Phylogenetic properties of Tertiary relict flora in the east Asian continental islands: imprint of climatic niche conservatism and in situ diversification

Understanding biodiversity patterns on islands has long been a central aim in ecology and conservation biology. Island‐specific biogeographical processes play substantial roles in the formation of endemic biota. Here, we examined how climate niche conservatism and geohistorical factors are interactively associated with in situ diversification of Tertiary relict flora in the east Asian continental islands. We generated two novel datasets for species distribution and phylogeny that included all of the known vascular plant species in Japan (5575). Then we tested phylogenetic signal of climatic tolerance, in terms of absolute minimum temperature and water balance, and explored environmental predictors of phylogenetic structure (evolutionary derivedness and clustering) of species assemblages. Although phylogenetic signal of climatic tolerance was significant across the phylogeny of most species, the strength of climatic niche conservatism differed among ferns, gymnosperms, angiosperm trees, and angiosperm herbs. For angiosperm trees, cold temperatures acted as environmental filters that generated phylogenetic derivedness/clustering of species assemblages. For fern and angiosperm herb species, however, phylogenetic properties were not associated with climatic harshness. These contrasting patterns among groups reflected climate niche evolution in vascular plants with different growth forms and traits; for example, diversification of angiosperm trees (but not fern and herb) occurred in response to historical climatic cooling. More importantly, geographical constraints contributed to evolutionary radiation that resulted from isolation by distance from the continent or by elevation. Quaternary climate change was also associated with clade‐specific radiation in refugial habitats. The degree to which geographical, geological, and palaeoclimatic variables explain the phylogenetic structure underscores the importance of isolation‐ and habitat‐stability‐related geohistorical processes in driving in situ diversification despite climatic niche conservatism. We propose that the highly endemic flora of the east Asian islands resulted from the interplay of idiosyncratic regional factors, and ecological and evolutionary processes, such as climate niche assembly and adaptive/nonadaptive radiation.     

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.     

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.     

Geographic patterns and environmental correlates of phylogenetic relatedness and diversity for freshwater fish assemblages in North America

The tropical niche conservatism hypothesis suggests that most groups should be most phylogenetically clustered in cold, dry environments. This idea has been well-tested in plants and some animal groups, but not for fishes. We assess the geographic patterns of freshwater fish phylogenetic structure and investigate the relationships between these patterns and environmental variables across North America and within two biogeographic realms. Phylogenetic relatedness and diversity of 360 freshwater fish assemblages across North America were quantified with three metrics based on a well-dated phylogeny, and were related to 15 environmental variables using correlation and regression analyses. Geographically, the data were analyzed for North America as well as for separate biogeographic realms. We found that cold temperatures are the strongest determinant of phylogenetic clustering overall. However, in the arid west, clustering is most pronounced in the driest regions. In eastern North America, phylogenetic clustering increases at higher latitudes, while the reverse is true in western North America. The strongest phylogenetic clustering for freshwater fish assemblages on the continent is found in the most arid, rather than the coldest, climate in North America. Our results highlight that patterns of phylogenetic structure of freshwater fishes in North America are driven by both ecological and evolutionary processes that differ regionally.     

The tortoise and the hare: ecology of angiosperm dominance and gymnosperm persistence

Gymnosperms, and conifers in particular, are sometimes very productive trees yet angiosperms dominate most temperate and tropical vegetation. Current explanations for angiosperm success emphasize the advantages of insect pollination and seed dispersal by animals for the colonization of isolated habitats. Differences between gymnosperm and angiosperm reproductive and vegetative growth rates have been largely ignored. Gymnosperms are all woody, perennial and usually have long reproductive cycles. Their leaves are not as fully vascularized as those of angiosperms and are more stereotyped in shape and size. Gymnosperm tracheids are generally more resistant to solute flow than angiosperm vessels. A consequence of the less efficient transport system is that maximum growth rates of gymnosperms are lower than maximum growth rates of angiosperms in well lit, well watered habitats. Gymnosperm seedlings may be particularly uncompetitive since their growth depends on a single cohort of relatively inefficient leaves. Later, some gymnosperms attain a higher productivity than co-occurring angiosperm trees by accumulating several cohorts of leaves with a higher total leaf area. These functional constraints on gymnosperm growth rates suggest that gymnosperms will be restricted to areas where growth of angiosperm competitors is reduced, for example, by cold or nutrient shortages. Biogeographic evidence supports this prediction since conifers are largely confined to high latitudes and elevations or nutrient-poor soils. Experimental studies show that competition in the regeneration niche (between conifer seedlings and angiosperm herbs and shrubs) is common and significantly affects conifer growth and survival, Fast-growing angiosperms, especially herbs and shrubs, may also change the frequency of disturbance regimes thereby excluding slower-growing gymnosperms. Shade-tolerant and early successional conifers share similar characteristics of slow initial growth and low plasticity to a change in resources. Shade-tolerant gymnosperms would be expected to occur only where forest openings are small or otherwise unsuitable for rapid filling by fast-growing angiosperm trees, lianas or shrubs. The limited evidence available suggests that shade-tolerant conifers are confined to forests with small gap sizes where large disturbances are very rare. The regeneration hypothesis for gymnosperm exclusion by angiosperms is consistent with several aspects of the fossil record such as the early disappearance of gymnosperms from early successional environments where competition with angiosperms would have been most severe. However there are unresolved difficulties in interpreting process from paleoecological pattern which prevent the testing of alternative hypotheses.     

Evolutionary history of the flora of Mexico: Dry forests cradles and museums of endemism

Mexico is considered an exceptional biogeographic area with a varied endemic flora, however spatial phylogenetic measures of biodiversity have not yet been estimated to understand how its flora assembled to form the current vegetation. Patterns of species richness, endemism, phylogenetic diversity, phylogenetic endemism and centers of neo‐ and paleo‐endemism were determined to examine differences and congruence among these measures, and their implications for conservation. Of 24 360 vascular plant species 10 235 (42%) are endemic. Areas of endemism and phylogenetic endemism were associated with dry forests in zones of topographic complexity in mountain systems, in deserts, and in isolated xeric vegetation. Every single locality where seasonally tropical dry forests have been reported in Mexico was identified as an area of endemism. Significant phylogenetic diversity was the most restricted and occurred in the Trans‐Mexican Volcanic Belt and in the Sierra de Chiapas. Notably, the highest degree of phylogenetic clustering comprising neo‐, paleo‐, and super‐endemism was identified in southernmost Mexico. Most vascular plant lineages diverged in the Miocene (5–20 mya) when arid environments expanded across the world. The location of Mexico between two very large landmasses and the fact that more than fifty percent of its surface is arid favored the establishment of tropical lineages adapted to extreme seasonality and aridity. These lineages were able to migrate from both North and South America across Central America presumably during the Miocene and to diversify, illustrating the signature of the flora of Mexico of areas of endemism with a mixture of neo‐ and paleo‐endemism.     

Tree height–diameter allometry across the United States

The relationship between tree height and diameter is fundamental in determining community and ecosystem structure as well as estimates of biomass and carbon storage. Yet our understanding of how tree allometry relates to climate and whole organismal function is limited. We used the Forest Inventory and Analysis National Program database to determine height–diameter allometries of 2,976,937 individuals of 293 tree species across the United States. The shape of the allometric relationship was determined by comparing linear and nonlinear functional forms. Mixed‐effects models were used to test for allometric differences due to climate and floristic (between angiosperms and gymnosperms) and functional groups (leaf habit and shade tolerance). Tree allometry significantly differed across the United States largely because of climate. Temperature, and to some extent precipitation, in part explained tree allometric variation. The magnitude of allometric variation due to climate, however, had a phylogenetic signal. Specifically, angiosperm allometry was more sensitive to differences in temperature compared to gymnosperms. Most notably, angiosperm height was more negatively influenced by increasing temperature variability, whereas gymnosperm height was negatively influenced by decreasing precipitation and increasing altitude. There was little evidence to suggest that shade tolerance influenced tree allometry except for very shade‐intolerant trees which were taller for any given diameter. Tree allometry is plastic rather than fixed and scaling parameters vary around predicted central tendencies. This allometric variation provides insight into life‐history strategies, phylogenetic history, and environmental limitations at biogeographical scales.     

Climatic correlates of phylogenetic relatedness of woody angiosperms in forest communities along a tropical elevational gradient in South America

Aims This study assesses the relationship between phylogenetic relatedness of angiosperm tree species and climatic variables in local forests distributed along a tropical elevational gradient in South America. In particular, this paper addresses two questions: Is phylogenetic relatedness of plant species in communities related to temperature variables more strongly than to water variables for tropical elevational gradients? Is phylogenetic relatedness of plant species in communities driven by extreme climatic conditions (e.g. minimum temperature (MT) and water deficit) more strongly than by climatic seasonal variability (e.g. temperature seasonality and precipitation seasonality)?Methods I used a set of 34 angiosperm woody plant assemblages along an elevational gradient in the Andes within less than 5 degrees of the equator. Phylogenetic relatedness was quantified as net relatedness index (NRI) and nearest taxon index (NTI) and was related to major climatic variables. Correlation analysis and structure equation modeling approach were used to assess the relationships between phylogenetic relatedness and climatic variables.Important findings Phylogenetic relatedness of angiosperm woody species in the local forest communities is more strongly associated with temperature-related variables than with water-related variables, is positively correlated with mean annual temperature (MAT) and MT, and is related with extreme cold temperature more strongly than with seasonal temperature variability. NTI was related with elevation, MAT and MT more strongly than was NRI. Niche convergence, rather than niche conservatism, has played a primary role in driving community assembly in local forests along the tropical elevational gradient examined. Negative correlations of phylogenetic relatedness with elevation and higher correlations of phylogenetic relatedness with elevation and temperature for NTI than for NRI indicate that evolution of cold tolerance at high elevations in tropical regions primarily occurred at recent (terminal) phylogenetic nodes widely distributed among major clades.     

The macroevolution of climatic niches and its role in ant diversification

1. Investigating how climatic niches change over evolutionary timescales is a necessary step to understanding the current distribution of lineages, yet few studies have addressed this issue using comprehensive datasets. In this study, the evolution of ant climatic niches is investigated at a global scale based on bioclimatic data associated with 163 481 ant occurrence records. The resulting dataset was subjected to principal component analysis, and the scores obtained were used to characterise the main axes of ant climatic niche evolution. 2. Principal component axis 1 (PC1) reflected variation in average temperature and seasonality – consistent with typical tropical/temperate gradients – whereas PC2 was associated with varying levels of aridity. Evolution along these two niche axes was markedly different: differences in the amount of explained variance between PC1 (65%) and PC2 (19%) suggest that climatic niche evolution was nearly three times more pronounced along a tropical–temperate climate axis. 3. There was statistically significant phylogenetic signal on PC1, with genera occupying more tropical conditions diversifying at a faster rate, yet neither of these results is significant on PC2. In addition, most of the ancient ant lineages are associated with conditions of low seasonality and high temperatures. 4. These results provide partial support for the tropical conservatism hypothesis as an explanation for geographical patterns of ant species richness.     

Postglacial colonization of Europe by the barbastelle bat: agreement between molecular data and past predictive modelling

The barbastelle (Barbastella barbastellus) is a rare forest bat with a wide distribution in Europe. Here, we combine results from the analysis of two mtDNA fragments with species distribution modelling to determine glacial refugia and postglacial colonization routes. We also investigated whether niche conservatism occurs in this species. Glacial refugia were identified in the three southern European peninsulas: Iberia, Italy and the Balkans. These latter two refugia played a major role in the postglacial colonization process, with their populations expanding to England and central Europe, respectively. Palaeo‐distribution models predicted that suitable climatic conditions existed in the inferred refugia during the last glacial maximum (LGM). Nevertheless, the overlap between the current and the LGM distributions was almost inexistent in Italy and in the Balkans, meaning that B. barbastellus populations were forced to shift range between glacial and interglacial periods, a process that probably caused some local extinctions. In contrast, Iberian populations showed a ‘refugia within refugium’ pattern, with two unconnected areas containing stable populations (populations that subsisted during both glacial and interglacial phases). Moreover, the match between LGM models and the refugial areas determined by molecular analysis supported the hypothesis of niche conservatism in B. barbastellus. We argue that geographic patterns of genetic structuring, altogether with the modelling results, indicate the existence of four management units for conservation: Morocco, Iberia, Italy and UK, and Balkans and central Europe. In addition, all countries sampled possessed unique gene pools, thus stressing the need for the conservation of local populations.     

Ecological traits influence the phylogenetic structure of bird species co‐occurrences worldwide

The extent to which species’ ecological and phylogenetic relatedness shape their co‐occurrence patterns at large spatial scales remains poorly understood. By quantifying phylogenetic assemblage structure within geographic ranges of >8000 bird species, we show that global co‐occurrence patterns are linked – after accounting for regional effects – to key ecological traits reflecting diet, mobility, body size and climatic preference. We found that co‐occurrences of carnivorous, migratory and cold‐climate species are phylogenetically clustered, whereas nectarivores, herbivores, frugivores and invertebrate eaters tend to be more phylogenetically overdispersed. Preference for open or forested habitats appeared to be independent from the level of phylogenetic clustering. Our results advocate for an extension of the tropical niche conservatism hypothesis to incorporate ecological and life‐history traits beyond the climatic niche. They further offer a novel species‐oriented perspective on how biogeographic and evolutionary legacies interact with ecological traits to shape global patterns of species coexistence in birds.     

Phylogenetic structure and phylogenetic diversity of angiosperm assemblages in forests along an elevational gradient in Changbaishan,China

Aims Understanding what drives the variation in species composition and diversity among local communities can provide insights into the mechanisms of community assembly. Because ecological traits are often thought to be phylogenetically conserved, there should be patterns in phylogenetic structure and phylogenetic diversity in local communities along ecological gradients. We investigate potential patterns in angiosperm assemblages along an elevational gradient with a steep ecological gradient in Changbaishan, China.Methods We used 13 angiosperm assemblages in forest plots (32×32 m) distributed along an elevational gradient from 720 to 1900 m above sea level. We used Faith's phylogenetic diversity metric to quantify the phylogenetic alpha diversity of each forest plot, used the net relatedness index to quantify the degree of phylogenetic relatedness among angiosperm species within each forest plot and used a phylogenetic dissimilarity index to quantify phylogenetic beta diversity among forest plots. We related the measures of phylogenetic structure and phylogenetic diversity to environmental (climatic and edaphic) factors.Important findings Our study showed that angiosperm assemblages tended to be more phylogenetically clustered at higher elevations in Changbaishan. This finding is consistent with the prediction of the phylogenetic niche conservatism hypothesis, which highlights the role of niche constraints in governing the phylogenetic structure of assemblages. Our study also showed that woody assemblages differ from herbaceous assemblages in several major aspects. First, phylogenetic clustering dominated in woody assemblages, whereas phylogenetic overdispersion dominated in herbaceous assemblages; second, patterns in phylogenetic relatedness along the elevational and temperature gradients of Changbaishan were stronger for woody assemblages than for herbaceous assemblages; third, environmental variables explained much more variations in phylogenetic relatedness, phylogenetic alpha diversity and phylogenetic beta diversity for woody assemblages than for herbaceous assemblages.     

Global angiosperm family richness revisited: linking ecology and evolution to climate

Aim The global richness gradient of angiosperm families is correlated with current climate, and it has been claimed that historical processes are not necessary to understand patterns of plant family richness. This claim has drawn criticism, and there have been doubts about the quality of the data used to quantify the pattern. We revisit this issue using the Angiosperm Phylogeny Group (APG) III classification and revised range maps, and we incorporate an evolutionary variable, family age, to explore covariation between evolution and ecology and their links to climate via the tropical conservatism hypothesis (TCH). Location Global. Methods The richness pattern for 408 families was derived from range maps, and family ages were derived from a dated angiosperm phylogeny. Patterns were generated for all families, 143 families composed of trees, and 149 families composed of herbs. We also examined family range size patterns to test the extent to which extratropical floras are nested subsets of tropical floras. Ordinary least squares (OLS) multiple and partial regressions were used to generate climate models for richness, mean range size and mean age for each plant dataset and to evaluate the covariation between contemporary climate and clade age as correlates of family richness. Results We confirmed the strong association between contemporary climate and family richness. Age patterns predicted by TCH were also found for families comprising trees. The richness of herbaceous families, in contrast, was correlated with climate but the age pattern was not as predicted by TCH. Floras in cold and dry areas are strongly nested within richer tropical floras. Main conclusions Phylogenetic niche conservatism at the family level offers a likely explanation for the global diversity gradient of trees, but not for non‐desert herbs, probably because of the faster evolutionary rates for herbs and less constrained evolutionary responses to climate change. Thus, it appears that multiple processes account for the overall angiosperm family gradient. Our analysis also demonstrates that even very strong associations of taxon richness and climate do not preclude evolutionary processes, as has been widely argued, and that climatic and evolutionary hypotheses for richness gradients are not mutually exclusive.     

Postglacial migration supplements climate in determining plant species ranges in Europe

The influence of dispersal limitation on species ranges remains controversial. Considering the dramatic impacts of the last glaciation in Europe, species might not have tracked climate changes through time and, as a consequence, their present-day ranges might be in disequilibrium with current climate. For 1016 European plant species, we assessed the relative importance of current climate and limited postglacial migration in determining species ranges using regression modelling and explanatory variables representing climate, and a novel species-specific hind-casting-based measure of accessibility to postglacial colonization. Climate was important for all species, while postglacial colonization also constrained the ranges of more than 50 per cent of the species. On average, climate explained five times more variation in species ranges than accessibility, but accessibility was the strongest determinant for one-sixth of the species. Accessibility was particularly important for species with limited long-distance dispersal ability, with southern glacial ranges, seed plants compared with ferns, and small-range species in southern Europe. In addition, accessibility explained one-third of the variation in species' disequilibrium with climate as measured by the realized/potential range size ratio computed with niche modelling. In conclusion, we show that although climate is the dominant broad-scale determinant of European plant species ranges, constrained dispersal plays an important supplementary role.     

Niche conservatism and the differences in species richness at the transition of tropical and subtropical climates in South America

Although detected long ago, latitudinal disparity in species richness lacks a consensus regarding its underlying mechanisms. We evaluated whether the main predictions derived from the tropical niche conservatism hypothesis help to explain differences regarding species richness and turnover of species and lineages between forests located in tropical and subtropical climates. If tropical niches are retained, we predict that only a subset of tropical lineages disperses and establishes outside the tropics; tip‐level phylogenetic clustering increases outside the tropics; and the climatic variation drives species richness indirectly via constraints to the distribution of lineages. We compiled 58 checklists along tropical and subtropical sites of riparian forests in southeastern South America. We tested the frequency of niches shifts for species and lineages and the abundance of taxa in each climate. Next, we checked the likelihood of pathways linking climatic and spatial predictors directly with species richness and via phylogenetic clustering estimates. Several lineages only occurred in the tropics, and the number of species and lineages that occurred in both climates was lower than expected by chance. Conversely, few lineages were exclusively subtropical and diversified in the subtropics. Phylogenetic clustering increased in subtropical sites and was correlated with decreasing species richness. An interaction between mean temperature of coldest quarter and precipitation seasonality explained most variation in species richness via increases in phylogenetic clustering. These results support an important contribution of climatic niche conservatism to explain richness disparities between tropics and subtropics, mainly because of the inability of most lineages to colonize the subtropics, which is very likely related to cold intolerance. Since niche conservatism likely drives most of the variation in tree species richness in the region, it provides a mechanistic interpretation of the observed patterns, thus fostering the understanding of richness disparities between these tropical and subtropical tree communities.     

Environmental filters shaping angiosperm tree assembly along climatic and geographic gradients

Question

Global‐scale forest censuses provide an opportunity to understand diversification processes in woody plant communities. Based on the climatic or geographic filtering hypotheses associated with tropical niche conservatism and dispersal limitation, we analysed phylogenetic community structures across a wide range of biomes and evaluated to what extent region‐specific processes have influenced large‐scale diversity patterns of tree species communities across latitude or continent.

Location

Global.

Methods

We generated a data set of species abundances for 21,379 angiosperm woody plants in 843 plots worldwide. We calculated net relatedness index (NRI) for each plot, based on a single global species pool and regional species pools, and phylogenetic β‐diversity (PBD) between plots. Then, we explored the correlations of NRI with climatic and geographic variables, and clarified phylogenetic dissimilarity along geographic and climatic differences. We also compared these patterns for South America, Africa, the Indo‐Pacific, Australia, the Nearctic, Western Palearctic and Eastern Palearctic.

Results

NRI based on a global‐scale species pool was negatively associated with precipitation and positively associated with Quaternary temperature change. PBD was positively associated with geographic distance and precipitation difference between plots across tropical and extratropical biomes. Moreover, phylogenetic dissimilarity was smaller in extratropical regions than in regions including the tropics, although temperate forests of the Eastern Palearctic showed a greater dissimilarity within extratropical regions.

Conclusions

Our findings support predictions of the climatic and geographic filtering hypotheses. Climatic filtering (climatic harshness and paleoclimatic change) relative to tropical niche conservatism played a role in sorting species from the global species pool and shaped the large‐scale diversity patterns, such as the latitudinal gradient observed across continents. Geographic filtering associated with dispersal limitation substantially contributed to regional divergence of tropical/extratropical biomes among continents. Old, long‐standing geographic barriers and recent climatic events differently influenced evolutionary diversification of angiosperm tree communities in tropical and extratropical biomes.     

Evolutionary constraints on regional faunas: whom, but not how many

The latitudinal diversity gradient has been hypothesized to reflect past evolutionary dynamics driven by climatic niche conservation during cladogenesis, i.e. the tropical conservatism hypothesis. Here we show that the species diversity of treefrogs (Hylidae) across the western hemisphere is actually independent of evolutionary niche dynamics. We evaluated three key predictions of the tropical conservatism hypothesis that relate to the relationships between climate, species richness and the phylogenetic structure of regional treefrog faunas across the continental Americas. Species composition was dependent on the inability of some lineages to evolve cold tolerance, but the actual number of species in a region was strongly predicted by precipitation, not temperature. Moreover, phylogenetic structure was independent of precipitation. Thus, species in low-richness areas were no more closely related than species in highly diverse regions. These results provide no support for the tropical conservatism hypothesis. Instead, they show that regional species composition and richness are constrained by different climatic components, demonstrating that global biodiversity gradients can be independent of niche stasis during cladogenesis.     

Climate refugia and migration requirements in complex landscapes

All of today's species have proven their ability to cope with climate change during the glacial‐interglacial cycles of the Quaternary, but future migration requirements may be different regarding speed, direction, geographic barriers, and availability of nearby climate refugia. Here, we contribute a landscape‐level climatic analysis of postglacial vs. projected future migration requirements for 24 common western North American tree species. Using a recently developed velocity of climate change algorithm, we quantify required migration velocities for all populations of species to track climate habitat, based on projections from general circulation models for the 2080s and the last glacial maximum, 21 000 yr ago. Specifically, we ask if nearby climate refugia exist for at least some populations within species ranges and whether the current landscape position of species imply qualitatively different migration requirements in the future compared to those during glacial‐interglacial cycles. Results showed that velocities to reach the nearest climate refugia in the future still exceed the fastest reconstructed post‐glacial migration requirements, but not by orders of magnitude. Regarding landscape positions, we find a low correlation among past and future migration requirements (r = 0.38), suggesting that qualitatively different migration patterns may emerge in the future for some species. Species identified as occupying landscape positions requiring disproportionally faster migration requirements in the future include whitebark pine, pinyon pine, and coast redwood. We discuss uncertainties of our analytical approach as well as implications for human‐assisted migration and conservation action to address climate change.     

Niche conservatism and phylogenetic clustering in a tribe of arid‐adapted marsupial mice,the Sminthopsini

The progressive expansion of the Australian arid zone during the last 20 Ma appears to have spurred the diversification of several families of plants, vertebrates and invertebrates, yet such taxonomic groups appear to show limited niche radiation. Here, we test whether speciation is associated with niche conservatism (constraints on ecological divergence) or niche divergence in a tribe of marsupial mice (Sminthopsini; 23 taxa) that includes the most speciose genus of living dasyurids, the sminthopsins. To that end, we integrated phylogenetic data with ecological niche modelling, to enable us to reconstruct the evolution of climatic suitability within Sminthopsini. Niche overlap among species was low‐moderate (but generally higher than expected given environmental background similarity), and the degree of phylogenetic clustering increased with aridity. Climatic niche reconstruction illustrates that there has been little apparent evolution of climatic tolerance within clades. Accordingly, climatic disparity tends to be accumulated among clades, suggesting considerable niche conservatism. Our results also indicate that evolution of climatic tolerances has been heterogeneous across different dimensions of climate (temperature vs. precipitation) and across phylogenetic clusters (Sminthopsis murina group vs. other groups). Although some results point to the existence of shifts in climatic niches during the speciation of sminthopsins, our study provides evidence for substantial phylogenetic niche conservatism in the group. We conclude that niche diversification had a low impact on the speciation of this tribe of small, but highly mobile marsupials.