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.     

Quaternary and pre‐Quaternary historical legacies in the global distribution of a major tropical plant lineage

Aim The relative importance of current climate and past historical legacies is hotly debated. Here, we assess their role in determining the global distribution and diversity patterns of palms (Arecaceae), a widespread, species‐rich group of keystone ecological importance in tropical ecosystems. Location Global. Methods We assembled country‐level species lists world‐wide and compiled associated data on potential contemporary environmental drivers (current climate, habitat heterogeneity, area, and insularity), Quaternary glacial–interglacial climate change and major biogeographic regions to evaluate to what extent the global distribution and species richness patterns in palms reflect Quaternary climatic oscillations or regional effects reflecting pre‐Quaternary legacies. We also assessed for the first time if historical legacies differ between continents and islands, providing novel insights into determinants of insular species richness. Results Palm species richness was significantly affected by Quaternary climate changes and further differed between biogeographic regions even when both current environmental conditions and Quaternary climate changes were accounted for. In contrast, global limits to the distribution of the palm family were best explained by current temperature while biogeographic regional differences were unimportant and Quaternary climate change caused only a small constraint. Historical legacies were weak on islands, with only a small regional effect and no effect of Quaternary climate changes. Main conclusions Strong historical legacies supplement current environment as determinants of palm species richness. These primarily comprise pre‐Quaternary historical effects, reflected in low African species richness (possibly linked to pre‐Quaternary extinctions) and outstandingly high Neotropical and Indomalayan palm species richness (possibly linked to these regions' long‐term climatic suitability for palms). In contrast to species richness, the global distribution of the family range is largely in equilibrium with current climate. The small historical effects on islands are consistent with climatic buffering from their oceanic environment.     

Interactive effects of elevation,species richness and extreme climatic events on plant–pollinator networks

Plant–pollinator interactions are essential for the functioning of terrestrial ecosystems, but are increasingly affected by global change. The risks to such mutualistic interactions from increasing temperature and more frequent extreme climatic events such as drought or advanced snow melt are assumed to depend on network specialization, species richness, local climate and associated parameters such as the amplitude of extreme events. Even though elevational gradients provide valuable model systems for climate change and are accompanied by changes in species richness, responses of plant–pollinator networks to climatic extreme events under different environmental and biotic conditions are currently unknown. Here, we show that elevational climatic gradients, species richness and experimentally simulated extreme events interactively change the structure of mutualistic networks in alpine grasslands. We found that the degree of specialization in plant–pollinator networks (H2′) decreased with elevation. Nonetheless, network specialization increased after advanced snow melt at high elevations, whereas changes in network specialization after drought were most pronounced at sites with low species richness. Thus, changes in network specialization after extreme climatic events depended on climatic context and were buffered by high species richness. In our experiment, only generalized plant–pollinator networks changed in their degree of specialization after climatic extreme events. This indicates that contrary to our assumptions, network generalization may not always foster stability of mutualistic interaction networks.     

Determinants of species richness in generalist and specialist Mediterranean butterflies: the negative synergistic forces of climate and habitat change

Although it is well established that butterfly richness is affected by climate and human factors (e.g. habitat disturbance and degradation) at different spatial scales, the drivers behind these changes vary greatly according to the geographical region and the ecology of the species concerned. It is essential that this variation be understood if trends in diversity are to be predicted with any degree of confidence under a scenario of global change. Here we examine patterns of butterfly species richness among groups differing in degree of habitat specialization, diet breadth and mobility in the north‐west Mediterranean Basin, a European hotspot for this taxon. We analyze a large number of butterfly communities and take into consideration the main potential drivers, that include climatic, geographic and resource variables, landscape structure and human environmental impact at different spatial scales. Our study shows that both climatic and anthropogenic factors play an important role in determining butterfly species richness in the north‐west Mediterranean Basin, but that their relative impact differs between specialist and generalist groups. At lower altitudes, water availability, a product of the interplay between temperature and rainfall, and negative effects of temperature appear as the most determinant factors. Maximum diversity was observed at mid‐altitudes, which reveals the importance from a conservation point of view of Mediterranean mountain ranges. The results suggest serious population declines in specialist species restricted to mountain areas as a result of climate warming in combination with habitat loss caused by the abandonment of grazing and mowing. They also suggest negative trends for generalist species due to an increase in aridity in combination with an increase in intensification of human land use in lowland areas. Such synergies are expected to lead to rapid declines in Mediterranean butterfly populations in the coming years, thereby posing a severe threat for the conservation of European biodiversity.     

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.     

The geographical variation of network structure is scale dependent: understanding the biotic specialization of host–parasitoid networks

Research on the structure of ecological networks suggests that a number of universal patterns exist. Historically, biotic specialization has been thought to increase towards the Equator. Yet, recent studies have challenged this view showing non‐conclusive results. Most studies analysing the geographical variation in biotic specialization focus, however, only on the local scale. Little is known about how the geographical variation of network structure depends on the spatial scale of observation (i.e. from local to regional spatial scales). This should be remedied, as network structure changes as the spatial scale of observation changes, and the magnitude and shape of these changes can elucidate the mechanisms behind the geographical variation in biotic specialization. Here we analyse four facets of biotic specialization in host–parasitoid networks along gradients of climatic constancy, classifying the networks according to their spatial extension (local or regional). Namely, we analyse network connectance, consumer diet overlap, consumer diet breadth, and resource vulnerability at both local and regional scales along the gradients of both current climatic constancy and historical climatic change. While at the regional scale none of the climatic variables are associated to biotic specialization, at the local scale, network connectance, consumer diet overlap, and resource vulnerability decrease with current climatic constancy, whereas consumer generalism increases (i.e. broader diet breadths in tropical areas). Similar patterns are observed along the gradient of historical climatic change. We provide an explanation based on different beta‐diversity for consumers and resources across the geographical gradients. Our results show that the geographical gradient of biotic specialization is not universal. It depends on both the facet of biotic specialization and the spatial scale of observation.     

Richness patterns, species distributions and the principle of extreme deconstruction

Aim To analyse the global patterns in species richness of Viperidae snakes through the deconstruction of richness into sets of species according to their distribution models, range size, body size and phylogenetic structure, and to test if environmental drivers explaining the geographical ranges of species are similar to those explaining richness patterns, something we called the extreme deconstruction principle. Location Global. Methods We generated a global dataset of 228 terrestrial viperid snakes, which included geographical ranges (mapped at 1° resolution, for a grid with 7331 cells world‐wide), body sizes and phylogenetic relationships among species. We used logistic regression (generalized linear model; GLM) to model species geographical ranges with five environmental predictors. Sets of species richness were also generated for large and small‐bodied species, for basal and derived species and for four classes of geographical range sizes. Richness patterns were also modelled against the five environmental variables through standard ordinary least squares (OLS) multiple regressions. These subsets are replications to test if environmental factors driving species geographical ranges can be directly associated with those explaining richness patterns. Results Around 48% of the total variance in viperid richness was explained by the environmental model, but richness sets revealed different patterns across the world. The similarity between OLS coefficients and the primacy of variables across species geographical range GLMs was equal to 0.645 when analysing all viperid snakes. Thus, in general, when an environmental predictor it is important to model species geographical ranges, this predictor is also important when modelling richness, so that the extreme deconstruction principle holds. However, replicating this correlation using subsets of species within different categories in body size, range size and phylogenetic structure gave more variable results, with correlations between GLM and OLS coefficients varying from –0.46 up to 0.83. Despite this, there is a relatively high correspondence (r = 0.73) between the similarity of GLM‐OLS coefficients and R² values of richness models, indicating that when richness is well explained by the environment, the relative importance of environmental drivers is similar in the richness OLS and its corresponding set of GLMs. Main conclusions The deconstruction of species richness based on macroecological traits revealed that, at least for range size and phylogenetic level, the causes underlying patterns in viperid richness differ for the various sets of species. On the other hand, our analyses of extreme deconstruction using GLM for species geographical range support the idea that, if environmental drivers determine the geographical distribution of species by establishing niche boundaries, it is expected, at least in theory, that the overlap among ranges (i.e. richness) will reveal similar effects of these environmental drivers. Richness patterns may be indeed viewed as macroecological consequences of population‐level processes acting on species geographical ranges.     

Plant species richness in Fennoscandia: evaluating the relative importance of climate and history

In Fennoscandia, the species richness of vascular plants in 75 × 75 km squares is highly correlated with geographical (latitude and longitude) and climatic variables (accumulated respiration sum, mean January temperature, and mean July temperature). When generalised additive models (GAM) are used, over 80% of the variation in richness can be statistically explained by geography and climate. Even though climate has such a high explanatory power we present several arguments for interpreting these results with care. Climate has no ecologically sound explanatory power when the variation due to latitude and longitude is accounted for, and the strongest latitudinal gradient in summer temperature is in an area where the latitudinal gradient in species richness is absent. We discuss the role that Holocene history might have on the variation in species richness, and argue that history and climate should be considered simultaneously when explaining the observed patterns in the geographical variation of species richness.     

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.     

Historical climate‐change influences modularity and nestedness of pollination networks

The structure of species interaction networks is important for species coexistence, community stability and exposure of species to extinctions. Two widespread structures in ecological networks are modularity, i.e. weakly connected subgroups of species that are internally highly interlinked, and nestedness, i.e. specialist species that interact with a subset of those species with which generalist species also interact. Modularity and nestedness are often interpreted as evolutionary ecological structures that may have relevance for community persistence and resilience against perturbations, such as climate‐change. Therefore, historical climatic fluctuations could influence modularity and nestedness, but this possibility remains untested. This lack of research is in sharp contrast to the considerable efforts to disentangle the role of historical climate‐change and contemporary climate on species distributions, richness and community composition patterns. Here, we use a global database of pollination networks to show that historical climate‐change is at least as important as contemporary climate in shaping modularity and nestedness of pollination networks. Specifically, on the mainland we found a relatively strong negative association between Quaternary climate‐change and modularity, whereas nestedness was most prominent in areas having experienced high Quaternary climate‐change. On islands, Quaternary climate‐change had weak effects on modularity and no effects on nestedness. Hence, for both modularity and nestedness, historical climate‐change has left imprints on the network structure of mainland communities, but had comparably little effect on island communities. Our findings highlight a need to integrate historical climate fluctuations into eco‐evolutionary hypotheses of network structures, such as modularity and nestedness, and then test these against empirical data. We propose that historical climate‐change may have left imprints in the structural organisation of species interactions in an array of systems important for maintaining biological diversity.     

Amphibian Beta Diversity in the Brazilian Atlantic Forest: Contrasting the Roles of Historical Events and Contemporary Conditions at Different Spatial Scales

Current patterns of biodiversity distribution result from a combination of historical and contemporary processes. Here, we compiled checklists of amphibian species to assess the roles of long-term climate stability (Quaternary oscillations), contemporary environmental gradients and geographical distance as determinants of change in amphibian taxonomic and phylogenetic composition in the Brazilian Atlantic Forest. We calculated beta diversity as both variation in species composition (CBD) and phylogenetic differentiation (PBD) among the assemblages. In both cases, overall beta diversity was partitioned into two basic components: species replacement and difference in species richness. Our results suggest that the CBD and PBD of amphibians are determined by spatial turnover. Geographical distance, current environmental gradients and long-term climatic conditions were complementary predictors of the variation in CBD and PBD of amphibian species. Furthermore, the turnover components between sites from different regions and between sites within the stable region were greater than between sites within the unstable region. On the other hand, the proportion of beta-diversity due to species richness difference for both CBD and PBD was higher between sites in the unstable region than between sites in the stable region. The high turnover components from CBD and PBD between sites in unstable vs stable regions suggest that these distinct regions have different biogeographic histories. Sites in the stable region shared distinct clades that might have led to greater diversity, whereas sites in the unstable region shared close relatives. Taken together, these results indicate that speciation, environmental filtering and limited dispersal are complementary drivers of beta-diversity of amphibian assemblages in the Brazilian Atlantic Forest.     

Role of climate and geohistorical factors in driving plant richness patterns and endemicity on the east Asian continental islands

We investigated the roles of climate, geography, and geology in plant diversity and endemicity on the east Asian continental islands, by testing predictions from contrasting hypotheses considering current climate, habitat stability, and isolation as major drivers of plant richness and endemicity. We created a fine‐resolution map of vascular plant richness (5614 species) with 10 × 10 km grid cells. Using this large dataset and regression models, we explored correlations between species richness/number of endemics and temperature, precipitation, Quaternary temperature/precipitation changes, Quaternary alluvial development and volcanic disturbances (presence of alluvial plains and of pyroclastic flows), distance from the continent, and elevation. We applied these analyses to the vascular plant assemblage as a whole and separately to trees, herbs, and ferns. Temperature and precipitation were associated with the richness of vascular plants overall and of their endemics. Quaternary temperature change was negatively associated with the richness of vascular plants overall and of their endemics. The presence of pyroclastic flows and of lowland alluvial plains was negatively associated with those. Distance from the continent and elevation were positively associated with endemic species richness, especially those of trees and herbs. While current climate was an important predictor of species richness (especially of ferns), geographical isolation and habitat stability were the main predictors of the endemic species richness of trees and herbs. The relative importance of current climate and historical factors may be related to the dispersal ability of functional groups. Our results illustrate that the diverse geographical conditions reflecting the characteristics of the island led to the various historical effects on biodiversity patterns. The highly endemic flora on the east Asian islands resulted from species accumulation and in situ diversification, suggesting that the climate and historical hypotheses are not mutually exclusive, but can be reconciled as the interplay between recent ecological and evolutionary processes.     

Palaeo-precipitation is a major determinant of palm species richness patterns across Madagascar: a tropical biodiversity hotspot

The distribution of rainforest in many regions across the Earth was strongly affected by Pleistocene ice ages. However, the extent to which these dynamics are still important for modern-day biodiversity patterns within tropical biodiversity hotspots has not been assessed. We employ a comprehensive dataset of Madagascan palms (Arecaceae) and climate reconstructions from the last glacial maximum (LGM; 21 000 years ago) to assess the relative role of modern environment and LGM climate in explaining geographical species richness patterns in this major tropical biodiversity hotspot. We found that palaeoclimate exerted a strong influence on palm species richness patterns, with richness peaking in areas with higher LGM precipitation relative to present-day even after controlling for modern environment, in particular in northeastern Madagascar, consistent with the persistence of tropical rainforest during the LGM primarily in this region. Our results provide evidence that diversity patterns in the World''s most biodiverse regions may be shaped by long-term climate history as well as contemporary environment.     

High proportion of smaller ranged hummingbird species coincides with ecological specialization across the Americas

Ecological communities that experience stable climate conditions have been speculated to preserve more specialized interspecific associations and have higher proportions of smaller ranged species (SRS). Thus, areas with disproportionally large numbers of SRS are expected to coincide geographically with a high degree of community-level ecological specialization, but this suggestion remains poorly supported with empirical evidence. Here, we analysed data for hummingbird resource specialization, range size, contemporary climate, and Late Quaternary climate stability for 46 hummingbird–plant mutualistic networks distributed across the Americas, representing 130 hummingbird species (ca 40% of all hummingbird species). We demonstrate a positive relationship between the proportion of SRS of hummingbirds and community-level specialization, i.e. the division of the floral niche among coexisting hummingbird species. This relationship remained strong even when accounting for climate, furthermore, the effect of SRS on specialization was far stronger than the effect of specialization on SRS, suggesting that climate largely influences specialization through species'' range-size dynamics. Irrespective of the exact mechanism involved, our results indicate that communities consisting of higher proportions of SRS may be vulnerable to disturbance not only because of their small geographical ranges, but also because of their high degree of specialization.     

Facilitative plant interactions and climate simultaneously drive alpine plant diversity

Interactions among species determine local‐scale diversity, but local interactions are thought to have minor effects at larger scales. However, quantitative comparisons of the importance of biotic interactions relative to other drivers are rarely made at larger scales. Using a data set spanning 78 sites and five continents, we assessed the relative importance of biotic interactions and climate in determining plant diversity in alpine ecosystems dominated by nurse‐plant cushion species. Climate variables related with water balance showed the highest correlation with richness at the global scale. Strikingly, although the effect of cushion species on diversity was lower than that of climate, its contribution was still substantial. In particular, cushion species enhanced species richness more in systems with inherently impoverished local diversity. Nurse species appear to act as a ‘safety net’ sustaining diversity under harsh conditions, demonstrating that climate and species interactions should be integrated when predicting future biodiversity effects of climate change.     

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.     

Relative importance of long‐term changes in climate and land‐use on the phenology and abundance of legume crop specialist and generalist aphids

Abstract Insect populations are prone to respond to global changes through shifts in phenology, distribution and abundance. However, global changes cover several factors such as climate and land-use, the relative importance of these being largely unknown. Here, we aim at disentangling the effects of climate, land-use, and geographical drivers on aphid abundance and phenology in France, at a regional scale and over the last 40 years. We used aerial data obtained from suction traps between 1978 and 2015 on five aphid species varying in their degree of specialization to legumes, along with climate, legume crop area and geographical data. Effects of environmental and geographical variables on aphid annual abundance and spring migration dates were analyzed using generalized linear mixed models. We found that within the last four decades, aphids have advanced their spring migration by a month, mostly due to the increase in temperature early in the year, and their abundance decreased by half on average, presumably in response to a combination of factors. The influence of legume crop area decreased with the degree of specialization of the aphid species to such crops. The effect of geographical variation was high even when controlling for environmental variables, suggesting that many other spatially structured processes act on aphid population characteristics. Multifactorial analyses helped to partition the effects of different global change drivers. Climate and land-use changes have strong effects on aphid populations, with important implications for future agriculture. Additionally, trait-based response variation could have major consequences at the community scale.     

Disentangling the Role of Climate,Topography and Vegetation in Species Richness Gradients

Environmental gradients (EG) related to climate, topography and vegetation are among the most important drivers of broad scale patterns of species richness. However, these different EG do not necessarily drive species richness in similar ways, potentially presenting synergistic associations when driving species richness. Understanding the synergism among EG allows us to address key questions arising from the effects of global climate and land use changes on biodiversity. Herein, we use variation partitioning (also know as commonality analysis) to disentangle unique and shared contributions of different EG in explaining species richness of Neotropical vertebrates. We use three broad sets of predictors to represent the environmental variability in (i) climate (annual mean temperature, temperature annual range, annual precipitation and precipitation range), (ii) topography (mean elevation, range and coefficient of variation of elevation), and (iii) vegetation (land cover diversity, standard deviation and range of forest canopy height). The shared contribution between two types of EG is used to quantify synergistic processes operating among EG, offering new perspectives on the causal relationships driving species richness. To account for spatially structured processes, we use Spatial EigenVector Mapping models. We perform analyses across groups with distinct dispersal abilities (amphibians, non-volant mammals, bats and birds) and discuss the influence of vagility on the partitioning results. Our findings indicate that broad scale patterns of vertebrate richness are mainly affected by the synergism between climate and vegetation, followed by the unique contribution of climate. Climatic factors were relatively more important in explaining species richness of good dispersers. Most of the variation in vegetation that explains vertebrate richness is climatically structured, supporting the productivity hypothesis. Further, the weak synergism between topography and vegetation urges caution when using topographic complexity as a surrogate of habitat (vegetation) heterogeneity.     

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.     

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.