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.     

Parasite diversity declines with host evolutionary distinctiveness: A global analysis of carnivores

Evolutionarily distinctive host lineages might harbor fewer parasite species because they have fewer opportunities for parasite sharing than hosts having extant close relatives, or because diverse parasite assemblages promote host diversification. We evaluate these hypotheses using data from 930 species of parasites reported to infect free‐living carnivores. We applied nonparametric richness estimators to estimate parasite diversity among well‐sampled carnivore species and assessed how well host evolutionary distinctiveness, relative to other biological and environmental factors, explained variation in estimated parasite diversity. Species richness estimates indicate that the current published literature captures less than 50% of the true parasite diversity for most carnivores. Parasite species richness declined with evolutionary distinctiveness of carnivore hosts (i.e., length of terminal ranches of the phylogeny) and increased with host species body mass and geographic range area. We found no support for the hypothesis that hosts from more diverse lineages support a higher number of generalist parasites, but we did find evidence that parasite assemblages might have driven host lineage diversification through mechanisms linked to sexual selection. Collectively, this work provides strong support for host evolutionary history being an essential predictor of parasite diversity, and offers a simple model for predicting parasite diversity in understudied carnivore species.     

Latitudinal gradient in species richness of the New World Triatominae (Reduviidae)

Aim To quantify the latitudinal gradient in species richness in the New World Triatominae and to explore the species‐energy and area hypotheses as possible causes. Location The gradient was studied for North and South America, between 43° N and 32° S. Methods A database was constructed containing the geographical distribution of the 118 New World Triatominae species based on data extracted from several published sources. Species richness was recorded as the number of species present within 5° latitudinal bands. We used univariate and multivariate models to analyse the relationship between area within each latitudinal belt, land surface temperature, and potential evapotranspiration as explanatory variables, and species richness. All variables were georeferenced and data were extracted using a GIS. Results Species richness of Triatominae increases significantly from the poles towards the Equator, peaking over the 5°?10 ° S latitudinal band. It increases according to a linear model, both north and south of the Equator, although a quadratic model fits better to southern hemisphere data. Richness correlates with habitable geographical area, when it is analysed through a nonlinear multiple regression factoring out latitude, only in the southern hemisphere. Regarding the species‐energy hypothesis, a multiple regression analysis controlling the effect of latitude shows a significant relationship between temperature and species richness. This effect is more pronounced in the southern hemisphere. Species richness shows a strong longitudinal trend south of the Equator (increasing to the east), but not north of the Equator. This differential pattern is reflected in significant interactions between longitude and both latitude and temperature in models of the species richness of the New World Triatominae. Main conclusions To our knowledge, this is the first time that a latitudinal gradient in species richness has been shown and analysed for obligate haematophagous organisms, and it shows that the species–energy hypothesis can account for this phenomenon. This relationship is stronger in the southern hemisphere.     

Determinants of local ant (Hymenoptera: Formicidae) species richness and activity density across Europe

1. Species richness is influenced by local habitat features and large‐scale climatic gradients. Usually, both influences are studied in isolation because of the divergent spatial scales at which they occur. Here, we compared the influence of large‐scale climate and local habitat type on European ants using a continent‐wide, standardised sampling programme. 2. We investigated species richness and activity density from pitfall traps distributed over four habitat types at 17 locations from northern Sweden to Spain and Greece. Species richness and activity density were analysed with respect to ambient energy [equilibrium evapotranspiration (EET)] and productive energy (net primary productivity). Furthermore, we compared ant richness and activity density between the four habitat types: arable land, scrubland, grassland, and forest. 3. Species richness and activity density of ants increased with equilibrium evapotranspiration (EET), explaining 30.2% of the total variation in species richness and 24.2% of activity density. Habitat type explained an additional 19.2% of the variation in species richness and 20.2% of activity density, and was not related to productivity. Species richness and activity density were highest in scrubland and significantly lower in forest and (marginally significant) in arable land. 4. The increase in EET and the decrease in forest confirms the pronounced thermophily of ants, whereas the decrease in arable land is probably caused by soil disturbance.     

Speciation dynamics during the global radiation of extant bats

Species richness varies widely across extant clades, but the causes of this variation remain poorly understood. We investigate the role of diversification rate heterogeneity in shaping patterns of diversity across families of extant bats. To provide a robust framework for macroevolutionary inference, we assemble a time‐calibrated, species‐level phylogeny using a supermatrix of mitochondrial and nuclear sequence data. We analyze the phylogeny using a Bayesian method for modeling complex evolutionary dynamics. Surprisingly, we find that variation in family richness can largely be explained without invoking heterogeneous diversification dynamics. We document only a single well‐supported shift in diversification dynamics across bats, occurring at the base of the subfamily Stenodermatinae. Bat diversity is phylogenetically imbalanced, but—contrary to previous hypotheses—this pattern is unexplained by any simple patterns of diversification rate heterogeneity. This discordance may indicate that diversification dynamics are more complex than can be captured using the statistical tools available for modeling data at this scale. We infer that bats as a whole are almost entirely united into one macroevolutionary cohort, with decelerating speciation through time. There is also a significant relationship between clade age and richness, suggesting that global bat diversity may still be expanding.     

Relative roles of ecological and energetic constraints,diversification rates and region history on global species richness gradients

Regions worldwide differ markedly in species richness. Here, for birds and mammals worldwide, we directly compare four sets of hypotheses regarding geographical richness gradients: (1) evolutionary, emphasising heterogeneity in diversification rates, (2) historical, related to differences in region ages and sizes, (3) energetic, associated with variation in productive or ambient energy and (4) ecological, reflecting differences in ecological niche diversity. Among highly independent regions, or ‘evolutionary arenas’, we find that richness is weakly influenced by richness‐standardised ecological niche diversity, questioning the significance of ecological constraints for producing large‐scale diversity gradients. In contrast, we find strong evidence for the importance of region area and its changes over time, together with a role for temperature. These predictors affect richness predominately directly without concomitant positive effects on diversification rates. This suggests that regional richness is governed by historical and evolutionary processes, which promote region‐specific accumulation of diversity through time or following asymmetrical dispersal.     

Regional variation in the historical components of global avian species richness

Aim Using a global data base of the distribution of extant bird species, we examine the evidence for spatial variation in the evolutionary origins of contemporary avian diversity. In particular, we assess the possible role of the timing of mountain uplift in promoting diversification in different regions. Location Global. Methods We mapped the distribution of avian richness at four taxonomic levels on an equal‐area 1° grid. We examined the relationships between richness at successive taxonomic levels (e.g. species richness vs. genus richness). We mapped the residuals from linear regressions of these relationships to identify areas that are exceptional in the number of lower taxa relative to the number of higher taxa. We use generalized least squares models to test the influence of elevation range and temperature on lower‐taxon richness relative to higher‐taxon richness. Results Peaks of species richness in the Neotropics were congruent with patterns of generic richness, whilst peaks in Australia and the Himalayas were congruent with patterns of both genus and family richness. Hotspots in the Afrotropics did not reflect higher‐taxon patterns. Regional differences in the relationship between richness at successive taxonomic levels revealed variation in patterns of taxon co‐occurrence. Species and genus co‐occurrence was positively associated with elevational range across much of the world. Taxon occurrence in the Neotropics was associated with a positive interaction between elevational range and temperature. Conclusions These results demonstrate that contemporary patterns of richness show different associations with higher‐taxon richness in different regions, which implies that the timing of historical effects on these contemporary patterns varies across regions. We suggest that this is due to dispersal limitation and phylogenetic constraints on physiological tolerance limits promoting diversification. We speculate that diversification rates respond to long‐term changes in the Earth's topography, and that the role of tropical mountain ranges is implicated as a correlate of contemporary diversity, and a source of diversification across avian evolutionary history.     

Neotropical savanna ants show a reversed latitudinal gradient of species richness,with climatic drivers reflecting the forest origin of the fauna

Aim

To evaluate the extent to which ant species richness in Neotropical savannas varies with macrogeographic variables, and to identify the potential climatic drivers of such variation.

Location

The Cerrado savanna biome of central Brazil, in a region spanning ca. 20° of latitude and 18°of longitude.

Methods

Standardized sampling of the arboreal and ground‐dwelling faunas was performed in 29 well‐preserved savanna sites using pitfall traps. Species were classified according to their habitat affinities: open‐savanna specialists, forest‐associated species or habitat generalists. We used generalized linear models to evaluate the importance of geographic (latitude, longitude and elevation) and climatic (mean temperature and three metrics of rainfall) variables as predictors of species richness.

Results

The total number of species recorded at each site varied more than twofold (from 59 to 144), and latitude was the best geographic correlate of overall species richness. However, contrary to the expected pattern, more species were found at higher than lower latitudes. This reversed latitudinal pattern of diversity occurred for both the arboreal and ground‐dwelling faunas, and for the habitat generalists and forest specialists. The savanna specialists showed a mid‐latitudinal peak in diversity. Overall, there was a significant positive association between rainfall and species richness, but the strength of this relationship varied with ant habitat affinity.

Main conclusions

The Cerrado ant fauna shows a reverse latitudinal gradient in species diversity, and this can be explained by increasing rainfall during the warmest months of the year (and therefore in plant productivity) with increasing latitude. The sensitivity of Cerrado ant diversity to declining rainfall contrasts with the high resilience to aridity of the Australian savanna ant fauna, and this reflects the contrasting evolutionary histories of these faunas. Our findings highlight the importance of historical processes as drivers of intercontinental contrasts in macroecological patterns.     

Patterns of distribution for southern Australian marine echinoderms and decapods

Aim To relate patterns of distribution of marine echinoderms and decapods around southern Australia to major ecological and historical factors. Location Shallow‐water (0–100 m) marine waters off southern Australia, south of 30° S. Methods (1) Record the presence/absence of known echinoderm and decapod species in cells of c. 1° latitude and longitude, along the coast of southern mainland Australia and Tasmania. (2) Describe patterns in species composition, species richness and endemism through gradient analysis, ordination and cluster analysis. (3) Relate these patterns to distance and temperature gradients, the area of continental shelf, the average size of species range, and known historical factors. Results Species composition varied with both latitude and longitude. Species richness was relatively constant from east to west but graded with latitude from high in the warm‐temperate regions around Perth and Sydney to low in cool‐temperate southern Tasmania. Species richness was not related to the area of continental shelf or average species range size. Species turnover was not correlated with rates of temperature change. It was problematic to separate distance from temperature gradients, but there was evidence that the southern distribution limits of some species are related to minimum sea surface temperature. Within the taxonomic groups surveyed, evolutionary radiation has been largely limited to a few cosmopolitan species‐rich genera. Main conclusions There are historical as well as ecological hypotheses explaining the latitudinal gradient of marine species richness in southern Australia: (1) the continual invasion and speciation of species of tropical origin as Australia has split from Gondwana and drifted northward; (2) progressive extinction of some Gondwanan cool‐temperate species at the limits of their range; (3) low level of immigration of additional cool‐temperate species; and (4) some in situ endemic speciation.     

Time and environment explain the current richness distribution of non‐marine turtles worldwide

Ecological, historical, and evolutionary hypotheses are important to explain geographical diversity gradients in many clades, but few studies have combined them into a single analysis allowing a comparison of their relative importance. This study aimed to evaluate the relative importance of ecological, historical, and evolutionary hypotheses in explaining the current global distribution of non‐marine turtles, a group whose distribution patterns are still poorly explored. We used data from distribution range maps of 336 species of non‐marine turtles, environmental layers, and phylogeny to obtain richness estimates of these animals in 2° × 2° cells and predictors related to ecological, evolutionary and historical hypotheses driving richness patterns. Then we used a path analysis to evaluate direct and indirect effects of the predictors on turtle richness. Ancestral area reconstruction was also performed in order to evaluate the influence of time‐for‐speciation in the current diversity of the group. We found that environmental variables had the highest direct effects on non‐marine turtle richness, whereas diversification rates and area available in the last 55 million yr minimally influenced turtle distributions. We found evidence for the time‐for‐speciation effect, since regions colonized early were generally richer than recently colonized regions. In addition, regions with a high number of colonization events had a higher number of turtle species. Our results suggested that ecological processes may influence non‐marine turtle richness independent of diversification rates, but they are probably related to dispersal abilities. However, colonization time was also an important component that must be taken into account. Finally, our study provided additional support for the importance of ecological (climate and productivity) and historical (time‐for‐speciation and dispersal) processes in shaping current biodiversity patterns.     

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.     

Evolutionary processes,dispersal limitation and climatic history shape current diversity patterns of European dragonflies

We investigated the effects of contemporary and historical factors on the spatial variation of European dragonfly diversity. Specifically, we tested to what extent patterns of endemism and phylogenetic diversity of European dragonfly assemblages are structured by 1) phylogenetic conservatism of thermal adaptations and 2) differences in the ability of post‐glacial recolonization by species adapted to running waters (lotic) and still waters (lentic). We investigated patterns of dragonfly diversity using digital distribution maps and a phylogeny of 122 European dragonfly species, which we constructed by combining taxonomic and molecular data. We calculated total taxonomic distinctiveness and mean pairwise distances across 4192 50 × 50 km equal‐area grid cells as measures of phylogenetic diversity. We compared species richness with corrected weighted endemism and standardized effect sizes of mean pairwise distances or residuals of total taxonomic distinctiveness to identify areas with higher or lower phylogenetic diversity than expected by chance. Broken‐line regression was used to detect breakpoints in diversity–latitude relationships. Dragonfly species richness peaked in central Europe, whereas endemism and phylogenetic diversity decreased from warm areas in the south‐west to cold areas in the north‐east and with an increasing proportion of lentic species. Except for species richness, all measures of diversity were consistently higher in formerly unglaciated areas south of the 0°C isotherm during the Last Glacial Maximum than in formerly glaciated areas. These results indicate that the distributions of dragonfly species in Europe were shaped by both phylogenetic conservatism of thermal adaptations and differences between lentic and lotic species in the ability of post‐glacial recolonization/dispersal in concert with the climatic history of the continent. The complex diversity patterns of European dragonflies provide an example of how integrating climatic and evolutionary history with contemporary ecological data can improve our understanding of the processes driving the geographical variation of biological diversity.     

A Dated Phylogeny Complements Macroecological Analysis to Explain the Diversity Patterns in Geonoma (Arecaceae)

Integrating phylogenetic data into macroecological studies of biodiversity patterns may complement the information provided by present‐day spatial patterns. In the present study, we used range map data for all Geonoma (Arecaceae) species to assess whether Geonoma species composition forms spatially coherent floristic clusters. We then evaluated the extent to which the spatial variation in species composition reflects present‐day environmental variation vs. nonenvironmental spatial effects, as expected if the pattern reflects historical biogeography. We also examined the degree of geographic structure in the Geonoma phylogeny. Finally, we used a dated phylogeny to assess whether species richness within the floristic clusters was constrained by a specific historical biogeographic driver, namely time‐for‐diversification. A cluster analysis identified six spatially coherent floristic clusters, four of which were used to reveal a significant geographic phylogenetic structure. Variation partitioning analysis showed that 56 percent of the variation in species composition could be explained by spatial variables alone, consistent with historical factors having played a major role in generating the Geonoma diversity pattern. To test for a time‐for‐diversification effect, we correlated four different species richness measures with the diversification time of the earliest large lineage that is characteristic of each cluster. In support of this hypothesis, we found that geographic areas with higher richness contained older radiations. We conclude that current geographic diversity patterns in Geonoma reflect the present‐day climate, but to a larger extent are related to nonenvironmental spatial constraints linked to colonization time, dispersal limitation, and geological history, followed by within‐area evolutionary diversification. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Multiregional comparison of the ecological and phylogenetic structure of butterfly species richness gradients

Aim To examine butterfly species richness gradients in seven regions/countries and to quantify geographic mean root distance (MRD) patterns. My primary goal is to determine the extent to which an explanation for butterfly richness patterns based on tropical niche conservatism and the evolution of cold tolerance, proposed for the fauna of Canada and the USA, applies to other parts of the world. Location USA/Canada, Mexico, Europe/NW Africa, Transbaikal Siberia, Chile, South Africa and Australia. Methods Digitized range maps for butterfly species in each region were used to map richness patterns in summer (for all areas) and winter (for USA/Canada, Europe/NW Africa and Australia). A phylogeny resolved to subfamily was used to map the geographic MRD patterns. Regression trees and general linear models examined climatic and vegetation correlates of species richness and MRD within and among regions. Results Various combinations of climate and vegetation were strong predictors of species richness gradients within regions, but unresolved ‘regional’ factors contributed to the multiregional pattern. Regionally based differences in phylogenetic structure also exist, but MRD is negatively correlated with temperature both within and across areas. MRD patterns consistent with tropical niche conservatism occur in most areas. With a possible partial exception of Mexico, faunas in cold climates and in mountains are more derived than faunas in lowlands and tropical/subtropical climates. In USA/Canada, Europe and Australia, winter faunas are more derived than summer faunas. Main conclusions The phylogenetic pattern previously found in the USA and Canada is widespread in both the Northern and Southern Hemispheres, and niche conservatism and the evolution of cold tolerance is the likely explanation for the development of the global butterfly species richness gradient over evolutionary time. Contemporary climate also influences species richness patterns but is unlikely to be a complete explanation globally. The importance of climate is also manifested in the seasonal loss of more basal butterfly elements outside the tropics in winter.     

Explaining disparities in species richness between Mediterranean floristic regions: a case study in Gladiolus (Iridaceae)

Aim The causes of geographical variation in species richness in clades that do not follow the latitudinal diversity gradient have rarely been investigated. Here, we examine spatial asymmetries of diversity in Gladiolus (Iridaceae), a large genus (> 260 species) that is present in two mediterranean climate biomes: the Cape of southern Africa (106 species) and the Mediterranean Basin (7 species). Despite convergence of climatic conditions between the two regions, the species density of Gladiolus is over one order of magnitude higher in the Cape than in the Mediterranean Basin. We investigate whether the diversity disparities observed in the genus are better explained by recent colonization of species‐poor areas (temporal hypothesis) or by differential rates of diversification (evolutionary hypothesis). Location Africa, Madagascar and Eurasia Methods We employ a recently developed Bayesian method for the estimation of diversification rates and a biogeographical optimization approach within a phylogenetic framework. Results In Gladiolus, the ‘diversity anomaly’ between the two Mediterranean climate regions cannot be explained solely by the time available for speciation in the Cape, but is also due to locally reduced rates of diversification in the Mediterranean Basin. Furthermore, high overall diversity in southern Africa stems from an ancient origin in the Cape allied with high rates of diversification in the summer‐rainfall region of the subcontinent. Main conclusions Both evolutionary and temporal hypotheses must be taken into account in order to explain the diversity anomaly between the Mediterranean Basin and the Cape. Our results suggest that regions at comparable latitudes and/or with similar climate may not converge in diversity levels due to heterogeneity of diversification rates and contrasting biogeographical histories.     

净多样化速率和进化时间对虎耳草目科间物种多样性差异的影响

不同生物类群包含的物种数目常存在巨大差异,这是生态学和生物学研究中普遍观察到的现象。然而,这一现象产生的原因仍然是未解之谜。从宏观进化的角度,进化时间假说和多样化速率假说是两个比较流行的假说。进化时间假说认为类群的演化时间越长,积累的物种丰富度越高; 而多样化速率假说认为类群的净多样化速率越快,则其物种丰富度越高。为验证这两个假说,该文以一棵包含1 539个物种化石定年的虎耳草目系统发育树为基础,通过宏观进化分析获取了虎耳草目内15个科的物种形成和灭绝速率,并计算了每个科的平均多样化速率。结果表明:(1)虎耳草目的物种多样化速率有着增加的趋势,并且多样化速率的增加主要出现在温带和高山类群,如茶藨子科、景天科和芍药科等。(2)采用系统发育广义最小二乘模型(PGLS)和线性回归模型(LM)结果表明,虎耳草目15个科的物种丰富度与科的分化时间和科内物种的最近共同祖先年龄都没有显著相关关系,而与净多样化速率显著正相关(R² =0.380,P<0.05)。该研究支持了多样化速率假说,认为不同科的净多样化速率的差异是导致虎耳草目科间物种数目差异的主要原因之一。全球气候变冷可能为虎耳草目中草本、落叶乔木和灌木等能够适应寒冷环境的类群提供了分布范围扩张和物种快速多样化的机会。该研究表明在温带和高山扩张类群中,物种净多样化速率可能是导致不同类群物种数目差异的主要原因。     

Global biodiversity and biogeography of mangrove crabs: Temperature,the key driver of latitudinal gradients of species richness

Mangroves are ideal habitat for a variety of marine species especially brachyuran crabs as the dominant macrofauna. However, the global distribution, endemicity, and latitudinal gradients of species richness in mangrove crabs remains poorly understood. Here, we assessed whether species richness of mangrove crabs decreases towards the higher latitudes and tested the importance of environmental factors such as Sea Surface Temperature (SST) in creating the latitudinal gradients in species richness of mangrove crabs. A total of 8262 distribution records of 481 species belonging to six families of mangrove crabs including Camptandriidae, Dotillidae, Macrophthalmidae, Ocypodidae, Sesarmidae, and Oziidae were extracted from open-access databases or collected by the authors, quality controlled, cleaned, and analyzed. Species richness was plotted against 5° latitudinal bands in relation to environmental factors. The R software and ArcGIS 10.6.1 were used to analyze the species latitudinal range and richness as well as to map the distribution of mangrove forest, endemic species, species geographical distribution records, and biogeographic regions. The Indo-West Pacific showed the highest species richness of mangrove crabs where more than 65% of species were found in the Indian Ocean and along the western Pacific Ocean. Our results showed that there are 11 significantly different biogeographic regions of mangrove crabs. The highest endemicity rate was observed in the NW Pacific Ocean (29%). Latitudinal patterns of species richness in Macrophthalmidae, Ocypodidae, and Sesarmidae showed an increasing trend from the poles toward the intermediate latitudes including one dip near the equator. However, latitudinal gradients in Camptandriidae, Dotillidae, and Oziidae were unimodal increasing from the higher latitudes towards the equator. Species richness per 5° latitudinal bands significantly increased following mean SST mean (°C), calcite, euphotic depth (m), and mangrove area (km²) across all latitudes, and tide average within each hemisphere. Species richness significantly decreased with dissolved O₂ (ml l⁻¹) and nitrate (μmol l⁻¹) over all latitudes and in the southern hemisphere. The climax of global latitudinal species richness for some mangrove was observed along latitudes 20° N and 15°–25° S, not at the equator. This can suggest that temperature is probably the key driver of latitudinal gradients of mangrove crabs’ species richness. Species richness and mangrove area were also highly correlated.     

Mutualistic mimicry enhances species diversification through spatial segregation and extension of the ecological niche space

Species richness varies among clades, yet the drivers of diversification creating this variation remain poorly understood. While abiotic factors likely drive some of the variation in species richness, ecological interactions may also contribute. Here, we examine one class of potential contributors to species richness variation that is particularly poorly understood: mutualistic interactions. We aim to elucidate large‐scale patterns of diversification mediated by mutualistic interactions using a spatially explicit population‐based model. We focus on mutualistic Müllerian mimicry between conspicuous toxic prey species, where convergence in color patterns emerges from predators' learning process. To investigate the effects of Müllerian mimicry on species diversification, we assume that some speciation events stem from shifts in ecological niches, and can also be associated with shift in mimetic color pattern. Through the emergence of spatial mosaics of mimetic color patterns, Müllerian mimicry constrains the geographical distribution of species and allows different species occupying similar ecological niches to exist simultaneously in different regions. Müllerian mimicry and the resulting spatial segregation of mimetic color patterns thus generate more balanced phylogenetic trees and increase overall species diversity. Our study sheds light on complex effects of Müllerian mimicry on ecological, spatial, and phylogenetic diversification.     

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

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