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Recent analyses using large-scale phylogenies suggest a radically different history for the evolution of live birth and egg laying in squamate reptiles (lizards and snakes) than traditionally understood. What is the ancestral condition for lizards and snakes? How frequently does live bearing evolve in egg-laying lineages? Can the eggshell ever re-evolve in live-bearing lineages? Answering these fundamental questions about the evolution of key physiological processes will require additional data from genomic, developmental, and fossil data. J. Exp. Zool. (Mol. Dev. Evol.) 324B:467-472, 2015. © 2015 Wiley Periodicals, Inc. How to cite this article: Pyron R, Burbrink FT. 2015. Contrasting models of parity-mode evolution in squamate reptiles. J. Exp. Zool. (Mol. Dev. Evol.) 324B:467-472. J. Exp. Zool. (Mol. Dev. Evol.) 324B:467-472, 2015

Time-calibrated phylogenies based on molecular data provide a framework for comparative studies. Calibration methods to combine fossil information with molecular phylogenies are, however, under active development, often generating disagreement about the best way to incorporate paleontological data into these analyses. This study provides an empirical comparison of the most widely used approach based on node-dating priors for relaxed clocks implemented in the programs BEAST and MrBayes, with two recently proposed improvements: one using a new fossilized birth–death process model for node dating (implemented in the program DPPDiv), and the other using a total-evidence or tip-dating method (implemented in MrBayes and BEAST). These methods are applied herein to tetraodontiform fishes, a diverse group of living and extinct taxa that features one of the most extensive fossil records among teleosts. Previous estimates of time-calibrated phylogenies of tetraodontiforms using node-dating methods reported disparate estimates for their age of origin, ranging from the late Jurassic to the early Paleocene (ca. 150– 59 Ma). We analyzed a comprehensive dataset with 16 loci and 210 morphological characters, including 131 taxa (95 extant and 36 fossil species) representing all families of fossil and extant tetraodontiforms, under different molecular clock calibration approaches. Results from node-dating methods produced consistently younger ages than the tip-dating approaches. The older ages inferred by tip dating imply an unlikely early-late Jurassic (ca. 185–119 Ma) origin for this order and the existence of extended ghost lineages in their fossil record. Node-based methods, by contrast, produce time estimates that are more consistent with the stratigraphic record, suggesting a late Cretaceous (ca. 86–96 Ma) origin. We show that the precision of clade age estimates using tip dating increases with the number of fossils analyzed and with the proximity of fossil taxa to the node under assessment. This study suggests that current implementations of tip dating may overestimate ages of divergence in calibrated phylogenies. It also provides a comprehensive phylogenetic framework for tetraodontiform systematics and future comparative studies.

We describe a new species, Aspidura ravanai sp. nov., representing the eighth species of the genus known from Sri Lanka. The new species is readily distinguished from all other congeners by its colour pattern, scalation, and genetic distinctiveness. Our molecular results indicated that Aspidura began diversifying in the Miocene approximately 18 Ma, and A. ravanai sp. nov. diverged from its likely sister lineage A. trachyprocta at least 6.2 Ma. The species is currently known only from the type locality, on the western slopes of Sri Pada Peak in the central highlands of Sri Lanka.

Human activities continue to erode the tree of life, requiring us to prioritize research and conservation. Amphibians represent key victims and bellwethers of global change, and the need for action to conserve them is drastically outpacing knowledge. We provide a phylogeny incorporating nearly all extant amphibians (7,238 species). Current amphibian diversity is composed of both older, depauperate lineages and extensive, more recent tropical radiations found in select clades. Frog and salamander diversification increased strongly after the Cretaceous-Palaeogene boundary, preceded by a potential mass-extinction event in salamanders. Diversification rates of subterranean caecilians varied little over time. Biogeographically, the Afro- and Neotropics harbour a particularly high proportion of Gondwanan relicts, comprising species with high evolutionary distinctiveness (ED). These high-ED species represent a large portion of the branches in the present tree: around 28% of all phylogenetic diversity comes from species in the top 10% of ED. The association between ED and imperilment is weak, but many species with high ED are now imperilled or lack formal threat status, suggesting opportunities for integrating evolutionary position and phylogenetic heritage in addressing the current extinction crisis. By providing a phylogenetic estimate for extant amphibians and identifying their threats and ED, we offer a preliminary basis for a quantitatively informed global approach to conserving the amphibian tree of life.

The purpose of this application, under Article 81 of the Code, is to conserve the name dicroglossidae Dubois, 1987 for a family of frogs. Analysis of the publication where the name dicroglossidae was first used (Anderson, 1871) showed that this was an incorrect subsequent spelling of discoglossidae Gunther, 1858 and therefore not an available name. This name would have been made available by Dubois (1987) except that Dicroglossus Gunther, 1860 was then considered a junior subjective synonym of Euphlyctis Fitzinger, 1843 and thus unable to be the type-genus of a new family-group name according to Article 11.7.1.1 of the Code. However, dicroglossidae is a widely used family name in the recent taxonomy of amphibians. It includes about 180 species distributed in sub-Saharan Africa and tropical Asia. We therefore ask the International Commission on Zoological Nomenclature to make the name dicroglossini Dubois, 1987, with the type genus Dicroglossus Gunther, 1860, available by original designation of ...

Phylogenetic niche conservatism (PNC) typically refers to the tendency of closely related species to be more similar to each other in terms of niche than they are to more distant relatives. This has been implicated as a potential driving force in speciation and other species-richness patterns, such as latitudinal gradients. However, PNC has not been very well defined in most previous studies. Is it a pattern or a process? What are the underlying endogenous (e.g. genetic) and exogenous (e.g. ecological) factors that cause niches to be conserved? What degree of similarity is necessary to qualify as PNC? Is it possible for the evolutionary processes causing niches to be conserved to also result in niche divergence in different habitats? Here, we revisit these questions, codifying a theoretical and operational definition of PNC as a mechanistic evolutionary process resulting from several factors. We frame this both from a macroevolutionary and population-genetic perspective. We discuss how different axes of physical (e.g. geographic) and environmental (e.g. climatic) heterogeneity interact with the fundamental process of PNC to produce different outcomes of ecological speciation. We also review tests for PNC, and suggest ways that these could be improved or better utilized in future studies. Ultimately, PNC as a process has a well-defined mechanistic basis in organisms, and future studies investigating ecological speciation would be well served to consider this, and frame hypothesis testing in terms of the processes and expected patterns described herein. The process of PNC may lead to patterns where niches are conserved (more similar than expected), constrained (divergent within a limited subset of available niches), or divergent (less similar than expected), based on degree of phylogenetic relatedness between species.

Amphibia comprises over 7000 extant species distributed in almost every ecosystem on every continent except Antarctica. Most species also show high specificity for particular habitats, biomes, or climatic niches, seemingly rendering long-distance dispersal unlikely. Indeed, many lineages still seem to show the signature of their Pangaean origin, approximately 300 Ma later. To date, no study has attempted a large-scale historical-biogeographic analysis of the group to understand the distribution of extant lineages. Here, I use an updated chronogram containing 3309 species (~45% of extant diversity) to reconstruct their movement between 12 global ecoregions. I find that Pangaean origin and subsequent Laurasian and Gondwanan fragmentation explain a large proportion of patterns in the distribution of extant species. However, dispersal during the Cenozoic, likely across land bridges or short distances across oceans, has also exerted a strong influence. Finally, there are at least three strongly supported instances of long-distance oceanic dispersal between former Gondwanan landmasses during the Cenozoic. Extinction from intervening areas seems to be a strong factor in shaping present-day distributions. Dispersal and extinction from and between ecoregions are apparently tied to the evolution of extraordinarily adaptive expansion-oriented phenotypes that allow lineages to easily colonize new areas and diversify, or conversely, to extremely specialized phenotypes or heavily relictual climatic niches that result in strong geographic localization and limited diversification. (Amphibians; caecilians; dispersal; frogs; historical biogeography; oceanic dispersal; salamanders; vicariance.)

Aim Many ecological and evolutionary hypotheses have been proposed to explain latitudinal diversity gradients. However, any mechanistic explanation must include factors that change at least one of the three processes that can directly affect species richness: speciation, extinction and dispersal. I hypothesize that higher extinction in temperate areas, higher speciation in tropical areas and reduced dispersal into temperate regions drives latitudinal diversity gradients in squamates, and that these processes may be common in other groups. Location World-wide. Methods I test for these processes using phylogenetic methods that can untangle speciation and extinction with respect to latitudinal position (GeoSSE), using a dated phylogeny containing c. 45% of all squamates (4161 species) with data on their geographic occurrence. Results I find that lineages in the tropics have high speciation and low extinction, but that temperate lineages have even higher speciation and extinction, leading to lower net diversification and higher turnover in temperate areas, with much higher rates of dispersal into the tropics from temperate areas than the reverse. Main conclusions Recent empirical studies using dated molecular phylogenies appear to be consistent in supporting a similar set of simple and intuitive results for processes driving latitudinal gradients in species richness: higher net diversification rates in the tropics, higher relative extinction fractions in temperate regions and reduced dispersal out of the tropics. I suggest that higher temperate extinction represents a dominant force for the origin and maintenance of latitudinal gradients, particularly in groups with ancient temperate clades.

Aim Island biogeography predicts positive relationships between area and species richness resulting from the ecological processes of colonization in smaller areas and the evolutionary process of in situ diversification in larger areas. However, the relative contribution of these ecological and evolutionary processes to diversity on islands over time is unclear without associated measures of phylogenetic diversity (PD), which account for the evolutionary distinctiveness of component taxa. Location Global. Methods We assemble a global dataset of island richness for snakes, along with a phylogeny for the group, to examine the origins of species richness and PD on islands. Results Ecological models driven by colonization explain a majority of species richness in island snake faunas, while in situ diversification is rare, restricted to a few large islands and rarely contributing significantly to island richness. Similarly, PD on islands is tied to variables that affect colonization probability such as isolation and climate, but not area, indicating that in situ diversification does not exert a significant influence on island richness. Main conclusions Even large islands with high in situ diversification do not produce phylogenetically distinct faunas, but are instead dependent on colonization of new lineages from the continent, as predicted by ecological theory. Evolutionary processes such as in situ diversification are relatively insignificant drivers of both species richness and PD compared with colonization from regional species pools, even over evolutionary time-scales on large islands.

Both gene-tree discordance and unrecognized diversity are sources of error for accurate estimation of species trees, and can affect downstream diversification analyses by obscuring the correct number of nodes, their density, and the lengths of the branches subtending them. Although the theoretical impact of gene-tree discordance on evolutionary analyses has been examined previously, the effect of unsampled and cryptic diversity has not. Here, we examine how delimitation of previously unrecognized diversity in the milksnake (Lampropeltis triangulum) and use of a species-tree approach affects both estimation of the Lampropeltis phylogeny and comparative analyses with respect to the timing of diversification. Coalescent species delimitation indicates that L. triangulum is not monophyletic and that there are multiple species of milksnake, which increases the known species diversity in the genus Lampropeltis by 40%. Both genealogical and temporal discordance occurs between gene trees and the species tree, with evidence that mitochondrial DNA (mtDNA) introgression is a main factor. This discordance is further manifested in the preferred models of diversification, where the concatenated gene tree strongly supports an early burst of speciation during the Miocene, in contrast to species-tree estimates where diversification follows a birth-death model and speciation occurs mostly in the Pliocene and Pleistocene. This study highlights the crucial interaction among coalescent-based phylogeography and species delimitation, systematics, and species diversification analyses. (Divergence-time estimation; diversification rates; Lampropeltini; gene-tree/species- tree discordance; mtDNA introgression; Pleistocene diversification.)

Phylogenies are used to estimate rates of speciation and extinction, reconstruct historical diversification scenarios, and link these to ecological and evolutionary factors, such as climate or organismal traits. Recent models can now estimate the effects of binary, multistate, continuous, and biogeographic characters on diversification rates. Others test for diversity dependence (DD) in speciation and extinction, which has become recognized as an important process in numerous clades. A third class incorporates flexible time-dependent functions, enabling reconstruction of major periods of both expanding and contracting diversity. Although there are some potential problems (particularly for estimating extinction), these methods hold promise for answering many classic questions in ecology and evolution, such as the origin of adaptive radiations, and the latitudinal gradient in species richness.

Viviparity has putatively evolved 115 times in squamates (lizards and snakes), out of only ~ 140 origins in vertebrates, and is apparently related to colder climates and other factors such as body size. Viviparity apparently evolves from oviparity via egg-retention, and such taxa may thus still have the machinery to produce thick-shelled eggs. Parity mode is also associated with variable diversification rates in some groups. We reconstruct ancestral parity modes accounting for state-dependent diversification in a large-scale phylogenetic analysis, and find strong support for an early origin of viviparity at the base of Squamata, and a complex pattern of subsequent transitions. Viviparous lineages have higher rates of speciation and extinction, and greater species turnover through time. Viviparity is associated with lower environmental and body temperatures in lizards and amphisbaenians, but not female mass. These results suggest that parity mode is a labile trait that shifts frequently in response to ecological conditions.

Based on a molecular phylogeny and a comparison of maxillary dentition and morphology, the relationship between the genera Lycodon and Dinodon was investigated. Bayesian Inference and Maximum Likelihood analysis of two mitochondrial genes (cyt b and ND4) and two nuclear genes (c-mos and Rag1) suggested that the two genera shared a most recent common ancestor. However, Dinodon was paraphyletic and Lycodon was polyphyletic, each with respect to the other. The results from counts of maxillary teeth indicated that the diagnostic characters used by previous authors to separate Dinodon and Lycodon were not reliable. Taking the molecular and morphological evidence together, we synonymized Dinodon with Lycodon. In addition, the validity of the species L. futsingensis was confirmed to be distinctly different from the other species of Dinodon and Lycodon.

Geographic barriers and elevational gradients have long been recognized as important in species diversification. Here, we illustrate an example where both mechanisms have shaped the genetic structure of the Neotropical rainfrog, Pristimantis ornatissimus, which has also resulted in speciation. This species was thought to be a single evolutionary lineage distributed throughout the Ecuadorian Choco and the adjacent foothills of the Andes. Based on recent sampling of P. ornatissimus sensu lato, we provide molecular and morphological evidence that support the validity of a new species, which we name Pristimantis ecuadorensis sp. nov. The sister species are elevational replacements of each other; the distribution of Pristimantis ornatissimus sensu stricto is limited to the Ecuadorian Choco ecoregion (< 1100 m), whereas the new species has only been found at Andean localities between 1450–1480 m. Given the results of the Multiple Matrix Regression with Randomization analysis, the genetic difference between P. ecuadorensis and P. ornatissimus is not explained by geographic distance nor environment, although environmental variables at a finer scale need to be tested. Therefore this speciation event might be the byproduct of stochastic historic extinction of connected populations or biogeographic events caused by barriers to dispersal such as rivers. Within P. ornatissimus sensu stricto, morphological patterns and genetic structure seem to be related to geographic isolation (e.g., rivers). Finally, we provide an updated phylogeny for the genus, including the new species, as well as other Ecuadorian Pristimantis.

Globally distributed groups may show regionally distinct rates of diversification, where speciation is elevated given timing and sources of ecological opportunity. However, for most organisms, nearly complete sampling at genomic-data scales to reduce topological error in all regions is unattainable, thus hampering conclusions related to biogeographic origins and rates of diversification. We explore processes leading to the diversity of global ratsnakes and test several important hypotheses related to areas of origin and enhanced diversification upon colonizing new continents. We estimate species trees inferred from phylogenomic scale data (304 loci) while exploring several strategies that consider topological error from each individual gene tree. With a dated species tree, we examine taxonomy and test previous hypotheses that suggest the ratsnakes originated in the Old World (OW) and dispersed to New World (NW). Furthermore, we determine if dispersal to the NW represented a source of ecological opportunity, which should show elevated rates of species diversification. We show that ratsnakes originated in the OW during the mid-Oligocene and subsequently dispersed to the NW by the mid-Miocene; diversification was also elevated in a subclade of NW taxa. Finally, the optimal biogeographic region-dependent speciation model shows that the uptick in ratsnake diversification was associated with colonization of the NW. We consider several alternative explanations that account for regionally distinct diversification rates.

Comparative phylogeography allow us to understand how shared historical circumstances have shaped the formation of lineages, by examining a broad spectrum of co-distributed populations of different taxa. However, these types of studies are scarce in the Neotropics, a region that is characterized by high diversity, complex geology, and poorly understood biogeography. Here, we investigate the diversification patterns of five lineages of amphibians and reptiles, co-distributed across the Choco and Andes ecoregions in northwestern Ecuador. Mitochondrial DNA and occurrence records were used to determine the degree of geographic genetic divergence within species. Our results highlight congruent patterns of parapatric speciation and common geographical barriers for distantly related taxa. These comparisons indicate similar biological and demographic characteristics for the included clades, and reveal the existence of two new species of Pristimantis previously subsumed under P. walkeri, which we describe herein. Our data supports the hypothesis that widely distributed Chocoan taxa may generally experience their greatest opportunities for isolation and parapatric speciation across thermal elevational gradients. Finally, our study provides critical information to predict which unstudied lineages may harbor cryptic diversity, and how geology and climate are likely to have shaped their evolutionary history.

Adaptive radiation (AR) theory predicts that groups sharing the same source of ecological opportunity (EO) will experience deterministic species diversification and morphological evolution. Thus, deterministic ecological and morphological evolution should be correlated with deterministic patterns in the tempo and mode of speciation for groups in similar habitats and time periods. We test this hypothesis using well-sampled phylogenies of four squamate groups that colonized the New World (NW) in the Late Oligocene. We use both standard and coalescent models to assess species diversification, as well as likelihood models to examine morphological evolution. All squamate groups show similar early pulses of speciation, as well as diversity-dependent ecological limits on clade size at a continental scale. In contrast, processes of morphological evolution are not easily predictable and do not show similar pulses of early and rapid change. Patterns of morphological and species diversification thus appear uncoupled across these groups. This indicates that the processes that drive diversification and disparification are not mechanistically linked, even among similar groups of taxa experiencing the same sources of EO. It also suggests that processes of phenotypic diversification cannot be predicted solely from the existence of an AR or knowledge of the process of diversification.

Adaptive radiations are typically triggered when a lineage encounters a significant range of open niche space (ecological opportunity), stemming from colonisation of new areas, extinction of competitors or key innovations. The most well-known of these is the colonisation of new areas, through either dispersal into new regions or the invasion of novel ecological regimes. One aspect of ecological opportunity that has rarely been studied, however, is the extent to which pre-existent competitors act to limit diversification in newly colonised adaptive zones. Herein, we show that in multiple geographically independent invasions of freshwaters by marine Sea Catfishes (Ariidae), rates of both morphological disparification and lineage diversification are inversely related to the presence and diversity of other freshwater fish lineages. Only in one region (Australia-New Guinea) with an otherwise depauperate freshwater fauna, has an ariid invasion gained any substantial traction. This is true at both regional and community scales, suggesting that competitive constraints may be an important factor regulating adaptive radiation.

The extant amphibians are one of the most diverse radiations of terrestrial vertebrates (>6800 species). Despite much recent focus on their conservation, diversification, and systematics, no previous phylogeny for the group has contained more than 522 species. However, numerous studies with limited taxon sampling have generated large amounts of partially overlapping sequence data for many species. Here, we combine these data and produce a novel estimate of extant amphibian phylogeny, containing 2871 species (∼40% of the known extant species) from 432 genera (∼85% of the ∼500 currently recognized extant genera). Each sampled species contains up to 12,712 bp from 12 genes (three mitochondrial, nine nuclear), with an average of 2563 bp per species. This data set provides strong support for many groups recognized in previous studies, but it also suggests non-monophyly for several currently recognized families, particularly in hyloid frogs (e.g., Ceratophryidae, Cycloramphidae, Leptodactylidae, Strabomantidae). To correct these and other problems, we provide a revised classification of extant amphibians for taxa traditionally delimited at the family and subfamily levels. This new taxonomy includes several families not recognized in current classifications (e.g., Alsodidae, Batrachylidae, Rhinodermatidae, Odontophrynidae, Telmatobiidae), but which are strongly supported and important for avoiding non-monophyly of current families. Finally, this study provides further evidence that the supermatrix approach provides an effective strategy for inferring large-scale phylogenies using the combined results of previous studies, despite many taxa having extensive missing data.

Aim We examine diversification in Caribbean alsophiine snakes and hypothesize that, given the ecological opportunity presented by colonization of the West Indies, alsophiines should show the signature of an early burst of diversification and associated low within-clade ecological and morphological disparification. We also test whether changes in morphology and ecology are associated with changes in diversification rate, as trait-dependent diversification is hypothesized to affect historical inferences of diversification and disparification. Finally, as replicated radiations are found across the West Indies in the anoles, we test for significant differences in ecological and morphological assemblages and rates among the major island groups. Location The West Indies. Methods A time-calibrated phylogeny produced from six genes using relaxed clock methods in beast was constructed to estimate ancestral areas using Lagrange. Maximum body size and ecological niche were scored for all species in the phylogeny, and comparative phylogenetic methods in R using geiger, laser, ape and our own code were used to examine diversification through time, disparification and trait-dependent diversification from this dated phylogeny. Results The pattern of species diversification did not differ significantly from the Yule model of diversification. Morphology and ecology fitted a Brownian and white noise model of diversification, respectively. Although not significantly different, morphological disparification was lower than the Brownian null model, whereas ecological disparification was significantly greater than the null. Traitdependent diversification analyses suggested that the constant null models provided the best fit to these data. There was no significant signal of rate variation among the major island groups for size, but moderate evidence for niche. Main conclusions Although ecological opportunity was similarly present for alsophiines as it was for anoles, the snakes fail to show an early burst of speciation. Potential reasons for this include the young age of the group, and staggered diversification due to waiting times between island colonization. In turn, ecological and morphological disparities do not necessarily follow predictable patterns related to species diversification. Thus, the presence of ecological opportunity alone is not necessarily sufficient to trigger replicated adaptive radiations in areas.

Snake diversity varies by at least two orders of magnitude among extant lineages, with numerous groups containing only one or two species, and several young clades exhibiting exceptional richness (>700 taxa). With a phylogeny containing all known families and subfamilies, we find that these patterns cannot be explained by background rates of speciation and extinction. The majority of diversity appears to derive from a radiation within the superfamily Colubroidea, potentially stemming from the colonization of new areas and the evolution of advanced venom-delivery systems. In contrast, negative relationships between clade age, clade size, and diversification rate suggest the potential for possible bias in estimated diversification rates, interpreted by some recent authors as support for ecologically mediated limits on diversity. However, evidence from the fossil record indicates that numerous lineages were far more diverse in the past, and that extinction has had an important impact on extant diversity patterns. Thus, failure to adequately account for extinction appears to prevent both rate- and diversity-limited models from fully characterizing richness dynamics in snakes. We suggest that clade-level extinction may provide a key mechanism for explaining negative or hump-shaped relationships between clade age and diversity, and the prevalence of ancient, species-poor lineages in numerous groups.

Were molecular data available for extinct taxa, questions regarding the origins of many groups could be settled in short order. As this is not the case, various strategies have been proposed to combine paleontological and neontological data sets. The use of fossil dates as node age calibrations for divergence time estimation from molecular phylogenies is commonplace. In addition, simulations suggest that the addition of morphological data from extinct taxa may improve phylogenetic estimation when combined with molecular data for extant species, and some studies have merged morphological and molecular data to estimate combined evidence phylogenies containing both extinct and extant taxa. However, few, if any, studies have attempted to estimate divergence times using phylogenies containing both fossil and living taxa sampled for both molecular and morphological data. Here, I infer both the phylogeny and the time of origin for Lissamphibia and a number of stem tetrapods using Bayesian methods based on a data set containing morphological data for extinct taxa, molecular data for extant taxa, and molecular and morphological data for a subset of extant taxa. The results suggest that Lissamphibia is monophyletic, nested within Lepospondyli, and originated in the late Carboniferous at the earliest. This research illustrates potential pitfalls for the use of fossils as post hoc age constraints on internal nodes and highlights the importance of explicit phylogenetic analysis of extinct taxa. These results suggest that the application of fossils as minima or maxima on molecular phylogenies should be supplemented or supplanted by combined evidence analyses whenever possible.

What explains the striking variation in local species richness across the globe and the remarkable diversity of rainforest sites in Amazonia? Here, we apply a novel phylogenetic approach to these questions, using treefrogs (Hylidae) as a model system. Hylids show dramatic variation in local richness globally and incredible local diversity in Amazonia. We find that variation in local richness is not explained primarily by climatic factors, rates of diversification (speciation and extinction) nor morphological variation. Instead, local richness patterns are explained predominantly by the timing of colonization of each region, and Amazonian megadiversity is linked to the long-term sympatry of multiple clades in that region. Our results also suggest intriguing interactions between clade diversification, trait evolution and the accumulation of local richness. Specifically, sympatry between clades seems to slow diversification and trait evolution, but prevents neither the accumulation of local richness over time nor the co-occurrence of similar species.

Molecular phylogenies are often used to test hypotheses about the tempo and mode of speciation and extinction. One commonly used statistic is Pybus and Harvey's γ, which measures the density of ordered internode distances on an ultrametric tree to infer earlier (negative γ) or later (positive γ) bursts of diversification. However, coalescent theory predicts that γ might be biased toward negative values (inferring early bursts of diversification) when using gene trees rather than species trees. Gene divergences predate species divergences, increasingly so at higher effective population sizes (N(e)), and proportionally more so toward the tips of the tree. Thus, gene trees will have a higher density of older nodes in many cases (particularly at higher N(e)), due to the disproportionate lengthening of terminal branches. This will yield an artifactual signature of early bursts of diversification when estimating γ from gene trees. We simulate gene trees within species trees under both Yule (pure-birth) and birth-death processes, and demonstrate support for these predictions. However, for most realistic estimates of θ in natural populations, gene trees provide relatively good estimates of γ, despite the disproportionate overestimation of younger node ages. This is corroborated with an empirical dataset of North American fence lizards (Sceloporus).

The superfamily Colubroidea (> 2500 species) includes the majority of snake species and is one of the most conspicuous and well-known radiations of terrestrial vertebrates. However, many aspects of the phylogeny of the group remain contentious, and dozens of genera have yet to be included in molecular phylogenetic analyses. We present a new, large-scale, likelihood-based phylogeny for the colubroids, including 761 species sampled for up to five genes: cytochrome b (93% of 761 species sampled), ND4 (69%), ND2 (28%), c-mos (54%), and RAG-1 (13%), totaling up to 5814bp per species. We also compare likelihood bootstrapping and a recently proposed ultra-fast measure of branch support (Shimodaira-Hasegawa-like [SHL] approximate likelihood ratio), and find that the SHL test shows strong support for several clades that were weakly-supported by bootstrapping in this or previous analyses (e.g., Dipsadinae, Lamprophiidae). We find that SHL values are positively related to branch lengths, but show stronger support for shorter branches than bootstrapping. Despite extensive missing data for many taxa (mean=67% per species), neither bootstrap nor SHL support values for terminal species are related to their incompleteness, and that most highly incomplete taxa are placed in the expected families from previous taxonomy, typically with very strong support. The phylogeny indicates that the Neotropical colubrine genus Scaphiodontophis represents an unexpectedly ancient lineage within Colubridae. We present a revised higher-level classification of Colubroidea, which includes a new subfamily for Scaphiodontophis (Scaphiodontophiinae). Our study provides the most comprehensive phylogeny of Colubroidea to date, and suggests that SHL values may provide a useful complement to bootstrapping for estimating support on likelihood-based trees.

Estimating divergence times using molecular sequence data has become a common application of statistical phylogenetics. However, disparities between estimated ages for clades produced in different studies have become equally commonplace. Here, I propose a method for the objective assessment of the likelihood of inferred divergence times to evaluate the placement of fossil constraints using information from the broader fossil record. The inclusion of nodes from the Tree of Life for which credible age ranges are known, in addition to the fossil constraints used in the ingroup, will allow for the comparison of alternate fossil placements when the phylogenetic affinity of a fossil is ambiguous as well as provide a heuristic assessment of the global likelihood of estimated divergence times. The use of these "likelihood checkpoints" will allow for the comparison of inferred dates across data sets and across taxonomic groups to place divergence time estimates into a broader evolutionary timescale. The method is illustrated with an example using an expanded phylogenetic estimate of the Gnathostomata, inferred with relaxed-clock molecular dating methods. (Divergence time estimation; fossil calibrations; maximum likelihood; molecular phylogenetics; relaxed clocks.)

Ecological adaptive radiation theory predicts an increase in both morphological and specific diversification when organisms colonize new environments. Accordingly, bursts of morphological diversification, characterized by low within-subclade morphological disparity, may be associated with these increases in speciation rates. Conversely, increasing species density, reduction in available habitat, or increasing extinction rates are expected to cause rates of diversification to decline. We test these hypotheses by examining the tempo and mode of speciation in the lampropeltinine snakes, a morphologically variable group that colonized the New World approximately 24 million years ago and radiated throughout the Miocene. We show that specific diversification increased early in the history of the group, and that most morphological variation is partitioned among, rather than within subclades. These patterns provide further evidence for the hypothesis that morphological variation tends to be strongly partitioned among lineages when clades undergo early bursts of species diversification. A reduction in speciation rates may be indicative of density dependent effects due to a saturation of available ecological opportunity, rather than increases in extinction rates at the onset of the Pleistocene/Pliocene glacial cycles. This evidence runs counter to the general Pleistocene species pump model.

Evolutionary correlations between functionally related character suites are expected as a consequence of coadaptation due to physiological relationships between traits. However, significant correlations may also exist between putatively unrelated characters due to shared relationships between those traits and underlying variables, such as body size. Although such patterns are often dismissed as simple body size scaling, this presumption may overlook important evolutionary patterns of diversification. If body size is the primary determinant of potential diversity in multiple unrelated characters, the observed differentiation of species may be governed by variability in body size, and any biotic or abiotic constraints on the diversification thereof. Here, we demonstrate that traits related to both predatory specialization (gape and diet preference) and predatory avoidance (the development of Batesian mimicry) are phylogenetically correlated in the North American snake tribe Lampropeltini. This is apparently due to shared relationships between those traits and adult body size, suggesting that size is the primary determinant of ecomorphological differentiation in the lampropeltinines. Diversification in body size is apparently not linked to climatic or environmental factors, and may have been driven by interspecific interactions such as competition. Additionally, we find the presence of a 'key zone' for the development of both rattle- and coral snake mimicry; only small snakes feeding primarily on ectothermic prey develop mimetic colour patterns, in or near the range of venomous model species.

3 Corresponding author Abstract We present a systematic revision of the Lampropeltis getula group, based on a recent range-wide phylogeographic analysis. We define our theoretical and operational concepts of species delimitation, and provide diagnoses based on mitochondrial DNA evidence, ecological niche modeling, morphology, and historical precedence. We find support for the recognition of five distinct species, which bear the name of the nominate subspecies found primarily within the range of each phylogeographic lineage: the Eastern lineage (Lampropeltis getula, Eastern Kingsnake), the Mississippi lineage (L. nigra, Black Kingsnake), the Central lineage (L. holbrooki, Speckled Kingsnake), the Desert lineage (L. splendida, Desert Kingsnake), and the Western lineage (L. californiae, California Kingsnake). Interestingly, all of these taxa had originally been described as distinct species and recognized as such for up to 101 years (in the case of L. californiae) before being demoted to subspecies. We discuss the impact that increasingly detailed genetic information from phylogeographic analyses may have on traditional taxonomy.

Niche conservatism and niche divergence are both important ecological mechanisms associated with promoting allopatric speciation across geographical barriers. However, the potential for variable responses in widely distributed organisms has not been fully investigated. For allopatric sister lineages, three patterns for the interaction of ecological niche preference and geographical barriers are possible: (i) niche conservatism at a physical barrier; (ii) niche divergence at a physical barrier; and (iii) niche divergence in the absence of a physical barrier. We test for the presence of these patterns in a transcontinentally distributed snake species, the common kingsnake (Lampropeltis getula), to determine the relative frequency of niche conservatism or divergence in a single species complex inhabiting multiple distinct ecoregions. We infer the phylogeographic structure of the kingsnake using a range-wide data set sampled for the mitochondrial gene cytochrome b. We use coalescent simulation methods to test for the presence of structured lineage formation vs. fragmentation of a widespread ancestor. Finally, we use statistical techniques for creating and evaluating ecological niche models to test for conservatism of ecological niche preferences. Significant geographical structure is present in the kingsnake, for which coalescent tests indicate structured population division. Surprisingly, we find evidence for all three patterns of conservatism and divergence. This suggests that ecological niche preferences may be labile on recent phylogenetic timescales, and that lineage formation in widespread species can result from an interaction between inertial tendencies of niche conservatism and natural selection on populations in ecologically divergent habitats.

The impact of climate change during the Pleistocene on organisms living in temperate areas of the world has been documented for more than fifty years (Rand, 1948). A major impact of Pleistocene glacial cycles on vertebrates in temperate zones is hypothesized to be the isolation of species into allopatric populations (Avise and Walker, 1998). Refugial isolation and subsequent lineage formation during the Pleistocene has been invoked as a primary mechanism resulting in modern species diversity (Johnson and Cicero, 2004). Under the Pleistocene speciation model, these glacial cycles acted as a ‘species pump,’ producing the majority of extant organisms inhabiting temperate areas. These arguments, usually focused on dating the origin of sister species using a fixed rate of divergence in birds, have suggested that the majority of speciation events occurred in the Pleistocene (Johnson and Cicero, 2004; Weir and Schluter, 2007). In a more limited study, Avise et al. (1998) suggested that the Pleistocene was important for speciation in reptiles, amphibians, and fish as well. In contrast to the Pleistocene diversification model, it has been shown that many speciation events in birds occurred much earlier than the Pleistocene (Zink and Slowinski, 1995; Klicka and Zink, 1997), potentially calling into question the effects of glacial cycles on the formation of species. Unfortunately, most of these studies have invoked a molecular clock using a fixed rate of mtDNA substitution over time, and thus did not consider the impact of molecular rate variation for assessing error in divergence dates. The effect of Pleistocene climatic cycles on speciation has not been investigated in other large groups of vertebrates, particularly those that may be more sensitive to the effects of glacial cycles, such as ectotherms like reptiles. We investigate these patterns in a major New World (NW) group of colubroid snakes, the tribe Lampropeltini, inferring a robust phylogeny for the group and estimating the timing of speciation and diversification. We use multiple nuclear and mitochon

Aim A latitudinal gradient in species richness, defined as a decrease in biodiversity away from the equator, is one of the oldest known patterns in ecology and evolutionary biology. However, there are also many known cases of increasing poleward diversity, forming inverse latitudinal biodiversity gradients. As only three processes (speciation, extinction and dispersal) can directly affect species richness in areas, similar factors may be responsible for both classical (high tropical diversity) and inverse (high temperate diversity) gradients. Thus, a modified explanation for differential species richness which accounts for both patterns would be preferable to one which only explains high tropical biodiversity.

Most phylogeographic studies examine organisms that do not have transcontinental distributions and therefore the genetic and temporal effects of barriers across an entire continent cannot be assessed with respect to a single species. We examined the phylogeographic structure, lineage age, and historical demography using sequences from the mtDNA cytochrome b gene of the widespread North American racer (Coluber constrictor), one of the few abundant transcontinental snakes that occurs throughout many diverse biomes. Our results indicate that this complex is comprised of six lineages differing greatly in geographic extent, with the largest (a central US clade) being approximately 26 times greater than the smallest (a lineage restricted to the Florida Panhandle and nearby portions of adjacent States). Most of the six lineages appear to be separated at previously identified genetic barriers for several vertebrates with similar ranges. Lineage diversification in this species began in the late Miocene, separating populations in the Florida Peninsula from the remainder of the US. Diversification of lineages continued throughout the Pliocene and early Pleistocene. Four of the six lineages occur east of the Mississippi River, with only two distinctly young ( approximately 1.5 mya) lineages found west of the Mississippi River (one occurs west of Continental Divide). All methods of demographic inference, including the mismatch distribution, Fu and Li's D and Tajima's D , and Bayesian skyline plots revealed population expansion occurring in the mid-to-late Pleistocene for every lineage, regardless of size or proximity to formerly glaciated areas. Population expansion for lineages found east of the Mississippi River occurred earlier and was much greater than those found west of the River.

A well-known issue in phylogenetics is discordance among gene trees, species trees, morphology, and other data types. Gene-tree discordance is often caused by incomplete lineage sorting, lateral gene transfer, and gene duplication. Multispecies-coalescent methods can account for incomplete lineage sorting and are believed by many to be more accurate than concatenation. However, simulation studies and empirical data have demonstrated that concatenation and species tree methods often recover similar topologies. We use three popular methods of phylogenetic reconstruction (one concatenation, two species tree) to evaluate relationships within Teiidae. These lizards are distributed across the United States to Argentina and the West Indies, and their classification has been controversial due to incomplete sampling and the discordance among various character types (chromosomes, DNA, musculature, osteology, etc.) used to reconstruct phylogenetic relationships. Recent morphological and molecular analyses of the group resurrected three genera and created five new genera to resolve non-monophyly in three historically ill-defined genera: Ameiva, Cnemidophorus, and Tupinambis. Here, we assess the phylogenetic relationships of the Teiidae using "next-generation" anchored-phylogenomics sequencing. Our final alignment includes 316 loci (488,656bp DNA) for 244 individuals (56 species of teiids, representing all currently recognized genera) and all three methods (ExaML, MP-EST, and ASTRAL-II) recovered essentially identical topologies. Our results are basically in agreement with recent results from morphology and smaller molecular datasets, showing support for monophyly of the eight new genera. Interestingly, even with hundreds of loci, the relationships among some genera in Tupinambinae remain ambiguous (i.e. low nodal support for the position of Salvator and Dracaena).

Predicting species presence and richness on islands is important for understanding the origins of communities and how likely it is that species will disperse and resist extinction. The equilibrium theory of island biogeography (ETIB) and, as a simple model of sampling abundances, the unified neutral theory of biodiversity (UNTB), predict that in situations where mainland to island migration is high, species-abundance relationships explain the presence of taxa on islands. Thus, more abundant mainland species should have a higher probability of occurring on adjacent islands. In contrast to UNTB, if certain groups have traits that permit them to disperse to islands better than other taxa, then phylogeny may be more predictive of which taxa will occur on islands. Taking surveys of 54 island snake communities in the Eastern Nearctic along with mainland communities that have abundance data for each species, we use phylogenetic assembly methods and UNTB estimates to predict island communities. Species richness is predicted by island area, whereas turnover from the mainland to island communities is random with respect to phylogeny. Community structure appears to be ecologically neutral and abundance on the mainland is the best predictor of presence on islands. With regard to young and proximate islands, where allopatric or cladogenetic speciation is not a factor, we find that simple neutral models following UNTB and ETIB predict the structure of island communities.

Amphibian diversity in the Western Ghats-Sri Lanka biodiversity hotspot is extremely high, especially for such a geo-graphically restricted area. Frogs in particular dominate these assemblages, and the family Rhacophoridae is chief among these, with hundreds of endemic species. These taxa continue to be described at a rapid pace, and several groups have recently been found to represent unique evolutionary clades at the genus level. Here, we report DNA sequences, larval and breeding data for two species of rhacophorid treefrog (Polypedates bijui and a new, hitherto undescribed species). Re-markably, they represent unique, independent clades which form successive sister groups to the Pseudophilautus (Sri Lan-ka) + Raorchestes (India, China & Indochina) clades. We place these species into two new genera (Beddomixalus gen. nov. and Mercurana gen. nov.). Both of these genera exhibit a distinct reproductive mode among Rhacophoridae of pen-insular India and Sri Lanka, with explosive breeding and semiterrestrial, unprotected, non-pigmented eggs oviposited in seasonal swamp pools, which hatch into exotrophic, free-living aquatic tadpoles. Relationships and representation of re-productive modes in sister taxa within the larger clade into which these novel genera are placed, is also discussed. These results suggest that more undescribed taxa may remain to be discovered in South Asia, and the crucial importance of con-serving remaining viable habitats.

The Asian keelback snakes (genus Amphiesma) are a widely distributed group of Old World natricines, inhabiting a variety of niches and exhibiting significant morphological variation. Recent molecular phylogenies suggest that this genus is not monophyletic, and that additional cryptic diversity is also likely present. We conducted a phylogenetic analysis of the group based on 3162 bp of one mitochondrial gene (Cyt. b) and three nuclear genes (C-mos, Rag1, NT3), sampling 18 species in addition to those sequenced in previous works. All analyses consistently show that Amphiesma consists of three distinct, monophyletic lineages with strong support. We divide Amphiesma into three genera, Amphiesma, Hebius, and Herpetoreas. The genus Amphiesma is monotypic, Herpetoreas contains three species, and Hebius comprises the remaining 39 species. On the basis of a combination of molecular analyses and external morphological comparisons, we describe a new species in the Herpetoreas group from China as H. burbrinki sp. nov. Several other species are shown to be non-monophyletic or contain significant levels of intraspecific genetic diversity. Another Old World natricine genera, Xenochrophis is also found to be non-monophyletic. Our results indicate that further taxonomic revisions are needed in Natricinae, at multiple levels.

When we submitted our manuscript (Pyron and Burbrink, 2014) on 2 May 2013, I did not anticipate how controversially it would be received, or the magnitude of the response. Articles in this issue (Blackburn, 2015a,b; Duchene and Lanfear, 2015; Griffith et al., 2015; King and Lee, 2015b; Shine, 2015; Stewart, 2015; Wright et al., 2015) and in other journals (King and Lee, 2015a) have re-examined our data, analyses, and conclusions, drawing a wide range of inferences. Our summary (Pyron and Burbrink, 2015) was written before any of the responses appeared, and merely gives an overview of our original conclusions, with some suggestions for future research. Here, I take a final opportunity to digest the recent responses to Pyron and Burbrink (2014), and offer a complementary perspective. Future studies of parity-mode evolution should consider the following four points

One of Darwin's most widely known conjectures is that prey are tame on remote islands, where mammalian predators are absent. Many species appear to permit close approach on such islands, but no comparative studies have demonstrated reduced wariness quantified as flight initiation distance (FID; i.e. predator–prey distance when the prey begins to flee) in comparison with mainland relatives. We used the phylogenetic comparative method to assess influence of distance from the mainland and island area on FID of 66 lizard species. Because body size and predator approach speed affect predation risk, we included these as independent variables. Multiple regression showed that FID decreases as distance from mainland increases and is shorter in island than mainland populations. Although FID increased as area increased in some models, collinearity made it difficult to separate effects of area from distance and island occupancy. FID increases as SVL increases and approach speed increases; these effects are statistically independent of effects of distance to mainland and island occupancy. Ordinary least-squares models fit the data better than phylogenetic regressions, indicating little or no phylogenetic signal in residual FID after accounting for the independent variables. Our results demonstrate that island tameness is a real phenomenon in lizards.

Next-generation genomic sequencing promises to quickly and cheaply resolve remaining contentious nodes in the Tree of Life, and facilitates species-tree estimation while taking into account stochastic genealogical discordance among loci. Recent methods for estimating species trees bypass full likelihood-based estimates of the multi-species coalescent, and approximate the true species-tree using simpler summary metrics. These methods converge on the true species-tree with sufficient genomic sampling, even in the anomaly zone. However, no studies have yet evaluated their efficacy on a large-scale phylogenomic dataset, and compared them to previous concatenation strategies. Here, we generate such a dataset for Caenophidian snakes, a group with >2500 species that contains several rapid radiations that were poorly resolved with fewer loci. We generate sequence data for 333 single-copy nuclear loci with ∼100% coverage (∼0% missing data) for 31 major lineages. We estimate phylogenies using neighbor joining, maximum parsimony, maximum likelihood, and three summary species-tree approaches (NJst, STAR, and MP-EST). All methods yield similar resolution and support for most nodes. However, not all methods support monophyly of Caenophidia, with Acrochordidae placed as the sister taxon to Pythonidae in some analyses. Thus, phylogenomic species-tree estimation may occasionally disagree with well-supported relationships from concatenated analyses of small numbers of nuclear or mitochondrial genes, a consideration for future studies. In contrast for at least two diverse, rapid radiations (Lamprophiidae and Colubridae), phylogenomic data and species-tree inference do little to improve resolution and support. Thus, certain nodes may lack strong signal, and larger datasets and more sophisticated analyses may still fail to resolve them.

Phylogenetic niche conservatism (PNC) typically refers to the tendency of closely related species to be more similar to each other in terms of niche than they are to more distant relatives. This has been implicated as a potential driving force in speciation and other species-richness patterns, such as latitudinal gradients. However, PNC has not been very well defined in most previous studies. Is it a pattern or a process? What are the underlying endogenous (e.g. genetic) and exogenous (e.g. ecological) factors that cause niches to be conserved? What degree of similarity is necessary to qualify as PNC? Is it possible for the evolutionary processes causing niches to be conserved to also result in niche divergence in different habitats? Here, we revisit these questions, codifying a theoretical and operational definition of PNC as a mechanistic evolutionary process resulting from several factors. We frame this both from a macroevolutionary and population-genetic perspective. We discuss how different axes of physical (e.g. geographic) and environmental (e.g. climatic) heterogeneity interact with the fundamental process of PNC to produce different outcomes of ecological speciation. We also review tests for PNC, and suggest ways that these could be improved or better utilized in future studies. Ultimately, PNC as a process has a well-defined mechanistic basis in organisms, and future studies investigating ecological speciation would be well served to consider this, and frame hypothesis testing in terms of the processes and expected patterns described herein. The process of PNC may lead to patterns where niches are conserved (more similar than expected), constrained (divergent within a limited subset of available niches), or divergent (less similar than expected), based on degree of phylogenetic relatedness between species.

Snake diversity in the island of Sri Lanka is extremely high, hosting at least 89 inland (i.e., non-marine) snake species, of which at least 49 are endemic. This includes the endemic genera Aspidura, Balanophis, Cercaspis, Haplocercus, and Pseudotyphlops, which are of uncertain phylogenetic affinity. We present phylogenetic evidence from nuclear and mitochondrial loci showing the relationships of 40 snake species from Sri Lanka (22 endemics) to the remaining global snake fauna. To determine the phylogenetic placement of these species, we create a molecular dataset containing 10 genes for all global snake genera, while also sampling all available species for genera with endemic species occurring in Sri Lanka. Our sampling comprises five mitochondrial genes (12S, 16S, cyt-b, ND2, and ND4) and five nuclear genes (BDNF, c-mos, NT3 RAG-1, and RAG-2), for a total of up to 9582bp per taxon. We find that the five endemic genera represent portions of four independent colonizations of Sri Lanka, with Cercaspis nested within Colubrinae, Balanophis in Natricinae, Pseudotyphlops in Uropeltidae, and that Aspidura+Haplocercus represents a distinct, ancient lineage within Natricinae. We synonymize two endemic genera that render other genera paraphyletic (Haplocercus with Aspidura, and Cercaspis with Lycodon), and discover that further endemic radiations may be present on the island, including a new taxon from the blindsnake family Typhlopidae, suggesting a large endemic radiation. Despite its small size relative to other islands such as New Guinea, Borneo, and Madagascar, Sri Lanka has one of the most phylogenetically diverse island snake faunas in the world, and more research is needed to characterize the island's biodiversity, with numerous undescribed species in multiple lineages.

Numerous taxa show ancient intertropical disjunct distributions. Many can be explained by well-known processes of historical vicariance, such as the breakup of Gondwanaland. Others, such as Asian-Neotropical divergences are not as well understood. To clarify the phylogenetic position and understand biogeographic and temporal origins of the geographically disjunct and morphologically unique genera of hinged-teeth snakes, Scaphiodontophis (n=1) and Sibynophis (n=9; Colubridae), we inferred a time-calibrated phylogeny with additional 107 taxa representing the superfamily Colubroidea using four genes (c-mos, cyt-b, ND2, RAG-1; 3085 bp). We used this tree to estimate ancestral areas for the group. The results show that Scaphiodontophis is sister to Sibynophis, both originated in the late Eocene/Oligocene in Asia and likely dispersed through Beringia to the New World, but unlike other snake groups left no extant species in temperate North America. Current recognition of Scaphiodontophiinae renders Colubrinae paraphyletic, and we resurrect the previously named subfamily Sibynophiinae to encompass both genera and use the tribes Sibynophiini (Sibynophis) and Scaphiodontophiini (Scaphiodontophis) to highlight the geographically distinct areas occupied by these taxa. These results suggest that intercontinental dispersal with extinction in intermediate areas can explain puzzling patterns of ancient intertropical disjunct distributions.

Snake diversity varies by at least two orders of magnitude among extant lineages, with numerous groups containing only one or two species, and several young clades exhibiting exceptional richness (700 taxa). With a phylogeny containing all known families and subfamilies, we find that these patterns cannot be explained by background rates of speciation and extinction. The majority of diversity appears to derive from a radiation within the superfamily Colubroidea, potentially stemming from the colonization of new areas and the evolution of advanced venom-delivery systems. In contrast, negative relationships between clade age, clade size, and diversification rate suggest the potential for possible bias in estimated diversification rates, interpreted by some recent authors as support for ecologically mediated limits on diversity. However, evidence from the fossil record indicates that numerous lineages were far more diverse in the past, and that extinction has had an important impact on extant diversity patterns. Thus, failure to adequately account for extinction appears to prevent both rate- and diversity-limited models from fully characterizing richness dynamics in snakes. We suggest that clade-level extinction may provide a key mechanism for explaining negative or hump-shaped relationships between clade age and diversity, and the prevalence of ancient, species-poor lineages in numerous groups.

Based on a combination of six mitochondrial gene fragments (12S RNA, cyt b, ND1, ND2, ND4 and CO1) and one nuclear gene (c-mos) from 22 genera we infer phylogenetic relationships among natricine snakes and examine the date and area of origin of these snakes. Our phylogenetic results indicate: (1) the subfamily Natricinae is strongly supported as monophyletic including a majority of extant genera, and a poorly known and previously unassigned species Trachischium monticola, (2) two main clades are inferred within Natricinae, one containing solely taxa from the Old World (OW) and the other comprising taxa from a monophyletic New World (NW) group with a small number of OW relatives. Within the first clade, the genera Xenochrophis and Amphiesma are apparently not monophyletic. Divergence dating and ancestral area estimation indicate that the natricines originated in tropical Asia during the later Eocene or the Oligocene. We recover two major dispersals events out of Asia, the first to Africa in the Oligocene (28 Ma) and the second to the Western Palearctic and the New World at 27 Ma. This date is consistent with the dispersal of numerous other OW groups into the NW.

Molecular phylogenies are often used to test hypotheses about the tempo and mode of speciation and extinction. One commonly used statistic is Pybus and Harvey's γ, which measures the density of ordered internode distances on an ultrametric tree to infer earlier (negative γ) or later (positive γ) bursts of diversification. However, coalescent theory predicts that γ might be biased toward negative values (inferring early bursts of diversification) when using gene trees rather than species trees. Gene divergences predate species divergences, increasingly so at higher effective population sizes (N(e)), and proportionally more so toward the tips of the tree. Thus, gene trees will have a higher density of older nodes in many cases (particularly at higher N(e)), due to the disproportionate lengthening of terminal branches. This will yield an artifactual signature of early bursts of diversification when estimating γ from gene trees. We simulate gene trees within species trees under both Yule (pure-birth) and birth-death processes, and demonstrate support for these predictions. However, for most realistic estimates of θ in natural populations, gene trees provide relatively good estimates of γ, despite the disproportionate overestimation of younger node ages. This is corroborated with an empirical dataset of North American fence lizards (Sceloporus).

Aim Three common patterns have emerged in comparative phylogeographic analyses at many barriers: (1) a potential geographic pseudocongruence of lineage divergences; (2) a disconnect between the inference of temporally clustered, relatively recent timing for observed speciation events, and dates spanning a broader, apparently random time-scale; and (3) an apparent prevalence of speciation with recent or continuing gene flow. It is unclear if there is a unifying explanation for these phenomena. We argue that the interaction between geographic barriers to dispersal and ecological limits on the distribution of species can explain these patterns. We suggest that these patterns can be explained by the presence of a continuum between two underlying processes, here termed ‘hard’ and ‘soft’ allopatric divergence, which result from the interplay between organismal ecology and the physioecological nature of geographic barriers. Location Examples from North America. Methods We examine comparative phylogeographic divergences in 18 groups of terrestrial vertebrates at two major biogeographic features in North America ‐ the Mississippi River Embayment and the Cochise Filter Barrier ‐ to test predictions made by this hypothesis. Results We find support for the two distinct processes of hard and soft allopatry, and note several examples exhibiting characteristics of both. Hard allopatry is caused by physical barriers promoting divergence as a function of consistent geographic isolation. Soft allopatry is caused by ecological processes that isolate populations geographically in allopatric refugia through niche conservatism, or across ecological transition zones through niche divergence, but which may be periodic or inconsistent through time.

The impact of barriers to dispersal and gene flow is often inferred to be the primary cause of lineage divergence and phylogeographic structure in terrestrial organisms. In particular, the Mississippi River has been implicated as a barrier to gene flow in many species, including aquatic taxa. However, if barriers are permeable to organisms, then phylogeographic structure may be difficult to detect due to gene flow between lineages. Using time-calibrated Bayesian phylogenetic analyses of mtDNA, and phylogeographic coalescent simulations, we determine if the Mississippi River operates as a barrier to gene flow in the aquatic diamond-backed watersnake (Nerodia rhombifer). The phylogenetic analyses support a basal division within N. rhombifer mtDNA lineages that coincides with populations generally east and west of the Mississippi River. These results, and that of the divergence dating analyses, therefore suggest that the river was a significant barrier to gene flow in the Pleistocene � 1.4 million years ago, presumably during an interglacial period when the river was much wider. However, we also detect western haplotypes in the eastern clade, and vice versa, thereby indicating that this barrier has not been complete. Nonetheless, the coalescent simulations that account for limited migration suggest that the Mississippi River was an important feature that shaped the phylogeographic history of this aquatic snake in the USA despite limited gene flow.

Ecological adaptive radiation theory predicts an increase in both morphological and specific diversification when organisms colonize new environments. Accordingly, bursts of morphological diversification, characterized by low within-subclade morphological disparity, may be associated with these increases in speciation rates. Conversely, increasing species density, reduction in available habitat, or increasing extinction rates are expected to cause rates of diversification to decline. We test these hypotheses by examining the tempo and mode of speciation in the lampropeltinine snakes, a morphologically variable group that colonized the New World approximately 24 million years ago and radiated throughout the Miocene. We show that specific diversification increased early in the history of the group, and that most morphological variation is partitioned among, rather than within subclades. These patterns provide further evidence for the hypothesis that morphological variation tends to be strongly partitioned among lineages when clades undergo early bursts of species diversification. A reduction in speciation rates may be indicative of density dependent effects due to a saturation of available ecological opportunity, rather than increases in extinction rates at the onset of the Pleistocene/Pliocene glacial cycles. This evidence runs counter to the general Pleistocene species pump model.