Evolutionary and ecological causes of species richness patterns in North American angiosperm trees

Climate and evolutionary factors (e.g. diversification, time‐for‐speciation, niche conservatism) are both thought to be major drivers of species richness in regional assemblages. However, few studies have simultaneously investigated the relative effects of climate and evolutionary factors on species richness across a broad geographical extent. Here, we assess their relative effects on species richness of angiosperm trees across North America. Species richness of angiosperm trees in 1175 regional assemblages were related to climate and phylogenetic structure using a structural equation modeling (SEM) approach. Climate was quantified based on the mean temperature of the coldest month and mean annual precipitation. Evolutionary factors (time‐for‐speciation vs diversification) were inferred from phylogeny‐based measures of mean root distance, phylogenetic species variability, and net relatedness index. We found that at the continental scale, species richness is correlated with temperature and precipitation with approximately similar strength. In the SEM with net relatedness index and phylogenetic species variability and with all the 1175 quadrats, the total direct effect size of phylogenetic structure on species richness is greater than the total direct effect size of climate on species richness by a factor of 3.7. The specific patterns of phylogenetic structure (i.e. greater phylogenetic distances in more species rich regions) are consistent with the idea that time and niche conservatism drive richness patterns in North American angiosperm trees. We conclude that angiosperm tree species richness in regional assemblages in North America is more strongly related to patterns of phylogenetic relatedness than to climatic variation. The results of the present study support the idea that climatic and evolutionary explanations for richness patterns are not in conflict, and that evolutionary processes explain both the relationship between climate and richness and substantial variation in richness that is independent of climate.     

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.     

Disparities between observed and predicted impacts of climate change on winter bird assemblages

Understanding how climate change affects the structure and function of communities is critical for gauging its full impact on biodiversity. To date, community-level changes have been poorly documented, owing, in part, to the paucity of long-term datasets. To circumvent this, the use of ‘space-for-time’ substitution—the forecasting of temporal trends from spatial climatic gradients—has increasingly been adopted, often with little empirical support. Here we examine changes from 1975 to 2001 in three community attributes (species richness, body mass and occupancy) for 404 assemblages of terrestrial winter avifauna in North America containing a total of 227 species. We examine the accuracy of space-for-time substitution and assess causal associations between community attributes and observed changes in annual temperature using a longitudinal study design. Annual temperature and all three community attributes increased over time. The trends for the three community attributes differed significantly from the spatially derived predictions, although richness showed broad congruence. Correlations with trends in temperature were found with richness and body mass. In the face of rapid climate change, applying space-for-time substitution as a predictive tool could be problematic with communities developing patterns not reflected by spatial ecological associations.     

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.     

Functional Resilience against Climate-Driven Extinctions – Comparing the Functional Diversity of European and North American Tree Floras

Future global change scenarios predict a dramatic loss of biodiversity for many regions in the world, potentially reducing the resistance and resilience of ecosystem functions. Once before, during Plio-Pleistocene glaciations, harsher climatic conditions in Europe as compared to North America led to a more depauperate tree flora. Here we hypothesize that this climate driven species loss has also reduced functional diversity in Europe as compared to North America. We used variation in 26 traits for 154 North American and 66 European tree species and grid-based co-occurrences derived from distribution maps to compare functional diversity patterns of the two continents. First, we identified similar regions with respect to contemporary climate in the temperate zone of North America and Europe. Second, we compared the functional diversity of both continents and for the climatically similar sub-regions using the functional dispersion-index (FDis) and the functional richness index (FRic). Third, we accounted in these comparisons for grid-scale differences in species richness, and, fourth, investigated the associated trait spaces using dimensionality reduction. For gymnosperms we find similar functional diversity on both continents, whereas for angiosperms functional diversity is significantly greater in Europe than in North America. These results are consistent across different scales, for climatically similar regions and considering species richness patterns. We decomposed these differences in trait space occupation into differences in functional diversity vs. differences in functional identity. We show that climate-driven species loss on a continental scale might be decoupled from or at least not linearly related to changes in functional diversity. This might be important when analyzing the effects of climate-driven biodiversity change on ecosystem functioning.     

The environmental basis of North American species richness patterns among Epicauta (Coleoptera: Meloidae)

Understanding regional variability in species richness is necessary for conservation efforts to succeed in the face of large-scale environmental deterioration. Several analyses of North American vertebrates have shown that climatic energy provides the best explanation of contemporary species richness patterns. The paucity of analyses of insect diversity patterns, however, remains a serious obstacle to a general hypothesis of spatial variation in diversity. We collected species distribution data on a North American beetle genus, Epicauta (Coleoptera: Meloidae) and tested several major diversity hypotheses. These beetles are generally grasshopper egg predators as larvae, and angiosperm herbivores as adults. Epicauta richness is highest in the hot, dry American southwest, and decreases north and east, consistent with the species richness-energy hypothesis. Potential evapotranspiration, which is also the best predictor of richness patterns among North American vertebrates, explains 80.2% of the variability in Epicauta species richness. Net primary productivity and variables measuring climatic heat energy only (such as PET) are not generally comparable, though they are sometimes treated as if they were equivalent. We conclude that the species richness-energy hypothesis currently provides a better overall explanation for Epicauta species richness patterns in North America than other major diversity hypotheses. The observed relationship between climatic energy and regional species richness may provide significant insight into the response of ecological communities to climate change.     

Spatial patterns of woody plant and bird diversity: functional relationships or environmental effects?

Aim To understand cross‐taxon spatial congruence patterns of bird and woody plant species richness. In particular, to test the relative roles of functional relationships between birds and woody plants, and the direct and indirect environmental effects on broad‐scale species richness of both groups. Location Kenya. Methods Based on comprehensive range maps of all birds and woody plants (native species > 2.5 m in height) in Kenya, we mapped species richness of both groups. We distinguished species richness of four different avian frugivore guilds (obligate, partial, opportunistic and non‐frugivores) and fleshy‐fruited and non‐fleshy‐fruited woody plants. We used structural equation modelling and spatial regressions to test for effects of functional relationships (resource–consumer interactions and vegetation structural complexity) and environment (climate and habitat heterogeneity) on the richness patterns. Results Path analyses suggested that bird and woody plant species richness are linked via functional relationships, probably driven by vegetation structural complexity rather than trophic interactions. Bird species richness was determined in our models by both environmental variables and the functional relationships with woody plants. Direct environmental effects on woody plant richness differed from those on bird richness, and different avian consumer guilds showed distinct responses to climatic factors when woody plant species richness was included in path models. Main conclusions Our results imply that bird and woody plant diversity are linked at this scale via vegetation structural complexity, and that environmental factors differ in their direct effects on plants and avian trophic guilds. We conclude that climatic factors influence broad‐scale tropical bird species richness in large part indirectly, via effects on plants, rather than only directly as often assumed. This could have important implications for future predictions of animal species richness in response to climate change.     

Preliminary global assessment of terrestrial biodiversity consequences of sea-level rise mediated by climate change

Considerable attention has focused on the climatic effects of global climate change on biodiversity, but few analyses and no broad assessments have evaluated effects of sea-level rise on biodiversity. Taking advantage of new maps of marine intrusion under scenarios of 1 and 6 m sea-level rise, we calculated areal losses for all terrestrial ecoregions globally, with areal losses for particular ecoregions ranging from nil to complete. Marine intrusion is a global phenomenon, but its effects are most prominent in Southeast Asia and nearby islands, eastern North America, northeastern South America, and western Alaska. Making assumptions regarding faunal responses to reduced distributional areas of species endemic to ecoregions, we estimated likely numbers of extinctions caused by sea-level rise, and found that marine-intrusion-caused extinctions of narrow endemics are likely to be most prominent in northeastern South America, although anticipated extinctions in smaller numbers are scattered worldwide. This assessment serves as a complement to recent estimates of losses owing to changing climatic conditions, considering a dimension of biodiversity consequences of climate change that has not previously been taken into account.     

Local‐ to continental‐scale variation in the richness and composition of an aquatic food web

Aim We investigated patterns of species richness and composition of the aquatic food web found in the liquid‐filled leaves of the North American purple pitcher plant, Sarracenia purpurea (Sarraceniaceae), from local to continental scales. Location We sampled 20 pitcher‐plant communities at each of 39 sites spanning the geographic range of S. purpurea– from northern Florida to Newfoundland and westward to eastern British Columbia. Methods Environmental predictors of variation in species composition and species richness were measured at two different spatial scales: among pitchers within sites and among sites. Hierarchical Bayesian models were used to examine correlates and similarities of species richness and abundance within and among sites. Results Ninety‐two taxa of arthropods, protozoa and bacteria were identified in the 780 pitcher samples. The variation in the species composition of this multi‐trophic level community across the broad geographic range of the host plant was lower than the variation among pitchers within host‐plant populations. Variation among food webs in richness and composition was related to climate, pore‐water chemistry, pitcher‐plant morphology and leaf age. Variation in the abundance of the five most common invertebrates was also strongly related to pitcher morphology and site‐specific climatic and other environmental variables. Main conclusions The surprising result that these communities are more variable within their host‐plant populations than across North America suggests that the food web in S. purpurea leaves consists of two groups of species: (1) a core group of mostly obligate pitcher‐plant residents that have evolved strong requirements for the host plant and that co‐occur consistently across North America, and (2) a larger set of relatively uncommon, generalist taxa that co‐occur patchily.     

One,two and three‐dimensional geometric constraints and climatic correlates of North American tree species richness

The ‘mid‐domain effect’ (MDE) has received much attention recently 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 randomly placed within a bounded geographic area (i.e. the domain). The most common terrestrial mid‐domain models published to date have been 1‐D latitude or elevation models and 2‐D latitude‐longitude models. Here, we test 1‐D, 2‐D and 3‐D mid‐domain models incorporating latitude, longitude and elevation, and assess independent and concurrent effects of geometric constraints and climatic variables on species richness of North American trees. We use both the traditional ‘global’ regression models as well as geographically weighted regressions (‘local’ models) to examine local variation in the contribution of MDE and climatic variables to species richness across the domain. Our results show that in some dimensions the contribution of MDE to patterns of species richness can be quite substantial, and we show that in most cases a combination of MDE and climate predicted empirical species richness best in both local and global models. For the North American domain, MDE in the elevation dimension is clearly important in describing patterns of empirical species richness. We also show that the assumption of stationarity in global models is not met in the North American domain and that results of these models mask complex patterns in both the effect of MDE on richness and the response of species richness to climate. In particular we show the increased explanatory role of MDE in predicting species richness as domain edges are approached. Our results support the hypothesis that geometric constraints contribute to species richness patterns and we suggest the mid‐domain effect should be considered alongside more traditional environmental correlates in understanding patterns of species diversity.     

Energy and interspecific body size patterns of amphibian faunas in Europe and North America: anurans follow Bergmann's rule, urodeles its converse

Aim To describe broad‐scale geographical patterns of body size for European and North American amphibian faunas and to explore possible processes underlying these patterns. Specifically, we propose a heat balance hypothesis, as both heat conservation and heat gain determine the heat balance of ectotherms, and test it along with five other hypotheses that have a possible influence on body size gradients: size dependence, migration ability, primary productivity, seasonality and water availability. Location Western Europe and North America north of Mexico. Methods We processed distribution maps for native amphibian species to estimate the mean body size in 110 × 110 km cells and calculated eight environmental predictors to explore the relationship between environmental gradients and the observed patterns. We used least squares regression modelling and model selection approaches based on information theory to evaluate the relative support for each hypothesis. Results We found consistent body size gradients and similar relationships to environmental variables within each amphibian group in Europe and North America. Annual potential evapotranspiration, a measure of environmental energy, was the strongest predictor of mean body size in both regions. However, the contrasting responses to ambient energy in each group resulted in opposite geographical patterns, i.e. anurans increased in size from high‐ to low‐energy areas in both continents and urodeles showed the opposite pattern. Main conclusions Our results support the heat balance hypothesis, suggesting that the thermoregulatory abilities of anurans would allow them to reach larger sizes in colder climates by optimizing the trade‐off between heating and cooling rates, whereas a lack of such strategies among urodele faunas would explain why these organisms tend to be smaller in cooler areas. These findings may also have implications for the role of climate warming on the global decline of amphibians.     

Contribution of disturbance to distribution and abundance in a fire‐adapted system

Species distribution modeling (SDM) is an essential tool in understanding species ranges, but models haven't incorporated disturbance‐related variables. This is true even for regions where long histories of disturbance have resulted in disturbance‐adapted species. Therefore, the degree to which including disturbance‐related variables in SDMs might improve their performance is unclear. We used hierarchical partitioning to determine how fire patterns contribute to variation in species abundance and presence, examining both the total variation disturbance‐related variables explained, and how much of this variation is independent of soil and climate variables. For 27 Proteaceae species in the fire‐adapted Cape Floristic Region of South Africa , we found that fire variability, frequency, and area burned tended to have explanatory power similar in size to that of soil and climate variables. Importantly, for SDMs of abundance, fire‐related variables explained additional variation not captured by climatic variables, resulting in markedly increased model performance. In systems with high disturbance rates, species are less likely to be in equilibrium with their environment, and SDMs including variables describing disturbance regimes may be better able to capture the probability of a species being present at a site. Finally, the differential effect of fire on species abundance and presence suggests functional differences between these responses, which could hamper attempts to make predictions about species abundances using models of presence.     

Tropical niche conservatism and the species richness gradient of North American butterflies

Aim We explore the potential role of the ‘tropical conservatism hypothesis’ in explaining the butterfly species richness gradient in North America. Its applicability can be derived from the tropical origin of butterflies and the presumed difficulties in evolving the cold tolerance required to permit the colonization and permanent occupation of the temperate zone. Location North America. Methods Digitized range maps for butterfly species north of Mexico were used to map richness for all species, species with distributions north of the Tropic of Capricorn (Extratropicals), and species that also occupy the tropics (Tropicals). A phylogeny resolved to subfamily was used to map the geographical pattern of mean root distance, a metric of the evolutionary development of assemblages. Regression models and general linear models examined environmental correlates of overall richness and for Extratropicals vs. Tropicals, patterns in summer vs. winter, and patterns in northern vs. southern North America. Results Species in more basal subfamilies dominate the south, whereas more derived clades occupy the north. There is also a ‘latitudinal’ richness gradient in Canada/Alaska, whereas in the conterminous USA richness primarily varies longitudinally. Overall richness is associated with broad‐ and mesoscale temperature gradients. The richness of Tropicals is strongly associated with temperature and distance from winter population sources. The richness of Extratropicals in the north is most strongly correlated with the pattern of glacial retreat since the more recent Ice Age, whereas in the south, richness is positively associated with the range of temperatures in mountains and the presence of forests but is negatively correlated with the broad‐scale temperature gradient. Main conclusions The tropical conservatism hypothesis provides a possible explanation for the complex structure of the species richness gradient. The Canada/Alaska fauna comprises temperate, boreal and tundra species that are nevertheless constrained by cold climates and limited vegetation, coupled with possible post‐Pleistocene recolonization lags. In the USA tropical species are constrained by temperature in winter as well as recolonization distances in summer, whereas temperate‐zone groups are richer in cooler climates in mountains and forests, where winter conditions are more suitable for diapause. The evolution of cold tolerance is key to both the evolutionary and ecological patterns.     

Relative influences of current and historical factors on mammal and bird diversity patterns in deglaciated North America

Aim To investigate the relative contributions of current vs. historical factors in explaining broad‐scale diversity gradients using a combination of contemporary factors and a quantitative estimate of the temporal accessibility of areas for recolonization created by glacial retreat following the most recent Ice Age. Location The part of the Nearctic region of North America that was covered by ice sheets during the glacial maximum 20 000 BP. Methods We used range maps to estimate the species richness of mammals and terrestrial birds in 48 400 km² cells. Current conditions in each cell were quantified using seven climatic and topographical variables. Historical conditions were estimated using the number of years before present when an area became exposed as the ice sheets retreated during the post‐Pleistocene climate warming. We attempted to tease apart contemporary and historical effects using multiple regression, partial regression and spatial autocorrelation analysis. Results A measure of current energy inputs, potential evapotranspiration, explained 76–82% of the variance in species richness, but time since deglaciation explained an additional 8–13% of the variance, primarily due to effects operating at large spatial scales. Because of spatial covariation between the historical climates influencing the melting of the ice sheet and current climates, it was not possible to partition their effects fully, but of the independent effects that could be identified, current climate explained two to seven times more variance in richness patterns than age. Main Conclusions Factors acting in the present appear to have the strongest influence on the diversity gradient, but an historical signal persisting at least 13 000 years is still detectable. This has implications for modelling changes in diversity patterns in response to future global warming.     

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.     

Biodiversity loss under existing land use and climate change: an illustration using northern South America

Aim Species richness depends on climate and land use. Maintaining locations with favourable climate and land‐use patterns is critical for protecting biodiversity because the loss of either can reduce the species richness that an area supports. Currently, the Guiana Shield (GS) receives abundant precipitation and has relatively light land use. For species richness this constitutes a good–good combination of climate and land use, respectively. In contrast, much of eastern Brazil receives low levels of precipitation and has heavy land use, which is a bad–bad combination for species richness. Thus, the current distribution of precipitation and land use in northern South America is relatively favourable for biodiversity. Palaeoclimate and model studies suggest, however, that the precipitation patterns for the two regions have switched before and could switch in response to greenhouse gas emissions. This paper examines the potential consequences of reconfiguring climate with respect to existing land‐use patterns using South America as an example. Location South America north of 20° S and east of the Andes. Methods Ecosystem structure and function are modelled under (1) historical climate and (2) altered precipitation following a shift in the location of the Inter‐Tropical Convergence Zone (ITCZ). The distribution of precipitation, biomes, net primary productivity (NPP) and land use are then used to predict levels of species richness under the two climate scenarios. Results Climate changes could shift the distribution of vegetation and NPP such that conditions favourable for species richness in the GS region disappear. If land‐use patterns were not prohibitive in eastern Brazil, the improved climate conditions there could compensate for the GS loss (assuming migratory barriers are overcome). Instead, existing land‐use patterns cause the combined species richness projected for the two regions to plummet. Main conclusions Human activities will alter current configurations of land use and climate throughout the world. For species richness, new configurations are likely to include both positive and negative combinations of climate and land use. However, the irreversibility of past extinctions due to land‐use patterns loads the dice against species richness.     

Irrigation as an important factor in species distribution models

Species distribution models (SDMs) that employ climatic variables are widely used to predict potential distribution of invasive species. However, climatic variables derived from climate datasets do not account for anthropogenic influences on microclimate. Irrigation is a major anthropogenic activity that influences microclimate conditions and alters the distribution of species in anthropogenic landuses. SDM-based studies appear to ignore the effects of irrigation on microclimatic conditions. This study incorporated irrigation as a correction to precipitation data, to improve the predictive capacity of SDM. As a case study, we examined a SDM of Wasmannia auropunctata, an invasive species that originates in South and Central America, which has invaded tropical and subtropical regions around the world. The potential distribution of W. auropunctata was predicted using Maxent. The model was built based on climatic variables and species records from non-irrigated sites in the native range and then projected on a global scale. Invasive species records were used to evaluate the performance of the model. Precipitation-related variables were modified to approximate actual water input in irrigated areas. Precipitation correction relied on an estimate of irrigation inputs. The model with irrigation correction performed better than the corresponding model without correction, on a global scale and when it was examined in five different geographical regions of the model. These results demonstrate the importance of irrigation correction for assessing the distribution of W. auropunctata in various geographical regions. Accounting for irrigation is expected to improve SDMs for a variety of species.     

Ice age legacies in the geographical distribution of tree species richness in Europe

Aim This study uses a high‐resolution simulation of the Last Glacial Maximum (LGM) climate to assess: (1) whether LGM climate still affects the geographical species richness patterns in the European tree flora and (2) the relative importance of modern and LGM climate as controls of tree species richness in Europe. Location The parts of Europe that were unglaciated during the LGM. Methods Atlas data on the distributions of 55 tree species were linked with data on modern and LGM climate and climatic heterogeneity in a geographical information system with a 60‐km grid. Four measures of species richness were computed: total richness, and richness of the 18 most restricted species, 19 species of medium incidence (intermediate species) and 18 most widespread species. We used ordinary least‐squares regression and spatial autoregressive modelling to test and estimate the richness–climate relationships. Results LGM climate constituted the best single set of explanatory variables for richness of restricted species, while modern climate and climatic heterogeneity was best for total and widespread species richness and richness of intermediate species, respectively. The autoregressive model with all climatic predictors was supported for all richness measures using an information‐theoretic approach, albeit only weakly so for total species richness. Among the strongest relationships were increases in total and intermediate richness with climatic heterogeneity and in restricted richness with LGM growing‐degree‐days. Partial regression showed that climatic heterogeneity accounted for the largest unique variation fraction for intermediate richness, while LGM climate was particularly important for restricted richness. Main conclusions LGM climate appears to still affect geographical patterns of tree species richness in Europe, albeit the relative importance of modern and LGM climate depends on range size. Notably, LGM climate is a strong richness control for species with a restricted range, which appear to still be associated with their glacial refugia.     

Patterns of woody plant species richness in the Iberian Peninsula: environmental range and spatial scale

Aim Climate‐based models often explain most of the variation in species richness along broad‐scale geographical gradients. We aim to: (1) test predictions of woody plant species richness on a regional spatial extent deduced from macro‐scale models based on water–energy dynamics; (2) test if the length of the climate gradients will determine whether the relationship with woody species richness is monotonic or unimodal; and (3) evaluate the explanatory power of a previously proposed ‘water–energy’ model and regional models at two grain sizes. Location The Iberian Peninsula. Methods We estimated woody plant species richness on grid maps with c. 2500 and 22,500 km² cell size, using geocoded data for the individual species. Generalized additive models were used to explore the relationships between richness and climatic, topographical and substrate variables. Ordinary least squares regression was used to compare regional and more general water–energy models in relation to grain size. Variation partitioning by partial regression was applied to find how much of the variation in richness was related to spatial variables, explanatory variables and the overlap between these two. Results Water–energy dynamics generate important underlying gradients that determine the woody species richness even over a short spatial extent. The relationships between richness and the energy variables were linear to curvilinear, whereas those with precipitation were nonlinear and non‐monotonic. Only a small fraction of the spatially structured variation in woody species richness cannot be accounted for by the fitted variables related to climate, substrate and topography. The regional models accounted for higher variation in species richness than the water–energy models, although the water–energy model including topography performed well at the larger grain size. Elevation range was the most important predictor at all scales, probably because it corrects for ‘climatic error’ due to the unrealistic assumption that mean climate values are evenly distributed in the large grid cells. Minimum monthly potential evapotranspiration was the best climatic predictor at the larger grain size, but actual evapotranspiration was best at the smaller grain size. Energy variables were more important than precipitation individually. Precipitation was not a significant variable at the larger grain size when examined on its own, but was highly significant when an interaction term between itself and substrate was included in the model. Main conclusions The significance of range in elevation is probably because it corresponds to several aspects that may influence species diversity, such as climatic variability within grid cells, enhanced surface area, and location for refugia. The relative explanatory power of energy and water variables was high, and was influenced by the length of the climate gradient, substrate and grain size of the analysis. Energy appeared to have more influence than precipitation, but water availability is also determined by energy, substrate and topographic relief.     

Modelling geographical patterns in species richness using eigenvector-based spatial filters

Aim To test the mechanisms driving bird species richness at broad spatial scales using eigenvector‐based spatial filtering. Location South America. Methods An eigenvector‐based spatial filtering was applied to evaluate spatial patterns in South American bird species richness, taking into account spatial autocorrelation in the data. The method consists of using the geographical coordinates of a region, based on eigenanalyses of geographical distances, to establish a set of spatial filters (eigenvectors) expressing the spatial structure of the region at different spatial scales. These filters can then be used as predictors in multiple and partial regression analyses, taking into account spatial autocorrelation. Autocorrelation in filters and in the regression residuals can be used as stopping rules to define which filters will be used in the analyses. Results Environmental component alone explained 8% of variation in richness, whereas 77% of the variation could be attributed to an interaction between environment and geography expressed by the filters (which include mainly broad‐scale climatic factors). Regression coefficients of environmental component were highest for AET. These results were unbiased by short‐scale spatial autocorrelation. Also, there was a significant interaction between topographic heterogeneity and minimum temperature. Conclusion Eigenvector‐based spatial filtering is a simple and suitable statistical protocol that can be used to analyse patterns in species richness taking into account spatial autocorrelation at different spatial scales. The results for South American birds are consistent with the climatic hypothesis, in general, and energy hypothesis, in particular. Habitat heterogeneity also has a significant effect on variation in species richness in warm tropical regions.