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Among direct-developing rain frogs of the genus Craugastor is a clade of 19 described species (bocourti series) that occur in Mexico and northern Central America. Many of these 19 species have been described based on subtle morphological differences and have never been examined using molecular data. Here, we used a multilocus dataset (one mitochondrial (mtDNA) and four nuclear (nDNA) gene fragments, totalling 3,048 concatenated base pairs) to investigate species limits and phylogenetic relationships among 60 northern rain frogs referable to 12 species, with a focus on species from Guatemala. We inferred phylogenies using maximum likelihood and Bayesian analyses on separate mtDNA and nDNA datasets. Concatenated and coalescent species-tree analyses support the monophyly of multiple species, with interspecific relationships mostly unresolved. These mtDNA and nDNA trees were often incongruent with morphology-based taxonomy. For example, two genetically shallow clades contained individuals referable to at least five described species, whereas a single described species contained deep divergences indicative of multiple cryptic species. These findings indicate that morphology-based taxonomy has both overestimated and underestimated actual species diversity (depending on the species), an interpretation supported by two molecular species-delimitation procedures. Based on these findings, we synonymise C. glaucus (Lynch, 1967) and C. stuarti (Lynch, 1967) with C. xucanebi (Stuart, 1941). We also synonymise C. nefrens (Smith, 2005) and C. cyanochthebius McCranie & Smith, 2006 with C. campbelli (Smith, 2005). The molecular data also support multiple undescribed species, notably within C. decoratus (Taylor, 1942). Overall, we show how morphology-based species delimitation can both underestimate and overestimate species richness in morphologically conservative groups. [ABSTRACT FROM AUTHOR]

Two major types of species richness patterns are spatial (e.g. the latitudinal diversity gradient) and clade-based (e.g. the dominance of angiosperms among plants). Studies have debated whether clade-based richness patterns are explained primarily by larger clades having faster rates of species accumulation (speciation minus extinction over time; diversification-rate hypothesis) or by simply being older (clade-age hypothesis). However, these studies typically compared named clades of the same taxonomic rank, such as phyla and families. This study design is potentially biased against the clade-age hypothesis, since clades of the same rank may be more similar in age than randomly selected clades. Here, we analyse the causes of clade-based richness patterns across the tree of life using a large-scale, time-calibrated, species-level phylogeny and random sampling of clades. We find that within major groups of organisms (animals, plants, fungi, bacteria, archaeans), richness patterns are most strongly related to clade age. Nevertheless, weaker relationships with diversification rates are present in animals and plants. These overall results contrast with similar large-scale analyses across life based on named clades, which showed little effect of clade age on richness. More broadly, these results help support the overall importance of time for explaining diverse types of species richness patterns. [ABSTRACT FROM AUTHOR]

Understanding the origins of species richness patterns is a fundamental goal in ecology and evolutionary biology. Much research has focused on explaining two kinds of species richness patterns: (i) spatial species richness patterns (e.g. the latitudinal diversity gradient), and (ii) clade‐based species richness patterns (e.g. the predominance of angiosperm species among plants). Here, I highlight a third kind of richness pattern: trait‐based species richness (e.g. the number of species with each state of a character, such as diet or body size). Trait‐based richness patterns are relevant to many topics in ecology and evolution, from ecosystem function to adaptive radiation to the paradox of sex. Although many studies have described particular trait‐based richness patterns, the origins of these patterns remain far less understood, and trait‐based richness has not been emphasised as a general category of richness patterns. Here, I describe a conceptual framework for how trait‐based richness patterns arise compared to other richness patterns. A systematic review suggests that trait‐based richness patterns are most often explained by when each state originates within a group (i.e. older states generally have higher richness), and not by differences in transition rates among states or faster diversification of species with certain states. This latter result contrasts with the widespread emphasis on diversification rates in species‐richness research. I show that many recent studies of spatial richness patterns are actually studies of trait‐based richness patterns, potentially confounding the causes of these patterns. Finally, I describe a plethora of unanswered questions related to trait‐based richness patterns. [ABSTRACT FROM AUTHOR]

How many species are there on Earth and to what groups do these species belong? These fundamental questions span systematics, ecology, and evolutionary biology. Yet, recent estimates of overall global biodiversity have ranged wildly, from the low millions to the trillions. Insects are a pivotal group for these estimates. Insects make up roughly half of currently described extant species (across all groups), with ~1 million described species. Insect diversity is also crucial because many other taxa have species that may be unique to each insect host species, including bacteria, apicomplexan protists, microsporidian fungi, nematodes, and mites. Several projections of total insect diversity (described and undescribed) have converged on ~6 million species. However, these projections have not incorporated the morphologically cryptic species revealed by molecular data. Here, we estimate the extent of cryptic insect diversity. We perform a systematic review of studies that used explicit species-delimitation methods with multilocus data. We estimate that each morphology-based insect species contains (on average) 3.1 cryptic species. We then use these estimates to project the overall number of species on Earth and their distribution among major groups. Our estimates suggest that overall global biodiversity may range from 563 million to 2.2 billion species. [Biodiversity; cryptic species; insects; species delimitation; species richness.] [ABSTRACT FROM AUTHOR]

Many sexually selected traits function as weapons, and these weapons can be incredibly diverse. However, the factors underlying weapon diversity among species remain poorly understood, and a fundamental hypothesis to explain this diversity remains untested. Although weapons can serve multiple functions, an undeniably important function is their role in fights. Thus, a crucial hypothesis is that weapon diversification is driven by the evolution of weapon modifications that provide an advantage in combat (e.g. causing more damage). Here, we test this fighting-advantage hypothesis using data from 17 species of coreid bugs. We utilize the fact that male–male combat in coreids often results in detectable damage, allowing us to link different weapon morphologies to different levels of damage among species. We find that certain weapon morphologies inflict much more damage than others, strongly supporting the fighting-advantage hypothesis. Moreover, very different weapon morphologies can inflict similarly severe amounts of damage, leading to a weapon performance landscape with multiple performance peaks. This multi-peak pattern could potentially drive different lineages towards divergent weapon forms, further increasing weapon diversity among species. Overall, our results may help explain how sexually selected weapons have evolved into the diversity of forms seen today. [ABSTRACT FROM AUTHOR]

A major goal of ecology and evolutionary biology is to explain geographic patterns of species richness. Richness is often correlated with climatic variables. However, the processes underlying these climate‐diversity relationships remain poorly understood. Two potential hypotheses to explain these relationships involve: (i) faster diversification rates (speciation minus extinction) in high‐richness climates and (ii) earlier colonization of high‐richness climates, allowing more time for speciation to build up richness. Few studies have tested these hypotheses directly, and most focused on animal clades with limited richness. In this study, we test these hypotheses in Chinese angiosperms, encompassing ~10% of Earth's plant species, using large‐scale phylogenetic, climatic, and distributional data including 26,977 species. We find that climatic zones that were colonized earlier have higher species richness. By contrast, relationships between diversification rates and richness of climatic zones are often nonsignificant or negative. Our study reveals that even when richness is strongly correlated with climate, the underlying explanation may still be rooted in phylogenetic history. Thus, climate may not be a competing explanation for richness patterns relative to colonization times and diversification rates. We also show that the timing of colonization can be crucial for explaining richness patterns. Yet, many recent studies have ignored this explanation and instead have focused solely on rates of speciation and diversification as drivers of diversity gradients. [ABSTRACT FROM AUTHOR]

Many biodiversity studies focus on explaining high tropical species richness, but an equally dramatic yet understudied pattern involves the divergent richness of land, sea and freshwater. Here, we reveal the origins of these richness differences among habitats across animals and plants. Most plant and animal species are terrestrial, although these habitats cover only ~28% of Earth's surface. Marine habitats have fewer species over a larger area (~70%). Freshwater habitats have relatively high richness and exceptional phylogenetic diversity given their tiny area (2%). The relative richness of habitats is related to variation in diversification rates. Based on ancestral reconstructions of habitat, we find that most marine species are descended from marine ancestors and most terrestrial species from freshwater ancestors. Yet, most extant animal richness in freshwater is derived from terrestrial ancestors. Overall, our results reveal the origins of fundamental but neglected biodiversity patterns, and highlight the conservation importance of freshwater habitats. [ABSTRACT FROM AUTHOR]

Aim: A major challenge in ecology and evolutionary biology is to explain the dramatic differences in species richness among clades. Much variation in richness is explained by the differences in diversification rates among clades, and variation in diversification rates is often linked to various traits. But what types of traits are most important for explaining diversification? Here, we compared the impacts of different types of traits on diversification rates among lizard and snake families and tested predictions about the relative importance of ecology vs. morphology, static vs. dynamic traits and alpha vs. beta niche traits. Location: Global. Taxon: Squamata. Methods: We compared the relative impacts of traits related to biogeography (range size, range expansion), climate, life history (viviparity), microhabitat and morphology (body‐size) on diversification rates among all 72 family‐level clades of squamates. We compiled data on traits and tested for relationships between traits and diversification rates using phylogenetic multiple regression models. Results: The best‐fitting model explained ~60% of the variation in diversification rates across squamate families. This model included only microhabitat (proportion of arboreal species) and a novel, dynamic, ecological/biogeographic beta‐niche trait (rate of range expansion), which explained most variance. Other variables had more variable or non‐significant contributions, including rates of climatic‐niche change. Rates of range expansion were related to species richness, larger body size and faster rates of climatic‐niche change. Main conclusions: Overall, we provide possibly the most comprehensive comparison of the types of traits that can drive diversification. We also help explain diversity patterns in one of the largest vertebrate clades. We show that the rate of range expansion is the most important variable for explaining diversification rates and richness patterns in squamates. We also identify traits that help explain variation in rates of range expansion among clades. [ABSTRACT FROM AUTHOR]

The paleotropics harbor many biodiversity hotspots and show many different species richness patterns. However, it remains unclear which factors are the most important in directly shaping richness patterns among regions in the paleotropics (i.e. diversification rates, colonization times, and dispersal frequency). Here we used Cycadaceae as a model system to test the causes of regional richness patterns in the paleotropics. Specifically, we tested the roles of dispersal frequency, colonization time, diversification rates, and their combined role in explaining richness patterns among regions. We generated a well‐sampled, time‐calibrated phylogeny and then used this to estimate dispersal events, colonization times, and diversification rates. Richness patterns were significantly associated with the timing of the first colonization of each region and were best explained by the combined effects of colonization time and diversification rates. The number of dispersal events into each region and the diversification rates of species in each region were not significantly related to richness. Ancestral‐area reconstructions showed frequent migrations across Wallace's line, with a higher diversification rate east of Wallace's line than west of it. Overall, our study shows that colonization time can be an important factor for explaining regional richness patterns in the paleotropics. [ABSTRACT FROM AUTHOR]

Across the Tree of Life, there are dramatic differences in species numbers among groups. However, the factors that explain the differences among the deepest branches have remained unknown. We tested whether multicellularity and sexual reproduction might explain these patterns, since the most species-rich groups share these traits. We found that groups with multicellularity and sexual reproduction have accelerated rates of species proliferation (diversification), and that multicellularity has a stronger effect than sexual reproduction. Patterns of species richness among clades are then strongly related to these differences in diversification rates. Taken together, these results help explain patterns of biodiversity among groups of organisms at the very broadest scales. They may also help explain the mysterious preponderance of sexual reproduction among species (the 'paradox of sex') by showing that organisms with sexual reproduction proliferate more rapidly. [ABSTRACT FROM AUTHOR]

A fundamental goal of ecology is to reveal generalities in the myriad types of interactions among species, such as competition, mutualism and predation. Another goal is to explain the enormous differences in species richness among groups of organisms. Here, we show how these two goals are intertwined: we find that different types of species interactions have predictable impacts on rates of species diversification, which underlie richness patterns. On the basis of a systematic review, we show that interactions with positive fitness effects for individuals of a clade (e.g. insect pollination for plants) generally increase that clade's diversification rates. Conversely, we find that interactions with negative fitness effects (e.g. predation for prey, competition) generally decrease diversification rates. The sampled clades incorporate all animals and land plants, encompassing 90% of all described species across life. Overall, we show that different types of local‐scale species interactions can predictably impact large‐scale patterns of diversification and richness. [ABSTRACT FROM AUTHOR]

Many sexually selected traits function as weapons, and these weapons can be incredibly diverse. However, the factors underlying weapon diversity among species remain poorly understood, and a fundamental hypothesis to explain this diversity remains untested. Although weapons can serve multiple functions, an undeniably important function is their role in fights. Thus, a crucial hypothesis is that weapon diversification is driven by the evolution of weapon modifications that provide an advantage in combat (e.g. causing more damage). Here, we test this fighting-advantage hypothesis using data from 17 species of coreid bugs. We utilize the fact that male–male combat in coreids often results in detectable damage, allowing us to link different weapon morphologies to different levels of damage among species. We find that certain weapon morphologies inflict much more damage than others, strongly supporting the fighting-advantage hypothesis. Moreover, very different weapon morphologies can inflict similarly severe amounts of damage, leading to a weapon performance landscape with multiple performance peaks. This multi-peak pattern could potentially drive different lineages towards divergent weapon forms, further increasing weapon diversity among species. Overall, our results may help explain how sexually selected weapons have evolved into the diversity of forms seen today. [ABSTRACT FROM AUTHOR]

Niche breadth is a unifying concept spanning diverse aspects of ecology, evolution, and conservation biology. Niche breadth usually refers to the diversity of resources used or environments tolerated by an individual, population, species, or clade. Here we review key research in ecology, evolution, and conservation biology in light of niche breadth. Namely, we explore the role of niche breadth in shaping geographic distributions and species richness from local to landscape scales, how niche breadth evolves and influences lineage diversification, and its use for understanding species invasions, responses to climate change, vulnerability to extinction, and ecosystem functioning. This diverse literature informs a research agenda that identifies focused needs for further progress: testing the hierarchical nature of niche breadth (e.g., of individuals, populations, and species); quantifying correlations in niche breadth among different niche axes and the role of environmental drivers and organismal constraints in generating these correlations; and evaluating the factors that decouple fundamental and realized niches. We describe how this research agenda could help unify disparate subdisciplines and shed light on key questions in ecology, evolution, and conservation. [ABSTRACT FROM AUTHOR]

What an animal eats is a fundamental aspect of its biology, but the evolution of diet has not been studied across animal phylogeny. Here, we performed a large‐scale phylogenetic analysis to address three unresolved questions about the evolution of animal diets. (i) Are diets conserved across animal phylogeny? (ii) Does diet influence rates of species proliferation (diversification) among animal phyla? (iii) What was the ancestral diet of animals and major animal clades? We analyzed diet data for 1087 taxa, proportionally sampled among animal phyla based on the relative species richness of phyla. Our survey suggests that across animals, carnivory is most common (∼63%), herbivory less common (∼32%), and omnivory relatively rare (∼3%). Despite considerable controversy over whether ecological traits are conserved or labile, we found strong conservatism in diet over extraordinarily deep timescales. We found that diet is unrelated to rates of species diversification across animal phyla, contrasting with previous studies showing that herbivory increased diversification within some important groups (e.g., crustaceans, insects, and mammals). Finally, we estimated that the ancestor of all animals was most likely carnivorous, as were many major phyla (e.g., arthropods, molluscs, and chordates). Remarkably, our results suggest that many carnivorous species living today may have maintained this diet through a continuous series of carnivorous ancestors for >800 million years. [ABSTRACT FROM AUTHOR]

Most groups of organisms occur in multiple regions and have different numbers of species in different regions. These richness patterns are directly explained by speciation, extinction, and dispersal. Thus, regional richness patterns may be explained by differences in when regions were colonized (more time for speciation in regions colonized earlier), differences in how often they were colonized, or differences in diversification rates (speciation minus extinction) among regions (with diversification rates potentially influenced by area, climate, and/or many other variables). Few studies have tested all three factors, and most that did examined them only in individual clades. Here, we analyze a diverse set of 15 clades of plants and animals to test the causes of regional species richness patterns within clades. We find that time was the sole variable significantly explaining richness patterns in the best-fitting models for most clades (10/15), whereas time combined with other factors explained richness in all others. Time was the most important factor explaining richness in 13 of 15 clades, and it explained 72% of the variance in species richness among regions across all 15 clades (on average). Surprisingly, time was increasingly important in older and larger clades. In contrast, the area of the regions was relatively unimportant for explaining these regional richness patterns. A systematic review yielded 15 other relevant studies, which also overwhelmingly supported time over diversification rates (13 to 1, with one study supporting both diversification rates and time). Overall, our results suggest that colonization time is a major factor explaining regional-scale richness patterns within clades (e.g., families). [ABSTRACT FROM AUTHOR]

In a previous paper, we used simulations and empirical data to show that BAMM (Bayesian Analysis of Macroevolutionary Mixtures) can give misleading estimates of rates and rate shifts. In simulations, BAMM underestimated rate shifts across every tree analyzed, and assigned incorrect rates to most clades in most trees. In empirical analyses, BAMM behaved as expected from simulations, and assigned different rates to clades when clades were analyzed alone versus across the tree (i.e., with rate heterogeneity). Rabosky recently criticized our paper, focusing primarily on the idea that our comparison of BAMM to another approach (method‐of‐moments estimators of Magallón and Sanderson, or MS estimators) was unfair to BAMM. Here, we provide further evidence that BAMM gives misleading rate estimates in empirical studies. We then describe how Rabosky's rown method comparisons were either acknowledged as being problematic or were described inaccurately (to favor BAMM). Finally, we show that the MS estimators can perform well when rates vary over time, despite untested assertions that they require constant rates to be accurate. Many other methods are available for analyzing diversification rates: we argue that BAMM should be avoided for estimating both diversification rates and rate shifts. [ABSTRACT FROM AUTHOR]

The Tropical Andes make up Earth's most species-rich biodiversity hotspot for both animals and plants. Nevertheless, the ecological and evolutionary processes underlying this extraordinary richness remain uncertain. Here, we examine the processes that generate high richness in the Tropical Andes relative to other regions in South America and across different elevations within the Andes, using frogs as a model system. We combine distributional data, a newly generated time-calibrated phylogeny for 2,318 frog species, and phylogenetic comparative methods to test the relative importance of diversification rates and colonization times for explaining Andean diversity at different scales. At larger scales (among regions and families), we find that faster diversification rates in Andean clades most likely explain high Andean richness. In contrast, at smaller temporal and spatial scales (within family-level clades within the Andes), diversification rates rarely explain richness patterns. Instead, we show that colonization times are important for shaping elevational richness patterns within the Andes, with more species found in habitats colonized earlier. We suggest that these scale-dependent patterns might apply to many other richness gradients. Recognition of this scale dependence may help to reconcile conflicting results among studies of richness patterns across habitats, regions, and organisms. [ABSTRACT FROM AUTHOR]

Patterns of species richness among clades can be directly explained by the ages of clades or their rates of diversification. The factors that most strongly influence diversification rates remain highly uncertain, since most studies typically consider only a single predictor variable. Here, we explore the relative impacts of macroclimate (i.e., occurring in tropical vs. temperate regions) and microhabitat use (i.e., terrestrial, fossorial, arboreal, aquatic) on diversification rates of squamate reptile clades (lizards and snakes). We obtained data on microhabitat, macroclimatic distribution, and phylogeny for >4000 species. We estimated diversification rates of squamate clades (mostly families) from a time-calibrated tree, and used phylogenetic methods to test relationships between diversification rates and microhabitat and macroclimate. Across 72 squamate clades, the best-fitting model included microhabitat but not climatic distribution. Microhabitat explained ∼37% of the variation in diversification rates among clades, with a generally positive impact of arboreal microhabitat use on diversification, and negative impacts of fossorial and aquatic microhabitat use. Overall, our results show that the impacts of microhabitat on diversification rates can be more important than those of climate, despite much greater emphasis on climate in previous studies. [ABSTRACT FROM AUTHOR]

Aims Patterns of species richness, such as the remarkable biodiversity of tropical regions, have been documented and studied for centuries. However, their underlying evolutionary and ecological causes are still incompletely understood. A commonly stated paradigm in the literature is that high richness in some habitats is directly caused by one of three competing explanations: (1) greater time-for-speciation (earlier colonization), (2) more rapid diversification rates (faster speciation relative to extinction) or (3) higher carrying capacity. However, these three explanations have been relatively little studied using theoretical approaches (especially in terms of comparing all three). Furthermore, empirical studies give conflicting results about their relative importance. Here, we use simulations to study the processes that drive richness patterns along environmental gradients. Location Globally applicable. Methods We use individual-based and trait-based modelling of eco-evolutionary dynamics to simulate the evolutionary radiation of a clade across five habitats with differing ecological conditions, and track patterns of species richness within and between habitats over time. We specifically address the roles of time and diversification rates in explaining richness patterns and the potential impact of carrying capacity. Main results and conclusions Contrary to the widespread paradigm, we find that variation in carrying capacity can underlie differences in diversification rates and time-for-speciation among habitats. Therefore, carrying capacity is not a competing, alternative explanation for richness patterns. We also find that the time-for-speciation effect dominates richness patterns over short time-scales, whereas diversification rates dominate over longer time-scales. These latter observations can help reconcile the seemingly conflicting results of many empirical studies, which find that some patterns are explained by time and others by differences in diversification rates. [ABSTRACT FROM AUTHOR]

A major goal of evolutionary biology and ecology is to understand why species richness varies among clades. Previous studies have suggested that variation in richness among clades might be related to variation in rates of morphological evolution among clades (e.g., body size and shape). Other studies have suggested that richness patterns might be related to variation in rates of climatic-niche evolution. However, few studies, if any, have tested the relative importance of these variables in explaining patterns of richness among clades. Here, we test their relative importance among major clades of Plethodontidae, the most species-rich family of salamanders. Earlier studies have suggested that climatic-niche evolution explains patterns of diversification among plethodontid clades, whereas rates of morphological evolution do not. A subsequent study stated that rates of morphological evolution instead explained patterns of species richness among plethodontid clades (along with 'ecological limits' on richness of clades, leading to saturation of clades with species, given limited resources). However, they did not consider climatic-niche evolution. Using phylogenetic multiple regression, we show that rates of climatic-niche evolution explain most variation in richness among plethodontid clades, whereas rates of morphological evolution do not. We find little evidence that ecological limits explain patterns of richness among plethodontid clades. We also test whether rates of morphological and climatic-niche evolution are correlated, and find that they are not. Overall, our results help explain richness patterns in a major amphibian group and provide possibly the first test of the relative importance of climatic niches and morphological evolution in explaining diversity patterns. [ABSTRACT FROM AUTHOR]

Terrestrial environments occupy ~ 30% of the Earth's surface yet contain ~ 80% of all species. The causes of this dramatic biodiversity gradient have remained relatively unstudied. Here, I test the fundamental prediction that predominantly non-marine clades have more rapid rates of diversification than marine clades, using a time-calibrated phylogeny of animal phyla. The results strongly support this hypothesis. This pattern helps explain the higher richness of terrestrial environments and the dramatic variation in species richness among animal phyla. The results show the importance of ecology in explaining large-scale patterns of clade richness and of diversification rates in explaining Earth's largest biodiversity patterns. The results also demonstrate remarkable niche conservatism in habitats, in some cases lasting > 800 million years. Finally, the results highlight the surprisingly high species richness of freshwater habitats, which are nearly equal to marine environments despite their much smaller area (~ 2% of Earth's surface vs. 70% for marine habitats). [ABSTRACT FROM AUTHOR]

Time-calibrated phylogenies have become essential to evolutionary biology. A recurrent and unresolved question for dating analyses is whether genes with missing data cells should be included or excluded. This issue is particularly unclear for the most widely used dating method, the uncorrelated lognormal approach implemented in BEAST. Here, we test the robustness of this method to missing data. We compare divergence-time estimates from a nearly complete dataset (20 nuclear genes for 32 species of squamate reptiles) to those from subsampled matrices, including those with 5 or 2 complete loci only and those with 5 or 8 incomplete loci added. In general, missing data had little impact on estimated dates (mean error of ∼5 Myr per node or less, given an overall age of ∼220 Myr in squamates), even when 80% of sampled genes had 75% missing data. Mean errors were somewhat higher when all genes were 75% incomplete (∼17 Myr). However, errors increased dramatically when only 2 of 9 fossil calibration points were included (∼40 Myr), regardless of missing data. Overall, missing data (and even numbers of genes sampled) may have only minor impacts on the accuracy of divergence dating with BEAST, relative to the dramatic effects of fossil calibrations. [ABSTRACT FROM AUTHOR]

Aim Diversification rates are critically important for understanding patterns of species richness, both among clades and among regions. However, the ecological correlates of variation in diversification rates remain poorly explored. Here, we test several hypotheses relating diversification rate and niche width across amphibian families (frogs and salamanders). Location Global. Methods We characterized climatic niches for 5784 amphibian species using databases for species distributions and climate. We estimated the niche width of each family using the range of values for climatic variables across all sampled species, and using the mean of species niche widths. We estimated diversification rates for families given their total number of described species and a time-calibrated phylogeny. We estimated relationships between variables using phylogenetic comparative methods. Results We found a significant positive relationship between family niche width and diversification rate, but a weak relationship between mean species niche width and diversification rate, despite both niche width variables being correlated. In fact, the deviation from this relationship (i.e. residuals of family niche width versus mean species niche width) was the best predictor of diversification rate. The observed relationship between niche width and diversification was independent of clade range size and niche position (e.g. whether clades occurred in tropical or temperate climates) and significantly different from null patterns derived from random sampling effects. Main conclusions Our results identify climatic niche width, and especially the relationship between family and species-level niche widths, as a major correlate of diversification rates among amphibian families. These results suggest that climatic niche divergence among species within clades can be important in explaining large-scale diversity patterns, possibly even more so than a clade's geographic area or whether it is primarily temperate or tropical. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Saturation is the idea that a community is effectively filled with species, such that no more can be added without extinctions. This concept has important implications for many areas of ecology, such as species richness, community assembly, invasive species and climate change. Here, we illustrate how biogeography can be used to test for community saturation, when combined with data on local species richness, phylogeny and climate. We focus on a clade of frogs (Terrarana) and the impact of the Great American Biotic Interchange on patterns of local richness in Lower Middle America and adjacent regions. We analyse data on species richness at 83 sites and a time-calibrated phylogeny for 363 species. We find no evidence for saturation, and show instead that biotic interchange dramatically increased local richness in the region. We suggest that historical biogeography offers thousands of similar long-term natural experiments that can be used to test for saturation. [ABSTRACT FROM AUTHOR]

The Tropical Andes are an important global biodiversity hotspot, harbouring extraordinarily high richness and endemism. Although elevational richness and speciation have been studied independently in some Andean groups, the evolutionary and ecological processes that explain elevational richness patterns in the Andes have not been analysed together. Herein, we elucidate the processes underlying Andean richness patterns using glassfrogs (Centrolenidae) as a model system. Glassfrogs show the widespread mid-elevation diversity peak for both local and regional richness. Remarkably, these patterns are explained by greater time (montane museum) rather than faster speciation at mid-elevations (montane species pump), despite the recency of the major Andean uplift. We also show for the first time that rates of climatic-niche evolution and elevational change are related, supporting the hypothesis that climatic-niche conservatism decelerates species' shifts in elevational distributions and underlies the mid-elevation richness peak. These results may be relevant to other Andean clades and montane systems globally. [ABSTRACT FROM AUTHOR]

Aim Climatic niche breadth (the range of climatic conditions that a species experiences over space and time) is a fundamental topic in ecology, biogeography and evolution. But what determines the climatic niche width of species? In 1967, Janzen suggested that climatic niche widths for temperature were determined by levels of seasonal fluctuation in temperature at each locality, such that niche breadths are narrow in tropical species and broad in temperate species. However, it is unclear whether climatic niche breadths of species are determined more by seasonal variability within sites as opposed to climatic variation between sites across the species' range. We address this question here. Location Global. Methods We analysed three vertebrate clades (plethodontid salamanders, hylid frogs and phrynosomatid lizards) for which we had phylogenetic information and climatic data from localities throughout each species' geographic range, collectively including 409 species. We tested how climatic niche breadths of localities (i.e. temporal variation) are related to overall species climatic niche breadths (i.e. temporal and spatial variation) using phylogenetic comparative methods, focusing both on temperature extremes and precipitation. Results Across the three clades, we find that niche breadths for single localities generally span most of the species' climatic niche breadth, and are strongly correlated with overall species niche breadths. However, species with wider climatic niches also tend to show greater climatic divergence between localities. Main conclusions The extent to which the climatic niche breadths of species are determined by variation within localities versus spatial variation between localities has been largely unexplored. Our results suggest that within-locality seasonal variation explains most variation in climatic niche breadths among species. However, between-locality variation and local adaptation may also play some role. These results require more general testing, but have several important implications. [ABSTRACT FROM AUTHOR]

Parallel evolutionary radiations in adjacent locations have been documented in many systems, but typically at limited geographical scales. Here, we compare patterns of evolutionary radiation at the global scale in iguanian lizards, the dominant clade of lizards. We generated a new time-calibrated phylogeny including 153 iguanian species (based on mitochondrial and nuclear data) and obtained data on morphology and microhabitats. We then compared patterns of species diversification, morphological disparity, and ecomorphological relationships in the predominantly Old World and New World clades ( Acrodonta and Pleurodonta, respectively), focusing on the early portions of these radiations. Acrodonts show relatively constant rates of species diversification and disparity over time. In contrast, pleurodonts show an early burst of species diversification and less-than-expected morphological disparity early in their history, and slowing diversification and increasing disparity more recently. Analyses including all species (with MEDUSA) suggest accelerated diversification rates in certain clades within both Acrodonta and Pleurodonta, which strongly influences present-day diversity patterns. We also find substantial differences in ecomorphological relationships between these clades. Our results demonstrate that sister clades in different global regions can undergo very different patterns of evolutionary radiation over similar time frames. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●●, ●●-●●. [ABSTRACT FROM AUTHOR]

The rate of climatic-niche evolution is important to many research areas in ecology, evolution, and conservation biology, including responses of species to global climate change, spread of invasive species, speciation, biogeography, and patterns of species richness. Previous studies have implied that clades with higher rates of climatic-niche evolution among species should have species with narrower niche breadths, but there is also evidence suggesting the opposite pattern. However, the relationships between rate and breadth have not been explicitly analyzed. Here, we examine the relationships between the rate of climatic-niche evolution and climatic-niche breadth using phylogenetic and climatic data for 250 species in the salamander family Plethodontidae, a group showing considerable variation in both rates of climatic-niche evolution and climatic-niche breadths. Contrary to some expectations, we find no general relationship between climatic-niche breadth and the rate of climatic-niche evolution. Climatic-niche breadths for some ecologically important climatic variables considered separately (temperature seasonality and annual precipitation) do show significant relationships with the rate of climatic-niche evolution, but rates are faster in clades in which species have broader (not narrower) niche breadths. In summary, our results show that narrower niche breadths are not necessarily associated with faster rates of niche evolution. [ABSTRACT FROM AUTHOR]

Abstract: The treefrogs (Hylidae) make up one of the most species-rich families of amphibians. With 885 species currently described, they contain >13% of all amphibian species. In recent years, there has been considerable progress in resolving hylid phylogeny. However, the most comprehensive phylogeny to date () included only 292 species, was based only on parsimony, provided only poor support for most higher-level relationships, and conflicted with previous hypotheses in several parts (including the monophyly and relationships of major clades of Hylinae). Here, we present an expanded phylogeny for hylid frogs, including data for 362 hylid taxa for up to 11 genes (4 mitochondrial, 7 nuclear), including 70 additional taxa and >270 sequences not included in the previously most comprehensive analysis. The new tree from maximum likelihood analysis is more well-resolved, strongly supported, and concordant with previous hypotheses, and provides a framework for future systematic, biogeographic, ecological, and evolutionary studies. [Copyright &y& Elsevier]

Information about several papers discussed at a symposium of the Society of Systematic Biologists on species delimitation is presented. Species delimitation, the process by which species boundaries are determined and new species are discovered, is emerging as a major topic in modern systematics, and new methods for species delimitation are being developed. The symposium featured several biologists including Keith Crandall, Paul Hebert, Marshal Hedin, Loren Rieseberg and Jack Sullivan.