predicted variation in community phylogenetic metrics. Using NTI, species rich communities were composed of more distantly related taxa for all mammal communities, as well as for communities of carnivorans or ungulates. Temperature seasonality predicted the phylogenetic structure of mammal, carnivoran, and ungulate communities, and annual rainfall predicted primate community structure. Additional climate variables related to temperature and rainfall also predicted the phylogenetic structure of ungulate communities. We suggest that both past interspecific competition and habitat filtering have shaped variation in tropical mammal communities. The significant effect of climatic factors on community structure has important implications for the diversity of mammal communities given current models of future climate change.     

The fate of European breeding birds under climate,land‐use and dispersal scenarios

Many species have already shifted their distributions in response to recent climate change. Here, we aimed at predicting the future breeding distributions of European birds under climate, land‐use, and dispersal scenarios. We predicted current and future distributions of 409 species within an ensemble forecast framework using seven species distribution models (SDMs), five climate scenarios and three emission and land‐use scenarios. We then compared results from SDMs using climate‐only variables, habitat‐only variables or both climate and habitat variables. In order to account for a species’ dispersal abilities, we used natal dispersal estimates and developed a probabilistic method that produced a dispersal scenario intermediate between the null and full dispersal scenarios generally considered in such studies. We then compared results from all scenarios in terms of future predicted range changes, range shifts, and variations in species richness. Modeling accuracy was better with climate‐only variables than with habitat‐only variables, and better with both climate and habitat variables. Habitat models predicted smaller range shifts and smaller variations in range size and species richness than climate models. Using both climate and habitat variables, it was predicted that the range of 71% of the species would decrease by 2050, with a 335 km median shift. Predicted variations in species richness showed large decreases in the southern regions of Europe, as well as increases, mainly in Scandinavia and northern Russia. The partial dispersal scenario was significantly different from the full dispersal scenario for 25% of the species, resulting in the local reduction of the future predicted species richness of up to 10%. We concluded that the breeding range of most European birds will decrease in spite of dispersal abilities close to a full dispersal hypothesis, and that given the contrasted predictions obtained when modeling climate change only and land‐use change only, both scenarios must be taken into consideration.     

Species richness of migratory birds is influenced by global climate change

Aim  Global climate change is increasingly influencing ecosystems. Long-term effects on the species richness and composition of ecological communities have been predicted using modelling approaches but, so far, hardly demonstrated in the field. Here, we test whether changes in the composition of bird communities have been influenced by recent climate change.
Location  Europe.
Methods  We focus on the proportion of migratory and resident bird species because these groups are expected to respond differently to climatic change. We used the spatial relationship between climatic factors and bird communities in Europe to predict changes in 21 European bird communities under recent climate change.
Results  Observed changes corresponded significantly to predicted changes and could not be explained by the effects of spatial autocorrelation. Alternative factors such as changes in land use were tested in a first approximation as well but no effects were found.
Main conclusions  This study demonstrates that global climate change has already influenced the species richness and composition of European bird communities.     

Climate induced increases in species richness of marine fishes

Climate change has been predicted to lead to changes in local and regional species richness through species extinctions and latitudinal ranges shifts. Here, we show that species richness of fish in the North Sea, a group of ecological and socio-economical importance, has increased over a 22-year period and that this rise is related to higher water temperatures. Over eight times more fish species displayed increased distribution ranges in the North Sea (mainly small-sized species of southerly origin) compared with those whose range decreased (primarily large and northerly species). This increase in species richness can be explained from the fact that fish species richness in general decreases with latitude. This observation confirms that the interaction between large-scale biogeographical patterns and climate change may lead to increasing species richness at temperate latitudes.     

How much do we overestimate future local extinction rates when restricting the range of occurrence data in climate suitability models?

Climate suitability models are used to make projections of species’ potential future distribution under climate change. When studying the species richness with such modeling methods, the extent of the study range is of particular importance, especially when the full range of occurrence is not considered for some species, often because of geographical or political limits. Here we examine biases induced by the use of range‐restricted occurrence data on predicted changes in species richness and predicted extinction rates, at study area margins. We compared projections of future suitable climate space for 179 bird species breeding in Iberia and North Africa (27 of them breeding only in North Africa though potential colonizers in Europe), using occurrence data from the full Western Palaearctic (WP) species range and from the often‐considered European‐restricted range. Current and future suitable climatic spaces were modeled using an ensemble forecast technique applied to five general circulation models and three climate scenarios, with eight climatic variables and eight modeling techniques. The use of range‐restricted compared to the full WP occurrence data of a species led to an underestimate of its suitable climatic space. The projected changes in species richness across the focus area (Iberia) varied considerably according to the occurrence data we used, with higher local extinction rates with European‐restricted data (on average 38 vs 12% for WP data). Modeling results for species currently breeding only in North Africa revealed potential colonization of the Iberian Peninsula (from a climatic point of view), which highlights the necessity to consider species outside the focus area if interested in forecasted changes in species richness. Therefore, the modeling of current and future species richness can lead to misleading conclusions when data from a restricted range of occurrence is used. Consequently, climate suitability models should use occurrence data from the complete distribution range of species, or at least within biogeographical areas.     

Genes on the edge: A framework to detect genetic diversity imperiled by climate change

Ongoing global warming is disrupting several ecological and evolutionary processes, spanning different levels of biological organization. Species are expected to shift their ranges as a response to climate change, with relevant implications to peripheral populations at the trailing and leading edges. Several studies have analyzed the exposure of species to climate change but few have explored exposure at the intraspecific level. We introduce a framework to forecast exposure to climate change at the intraspecific level. We build on existing methods by combining correlative species distribution models, a model of species range dynamics, and a model of phylogeographic interpolation. We demonstrate the framework by applying it to 20 Iberian amphibian and reptile species. Our aims were to: (a) identify which species and intraspecific lineages will be most exposed to future climate change; (b) test if nucleotide diversity at the edges of species ranges are significantly higher or lower than on the overall range; and (c) analyze if areas of higher species gain, loss, and turnover coincide with those predicted for lineages richness and nucleotide diversity. We found that about 80% of the studied species are predicted to contract their range. Within each species, some lineages were predicted to contract their range, while others were predicted to maintain or expand it. Therefore, estimating the impacts of climate change at the species level only can underestimate losses at the intraspecific level. Some species had significant high amount of nucleotide at the trailing or leading edge, or both, but we did not find a consistent pattern across species. Spatial patterns of species richness, gain, loss, and turnover were fairly concurrent with lineages richness and nucleotide diversity. Our results support the need for increased attention to intraspecific diversity regarding monitoring and conservation strategies under climate change.     

Climate‐based empirical models show biased predictions of butterfly communities along environmental gradients

A better understanding of the factors that mould ecological community structure is required to accurately predict community composition and to anticipate threats to ecosystems due to global changes. We tested how well stacked climate‐based species distribution models (S‐SDMs) could predict butterfly communities in a mountain region. It has been suggested that climate is the main force driving butterfly distribution and community structure in mountain environments, and that, as a consequence, climate‐based S‐SDMs should yield unbiased predictions. In contrast to this expectation, at lower altitudes, climate‐based S‐SDMs overpredicted butterfly species richness at sites with low plant species richness and underpredicted species richness at sites with high plant species richness. According to two indices of composition accuracy, the Sorensen index and a matching coefficient considering both absences and presences, S‐SDMs were more accurate in plant‐rich grasslands. Butterflies display strong and often specialised trophic interactions with plants. At lower altitudes, where land use is more intense, considering climate alone without accounting for land use influences on grassland plant richness leads to erroneous predictions of butterfly presences and absences. In contrast, at higher altitudes, where climate is the main force filtering communities, there were fewer differences between observed and predicted butterfly richness. At high altitudes, even if stochastic processes decrease the accuracy of predictions of presence, climate‐based S‐SDMs are able to better filter out butterfly species that are unable to cope with severe climatic conditions, providing more accurate predictions of absences. Our results suggest that predictions should account for plants in disturbed habitats at lower altitudes but that stochastic processes and heterogeneity at high altitudes may limit prediction success of climate‐based S‐SDMs.     

Phylogenetic conservatism and biogeographic affinity influence woody plant species richness–climate relationships in eastern Eurasia

Mechanisms underlying species richness patterns remain a central yet controversial issue in biology. Climate has been regarded as a major determinant of species richness. However, the relative influences of different evolutionary processes, (i.e. niche conservatism, diversification rate and time for speciation) on species richness–climate relationships remain to be tested. Here, using newly compiled distribution maps for 11 422 woody plant species in eastern Eurasia, we estimated species richness patterns for all species and for families with tropical and temperate affinities separately, and explored the phylogenetic signals in species richness patterns of different families and their relationships with contemporary climate and climate change since the Last Glacial Maximum (LGM). We further compared the effects of niche conservatism (represented by contemporary-ancestral climatic niches differences), diversification rate and time for speciation (represented by family age) on variation in the slopes of species richness–climate relationships. We found that winter coldness was the best predictor for species richness patterns of most tropical families while Quaternary climate change was the best predictor for those of most temperate families. Species richness patterns of closely-related families were more similar than those of distantly-related families within eudicots, and significant phylogenetic signals characterized the slopes of species richness–climate relationships across all angiosperm families. Contemporary-ancestral climatic niche differences dominated variation in the relationships between family-level species richness and most climate variables. Our results indicate significant phylogenetic conservatism in family-level species richness patterns and their relationships with contemporary climate within eudicots. These findings shed light on the mechanisms underlying large-scale species richness patterns and suggest that ancestral climatic niche may influence the evolution of species richness–climate relationships in plants through niche conservatism.     

Projected Effects of Climate Change on Patterns of Vertebrate and Tree Species Richness in the Conterminous United States

General circulation models (GCM) predict that increasing levels of atmospheric carbon dioxide (CO₂) and other greenhouse gases will lead to dramatic changes in climate. It is known that the spatial variability of species richness over continental spatial scales is strongly correlated with contemporary climate. Assuming that this relationship between species richness and climate persists under conditions of increased CO₂, what changes could we expect to occur in terms of species richness? To address this question, I used observed relationships between contemporary richness and climate, coupled with climate projections from five GCM, to project these future changes. These models predict that the richness of vertebrate ectotherms will increase over most of the conterminous United States. Mammal and bird richness are predicted to decrease in much of the southern US and to increase in cool, mountainous areas. Woody plant richness is likely to increase throughout the North and West and to decrease in the southwestern deserts. These projections represent changes that are likely to occur over long time scales (millennia); short-term changes are expected to be mainly negative.     

Soil warming increases plant species richness but decreases germination from the alpine soil seed bank

Global warming is occurring more rapidly above the treeline than at lower elevations and alpine areas are predicted to experience above average warming in the future. Temperature is a primary factor in stimulating seed germination and regulating changes in seed dormancy status. Thus, plant regeneration from seed will be crucial to the persistence, migration and post disturbance recruitment of alpine plants in future climates. Here, we present the first assessment of the impact of soil warming on germination from the persistent alpine soil seed bank. Contrary to expectations, soil warming lead to reduced overall germination from the soil seed bank. However, germination response to soil temperature was species specific such that total species richness actually increased by nine with soil warming. We further explored the system by assessing the prevalence of seed dormancy and germination response to soil disturbance, the frequency of which is predicted to increase under climate change. Seeds of a significant proportion of species demonstrated physiological dormancy mechanisms and germination of several species appeared to be intrinsically linked to soil disturbance. In addition, we found no evidence of subalpine species and little evidence of exotic weed species in the soil, suggesting that the soil seed bank will not facilitate their invasion of the alpine zone. In conclusion, changes in recruitment via the alpine soil seed bank can be expected under climate change, as a result of altered dormancy alleviation and germination cues. Furthermore, the alpine soil seed bank, and the species richness therein, has the potential to help maintain local species diversity, support species range shift and moderate species dominance. Implications for alpine management and areas for further study are also discussed.     

Patterns of herbivore species richness in Kenya and current ecoclimatic stability

The increased attention to biodiversity worldwide has stimulatedinterest in understanding biophysical factors associated with indicators ofbiodiversity such as species richness. Although levels of biodiversity may seemto be equivalent in different areas, high species richness may be caused byaccumulation of species over a long time in places where environmentalconditions remained stable and predictable. The advanced very high resolutionradiometer (AVHRR)–normalized difference vegetation index (NDVI) has beenestablished to be a good proxy for studying interannual climate variability aswell as regional drought condition. In this study, I examined the relationshipbetween large herbivore species richness and AVHRR–NDVI derivedclimatic-variability indices, interannual average NDVI and coefficient ofvariation of NDVI at a regional spatial scale in Kenya. Regions with a relativelylow coefficient of variation of NDVI and high interannual average NDVIcharacterize current ecoclimatic stability. By contrast, a high coefficient ofvariation of NDVI and relatively low interannual average NDVI characterizeecoclimatic instability (drought risk). Statistical analyses revealed that a highinterannual average NDVI increases species richness, whereas a high coefficient ofvariation of NDVI lowers species richness. This indicates that maximum numbers ofspecies are found in regions with current ecoclimatic stability. Understandingsuch relationships can help in explaining spatial distribution of speciesrichness and predicting global changes resulting from human impacts on theenvironment.     

Modelling the species richness distribution of French dung beetles (Coleoptera, Scarabaeidae) and delimiting the predictive capacity of different groups of explanatory variables

Aim To predict French Scarabaeidae dung beetle species richness distribution, and to determine the possible underlying causal factors. Location The entire French territory has been studied by dividing it into 301 grid cells of 0.72 × 0.36 degrees. Method Species richness distribution was predicted using generalized linear models to relate the number of species with spatial, topographic and climate variables in grid squares previously identified as well sampled (n = 66). The predictive function includes the curvilinear relationship between variables, interaction terms and the significant third‐degree polynomial terms of latitude and longitude. The final model was validated by a jack‐knife procedure. The underlying causal factors were investigated by partial regression analysis, decomposing the variation in species richness among spatial, topographic and climate type variables. Results The final model accounts for 86.2% of total deviance, with a mean jack‐knife predictive error of 17.7%. The species richness map obtained highlights the Mediterranean as the region richest in species, and the less well‐explored south‐western region as also being species‐rich. The largest fraction of variability (38%) in the number of species is accounted for by the combined effect of the three groups of explanatory variables. The spatially structured climate component explains 21% of variation, while the pure climate and pure spatial components explain 14% and 11%, respectively. The effect of topography was negligible. Conclusions Delimiting the adequately inventoried areas and elaborating forecasting models using simple environmental variables can rapidly produce an estimate of the species richness distribution. Scarabaeidae species richness distribution seems to be mainly influenced by temperature. Minimum mean temperature is the most influential variable on a local scale, while maximum and mean temperature are the most important spatially structured variables. We suggest that species richness variation is mainly conditioned by the failure of many species to go beyond determined temperature range limits.     

Warmer and richer? Predicting the impact of climate warming on species richness in small temperate waterbodies

Climate change is expected to affect communities worldwide. Many studies focus on responses at the regional level and show an increase in species richness. However, less is known about the consequences of climate change at the local scale (in ecosystems). Small waterbodies, such as ponds, could play an important role for the assessment of the impact of future changes in climate at the local level. We evaluated here the potential changes due to climate warming in the species richness for various groups (plants, snails, beetles, dragonflies, amphibians) across 113 lowland and high altitude ponds in Switzerland. We modelled the relationships between species richness and environmental variables (including temperature) and predicted species richness changes for the end of the century (2090–2100; using the A2 IPCC scenario). Temperature rise could significantly increase pond species richness. For the five taxonomic groups pooled, species richness would potentially increase from 41 to 75 (+83%) in lowland ponds. In presently species‐poor high altitude ponds, the potential increase would be particularly marked, with a proportional increase (+150%; from 14 to 35 species) almost double that in lowland areas. A strong increase in species richness also resulted from models including changes in additional variables, such as landuse or water quality. Future reductions in water quality (e.g. increase in nutrients) may limit the predicted increase in lowland species richness or, conversely, result in a greater increase in species richness in high altitude areas. Nutrient enrichment is shown to affect the taxonomic groups differentially, with plant species richness the most negatively influenced. Climate warming could therefore affect species richness of temperate ponds not only regionally, but also at the local, within ecosystems‐scale; species richness could increase markedly in temperate regions, and especially so at higher altitude.     

Disentangling the drivers of continental mammalian endemism

Endemic species and species with small ranges are ecologically and evolutionarily distinct and are vulnerable to extinction. Determining which abiotic and biotic factors structure patterns of endemism on continents can advance our understanding of global biogeographic processes, but spatial patterns of mammalian endemism have not yet been effectively predicted and reconstructed. Using novel null model techniques, we reconstruct trends in mammalian endemism and describe the isolated and combined effects of physiographic, ecological, and evolutionary factors on endemism. We calculated weighted endemism for global continental ecoregions and compared the spatial distribution of endemism to niche-based, geographic null models of endemism. These null models distribute species randomly across continents, simulating their range sizes from their degree of climatic specialization. They isolate the effects of physiography (topography and climate) and species richness on endemism. We then ran linear and structural models to determine how topography and historical climate stability influence endemism. The highest rates of mammalian endemism were found in topographically rough, climatically stable ecoregions with many species. The null model that isolated physiography did not closely approximate the observed distribution of endemism (r² = .09), whereas the null model that incorporated both physiography and species richness did (r² = .59). The linear models demonstrate that topography and climatic stability both influenced endemism values, but that average climatic niche breadth was not highly correlated with endemism. Climate stability and topography both influence weighted endemism in mammals, but the spatial distribution of mammalian endemism is driven by a combination of physiography and species richness. Despite its relationship to individual range size, average climate niche breadth has only a weak influence on endemism. The results highlight the importance of historical biogeographic processes (e.g. centers of speciation) and geography in driving endemism patterns, and disentangle the mechanisms structuring species ranges worldwide.     

Wildfire–vegetation dynamics affect predictions of climate change impact on bird communities

Community‐level climate change indicators have been proposed to appraise the impact of global warming on community composition. However, non‐climate factors may also critically influence species distribution and biological community assembly. The aim of this paper was to study how fire–vegetation dynamics can modify our ability to predict the impact of climate change on bird communities, as described through a widely‐used climate change indicator: the community thermal index (CTI). Potential changes in bird species assemblage were predicted using the spatially‐explicit species assemblage modelling framework – SESAM – that applies successive filters to constrained predictions of richness and composition obtained by stacking species distribution models that hierarchically integrate climate change and wildfire–vegetation dynamics. We forecasted future values of CTI between current conditions and 2050, across a wide range of fire–vegetation and climate change scenarios. Fire–vegetation dynamics were simulated for Catalonia (Mediterranean basin) using a process‐based model that reproduces the spatial interaction between wildfire, vegetation dynamics and wildfire management under two IPCC climate scenarios. Net increases in CTI caused by the concomitant impact of climate warming and an increasingly severe wildfire regime were predicted. However, the overall increase in the CTI could be partially counterbalanced by forest expansion via land abandonment and efficient wildfire suppression policies. CTI is thus strongly dependent on complex interactions between climate change and fire–vegetation dynamics. The potential impacts on bird communities may be underestimated if an overestimation of richness is predicted but not constrained. Our findings highlight the need to explicitly incorporate these interactions when using indicators to interpret and forecast climate change impact in dynamic ecosystems. In fire‐prone systems, wildfire management and land‐use policies can potentially offset or heighten the effects of climate change on biological communities, offering an opportunity to address the impact of global climate change proactively.     

Species–energy relationship in the deep sea: a test using the Quaternary fossil record

Little is known about the processes regulating species richness in deep‐sea communities. Here we take advantage of natural experiments involving climate change to test whether predictions of the species–energy hypothesis hold in the deep sea. In addition, we test for the relationship between temperature and species richness predicted by a recent model based on biochemical kinetics of metabolism. Using the deep‐sea fossil record of benthic foraminifera and statistical meta‐analyses of temperature‐richness and productivity‐richness relationships in 10 deep‐sea cores, we show that temperature but not productivity is a significant predictor of species richness over the past c. 130 000 years. Our results not only show that the temperature‐richness relationship in the deep‐sea is remarkably similar to that found in terrestrial and shallow marine habitats, but also that species richness tracks temperature change over geological time, at least on scales of c. 100 000 years. Thus, predicting biotic response to global climate change in the deep sea would require better understanding of how temperature regulates the occurrences and geographical ranges of species.     

Quaternary climate changes explain diversity among reptiles and amphibians

It is widely believed that contemporary climate determines large-scale patterns of species richness. An alternative view proposes that species richness reflects biotic responses to historic climate changes. These competing "contemporary climate" vs "historic climate" hypotheses have been vigorously debated without reaching consensus. Here, we test the proposition that European species richness of reptiles and amphibians is driven by climate changes in the Quaternary. We find that climate stability between the Last Glacial Maximum (LGM) and the present day is a better predictor of species richness than contemporary climate; and that the 0°C isotherm of the LGM delimits the distributions of narrow-ranging species, whereas the current 0°C isotherm limits the distributions of wide-ranging species. Our analyses contradict previous studies of large-scale species richness patterns and support the view that "historic climate" can contribute to current species richness independently of and at least as much as contemporary climate.     

Effects of historical and contemporary factors on global patterns in avian species richness

Aim  The aim of this study was to determine how regional and historical factors influence global patterns in avian species richness.
Location  Global.
Methods  Using a comprehensive data set including 710 World Wildlife Fund terrestrial ecoregions covering nearly all the land surface of the Earth, avian species richness was compared among six biogeographical regions after accounting for sample area, elevational range and climate. Analysis of variance and multiple regressions were used. Spatial autocorrelation in model residuals was accounted for.
Results  Significant effects of region on avian species richness were found in nearly all comparisons between biogeographical regions.
Main conclusions  Regional and historical processes have played a role in regulating large-scale avian species richness patterns across the globe. Avian species richness in different regions of the world cannot be accurately predicted by a single global model. Avian species richness in areas of similar environments may differ substantially between regions, and thus avian species richness in one biogeographical region cannot be predicted using the richness–environment relationship derived from the data of another biogeographical region, even one with similar environments.