Phylogenetics of Cancer crabs (Crustacea: Decapoda: Brachyura).

We used morphological, mitochondrial DNA sequence, paleontological, and biogeographical information to examine the evolutionary history of crabs of the genus Cancer. Phylogenies inferred from adult morphology and DNA sequence of the cytochrome oxidase I (COI) gene were each well resolved and well supported, but differed substantially in topology. Four lines of evidence suggested that the COI data set accurately reflected Cancer phylogeny: (1) in the phylogeny inferred from morphological data, each Atlantic species was sister taxon to an ecologically similar Pacific species, suggesting convergence in morphology; (2) a single trans-Arctic dispersal event, as indicated by the phylogeny inferred from COI, is more parsimonious than two such dispersal events, as inferred from morphology; (3) test and application of a maximum likelihood molecular clock to the COI data yielded estimates of origin and speciation times that fit well with the fossil record; and (4) the tree inferred from the combined COI and morphology data was closely similar to the trees inferred from COI, although notably less well supported by the bootstrap. The phylogeny inferred from maximum likelihood analysis of COI suggested that Cancer originated in the North Pacific in the early Miocene, that the Atlantic species arose from a North Pacific ancestor, and that Cancer crabs invaded the Atlantic from the North Pacific 6-12 mya. This inferred invasion time is notably prior to most estimates of the date of submergence of the Bering Strait and the trans-Arctic interchange, but it agrees with fossil evidence placing at least one Cancer species in the Atlantic about 8 mya.     

Isolation and characterization of microsatellite loci in the marine gastropod Nucella lapillus

Our paper deals the cloning and characterization of microsatellites from Nucella lapillus, and tests cross‐species amplification in a congener and in two species of the confamilial genus Thais. Fourteen of 31 microsatellite loci tested were polymorphic, with 4–9 (mean 5.93) alleles per locus. The observed heterozygosity per locus varied from 0.10 to 0.85 (mean 0.37) and expected heterozygosity from 0.48 to 0.85 (mean 0.65). Most primer pairs were successfully amplified in N. freycineti, although only one primer pair was successfully amplified in both species of Thais. The markers are potentially useful for other species of Nucella.     

Evolutionary and biogeographical patterns within the smelt genus Hypomesus in the North Pacific Ocean

Aim We address questions about trans‐Pacific distributions of marine organisms and the North Pacific Ocean as a centre of marine biodiversity through a phylogenetic and biogeographical study of a pan‐Pacific genus of Northern Hemisphere smelts (Hypomesus, Pisces: Osmeridae). Location North Pacific Ocean. Methods Relationships of the five species of Hypomesus from throughout the North Pacific were reconstructed through maximum likelihood and Bayesian phylogenetic analyses of sequence data from two mitochondrial (cytb, 16S) and three nuclear (ITS2, S71, RAG1) gene regions of five to 25 individuals per species, totalling 3588 characters. The resulting phylogenies were used to test hypotheses of species relationships and geographical origins using both dispersal‐based and maximum likelihood methods for inferring ancestral areas (lagrange ). Cytb sequence divergence and a Bayesian approach (beast ) were used to estimate the timeframe of Hypomesus evolution, which was compared with work on similarly distributed taxa. Results Hypothesized trans‐Pacific Ocean relationships based on lateral line scale counts were not supported by the phylogeny, suggesting parallel evolution of this phenotype, although we found one such relationship between the western H. japonicus and the two eastern Pacific species (H. pretiosus and H. transpacificus). Dispersalist approaches rejected an early proposal of a double‐compression vicariant mechanism as well as an eastern Pacific origin. Results from the lagrange analysis suggested a more widespread ancestor, although also supporting a role for the western Pacific. Divergence estimates suggested that most splits between species occurred in the mid‐Miocene, and the most recent speciation event, between the eastern Pacific species, occurred in the Pliocene to early Pleistocene. Main conclusions Our molecular data indicate that the character historically used to define relationships within Hypomesus, lateral line scale count, does not reflect ancestry within the genus. Biogeographical reconstructions suggest an important role for the western North Pacific in the diversification of Hypomesus. While uncertainty remains over the date of origin for this genus, estimates place the divergences during periods of climatic cooling that have been important in generating diversity in a number of similarly distributed organisms. Additional comparative data will provide further insight into the relative importance of the western region in generating diversity in the North Pacific Ocean.     

Direct and indirect effects of predation,herbivory and surface rugosity on mussel recruitment

Summary The predatory gastropod Nucella lapillus, commonly preys upon the mussel, Mytilus edulis, and is thought to control the distribution and abundance of mussels on the rocky shores of New England, USA. In this study, done in Maine, USA, not only the presence of Nucella lapillus but also the roughness of the experimental surface and the presence of the herbivorous gastropod, Littorina littorea, were manipulated. Four types of surfaces were used as recruitment substrata for mussels: smooth bare granite, aggregations of the barnacle, Semibalanus balanoides, fiberglass resin castings of smooth bare granite and resin castings of aggregations of S. balanoides. To ensure that caged N. lapillus were not starving, barnacles were provided as alternative prey. Experiments showed no detectable effect of N. lapillus on the recruitment of M. edulis. Mussel recruitment was enhanced by surface rugosity and depressed by the activities of L. littorea. Analysis of covariance, using the number of algal species as the covariate, suggested that L. littorea reduced the number of newlyrecruited mussels by removing algae that provided recruitment sites, but no manipulations were done to test this conjecture. It is likely that previous reports of N. lapillus controlling mussel abundance are attributable to N. lapillus preying upon barnacles, which increase surface rugosity and enhance mussel recruitment. Review of literature on feeding preferences of N. lapillus supports this view. When handling times and prey availability are taken into account, Nucella shows a clear preference for barnacles over mussels.     

Genetic Variation in a Cline in a Living Intertidal Snail Arose in the Neogastropoda over 100 Million Years Ago

To understand how species adapt and evolve it is necessary to appreciate the relationship between genetic variation and the environment. Here, the fossil record and molecular data from different lineages of the marine Gastropoda are used to understand the evolution of genetic variations found in the nuclear gene for mitochondrial malate dehydrogenase (mMDH; EC 1.1.1.37) in the living intertidal Muricid snail, Nucella lapillus. Assuming a molecular clock, DNA sequences of mMDH indicate that two variants found in N. lapillus, mMDH⁹ and mMDH¹⁰, may have arisen as long as 144 MY (million years) ago and at least prior to the evolution of the Muricidae approximately 112–90 MY ago. The Muricidae contain by far the greatest majority of the Neogastropoda specialized for life in the intertidal habitat. In N. lapillus the mMDH⁹ and mMDH¹⁰ variants covary with variations in other biochemically defined loci, inherited phenotypic traits (shell shape and physiology) and karyotype frequencies to differentiate two distinct nuclear haplotypes that are associated with different temperature environments. The variations in shell shape that are associated with the haplotypes of N. lapillus represent adaptations to temperature stress and similar variations occur in other related intertidal molluscs whose lineages are much older than Nucella, which arose around 25 million years ago. It is suggested that the divergence of the mMDH variants found in N. lapillus may reflect an ancient genetic event, such as a chromosomal mutation, perhaps involving variation in other linked traits that together became important in the subsequent evolution of the marine Gastropoda.     

Global mitochondrial DNA phylogeography and biogeographic history of the antitropically and longitudinally disjunct marine bryozoan Membranipora membranacea L. (Cheilostomata): another cryptic marine sibling species complex?

The origin of disjunct distributions in high dispersal marine taxa remains an important evolutionary question as it relates to the formation of new species in an environment where barriers to gene flow are not always obvious. To reconstruct the relationships and phylogeographic history of the antitropically and longitudinally disjunct bryozoan Membranipora membranacea populations were surveyed with mtDNA cytochrome oxidase 1 (COI) sequences across its cosmopolitan range. Maximum parsimony, maximum likelihood and Bayesian genealogies revealed three deep clades in the North Pacific and one monophyletic clade each in the southeast Pacific (Chile), southwest Pacific (Australia/New Zealand), North Atlantic and southeast Atlantic (South Africa). Human-mediated dispersal has not impacted M. membranacea’s large-scale genetic structure. M. membranacea did not participate in the trans-arctic interchange. Episodic long-distance dispersal, combined with climatic vicariance can explain the disjunct distribution. Dispersal led southward across the tropics perhaps 13 mya in the East Pacific and again northwards perhaps 6 mya in the Eastern Atlantic to colonize the North Atlantic from the south, and along the West Wind Drift to colonize Australia. The clades differentiated over evolutionary time in their respective ocean region, potentially forming a sibling species complex. The taxonomic status of the clades is discussed.     

Tethyan vicariance, relictualism and speciation: evidence from a global molecular phylogeny of the opisthobranch genus Bulla

Aim Our aims were: (1) to reconstruct a molecular phylogeny of the cephalaspidean opisthobranch genus Bulla, an inhabitant of shallow sedimentary environments; (2) to test if divergence times are consistent with Miocene and later vicariance among the four tropical marine biogeographical provinces; (3) to examine the phylogenetic status of possible Tethyan relict species; and (4) to infer the timing and causes of speciation events. Location Tropical and warm‐temperate regions of the Atlantic, Indo‐West Pacific, Australasia and eastern Pacific. Methods Ten of the 12 nominal species of Bulla were sampled, in a total sample of 65 individuals, together with cephalaspidean outgroups. Phylogenetic relationships were inferred by Bayesian analysis of partial sequences of the mitochondrial cytochrome c oxidase I (COI) and 16S rRNA and nuclear 28S rRNA genes. Divergence times and rates of evolution were estimated using uncorrelated relaxed‐clock Bayesian methods with fossil calibrations (based on literature review and examination of fossil specimens), implemented in beast . The geographical pattern of speciation was assessed by estimating the degree of overlap between sister lineages. Results Four clades were supported: Indo‐West Pacific (four species), Australasia (one species), Atlantic plus eastern Pacific (three species) and Atlantic (two species), with estimated mean ages of 35–46 Ma. Nominal species were monophyletic, but deep divergences were found within one Indo‐West Pacific and one West Atlantic species. Species‐level divergences occurred in the Miocene or earlier. The age of a sister relationship across the Isthmus of Panama was estimated at 7.9–32.1 Ma, and the divergence of a pair of sister species on either side of the Atlantic Ocean occurred 20.4–27.2 Ma. Main conclusions Fossils suggest that Bulla originated in the Tethys realm during the Middle Eocene. Average ages of the four main clades fall in the Eocene, and far pre‐date the 18–19 Ma closure of the Tethys Seaway. This discrepancy could indicate earlier vicariant events, selective extinction or errors of calibration. Similarly, the transisthmian divergence estimate far pre‐dates the uplift of the Panamanian Isthmus at about 3 Ma. Speciation events occurred in the Miocene, consistent with tectonic events in the central Indo‐West Pacific, isolation of the Arabian Sea by upwelling and westward trans‐Atlantic dispersal. Differences in habitat between sister species suggest that ecological speciation may also have played a role. The basal position of the Australasian species supports its interpretation as a Tethyan relict.     

Phylogeography of the Indo‐West Pacific maskrays (Dasyatidae,Neotrygon): a complex example of chondrichthyan radiation in the Cenozoic

Maskrays of the genus Neotrygon (Dasyatidae) have dispersed widely in the Indo‐West Pacific being represented largely by an assemblage of narrow‐ranging coastal endemics. Phylogenetic reconstruction methods reproduced nearly identical and statistically robust topologies supporting the monophyly of the genus Neotrygon within the family Dasyatidae, the genus Taeniura being consistently basal to Neotrygon, and Dasyatis being polyphyletic to the genera Taeniurops and Pteroplatytrygon. The Neotrygon kuhlii complex, once considered to be an assemblage of color variants of the same biological species, is the most derived and widely dispersed subgroup of the genus. Mitochondrial (COI, 16S) and nuclear (RAG1) phylogenies used in synergy with molecular dating identified paleoclimatic fluctuations responsible for periods of vicariance and dispersal promoting population fragmentation and speciation in Neotrygon. Signatures of population differentiation exist in N. ningalooensis and N. annotata, yet a large‐scale geological event, such as the collision between the Australian and Eurasian Plates, coupled with subsequent sea‐level falls, appears to have separated a once homogeneous population of the ancestral form of N. kuhlii into southern Indian Ocean and northern Pacific taxa some 4–16 million years ago. Repeated climatic oscillations, and the subsequent establishment of land and shallow sea connections within and between Australia and parts of the Indo‐Malay Archipelago, have both promoted speciation and established zones of secondary contact within the Indian and Pacific Ocean basins.     

Phylogeny,biogeography, and morphological ancestral character reconstruction in the Mediterranean genus Fumana (Cistaceae)

Fumana is a diverse genus of the Cistaceae family, consisting of 21 currently accepted species. In this study, nuclear (ITS) and plastid (matK, trnT‐L) molecular markers were used to reconstruct the phylogeny and to estimate divergence times, including 19 species of Fumana. Phylogenetic analyses (Bayesian Inference, Maximum Parsimony and Maximum Likelihood) confirmed the monophyly of Fumana and did not support the infrageneric divisions previously established. The results support four main clades that group species that differ in vegetative and reproductive characters. Given the impossibility to define morphological characters common to all species within the clades, our proposal is to reject infrageneric divisions. Molecular dating and ancestral area analyses provide evidence for a Miocene diversification of the genus in the north‐western Mediterranean. Ancestral state reconstructions revealed ancestral character states for some traits related to xeric and arid habitats, suggesting a preadaptation to the Mediterranean climate.     

The systematics of right whales (Mysticeti: Balaenidae)

Balaenidae (right whales) are large, critically endangered baleen whales represented by four living species. The evolutionary relationships of balaenids are poorly known, with the number of genera, relationships to fossil taxa, and position within Mysticeti in contention. This study employs a comprehensive set of morphological characters to address aspects of balaenid phylogeny. A sister‐group relationship between neobalaenids and balaenids is strongly supported, although this conflicts with molecular evidence, which may be an artifact of long‐branch attraction (LBA). Monophyly of Balaenidae is supported, and three major clades are recognized: (1) extinct genus Balaenula, (2) extant and extinct species of the genus Eubalaena, and (3) extant and extinct species of the genus Balaena plus the extinct taxon, Balaenella. The relationships of these clades to one another, as well as to the early Miocene stem balaenid, Morenocetus parvus, remain unresolved. Pliocene taxa, Balaenula astensis and Balaenula balaenopsis, form a clade that is the sister group to the Japanese Pliocene Balaenula sp. Eubalaena glacialis and Pliocene Eubalaena belgica, are in an unresolved polytomy with a clade including E. japonica and E. australis. Extant and fossil species of Balaena form a monophyletic group that is sister group to the Dutch Pliocene Balaenella, although phylogenetic relationships within Balaena remain unresolved.     

Molecular lineage diversity and inter‐oceanic biogeographical history in Hiatella (Mollusca,Bivalvia)

Hiatella is one of the most widespread marine bivalve genera, occurring in diverse habitats from the temperate to polar latitudes in both hemispheres, and in fossil strata since almost 150 Myr ago. Despite variation in some biological and morphological traits, characters to resolve the current systematic structure consistently across the range of the genus are not known: all samples are often referred to a single species, Hiatella arctica (L.). Exploring the systematics of Hiatella using partial sequences of three genes (mitochondrial COI, and the nuclear ANT and 28S rRNA), we find high diversity of deep lineages (11–22% p‐distance in COI), and identify at least 13 distinct taxa both by heuristic criteria (congruence of the nuclear and mtDNA data) and by coalescence‐based analyses. At several localities, two or three of these cryptic species were found in sympatry. In the framework of previous fossil evidence and of hypotheses of paleoceanographical connections, scenarios of the phylogeny and biogeographical history of the identified species at a range of different time scales are outlined. The distinction between the main North Pacific and North Atlantic Hiatella clades and systematic diversification within each of them seem to have followed a Miocene trans‐Panamanian invasion. Apart from such earlier intra‐basin diversification, the data suggest that three successive counter‐invasions from the Pacific to the Atlantic via the Arctic Ocean route have later contributed to the current North Atlantic Hiatella diversity. These invasions probably took place in connection with (i) the Great Trans‐Arctic Biotic Interchange in the Pliocene, (ii) the last interglacial period c. 120 kya and (iii) the Holocene, postdating the last glaciation. This sequence of trans‐Arctic invasions is largely analogous to that hypothesized for some other boreal‐arctic bivalves.     

Recent origin,active speciation and dispersal for the lichen genus Nephroma (Peltigerales) in Macaronesia

Aim We reconstructed the phylogeny of the lichen genus Nephroma (Peltigerales) to assess the relationships of species endemic to Macaronesia. We estimated dates of divergences to test the hypothesis that the species arose in Macaronesia (neo‐endemism) versus the oceanic archipelagos serving as refugia for formerly widespread taxa (palaeo‐endemism). Location Cosmopolitan with a special focus on the archipelagos of the Azores, Madeira and the Canary Islands. Methods DNA sequences were obtained from 18 species for three loci and analysed using maximum parsimony, maximum likelihood and Bayesian inferences. Divergence dates were estimated for the internal transcribed spacer (ITS)‐based phylogeny using a relaxed molecular clock. Reconstruction of the ancestral geographical range was conducted using the Bayesian 50% majority rule consensus tree under a parsimony method. Results The backbone phylogenetic tree was fully supported, with Nephroma plumbeum as sister to all other species. Four strongly supported clades were detected: the Nephroma helveticum, the N. bellum, the N. laevigatum and the N. parile clades. The latter two share a common ancestor and each includes a widespread Holarctic species (N. laevigatum and N. parile, respectively) and all species endemic to Macaronesia. The data suggest a neo‐endemic origin of Macaronesian taxa, a recent range expansion from Macaronesia of both widespread species, a range expansion limited to the Mediteranean Basin and south‐western Europe for another taxon, and a long dispersal event that resulted in a speciation event in the western parts of North America. Main conclusions The Macaronesian endemic species belong to two sister clades and originated from a most recent common ancestor (MRCA) shared with one widely distributed taxon, either N. parile or N. laevigatum. Estimates of the mean divergence dates suggest that the endemics originated in the archipelagos after the rise of the volcanic islands, along with the ancestor to the now widespread species, which probably expanded their range beyond Macaronesia via long‐distance dispersal. This study provides the first phylogenetic evidence of Macaronesian neo‐endemism in lichenized fungi and provides support for the hypothesis that oceanic islands may serve as a source for the colonization of continents. However, further data are needed to properly assess the alternative hypothesis, namely colonization from western North America.     

FURTHER OBSERVATIONS ON THE GROWTH OF THE COMMON DOG-WHELK, NUCELLA LAPILLUS (L.), IN THE LABORATORY

Juvenile Nueella lapillus, reared in aquaria from adults collectedat Hele in North Devon, did not change shape during the firsttwo years of life. In an earlier paper, Crothers (1977b)foundthat juveniles, reared under identical conditions from adultscollected at Porlock Weir in West Somerset, grew allometricallyduring their second year. The different pattern of growth isyet another feature, to add to differences in size and shape,of the unusual populations of Nucella found in the Severn Estuary. (Received 11 June 1979;     

狭义蛇葡萄属东亚支系的系统发生与生物地理演化

狭义蛇葡萄属(Ampelopsis s. str.)是葡萄科的落叶木质藤本植物,主要分布在北半球温带地区,特别是东亚为其重要的分布和演化中心。该研究选取了狭义蛇葡萄属15个种的37个样本,对其5个叶绿体基因片段(trnL F、rps16、psbA trnH、atpB rbcL和trnK petN)和2个核基因标记(ITS和GAI1)进行了分子测序;利用测序获得的分子数据探讨属内的系统发生关系,叶的性状演化以及生物地理起源演化。结果表明：(1)狭义蛇葡萄属是一个单系类群,属内东亚地区的物种聚成一支。(2)叶绿体基因数据分析结果表明,狭义蛇葡萄属东亚支系分为两支系,这两支系的分布范围大致与东亚植物区系的中国 日本森林和中国 喜马拉雅森林亚区范围相一致。(3)形态演化分析表明,掌状复叶为祖征,叶形态性状存在多次的独立起源和演化,其变化与系统进化没有明显的关系。(4)生物地理分析结果表明,狭义蛇葡萄属起源于北美,渐新世晚期至中新世早期迁移扩散至欧洲,中新世中期随着全球气温回暖迁移至东亚,并进一步在东亚地区快速分化形成多样化中心,这可能与中新世时期的气候温暖、受第四纪冰川影响较少以及东亚地区复杂的地形地貌有关。     

Phylogenomics,biogeography, and evolution of morphology and ecological niche of the eastern Asian–eastern North American Nyssa (Nyssaceae)

Nyssa (Nyssaceae, Cornales) represents a classical example of the well‐known eastern Asian–eastern North American floristic disjunction. The genus consists of three species in eastern Asia, four species in eastern North America, and one species in Central America. Species of the genus are ecologically important trees in eastern North American and eastern Asian forests. The distribution of living species and a rich fossil record of the genus make it an excellent model for understanding the origin and evolution of the eastern Asian–eastern North American floristic disjunction. However, despite the small number of species, relationships within the genus have remained unclear and have not been elucidated using a molecular approach. Here, we integrate data from 48 nuclear genes, fossils, morphology, and ecological niche to resolve species relationships, elucidate its biogeographical history, and investigate the evolution of morphology and ecological niches, aiming at a better understanding of the well‐known EA–ENA floristic disjunction. Results showed that the Central American (CAM) Nyssa talamancana was sister to the remaining species, which were divided among three, rapidly diversified subclades. Estimated divergence times and biogeographical history suggested that Nyssa had an ancestral range in Eurasia and western North America in the late Paleocene. The rapid diversification occurred in the early Eocene, followed by multiple dispersals between and within the Erasian and North American continents. The genus experienced two major episodes of extinction in the early Oligocene and end of Neogene, respectively. The Central American N. talamancana represents a relic lineage of the boreotropical flora in the Paleocene/Eocene boundary that once diversified in western North America. The results supported the importance of both the North Atlantic land bridge and the Bering land bridge (BLB) for the Paleogene dispersals of Nyssa and the Neogene dispersals, respectively, as well as the role of Central America as refugia of the Paleogene flora. The total‐evidence‐based dated phylogeny suggested that the pattern of macroevolution of Nyssa coincided with paleoclimatic changes. We found a number of evolutionary changes in morphology (including wood anatomy and leaf traits) and ecological niches (precipitation and temperature) between the EA–ENA disjunct, supporting the ecological selection driving trait evolutions after geographic isolation. We also demonstrated challenges in phylogenomic studies of lineages with rapid diversification histories. The concatenation of gene data can lead to inference of strongly supported relationships incongruent with the species tree. However, conflicts in gene genealogies did not seem to impose a strong effect on divergence time dating in our case. Furthermore, we demonstrated that rapid diversification events may not be recovered in the divergence time dating analysis using BEAST if critical fossil constraints of the relevant nodes are not available. Our study provides an example of complex bidirectional exchanges of plants between Eurasia and North America in the Paleogene, but “out of Asia” migrations in the Neogene, to explain the present disjunct distribution of Nyssa in EA and ENA.     

Phylogeny and biogeography of the American live oaks (Quercus subsection Virentes): a genomic and population genetics approach

The nature and timing of evolution of niche differentiation among closely related species remains an important question in ecology and evolution. The American live oak clade, Virentes, which spans the unglaciated temperate and tropical regions of North America and Mesoamerica, provides an instructive system in which to examine speciation and niche evolution. We generated a fossil‐calibrated phylogeny of Virentes using RADseq data to estimate divergence times and used nuclear microsatellites, chloroplast sequences and an intron region of nitrate reductase (NIA‐i3) to examine genetic diversity within species, rates of gene flow among species and ancestral population size of disjunct sister species. Transitions in functional and morphological traits associated with ecological and climatic niche axes were examined across the phylogeny. We found the Virentes to be monophyletic with three subclades, including a southwest clade, a southeastern US clade and a Central American/Cuban clade. Despite high leaf morphological variation within species and transpecific chloroplast haplotypes, RADseq and nuclear SSR data showed genetic coherence of species. We estimated a crown date for Virentes of 11 Ma and implicated the formation of the Sea of Cortés in a speciation event ~5 Ma. Tree height at maturity, associated with fire tolerance, differs among the sympatric species, while freezing tolerance appears to have diverged repeatedly across the tropical–temperate divide. Sympatric species thus show evidence of ecological niche differentiation but share climatic niches, while allopatric and parapatric species conserve ecological niches, but diverge in climatic niches. The mode of speciation and/or degree of co‐occurrence may thus influence which niche axis plants diverge along.     

Molecular systematics of the Middle American genus Hypopachus (Anura: Microhylidae)

We present the first phylogenetic study on the widespread Middle American microhylid frog genus Hypopachus. Partial sequences of mitochondrial (12S and 16S ribosomal RNA) and nuclear (rhodopsin) genes (1275 bp total) were analyzed from 43 samples of Hypopachus, three currently recognized species of Gastrophryne, and seven arthroleptid, brevicipitid and microhylid outgroup taxa. Maximum parsimony (PAUP), maximum likelihood (RAxML) and Bayesian inference (MrBayes) optimality criteria were used for phylogenetic analyses, and BEAST was used to estimate divergence dates of major clades. Population-level analyses were conducted with the programs NETWORK and Arlequin. Results confirm the placement of Hypopachus and Gastrophryne as sister taxa, but the latter genus was strongly supported as paraphyletic. The African phrynomerine genus Phrynomantis was recovered as the sister taxon to a monophyletic Chiasmocleis, rendering our well-supported clade of gastrophrynines paraphyletic. Hypopachus barberi was supported as a disjunctly distributed highland species, and we recovered a basal split in lowland populations of Hypopachus variolosus from the Pacific versant of Mexico and elsewhere in the Mesoamerican lowlands. Dating analyses from BEAST estimate speciation within the genus Hypopachus occurred in the late Miocene/early Pliocene for most clades. Previous studies have not found bioacoustic or morphological differences among these lowland clades, and our molecular data support the continued recognition of two species in the genus Hypopachus.     

Biogeography of Asterias: North Atlantic climate change and speciation

Fossil evidence suggests that the seastar genus Asterias arrived in the North Atlantic during the trans-Arctic interchange around 3.5 Ma. Previous genetic and morphological studies of the two species found in the Atlantic today suggested two possible scenarios for the speciation of A. rubens and A. forbesi. Through phylogenetic and population genetic analysis of data from a portion of the cytochrome oxidase I mitochondrial gene and a fragment of the ribosomal internal transcribed spacer region, I show that the formation of the Labrador Current 3.0 Ma was probably responsible for the initial vicariance of North Atlantic Asterias populations. Subsequent adaptive evolution in A. forbesi was then possible in isolation from the European species A. rubens. The contact zone between these two species formed recently, possibly due to a Holocene founding event of A. rubens in New England and the Canadian Maritimes.