Phylogeography of West Indies Coral snakes (Micrurus): Island colonisation and banding patterns

In this study, we analyse New World coral snakes in a phylogenetic framework based upon an increased molecular data set, including novel sequences for the only two sympatric species known from an island (Trinidad, West Indies). Their presence in Trinidad and absence in Tobago offers a unique system to study the phylogeography of the region. We assess the tempo and mode of colonisation of Micrurus on the island, in addition to discussing the phylogenetic relationships for the genus Micrurus concerning two phenotypic traits, body and tail banding patterns. These relationships are analysed for the first time on statistical coalescent phylogeographic discrete ancestral reconstruction. We find a robust phylogenetic component in these characteristics, where strongly supported clades are recovered: prior to the onset of the Early Miocene, a triadal and tricolour tail clade composed of species from South America, and a second clade dating to the Middle‐Late‐ Miocene with monadal and bicolour tails widely distributed from North to South America. The divergence between clades dates to the Oligocene and suggests an ancient pre‐isthmus divergence supporting the arrival of the triadal clade into South America, before the connection between Central and South America was established. We find the two coral snakes present in the West Indies, M. diutius and M. circinalis, belong to the triadal and monadal clades, respectively. Guyana and Trinidad Micrurus diutius share the same haplotypes suggesting a Late Pleistocene–Holocene vicariance when sea level rises separated Trinidad from the mainland. A second lineage of diutius‐like snakes is present in Guyana and is confirmed as M. lemniscatus which is assigned as a voucher and restricts the type locality for M. lemniscatus.     

Phylogenetics,delimitation and historical biogeography of the pantropical tree genus Thespesia (Malvaceae,Gossypieae)

Thespesia consists of 16 species of trees and shrubs from Southeast Asia–Oceania, Africa and America, the most well known being T. populnea, a small tree of tropical coastal areas around the world. Phylogenetic relationships in the genus and among its allies in tribe Gossypieae were inferred using three plastid and two nuclear regions to ascertain its generic delimitation and explore its biogeographical history. Maximum‐likelihood and Bayesian analyses confirmed that Thespesia is not monophyletic and, based on these results, Azanza is reinstated to accommodate the two species previously placed in Thespesia section Lampas. Dating analyses and ancestral range estimation indicated that Thespesia s.s. most likely originated in Southeast Asia–Oceania c. 30 Mya, but extant species did not begin to differentiate until the late Miocene. Two dispersal events, one into Africa c. 11 Mya and another into America (Antilles) c. 9 Mya, gave rise to the African and the Greater Antillean endemics, respectively. The two most widespread hydrochorous species, T. populnea and T. populneoides, originated in Southeast Asia–Oceania from where they spread to other parts of the world. Our analysis also indicated a much earlier origin than previously reported for the eumalvoid clade and its tribes Gossypieae, Malveae and Hibisceae suggesting that vicariance might have had an important role early in the history of these groups.     

Evolution of the African slippery frogs (Anura: Conraua), including the world’s largest living frog

Although Conraua goliath is well known as the largest living frog species, the diversity and evolution of the genus Conraua across sub-Saharan Africa remain poorly understood. We present multilocus phylogenetic analyses of the six currently recognized species that provide insights into divergence times, biogeography, body size evolution and undescribed species. An analysis of divergence times demonstrates that crown-group Conraua arose some time during the latest Oligocene to mid-Miocene followed by divergence into major lineages in the mid-Miocene that may reflect the fragmentation of widespread tropical forests in Africa that began at this time. We find three pairs of sister species, C. crassipes + C. beccarii, C. alleni + C. derooi and C. goliath + C. robusta, each of which diverged during the Miocene. These relationships reject phylogenetic hypotheses based solely on biogeography as the geographically peripheral C. beccarii from north-eastern Africa is nested within western African species and the Central African species do not form a clade. Our species delimitation analyses provide support for undescribed species in C. alleni, C. beccarii and C. derooi, and possibly C. crassipes, suggesting that the current taxonomy substantially underestimates species diversity. There is no clear directional trend of either increasing or decreasing body size in Conraua and the three largest species do not form a clade. With a robust phylogenetic hypothesis in hand, further field-based studies are needed to understand the evolution of morphology and life history in this charismatic African anuran clade.     

Phylogeny and species delimitation of North European Lumbricillus (Clitellata,Enchytraeidae)

The enchytraeid genus Lumbricillus comprises about 80 described species of clitellate worms, which are up to a few centimetres long, and they mostly inhabit the littoral zone of non‐tropical marine and brackish waters world‐wide. The phylogeny of this genus is poorly studied, but previous work has suggested that Lumbricillus is a non‐monophyletic group. In this study, species boundaries and the phylogeny of this genus is re‐assessed using more than 300 DNA‐barcoded specimens (using COI mtDNA), part of which was also sequenced for two additional mitochondrial and four nuclear molecular markers. Statistical and coalescent based applications were used for the delimitation of a total of 24 species, of which 20 were identified as belonging to 17 described morphospecies; one morphospecies was found to be a complex of four delimited species, and another four delimited species could not be matched with any described species. Furthermore, gene trees, concatenation and multispecies coalescent based species trees were estimated using Bayesian inference. The estimated phylogenies confirm a non‐monophyletic Lumbricillus as L. semifuscus is clearly excluded from the genus. Furthermore, the placement of a monophyletic clade consisting of L. arenarius, L. dubius, and an unidentified species varies between analyses; they are either found as the sister‐group to the genus Grania or as sister‐group to the remaining Lumbricillus, where the latter relationship is supported by the multispecies coalescent, which we consider as the most reliable method.     

Species delimitation in the Aleochara fucicola species complex (Coleoptera: Staphylinidae: Aleocharinae) and its phylogenetic relationships

To assess the accuracy of species delimitation and phylogenetic relationships of the Aleochara fucicola species complex, we performed molecular phylogenetic analyses. Detailed micromorphological characters were also examined using scanning electron microscopy (SEM). The molecular data set included two mitochondrial (COI and COII) and three nuclear protein‐coding genes (CAD, EF1‐α and wg) for 19 populations of five species. Significant discordance was found between mitochondrial and nuclear gene trees. Two species, A. puetzi (Assing) and A. segregata Yamamoto and Maruyama, were not separated in the mitochondrial gene trees, but clearly were differentiated in the nuclear and concatenated gene trees. The SEM data also supported the separation of these two species, but an analysis of genetic divergence data did not. Adaptation to extremely harsh environments might reduce morphological variation in the A. fucicola species complex during the colonization of seashores. We present a multilocus phylogeny of the species complex. It suggests that the ancestor of the A. fucicola species complex occurred along the southern coasts of Northeast Asia, followed by dispersals to northern coasts.     

Deep phylogeographic divergence of a migratory passerine in Sino‐Himalayan and Siberian forests: the Red‐flanked Bluetail (Tarsiger cyanurus) complex

Enormous mountainous forests in Sino‐Himalayans and Siberia harbor important avian biodiversity in the Northern Hemisphere. Numerous studies in last two decades have been contributed to systematics and taxonomy of passerines birds in these regions and have revealed various and complex phylogeographic patterns. A passerine species Red‐flanked Bluetail Tarsiger cyanurus provided a good system to manifest such evolutionary complexity. The subspecies T. c. cyanurus and T. c. rufilatus (or/and T. c. pallidior), divergent in morphology, acoustics, and migratory strategies are allopatric in Siberia and Sino‐Himalayan forests, respectively. The two taxa most likely deserve full species status but rigorous genetic analysis is missing. In this study, multilocus phylogeography based on mitochondrial DNA and Z‐linked DNA reveals that T. c. cyanurus and T. c. rufilatus are reciprocally monophyletic with significant statistical support and differ with a large number of diagnostic nucleotide sites resulting substantial genetic divergence. Our finding supports the proposed split of Tarsiger cyanurus s.l. that T. cyanurus and T. rufilatus should be treated as two full species. Whether “pallidior” is a subspecies or geographical form of T. rufilatus is still uncertain. Additionally, these two forest passerine species may have diverged 1.88 (3.25–1.30) Mya, which might be shaped by geographical vicariance due to grassland and desert steppe on the central Loess Plateau during the Pliocene. Taken together, this study and further suggests another independent example of North Palearctic–Sino‐Himalayan phylogeographic pattern in Palearctic birds.     

Evolutionary history of spiny‐tailed lizards (Agamidae: Uromastyx) from the Saharo‐Arabian region

The subfamily Uromastycinae within the Agamidae is comprised of 18 species: three within the genus Saara and 15 within Uromastyx. Uromastyx is distributed in the desert areas of North Africa and across the Arabian Peninsula towards Iran. The systematics of this genus has been previously revised, although incomplete taxonomic sampling or weakly supported topologies resulted in inconclusive relationships. Biogeographic assessments of Uromastycinae mostly agree on the direction of dispersal from Asia to Africa, although the timeframe of the cladogenesis events has never been fully explored. In this study, we analysed 129 Uromastyx specimens from across the entire distribution range of the genus. We included all but one of the recognized taxa of the genus and sequenced them for three mitochondrial and three nuclear markers. This enabled us to obtain a comprehensive multilocus time‐calibrated phylogeny of the genus, using the concatenated data and species trees. We also applied coalescent‐based species delimitation methods, phylogenetic network analyses and model‐testing approaches to biogeographic inferences. Our results revealed Uromastyx as a monophyletic genus comprised of five groups and 14 independently evolving lineages, corresponding to the 14 currently recognized species sampled. The onset of Uromastyx diversification is estimated to have occurred in south‐west Asia during the Middle Miocene with a later radiation in North Africa. During its Saharo‐Arabian colonization, Uromastyx underwent multiple vicariance and dispersal events, hypothesized to be derived from tectonic movements and habitat fragmentation due to the active continental separation of Arabia from Africa and the expansion and contraction of arid areas in the region.     

Lineage divergence and speciation in the Web‐toed Salamanders (Plethodontidae: Hydromantes) of the Sierra Nevada,California

Peripatric speciation and the importance of founder effects have long been controversial, and multilocus sequence data and coalescent methods now allow hypotheses of peripatric speciation to be tested in a rigorous manner. Using a multilocus phylogeographical data set for two species of salamanders (genus Hydromantes) from the Sierra Nevada of California, hypotheses of recent divergence by peripatric speciation and older, allopatric divergence were tested. Phylogeographical analysis revealed two divergent lineages within Hydromantes platycephalus, which were estimated to have diverged in the Pliocene. By contrast, a low‐elevation species, Hydromantes brunus, diverged from within the northern lineage of H. platycephalus much more recently (mid‐Pleistocene), during a time of major climatic change in the Sierra Nevada. Multilocus species tree estimation and coalescent estimates of divergence time, migration rate, and growth rate reject a scenario of ancient speciation of H. brunus with subsequent gene flow and introgression from H. platycephalus, instead supporting a more recent divergence with population expansion. Although the small, peripheral distribution of H. brunus suggests the possibility of peripatric speciation, the estimated founding population size of the species was too large to have allowed founder effects to be important in its divergence. These results provide evidence for both recent speciation, most likely tied to the climatic changes of the Pleistocene, and older lineage divergence, possibly due to geological events, and add to evidence that Pleistocene glacial cycles were an important driver of diversification in the Sierra Nevada.     

Evolutionary history of Gymnocarpos (Caryophyllaceae) in the arid regions from North Africa to Central Asia

Gymnocarpos has only about ten species distributed in the arid regions of Asia and Africa, but it exhibits a geographical disjunction between eastern Central Asia and western North Africa and Minor Asia. We sampled eight species of the genus and sequenced two chloroplast regions (rps16 and psbB–psbH), and the nuclear rDNA (ITS) to study the phylogeny and biogeography. The results of the phylogenetic analyses corroborated that Gymnocarpos is monophyletic, in the phylogenetic tree two well supported clades are recognized: clade 1 includes Gymnocarpos sclerocephalus and G. decandrus, mainly the North African group, whereas clade 2 comprises the remaining species, mainly in the Southern Arabian Peninsula. Molecular dating analysis revealed that the divergence age of Gymnocarpos was c. 31.33 Mya near the Eocene and Oligocene transition boundary, the initial diversification within Gymnocarpos dated to c. 6.69 Mya in the late Miocene, and the intraspecific diversification mostly occurred during the Quaternary climate oscillations. Ancestral area reconstruction suggested that the Southern Arabian Peninsula was the ancestral area for Gymnocarpos. Our conclusions revealed that the aridification since mid‐late Miocene significantly affected the diversification of the genus in these areas.     

Taxonomy and biogeography of Bunopus spatalurus (Reptilia; Gekkonidae) from the Arabian Peninsula

In the last decade, taxonomic studies have drastically increased the number of species known to inhabit the Arabian deserts. While ongoing phylogenetic studies continue to identify new species and high levels of intraspecific genetic diversity, few studies have yet explored the biogeographic patterns in this arid region using an integrative approach. In the present work, we apply different phylogenetic methods to infer relationships within the Palearctic naked‐toed geckos. We specifically address for the first time the taxonomy and biogeography of Bunopus spatalurus Anderson, 1901, from Arabia using multilocus concatenated and species tree phylogenies, haplotype networks and morphology. We also use species distribution modelling and phylogeographic interpolation to explore the phylogeographic structure of Bunopus spatalurus hajarensis in the Hajar Mountains and the roles of climatic stability and possible biogeographic barriers on lineage occurrence and contact zones in this arid mountain endemism hot spot. According to the inferred topology recovered using concatenated and species tree methods, the genus ‘Bunopus’ is polyphyletic. Bunopus tuberculatus and B. blanfordii form a highly supported clade closely related to Crossobamon orientalis, while the two subspecies of ‘Bunopus’ spatalurus branch together as an independent highly supported clade that diverged during the Miocene according to our estimations. Within B. s. hajarensis, three geographically structured clades can be recognized that according to our estimations diverged during the Late Miocene to Pliocene. The paleodistribution models indicate climatic stability during the Late Pleistocene and the lineage occurrence, and predicted contact zones obtained from phylogeographic interpolation therefore probably result from the older splits of the groups when these lineages originated in allopatry. As demonstrated by the results of the multilocus molecular phylogenetic analyses and the topological test carried out in this study, the genus ‘Bunopus’ is not monophyletic. To resolve this, we resurrect the genus Trachydactylus Haas and Battersby, 1959; for the species formerly referred to as Bunopus spatalurus. Considering the morphological differences, the high level of genetic differentiation in the 12S mitochondrial gene and the results of the phylogenetic and the cmos haplotype network analysis, we elevate Trachydactylus spatalurus hajarensis to the species level Trachydactylus hajarensis (Arnold, 1980).     

Miocene diversification of an open‐habitat predatorial passerine radiation,the shrikes (Aves: Passeriformes: Laniidae)

Diversification of avifaunas associated with savannah and steppes appears to correlate with open habitats becoming available, starting in the Miocene. Few comparative analyses exist for families for which all species are predominantly adapted to these habitats. One such group is Laniidae (Passeriformes), which are small‐ to medium‐sized predatory passerines known for their distinctive behaviour of impaling prey. We used multispecies coalescent‐based and concatenation methods to provide the first complete species‐level phylogeny for this group, as well as an estimate of the timing of diversification. Our analyses indicate that Laniidae as currently delimited is not monophyletic, as the genus Eurocephalus is not closely related to the remaining species. The two species currently assigned to the monotypic genera Urolestes and Corvinella are part of the same clade as the Lanius species, and we propose that they are included in the genus Lanius, making Laniidae monogeneric. The initial diversification of the clade is inferred to have occurred very rapidly, starting about 7.2–9.1 million years ago, timing depending on calibration method, but in either case coinciding with the expansion of C4 grasses. An African origin is inferred in the biogeographic analysis. In the redefined Laniidae, cooperative breeding is inferred to be restricted to a single clade, characterized by gregarious behaviour and rallying. Migratory behaviour evolved multiple times within the family.     

Phylogenomics unravels Quaternary vicariance and allopatric speciation patterns in temperate‐montane plant species: A case study on the Ranunculus auricomus species complex

The time frame and geographical patterns of diversification processes in European temperate‐montane herbs are still not well understood. We used the sexual species of the Ranunculus auricomus complex as a model system to understand how vicariance versus dispersal processes in the context of Pleistocene climatic fluctuations have triggered speciation in temperate‐montane plant species. We used target enrichment sequence data from about 600 nuclear genes and coalescent‐based species tree inference methods to resolve phylogenetic relationships among the sexual taxa of the complex. We estimated absolute divergence times and, using ancestral range reconstruction, we tested if speciation was enhanced by vicariance or by dispersal processes. Phylogenetic relationships among taxa were fully resolved with some incongruence in the position of the tetraploid R. marsicus. Speciation events took place in a very short time at the end of the Mid‐Pleistocene Transition (830–580 thousand years ago [ka]). A second wave of intraspecific geographical differentiation occurred at the end of the Riss glaciation or during the Eemian interglacial between 200 and 100 ka. Ancestral range reconstruction suggests a widespread European ancestor of the R. auricomus complex. Vicariance has triggered allopatric speciation in temperate‐montane plant species during the climatic deterioration that occurred during the last phase of the Mid‐Pleistocene Transition. Vegetation restructuring from forest into tundra could have confined these forest species into isolated glacial macro‐ and microrefugia. During subsequent warming periods, range expansions of these species could have been hampered by apomictic derivatives and by other congeneric competitors in the same habitat.     

Speciation,population structure,and demographic history of the Mojave Fringe‐toed Lizard (Uma scoparia), a species of conservation concern

The North American deserts were impacted by both Neogene plate tectonics and Quaternary climatic fluctuations, yet it remains unclear how these events influenced speciation in this region. We tested published hypotheses regarding the timing and mode of speciation, population structure, and demographic history of the Mojave Fringe‐toed Lizard (Uma scoparia), a sand dune specialist endemic to the Mojave Desert of California and Arizona. We sampled 109 individual lizards representing 22 insular dune localities, obtained DNA sequences for 14 nuclear loci, and found that U. scoparia has low genetic diversity relative to the U. notata species complex, comparable to that of chimpanzees and southern elephant seals. Analyses of genotypes using Bayesian clustering algorithms did not identify discrete populations within U. scoparia. Using isolation‐with‐migration (IM) models and a novel coalescent‐based hypothesis testing approach, we estimated that U. scoparia diverged from U. notata in the Pleistocene epoch. The likelihood ratio test and the Akaike Information Criterion consistently rejected nested speciation models that included parameters for migration and population growth of U. scoparia. We reject the Neogene vicariance hypothesis for the speciation of U. scoparia and define this species as a single evolutionarily significant unit for conservation purposes.     

Phylogenetic and diversity patterns of Blanus worm lizards (Squamata: Amphisbaenia): insights from mitochondrial and nuclear gene genealogies and species tree

Phylogenetic and phylogeographic patterns of amphisbaenians are poorly known. Molecular data from mitochondrial and nuclear loci are particularly needed for amphisbaenian phylogeny and taxonomy because their specializations to subterranean habits make morphology poorly informative and the occurrence of cryptic species probable. The Mediterranean genus Blanus includes five species – three of them have been recently studied mainly at the mitochondrial level. In this study, we collected mitochondrial (16S and nd4) and nuclear (mc1r and pomc) sequences from 49 specimens, including multiple individuals for each of the five species. We used multilocus coalescent‐based species‐tree inference and single‐gene analyses to estimate phylogenetic relationships among Blanus and to assess patterns of intraspecific differentiation within all the five species. Species‐tree and single‐gene phylogenies provided strong support for the Anatolian worm lizard B. strauchi lying outside a clade comprising all other congeners, with a sister relationship between the Iberian clade (B. cinereus and B. mariae) and the North African clade (B. tingitanus and B. mettetali). Mitochondrial and nuclear data supported the genetic distinctiveness of the recently described B. mariae and also indicated that the distribution of this species is wider than previously known and overlaps with B. cinereus in central Portugal. Blanus tingitanus showed two phylogeographic groups, from the northern and the southern portion of the range, respectively, having high mitochondrial and nuclear divergence and a possible contact zone in northwestern Morrocco. Finally, high genetic variation was found within B. mettetali and B. strauchi, suggesting in the latter case, the occurrence of cryptic taxa to be tested by further research.     

Systematics and phylogeny of Vasseuromys (Mammalia,Rodentia, Gliridae) with a description of a new species from the late Miocene of eastern Europe

Vasseuromys is a species‐rich genus of small‐ to medium‐sized glirids spanning the latest Oligocene to late Miocene of Europe and western Asia. Despite extensive discoveries over the past 50 years, little phylogenetic work has been done on Vasseuromys. This study presents the first phylogenetic analysis of the genus that includes all the described species and a new taxon Vasseuromys tectus sp. nov. from the late Miocene of eastern Europe, providing the first insights into the evolutionary relationships within the clade. Results suggest that the genus is clearly paraphyletic. Two strongly supported genus‐level clades are recognized within ‘Vasseuromys’: a restricted Vasseuromys clade (containing the three species, V. pannonicus, V. rugosus and V. tectus) and the Glirulus clade that includes ‘Vasseuromys’ duplex. The remaining ‘Vasseuromys’ species are found to constitute a set of paraphyletic taxa, with the polyphyletic ‘Ramys’ nested within it. The genus Gliruloides is synonymized with Glirulus. Vasseuromys tectus sp. nov. is the most derived member of the genus in having a greater number of cheek teeth ridges including constantly present anterotrope, centrotrope, second prototrope on M1–2, third metatrope on M2, two to three posterotropids on p4 and strong ectolophids on lower molars. The results of the study confirm a European origin for Vasseuromys while suggesting that the late Miocene species of the genus dispersed from the east in the early Turolian.     

Timing of clade divergence and discordant estimates of genetic and morphological diversity in the Slender Madtom,Noturus exilis (Ictaluridae)

Phylogeographic relationships, the timing of clade diversification, and the potential for cryptic diversity in the Slender Madtom, Noturus exilis, was investigated using mitochondrial Cyt b, nuclear RAG2, shape analysis, and meristic and pigmentation data. Three well-supported and deeply divergent clades were recovered from analyses of genetic data: Little Red River (White River drainage) clade, Arkansas + Red River (Mississippi River) clade, and a large clade of populations from the rest of the range of the species. Recovered clades showed little to no diagnostic morphological differences, supporting previous hypotheses of morphological conservatism in catfishes, and indicating morphology may commonly underestimate diversity in this group of fishes. The Little Red River clade is the most distinct lineage of N. exilis with 11 POM pores (vs. 10 in other populations) and unique Cyt b haplotypes and RAG 2 alleles. However, treating it as a species separate from N. exilis would imply that the other major clades of N. exilis are more closely related to one another than they are to the Little Red River clade, which was not supported.The UCLN age estimate for Noturus was 23.9 mya (95% HPD: 13.49, 35.43), indicating a late Oligocene to early Miocene origin. The age of N. exilis was estimated as late Miocene at 9.7 mya (95% HPD: 5.32, 14.93). Diversification within the species spanned the late Miocene to mid-Pleistocene. The largest clade of N. exilis, which dates to the late Miocene, includes populations from the unglaciated Eastern and Interior Highlands as well as the previously glaciated Central Lowlands. Diversification of this clade coincides with a drastic drop in sea-level and diversification of other groups of Central Highlands fishes (Centrarchidae and Cyprinidae). Sub-clades dating to the Pleistocene show that northern populations occurring in previously glaciated regions resulted from dispersal from populations in the Ozarks up the Mississippi River following retreat of the Pleistocene glaciers. Pre-Pleistocene vicariance, such as drainage pattern changes of the Mississippi River, also played a prominent role in the history of the species. The incorporation of a temporal estimate of clade diversification revealed that in some instances, phylogeographic breaks shared with other aquatic species were best explained by different or persistent vicariant events through time, rather than a single shared event.     

CALIBRATING DIVERGENCE TIMES ON SPECIES TREES VERSUS GENE TREES: IMPLICATIONS FOR SPECIATION HISTORY OF APHELOCOMA JAYS

Estimates of the timing of divergence are central to testing the underlying causes of speciation. Relaxed molecular clocks and fossil calibration have improved these estimates; however, these advances are implemented in the context of gene trees, which can overestimate divergence times. Here we couple recent innovations for dating speciation events with the analytical power of species trees, where multilocus data are considered in a coalescent context. Divergence times are estimated in the bird genus Aphelocoma to test whether speciation in these jays coincided with mountain uplift or glacial cycles. Gene trees and species trees show general agreement that diversification began in the Miocene amid mountain uplift. However, dates from the multilocus species tree are more recent, occurring predominately in the Pleistocene, consistent with theory that divergence times can be significantly overestimated with gene‐tree based approaches that do not correct for genetic divergence that predates speciation. In addition to coalescent stochasticity, Haldane's rule could account for some differences in timing estimates between mitochondrial DNA and nuclear genes. By incorporating a fossil calibration applied to the species tree, in addition to the process of gene lineage coalescence, the present approach provides a more biologically realistic framework for dating speciation events, and hence for testing the links between diversification and specific biogeographic and geologic events.     

Northern Hemisphere origin,transoceanic dispersal,and diversification of Ranunculeae DC. (Ranunculaceae) in the Cenozoic

Aim The role of dispersal versus vicariance for plant distribution patterns has long been disputed. We study the temporal and spatial diversification of Ranunculeae, an almost cosmopolitan tribe comprising 19 genera, to understand the processes that have resulted in the present inter‐continental disjunctions. Location All continents (except Antarctica). Methods Based on phylogenetic analyses of nuclear and chloroplast DNA sequences for 18 genera and 89 species, we develop a temporal–spatial framework for the reconstruction of the biogeographical history of Ranunculeae. To estimate divergence dates, Bayesian uncorrelated rates analyses and four calibration points derived from geological, fossil and external molecular information were applied. Parsimony‐based methods for dispersal–vicariance analysis (diva and Mesquite ) and a maximum likelihood‐based method (Lagrange ) were used for reconstructing ancestral areas. Six areas corresponding to continents were delimited. Results The reconstruction of ancestral areas is congruent in the diva and maximum likelihood‐based analyses for most nodes, but Mesquite reveals equivocal results at deep nodes. Our study suggests a Northern Hemisphere origin for the Ranunculeae in the Eocene and a weakly supported vicariance event between North America and Eurasia. The Eurasian clade diversified between the early Oligocene and the late Miocene, with at least three independent migrations to the Southern Hemisphere. The North American clade diversified in the Miocene and dispersed later to Eurasia, South America and Africa. Main conclusions Ranunculeae diversified between the late Eocene and the late Miocene. During this time period, the main oceanic barriers already existed between continents and thus dispersal is the most likely explanation for the current distribution of the tribe. In the Southern Hemisphere, a vicariance model related to the break‐up of Gondwana is clearly rejected. Dispersals between continents could have occurred via migration over land bridges, such as the Bering Land Bridge, or via long‐distance dispersal.     

Recent rapid speciation and ecomorph divergence in Indo‐Australian sea snakes

The viviparous sea snakes (Hydrophiinae) are a young radiation of at least 62 species that display spectacular morphological diversity and high levels of local sympatry. To shed light on the mechanisms underlying sea snake diversification, we investigated recent speciation and eco‐morphological differentiation in a clade of four nominal species with overlapping ranges in Southeast Asia and Australia. Analyses of morphology and stomach contents identified the presence of two distinct ecomorphs: a ‘macrocephalic’ ecomorph that reaches >2 m in length, has a large head and feeds on crevice‐dwelling eels and gobies; and a ‘microcephalic’ ecomorph that rarely exceeds 1 m in length, has a small head and narrow fore‐body and hunts snake eels in burrows. Mitochondrial sequences show a lack of reciprocal monophyly between ecomorphs and among putative species. However, individual assignment based on newly developed microsatellites separated co‐distributed specimens into four significantly differentiated clusters corresponding to morphological species designations, indicating limited recent gene flow and progress towards speciation. A coalescent species tree (based on mitochondrial and nuclear sequences) and isolation‐migration model (mitochondrial and microsatellite markers) suggest between one and three transitions between ecomorphs within the last approximately 1.2 million to approximately 840 000 years. In particular, the macrocephalic ‘eastern’ population of Hydrophis cyanocinctus and microcephalic H. melanocephalus appear to have diverged very recently and rapidly, resulting in major phenotypic differences and restriction of gene flow in sympatry. These results highlight the viviparous sea snakes as a promising system for speciation studies in the marine environment.