Variation In Diversification Rates Research Articles

Imbalanced speciation pulses sustain the radiation of mammals.

The evolutionary histories of major clades, including mammals, often comprise changes in their diversification dynamics, but how these changes occur remains debated. We combined comprehensive phylogenetic and fossil information in a new "birth-death diffusion" model that provides a detailed characterization of variation in diversification rates in mammals. We found an early rising and sustained diversification scenario, wherein speciation rates increased before and during the Cretaceous-Paleogene (K-Pg) boundary. The K-Pg mass extinction event filtered out more slowly speciating lineages and was followed by a subsequent slowing in speciation rates rather than rebounds. These dynamics arose from an imbalanced speciation process, with separate lineages giving rise to many, less speciation-prone descendants. Diversity seems to have been brought about by these isolated, fast-speciating lineages, rather than by a few punctuated innovations.

Tropical Origin, Global Diversification and Dispersal in the Pond Damselflies (Coenagrionoidea) Revealed by a New Molecular Phylogeny.

The processes responsible for the formation of Earth's most conspicuous diversity pattern, the latitudinal diversity gradient (LDG), remain unexplored for many clades in the Tree of Life. Here, we present a densely-sampled and dated molecular phylogeny for the most speciose clade of damselflies worldwide (Odonata: Coenagrionoidea), and investigate the role of time, macroevolutionary processes and biome-shift dynamics in shaping the LDG in this ancient insect superfamily. We used process-based biogeographic models to jointly infer ancestral ranges and speciation times, and to characterise within-biome dispersal and biome-shift dynamics across the cosmopolitan distribution of Coenagrionoidea. We also investigated temporal and biome-dependent variation in diversification rates. Our results uncover a tropical origin of pond damselflies and featherlegs ~ 105 Ma, while highlighting uncertainty of ancestral ranges within the tropics in deep time. Even though diversification rates have declined since the origin of this clade, global climate change and biome-shifts have slowly increased diversity in warm- and cold-temperate areas, where lineage turnover rates have been relatively higher. This study underscores the importance of biogeographic origin and time to diversify as important drivers of the LDG in pond damselflies and their relatives, while diversification dynamics have instead resulted in the formation of ephemeral species in temperate regions. Biome-shifts, although limited by tropical niche conservatism, have been the main factor reducing the steepness of the LDG in the last 30 Myr. With ongoing climate change and increasing northward range expansions of many damselfly taxa, the LDG may become less pronounced. Our results support recent calls to unify biogeographic and macroevolutionary approaches to increase our understanding of how latitudinal diversity gradients are formed and why they vary across time and among taxa.

All about being old and shooting hairs: clade age and urticating hair explain the patterns of diversification in tarantulas.

The extreme asymmetry of species richness distribution across the tree of life has always intrigued evolutionary biologists. Two competing explanations have been proposed to explain this pattern-the clade age hypothesis and diversification rate variation. While these two scenarios may not be mutually exclusive, to what extent time and diversification rates interact to explain species richness patterns remains understudied. Here, we investigate the relative influence of these two scenarios using tarantulas (Family: Theraphosidae) as a model. Tarantulas represent a speciose group of spiders found worldwide but exceptionally diverse in South America. These spiders show two distinct patterns of microhabitat use (ground-dwelling or arboreal) and defense strategies (presence or absence of urticating hairs). Using various trait-independent and dependent diversification models, we test the clade age hypothesis, the role of microhabitat, antipredator defense strategy, and geography in influencing diversification rates. Our results suggest that clade age is the primary predictor of species richness distribution across the tarantula subfamilies. However, the presence of urticating hair probably disrupted this pattern in some clades by increasing the net diversification rates, not by increasing the speciation rate but by reducing the extinction rate.

Paleoenvironmental models for Australia and the impact of aridification on blindsnake diversification

AbstractAimShifts in diversification rates of Australian flora and fauna have been associated with aridification, but the relationship between diversification rates and aridity has never been quantified. We employed multiple approaches to reconstruct paleoenvironments of Australia for the first time. We used this information, and phylogenetic‐based analyses, to explore how changes in temperature and increasing aridity during the Neogene influenced the diversification of the Australian blindsnakes. We tested whether diversification rates differ between arid‐adapted and mesic‐adapted lineages.TaxonTyphlopidae, Anilios blindsnakes.LocationAustralia.Materials and MethodsWe estimated the historical biogeography of blindsnakes using BioGeoBEARS. We synthesised multiple approaches to reconstruct paleotemperature and paleoaridity of Australia during the Neogene. We fitted several birth‐death models and estimated diversification rates under paleoenvironmental conditions using RPANDA. We further compared diversification rates between arid‐adapted lineages versus mesic‐adapted lineages using ClaDS and GeoHiSSE.ResultsAncestral area estimation indicated Australian blindsnakes have tropical grassland origins. We found that Australia‐specific regional paleotemperature and paleoaridity provided a better explanation for diversification rate variation than global paleotemperature. Specifically, our best‐fitting model indicated that speciation rates of blindsnakes decreased with increasing aridity. We found no difference in diversification rates between arid‐ and mesic‐adapted lineages.Main ConclusionsSoon after dispersing to Australia, the common ancestors of Australian blindsnakes diversified rapidly in mesic habitats during the early Miocene. However, as the continent became increasingly arid, diversification rates decreased. We found that shifts in the environment led to the emergence of two major clades: one remaining in primarily mesic habitats and the other adapting to the expanding arid biome. Our results emphasise the importance of both arid and tropical biomes as sources and sinks of diversification.

Combining palaeontological and neontological data shows a delayed diversification burst of carcharhiniform sharks likely mediated by environmental change

Estimating deep-time species-level diversification processes remains challenging. Both the fossil record and molecular phylogenies allow the estimation of speciation and extinction rates, but each type of data may still provide an incomplete picture of diversification dynamics. Here, we combine species-level palaeontological (fossil occurrences) and neontological (molecular phylogenies) data to estimate deep-time diversity dynamics through process-based birth–death models for Carcharhiniformes, the most speciose shark order today. Despite their abundant fossil record dating back to the Middle Jurassic, only a small fraction of extant carcharhiniform species is recorded as fossils, which impedes relying only on the fossil record to study their recent diversification. Combining fossil and phylogenetic data, we recover a complex evolutionary history for carcharhiniforms, exemplified by several variations in diversification rates with an early low diversity period followed by a Cenozoic radiation. We further reveal a burst of diversification in the last 30 million years, which is partially recorded with fossil data only. We also find that reef expansion and temperature change can explain variations in speciation and extinction through time. These results pinpoint the primordial importance of these environmental variables in the evolution of marine clades. Our study also highlights the benefit of combining the fossil record with phylogenetic data to address macroevolutionary questions.

Diversification dynamics of chameleons (Chamaeleonidae)

AbstractChameleons are charismatic and common lizards across Madagascar, Africa, and some surrounding regions. Little is known about their diversification dynamics and how this relates to their ecology, so we estimated diversification rate variation and consider this in the context of three hypotheses previously proposed in the literature. First, that the transoceanic dispersal from Africa to Madagascar on two separate occasions has resulted in fast radiation of Malagasy chameleons. Second, that the substantial floral turnover in their distributions within South Africa has resulted in rapid radiations of the endemic dwarf chameleons (Bradypodion). Finally, that the evolution of distinct ecomorphs of chameleon has fuelled fast diversification via adaptive radiations. We use the most recent and complete phylogeny of chameleons to estimate the diversification dynamics of the group using three methods: BAMM (which estimates constant or gradually changing diversification regimes and tests for shifts in these), MEDUSA (which tests for rate shifts in particular clades), and ClaDS (which estimates branch‐specific diversification rates). Our results from all analyses estimate a diversification rate increase in a clade containing most of the genusBradypodion, a group containing the South African dwarf chameleons which occur in recognized biodiversity hotspots in diverse habitats. We find no evidence for shifts resulting from dispersal events to Madagascar or related to the strong ecomorphological divergence of short‐tailed chameleon lineages (Brookesia,Palleon,Rhampholeon, andRieppeleon). The single burst of diversification within chameleons was in a clade which was associated with geographic areas which have experienced rapid habitat turnover and vicariance over the last ~10 million years. This suggests that ‘habitat vicariance’ resulting from ecological changes in vegetation has contributed to the diversity of species in this area by increasing diversification rates.

Global variation in diversification rate and species richness are unlinked in plants

Species richness varies immensely around the world. Variation in the rate of diversification (speciation minus extinction) is often hypothesized to explain this pattern, while alternative explanations invoke time or ecological carrying capacities as drivers. Focusing on seed plants, the world's most important engineers of terrestrial ecosystems, we investigated the role of diversification rate as a link between the environment and global species richness patterns. Applying structural equation modeling to a comprehensive distribution dataset and phylogenetic tree covering all circa 332,000 seed plant species and 99.9% of the world's terrestrial surface (excluding Antarctica), we test five broad hypotheses postulating that diversification serves as a mechanistic link between species richness and climate, climatic stability, seasonality, environmental heterogeneity, or the distribution of biomes. Our results show that the global patterns of species richness and diversification rate are entirely independent. Diversification rates were not highest in warm and wet climates, running counter to the Metabolic Theory of Ecology, one of the dominant explanations for global gradients in species richness. Instead, diversification rates were highest in edaphically diverse, dry areas that have experienced climate change during the Neogene. Meanwhile, we confirmed climate and environmental heterogeneity as the main drivers of species richness, but these effects did not involve diversification rates as a mechanistic link, calling for alternative explanations. We conclude that high species richness is likely driven by the antiquity of wet tropical areas (supporting the "tropical conservatism hypothesis") or the high ecological carrying capacity of warm, wet, and/or environmentally heterogeneous environments.

Diversification in the Rosales is influenced by dispersal, geographic range size, and pre‐existing species richness

Biodiversity results from origination and extinction, justifying interest in identifying traits that influence this balance. Traits implicated in the success or failure of lineages include dispersal, colonization ability, and geographic range size. We investigated the impact of dispersal and range size on contemporary diversity in the Rosales. We used the multiple-state speciation and extinction (MuSSE) method to explore the effects on genus-level diversification of two genus-level traits (geographic range size and within-genus proclivity to speciate) and two species traits (seed dispersal and growth habit) and the multiple hidden-state speciation and extinction (MuHiSSE) method for species-level associations. Finally, we conducted a PGLS (phylogenetic least-squares) analysis to distinguish between speciation within genera versus origination of new genera. At the species level, animal dispersal enhances diversification rate in both woody and herbaceous lineages, while woody lineages without animal dispersal have higher extinction rates than speciation rates. At the genus level, herbaceous taxa have positive diversification rates regardless of other character states. Diversification rate variation is also explained by two interactions: (1) a three-way interaction between large geographic range, animal-mediated dispersal, and high within-genus species richness, whereby genera possessing all three traits have high diversification rates, and (2) a four-way interaction by which the three-way interaction is stronger in woody genera than in herbaceous genera. Colonization ability may underlie the relationship between dispersal type and range size and may influence past diversification rates by decreasing extinction rates during late Cenozoic climate volatility. Thus, colonization ability could be used to predict future extinction risk to aid conservation.

Biogeography of a neotropical songbird radiation reveals similar diversification dynamics between montane and lowland clades

AbstractAimContinental evolutionary radiations provide opportunities to understand how landscape evolution and biotic factors interact to generate species diversity. Additionally, understanding whether diversification dynamics differ between montane and lowland environments is a long‐standing question with few comparative analyses in the Neotropics. To address these questions, we investigated the biogeographical patterns and the evolutionary processes underlying the diversification of a songbird genus, and compared diversification dynamics of clades occurring in lowland and montane Neotropical habitats.LocationNeotropical montane and lowland forests.TaxonArremon(Aves: Passerellidae).MethodsWe sequenced genomic data (ultra‐conserved elements, UCEs) of 92 individuals (including historical skin specimens) comprising 47 of 50 currently recognized subspecies in the genus and collected habitat association data to (1) build the most complete phylogenetic hypothesis for the genus to date using maximum likelihood and Bayesian methods with a concatenated matrix, and a multi‐species coalescent method based on quartets; and (2) reconstruct the evolution of their ancestral ranges, habitat association and diversification rates.ResultsAll phylogenetic methods recovered essentially the same topology with strong support values for most interspecific nodes revealing relationships among species. We found evidence for a montane and humid ancestral range in Central America in the late Miocene and a later expansion into the lowlands of Central America, as well as into the lowlands and mountains of South America. Despite some temporal variation in diversification rate, we found overall similar diversification dynamics between montane and lowland clades.Main conclusionSpecies diversity within the genus is likely underestimated by the current taxonomic arrangement. The colonization of lowlands and dry forests, and expansion across South America, may have provided new geographical and ecological opportunities for speciation resulting in high species diversification. Overall diversification dynamics were comparable between montane and lowland clades, contrasting with previous studies focused on such comparisons for Neotropical birds.

Exceptional evolutionary lability of flower-like inflorescences (pseudanthia) in Apiaceae subfamily Apioideae.

PremisePseudanthia are widespread and have long been postulated to be a key innovation responsible for some of the angiosperm radiations. The aim of our study was to analyze macroevolutionary patterns of these flower‐like inflorescences and their potential correlation with diversification rates in Apiaceae subfamily Apioideae. In particular, we were interested to investigate evolvability of pseudanthia and evaluate their potential association with changes in the size of floral display.MethodsThe framework for our analyses consisted of a time‐calibrated phylogeny of 1734 representatives of Apioideae and a morphological matrix of inflorescence traits encoded for 847 species. Macroevolutionary patterns in pseudanthia were inferred using Markov models of discrete character evolution and stochastic character mapping, and a principal component analysis was used to visualize correlations in inflorescence architecture. The interdependence between net diversification rates and the occurrence of pseudocorollas was analyzed with trait‐independent and trait‐dependent approaches.ResultsPseudanthia evolved in 10 major clades of Apioideae with at least 36 independent origins and 46 reversals. The morphospace analysis recovered differences in color and compactness between floral and hyperfloral pseudanthia. A correlation between pseudocorollas and size of inflorescence was also strongly supported. Contrary to our predictions, pseudanthia are not responsible for variation in diversification rates identified in this subfamily.ConclusionsOur results suggest that pseudocorollas evolve as an answer to the trade‐off between enlargement of floral display and costs associated with production of additional flowers. The high evolvability and architectural differences in apioid pseudanthia may be explained on the basis of adaptive wandering and evolutionary developmental biology.

What drives diversification? Range expansion tops climate, life history, habitat and size in lizards and snakes

AbstractAimA 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.LocationGlobal.TaxonSquamata.MethodsWe 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.ResultsThe 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 conclusionsOverall, 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.

Radiating pain: venom has contributed to the diversification of the largest radiations of vertebrate and invertebrate animals

BackgroundUnderstanding drivers of animal biodiversity has been a longstanding aim in evolutionary biology. Insects and fishes represent the largest lineages of invertebrates and vertebrates respectively, and consequently many ideas have been proposed to explain this diversity. Natural enemy interactions are often important in diversification dynamics, and key traits that mediate such interactions may therefore have an important role in explaining organismal diversity. Venom is one such trait which is intricately bound in antagonistic coevolution and has recently been shown to be associated with increased diversification rates in tetrapods. Despite ~ 10% of fish families and ~ 16% of insect families containing venomous species, the role that venom may play in these two superradiations remains unknown.ResultsIn this paper we take a broad family-level phylogenetic perspective and show that variation in diversification rates are the main cause of variations in species richness in both insects and fishes, and that venomous families have diversification rates twice as high as non-venomous families. Furthermore, we estimate that venom was present in ~ 10% and ~ 14% of the evolutionary history of fishes and insects respectively.ConclusionsConsequently, we provide evidence that venom has played a role in generating the remarkable diversity in the largest vertebrate and invertebrate radiations.

Limited evidence for a positive relationship between hybridization and diversification across seed plant families.

Hybridization has experimental and observational ties to evolutionary processes and outcomes such as adaptation, speciation, and radiation. Although it has been hypothesized that hybridization and diversification are positively correlated, this idea remains largely untested empirically, and hybridization can also potentially reduce diversity. Here, we use a hybridization database on 170 seed plant families, life history information, and a time-calibrated phylogeny to test for phylogenetically-corrected associations between hybridization and diversification rates, while also taking into account life-history traits that may be correlated with both processes. We use three methods to estimate diversification rates and two metrics of hybridization. Although hybridization explains only a small amount of overall variation in diversification rates, we show that diversification and hybridization are sometimes positively correlated, although the effect sizes are very small. Moreover, the relationship remains detectable when incorporating the correlations between diversification and two other life history characteristics, perenniality and woodiness. We discuss potential mechanisms for this association under four different scenarios: hybridization may drive diversification, diversification may drive hybridization, both hybridization and diversification may jointly be driven by other factors, or, as an alternative, that there is in fact no relationship between the two. We suggest future studies to disentangle the causal structure.

Empirical and Methodological Challenges to the Model-Based Inference of Diversification Rates in Extinct Clades.

Changes in speciation and extinction rates are key to the dynamics of clade diversification, but attempts to infer them from phylogenies of extant species face challenges. Methods capable of synthesizing information from extant and fossil species have yielded novel insights into diversification rate variation through time, but little is known about their behavior when analyzing entirely extinct clades. Here, we use empirical and simulated data to assess how two popular methods, PyRate and Fossil BAMM, perform in this setting. We inferred the first tip-dated trees for ornithischian dinosaurs and combined them with fossil occurrence data to test whether the clade underwent an end-Cretaceous decline. We then simulated phylogenies and fossil records under empirical constraints to determine whether macroevolutionary and preservation rates can be teased apart under paleobiologically realistic conditions. We obtained discordant inferences about ornithischian macroevolution including a long-term speciation rate decline (BAMM), mostly flat rates with a steep diversification drop (PyRate) or without one (BAMM), and episodes of implausibly accelerated speciation and extinction (PyRate). Simulations revealed little to no conflation between speciation and preservation, but yielded spuriously correlated speciation and extinction estimates while time-smearing tree-wide shifts (BAMM) or overestimating their number (PyRate). Our results indicate that the small phylogenetic data sets available to vertebrate paleontologists and the assumptions made by current model-based methods combine to yield potentially unreliable inferences about the diversification of extinct clades. We provide guidelines for interpreting the results of the existing approaches in light of their limitations and suggest how the latter may be mitigated. [BAMM; diversification; fossils; macroevolutionary rates; Ornithischia; PyRate.].

Macroevolutionary analyses of ciliates associated with hosts support high diversification rates

Ciliophora is a phylum that is comprised of extremely diverse microorganisms with regard to their morphology and ecology. They may be found in various environments, as free-living organisms or associated with metazoans. Such associations range from relationships with low metabolic dependence such as epibiosis, to more intimate relationships such as mutualism and parasitism. We know that symbiotic relationships occur along the whole phylogeny of the group, however, little is known about their evolution. Theoretical studies show that there are two routes for the development of parasitism, yet few authors have investigated the evolution of these characteristics using molecular tools. In the present study, we inferred a wide dated molecular phylogeny, based on the 18S rDNA gene, for the entire Ciliophora phylum, mapped life habits throughout the evolutionary time, and evaluated whether symbiotic relationships were linked to the variation in diversification rates and to the mode of evolution of ciliates. Our results showed that the last common ancestor for Ciliophora was likely a free-living organism, and that parasitism is a recent adaptation in ciliates, emerging more than once and independently via two distinct routes: (i) a free-living ciliate evolved into a mutualistic organism and, later, into a parasitic organism, and (ii) a free-living ciliate evolved directly into a parasitic organism. Furthermore, we have found a significant increase in the diversification rate of parasitic and mutualistic ciliates compared with their free-living conspecifics. The evolutionary success in different lineages of symbiont ciliates may be associated with many factors including type and colonization placement on their host, as well as physical and physiological conditions made available by the hosts.

Macroevolution and shifts in the feeding biology of the New World scarab beetle tribe Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae)

Abstract The New World scarab beetle tribe Phanaeini contains coprophagous, necrophagous, mycetophagous and suspected myrmecophilous species. We analyse the largest tribal molecular dataset assembled, incorporating, for the first time, the enigmatic monobasic genus Megatharsis, the thalassinus group of the subgenus Coprophanaeus (Metallophanaeus), and the subgenus Dendropaemon (Eurypodea) (formerly Tetramereia), unveiling their macroevolutionary and biogeographical history in light of Cenozoic abiotic changes and inferring shifts in feeding biology through time. We recover the contentious genus Gromphas outside an otherwise monophyletic Phanaeini. We infer Megatharsis in a clade containing the apparent myrmecophilous genus Dendropaemon, within the Coprophanaeus clade, and demonstrate that the subgenus Coprophanaeus (Metallophanaeus) is polyphyletic, whilst species groups within the subgenus Coprophanaeus (Coprophanaeus) are monophyletic. Our divergence time analyses and ancestral range estimation indicate an eastern South American origin for Phanaeini in the early Eocene, with subsequent colonization of Central America and the Nearctic during the Oligocene, long before a Panamanian land bridge. A shift to necrophagy in Coprophanaeus is possibly linked to increasing Neotropical small vertebrate diversity since the Eocene and, astonishingly, myrmecophily evolved from necrophagy 35 Mya. These drastic shifts in lifestyle are not concordant with variations in diversification rates and appear unlinked to Quaternary extinction of large mammals.

A model with many small shifts for estimating species-specific diversification rates

Understanding how and why diversification rates vary through time and space and across species groups is key to understanding the emergence of today's biodiversity. Phylogenetic approaches aimed at identifying variations in diversification rates during the evolutionary history of clades have focused on exceptional shifts subtending evolutionary radiations. While such shifts have undoubtedly affected the history of life, identifying smaller but more frequent changes is important as well. We developed ClaDS-a new Bayesian approach for estimating branch-specific diversification rates on a phylogeny that relies on a model with changes in diversification rates at each speciation event. We show, using Monte Carlo simulations, that the approach performs well at inferring both small and large changes in diversification. Applying our approach to bird phylogenies covering the entire avian radiation, we find that diversification rates are remarkably heterogeneous within evolutionarily restricted species groups. Some groups such as Accipitridae (hawks and allies) cover almost the full range of speciation rates found across the entire bird radiation. As much as 76% of the variation in branch-specific rates across this radiation is due to intraclade variation, suggesting that a large part of the variation in diversification rates is due to many small, rather than few large, shifts.

Tip rates, phylogenies and diversification: What are we estimating, and how good are the estimates?

Abstract Species‐specific diversification rates, or ‘tip rates’, can be computed quickly from phylogenies and are widely used to study diversification rate variation in relation to geography, ecology and phenotypes. These tip rates provide a number of theoretical and practical advantages, such as the relaxation of assumptions of rate homogeneity in trait‐dependent diversification studies. However, there is substantial confusion in the literature regarding whether these metrics estimate speciation or net diversification rates. Additionally, no study has yet compared the relative performance and accuracy of tip rate metrics across simulated diversification scenarios. We compared the statistical performance of three model‐free rate metrics (inverse terminal branch lengths; node density metric; DR statistic) and a model‐based approach (Bayesian analysis of macroevolutionary mixtures [BAMM]). We applied each method to a large set of simulated phylogenies that had been generated under different diversification processes. We summarized performance in relation to the type of rate variation, the magnitude of rate heterogeneity and rate regime size. We also compared the ability of the metrics to estimate both speciation and net diversification rates. We show decisively that model‐free tip rate metrics provide a better estimate of the rate of speciation than of net diversification. Error in net diversification rate estimates increases as a function of the relative extinction rate. In contrast, error in speciation rate estimates is low and relatively insensitive to extinction. Overall, and in particular when relative extinction was high, BAMM inferred the most accurate tip rates and exhibited lower error than non‐model‐based approaches. DR was highly correlated with true speciation rates but exhibited high error variance, and was the best metric for very small rate regimes. We found that, of the metrics tested, DR and BAMM are the most useful metrics for studying speciation rate dynamics and trait‐dependent diversification. Although BAMM was more accurate than DR overall, the two approaches have complementary strengths. Because tip rate metrics are more reliable estimators of speciation rate, we recommend that empirical studies using these metrics exercise caution when drawing biological interpretations in any situation where the distinction between speciation and net diversification is important.

Evolution of the latitudinal diversity gradient in the hyperdiverse ant genus Pheidole

AbstractAimThe latitudinal diversity gradient is the dominant geographic pattern of life on Earth, but a consensus understanding of its origins has remained elusive. The analysis of recently diverged, hyper‐rich invertebrate groups provides an opportunity to investigate latitudinal patterns with the statistical power of large trees while minimizing potentially confounding variation in ecology and history. Here, we synthesize global phylogenetic and macroecological data on a hyperdiverse (> 1,100 species) ant radiation, Pheidole and test predictions of three general explanations for the latitudinal gradient: variation in diversification rates, tropical conservatism and ecological regulation.LocationGlobal.Time periodThe past 35 million years.Major taxa studiedThe hyperdiverse ant genus Pheidole Westwood.MethodsWe assembled geographic data for 1,499 species and morphospecies, and inferred a dated phylogeny for 449 species of Pheidole, including 167 species newly sequenced for this study. We tested for correlations between diversification rate and latitude with Bayesian analysis of macroevolutionary mixtures (BAMM), hidden state speciation and extinction (HiSSE), geographic state speciation and extinction (GeoSSE), and a non‐parametric method (FiSSE), evaluated evidence for richness steady state, and examined patterns of diversification as Pheidole spread around the globe.ResultsThere was no evidence of systematic variation of net diversification rates with latitude across any of the methods. We found that Pheidole diversification occurred in bursts when new continents were colonized, followed by a slowdown in each region, but there is no evidence richness has saturated at an equilibrium in any region. Additionally, we found latitudinal affinity is moderately conserved with a Neotropical ancestor and simulations show that phylogenetic inertia alone is sufficient to produce the gradient pattern.Main conclusionsOur results provide no evidence that diversification rates vary systematically with latitude. Richness is far from steady state in each region, contrary to the ecological regulation hypothesis, although there is evidence that ecological opportunity promotes diversification after colonization of new areas. The fact that niche conservatism is strong enough to produce the gradient pattern is in accord with the tropical conservatism hypothesis. Overall, these results shed light on the mechanisms underlying the emergence of the diversity gradient within the past 34 million years, complementing recent work on deeper time‐scales, and more generally contribute toward much‐needed invertebrate perspective on global biodiversity dynamics.

Why are red flowers so rare? Testing the macroevolutionary causes of tippiness.

Traits that have arisen multiple times yet still remain rare present a curious paradox. A number of these rare traits show a distinct tippy pattern, where they appear widely dispersed across a phylogeny, are associated with short branches and differ between recently diverged sister species. This phylogenetic pattern has classically been attributed to the trait being an evolutionary dead end, where the trait arises due to some short-term evolutionary advantage, but it ultimately leads species to extinction. While the higher extinction rate associated with a dead end trait could produce such a tippy pattern, a similar pattern could appear if lineages with the trait speciated slower than other lineages, or if the trait was lost more often that it was gained. In this study, we quantify the degree of tippiness of red flowers in the tomato family, Solanaceae, and investigate the macroevolutionary processes that could explain the sparse phylogenetic distribution of this trait. Using a suite of metrics, we confirm that red-flowered lineages are significantly overdispersed across the tree and form smaller clades than expected under a null model. Next, we fit 22 alternative models using HiSSE (Hidden State Speciation and Extinction), which accommodates asymmetries in speciation, extinction and transition rates that depend on observed and unobserved (hidden) character states. Results of the model fitting indicated significant variation in diversification rates across the family, which is best explained by the inclusion of hidden states. Our best fitting model differs between the maximum clade credibility tree and when incorporating phylogenetic uncertainty, suggesting that the extreme tippiness and rarity of red Solanaceae flowers makes it difficult to distinguish among different underlying processes. However, both of the best models strongly support a bias towards the loss of red flowers. The best fitting HiSSE model when incorporating phylogenetic uncertainty lends some support to the hypothesis that lineages with red flowers exhibit reduced diversification rates due to elevated extinction rates. Future studies employing simulations or targeting population-level processes may allow us to determine whether red flowers in Solanaceae or other angiosperms clades are rare and tippy due to a combination of processes, or asymmetrical transitions alone.