Stratigraphy,facies and synsedimentary tectonics in the Jurassic Rosso Ammonitico Veronese (Altopiano di Asiago,NE Italy)

Summary The red, pelagic limestones of the Rosso Ammonitico Veronese (Upper Bajocian-Tithonian) of the Altopiano di Asiago area can be subdivided into eight facies. They differ from each other in structure (bedding style, presence and type of nodularity) and texture (nature of components, grain-vs mud-support, compactional features). Several discontinuities could be recognized, based on sedimentological or biostratigraphic evidence. In the context of a drowned platform, where sediments essentially consist of skeletal remains of both planktonic and benthic organism, the different facies are interpreted as reflecting a varying influence of currents on the winnowing of micrite and on triggering early cementation. Early cementation in turn, controlled the patterns of bioturbation and the response of sediments to later compaction and pressure-dissolution. At times, microbial mats, of unidentified nature, were important in trapping and binding sediment, giving rise to early lithified nodules and layers of stromatolitic aspect. The Rosso Ammonitico Veronese can be subdivided into three units: lower Rosso Ammonitico (RAI: Upper Bajocian-Lower Callovian), middle Rosso Ammonitico (RAM: Upper Callovian-Middle Oxfordian), and upper Rosso Ammonitico (RAS: Lower Kimmeridgian-Tithonian). Frequent ammonite moulds allow the precise dating of the base and top of each unit, and the documentation of facies heteropies and hiatusses in the more fossiliferous RAS. The lower unit (RAM) is massive and essentially nodular; the middle unit (RAM) is well bedded, non-nodular, and cherty; the upper unit (RAS) is richer in clay and typically nodular. The RAI and the RAS are present everywhere, though significant facies and thickness changes affect particularly the RAS; the RAM is much more variable, ranging from 0 to 10 metres. These variations, that may be gradual or abrupt, are inter-preted as the result of Middle-Late Callovian block-faulting which generated an irregular sea floor topography where the swells were more exposed to currents that continuously removed sediments, inducing long-lasting periods of nondeposition. Sediments preferentially accumulated in the adjacent lows. A confirmation of this hypothesis is provided by evidence of synsedimentary tectonics, described for the first time in the Rosso Ammonitico Veronese. Neptunian dykes, both vertical and horizontal, are developed at the top of the RAI and are filled with laminated sediments or collapse breccias. Glides of metre-size blocks and slumps are present at the top of the RAI and at the base of the RAM, respectively. Cm-thick layers of mud supported breccias are intercalated in the upper part of the RAI and within the RAM: they are interpreted as seismites. All these features document a tensional regime that generated fractures in more or less lithified sediments and failure along steep fault scarps or gently dipping slopes of tilted fault blocks. Recognition of this Callovian-Oxfordian tectonic activity shows that, after the Bajocian drowning, the Trento Plateau did not simply experience a uniform and general foundering: a small-scale block-faulting was still active and affected the pattern of facies distribution.     

Triassic nannoplankton limestones of deep basin origin in the central mediterranean region

Samples of Triassic pelagic limestones collected in several regions of the Central Mediterranean area were examined with the electron microscope. They show calcareous nannoplankton existing already from Middle Triassic time. The Middle Triassic nannofossils are contained in nodular limestones such as the “Ammonitico Rosso” of Epidaurus (Greece); the Upper Triassic ones are contained in nodular limestones like the Ammonitico Rosso of Hallstatt (Austria), as well as in Halobia cherty limestones of the Pindos basin in Italy, Yugoslavia and Greece. In the Halobia limestones it is possible, in spite of the recrystallization effects, to recognize such an abundance of organic forms — even if in fragments — that we can suppose the original sediment to be a kind of nannoplankton ooze.Because of the very low accumulation rate (few metres/m.y.) of the Middle Triassic Ammonitico Rosso limestones, the coccoliths cannot be considered a significant factor of carbonate pelagic sedimentation at this time; the very high accumulation rate (up to 25 metres/m.y.) of the Upper Triassic cherty limestones in the Pindos basin, on the contrary, allows the nannoplankton to be considered a very important agent of calcareous pelagic sedimentation. The authors hypothesize that the Dogger-Malm, mainly represented by radiolarites, was only a temporary interlude in the Mesozoic pelagic carbonate sedimentation.     

Cryptic diversity in the lizard genus Plica (Squamata): phylogenetic diversity and Amazonian biogeography

Intra‐ and interspecific genetic diversity of the lizard species Plica plica (9 localities) and Plica umbra (19 localities) from the Brazilian Amazon was analysed using two mitochondrial (16S rDNA and CO1) and one nuclear (prolactin receptor – PRLR) genes. We generated a maximum‐likelihood and Bayesian hypotheses of phylogenetic relationships, and using the bPTP and ABGD lineage delimiting methods inferred the most likely number of lineages within each species. Both methods delimited five distinct lineages in Plica plica and six lineages within Plica umbra. The nominal subspecies of Plica umbra was comprised of one lineage, while Plica umbra ochrocollaris was comprised of five lineages. In majority of the cases, lineages were restricted to the interfluves of major Amazonian rivers, and different lineages occupied distinct areas of endemism. Phylogenetic relationships of the lineages are largely concordant with the hypothesized formation of the areas of endemism. The geographic structuring of the clades and the delimitation of these clades as distinct lineages suggest the possibility that these lineages represent species. If the observed diversity of lineages within the genus Plica is characteristic of squamate reptiles of the Amazon region, the diversity of squamates is grossly underestimated.     

New information on Bobosaurus forojuliensis (Reptilia: Sauropterygia): implications for plesiosaurian evolution

Preparation of the holotype specimen of Bobosaurus forojuliensis, a large sauropterygian from the lower Carnian of northeastern Italy, revealed new morphological data relevant in establishing its phylogenetic affinities among pistosauroid taxa and its relationships with plesiosaurians. Inclusion of B. forojuliensis in two phylogenetic analyses focusing, respectively, on sauropterygians and pistosauroids agreed in placing the Italian taxon as closer to plesiosaurians than to other pistosauroids. The phylogenetic interpretation of Bobosaurus was not biased by assumptions on character weighting, is consistent with its relatively younger age compared to most pistosauroids, extends the fossil record of the plesiosaurian basal lineage back to the Carnian and supports the earliest diversification of the clade during the Late Triassic in agreement with the record of several distinct lineages of rhomaleosaurids, plesiosauroids and pliosauroids in the lowermost Jurassic. Bobosaurus shows that the evolution of the plesiosaurian body plan from the ancestral pistosauroid grade was a step-wise process, and that some of the vertebral and appendicular specialisations of Jurassic and Cretaceous plesiosaurians had already developed in the earliest Late Triassic.     

Origins and patterns of endemic diversity in two specialized lizard lineages from the Australian Monsoonal Tropics (Oedura spp.)

Aim

Savanna biomes cover around 20% of land surfaces, yet the origins and processes that have shaped their biodiversity remain understudied. Here, we assess the timing of diversification and how patterns of genetic diversity vary along an aridity gradient in specialized saxicoline gecko clades (Oedura spp.) from the tropical savannas of northern Australia.

Location

Australian Monsoonal Tropics (AMT), Kimberley region (Western Australia).

Methods

We compiled mitochondrial and nuclear data for two Kimberley endemic lizard clades (Oedura filicipoda/murrumanu and O. gracilis), and allied non‐Kimberley taxa (O. marmorata complex). Species delimitation methods were used to identify evolutionary lineages, Maximum‐likelihood and Bayesian phylogenetic methods were employed to assess relationships and diversification timeframes, and rainfall data and range sizes were tested for correlations.

Results

Phylogenetic analyses of cryptic or recently discovered lineage diversity revealed late‐Miocene to early‐Pliocene crown ages. Microendemism and diversity were highest in high‐rainfall regions, while the most widespread lineages occurred in the central and south‐east Kimberley, and showed evidence of introgression with parapatric lineages.

Main conclusions

The initial diversification in both clades was broadly concordant with global climatic events linked to the expansion of savanna biomes in the lateMiocene. Higher endemism in mesic and refugial areas suggests long histories of localized persistence, while wider distributions and evidence of introgression suggest a dynamic history at the arid‐monsoonal interface.     

Molecular phylogeny and intraspecific differentiation of the Eremias velox complex of the Iranian Plateau and Central Asia (Sauria,Lacertidae)

The Central Asian racerunner, Eremias velox, is a widely distributed lizard of the Eurasian lacertid genus Eremias. Nucleotide sequences of mitochondrial genes, cyt b and 12S rDNA from 13 geographically distant localities in Iran and Central Asia, were analysed. Phylogenetic analyses of the sequence data unambiguously recovered five major clades within the E. velox complex with a high level of genetic divergence, indicating long periods of isolation. The basal position of the Iranian clades in the phylogenetic trees suggests that the E. velox clade originated on the Iranian plateau in the Middle Miocene. According to our calibrations, the northern Iranian clade diverged first some 10–11 Ma and that the Central Asian lineages split from the northeastern Iranian lineage approximately 6 Ma, most likely as a result of uplifting of the Kopet‐Dagh Mountains in the northern margin of the Iranian plateau. Topology of the phylogenetic trees, combined with the degree of the genetic distances among the independent lineages recovered in this study, provide a solid foundation for a fundamental revision of the taxonomic status of the major clades within this species complex.     

Global interrelationships of Plesiosauria (Reptilia,Sauropterygia) and the pivotal role of taxon sampling in determining the outcome of phylogenetic analyses

Previous attempts to resolve plesiosaurian phylogeny are reviewed and a new phylogenetic data set of 66 taxa (67% of ingroup taxa examined directly) and 178 characters (eight new) is presented. We recover two key novel results: a monophyletic Plesiosauridae comprising Plesiosaurus dolichodeirus, Hydrorion brachypterygius, Microcleidus homalospondylus, Occitanosaurus tournemirensis and Seeleyosaurus guilelmiimperatoris; and five plesiosaurian taxa recovered outside the split between Plesiosauroidea and Pliosauroidea. These taxa are Attenborosaurus conybeari, ‘Plesiosaurus’macrocephalus and a clade comprising Archaeonectrus rostratus, Macroplata tenuiceps and BMNH 49202. Based on this result, a new name, Neoplesiosauria, is erected for the clade comprising Plesiosauroidea and Pliosauroidea. Taxon subsamples of the new dataset are used to simulate previous investigations of global plesiosaurian relationships. Based on these simulations, most major differences between previous global phylogenetic hypotheses can be attributed to differences in taxon sampling. These include the position of Leptocleididae and Polycotylidae and the monophyly or paraphyly of Rhomaleosauridae. On this basis we favour the results recovered by our, larger analysis. Leptocleididae and Polycotylidae are sister taxa, forming a monophyletic clade within Plesiosauroidea, indicating that the large‐headed, short‐necked ‘pliosauromorph’ body plan evolved twice within Plesiosauria. Rhomaleosauridae forms the monophyletic sister taxon of Pliosauridae within Pliosauroidea. Problems are identified with previous phylogenetic definitions of plesiosaurian clades and new, stem‐based definitions are presented that should maintain their integrity over a range of phylogenetic hypotheses. New, rank‐free clade names Cryptoclidia and Leptocleidia are erected to replace the superfamilies Cryptoclidoidea and Leptocleidoidea. These were problematic as they were nested within the superfamily Plesiosauroidea. The incongruence length difference test indicates no significant difference in levels of homoplasy between cranial and postcranial characters.     

Genetic diversity of the naked mole‐rat (Heterocephalus glaber)

The naked mole‐rat (Heterocephalus glaber) is used as an animal model in various studies, but not much is known on the genetic diversity of this animal. Here, on the basis of dataset collected from the most part of the distribution range of the naked mole‐rat, we reconstruct phylogenetic relationships between its different lineages using mitochondrial and nuclear markers. We also mapped the distribution of the main genetic lineages, dated the divergence using different Bayesian tree‐calibration techniques, and modeled the distribution of ecological niches for the period of last glacial maximum. Our results show the existence of two deeply divergent clades designated as the eastern clade (East Ethiopia) and the southern clade (South Ethiopia and North Kenya). Additional phylogeographic structure was demonstrated for each of these two clades. Divergence between these two main lineages dated back to the Middle Pleistocene (ca. 1.4–0.8 Mya) and may have been related to climate changes in Africa during the Mid‐Pleistocene Revolution. In light of substantial genetic differences between the eastern and southern lineages of the naked mole‐rat, these two clades can be considered as two deeply divergent subspecies or even as distinct species.     

Comparative phylogeography of livebearing fishes in the genera Poeciliopsis and Poecilia (Poeciliidae: Cyprinodontiformes) in central Mexico

Aim To test the hypothesis that the vicariant event responsible for north–south divergences in two clades of the fish genus Poeciliopsis Regan was also responsible for north–south divergences in the fish Poecilia butleri Jordan. Location Central Mexico. Methods Parsimony, distance, maximum likelihood and Bayesian phylogenetic analyses of a mitochondrial gene. Molecular clock test and Bayesian analyses of divergence time. Results We report concordant phylogeographical patterns between two clades in the genus Poeciliopsis (i.e. the other formed by P. latidens Garman and P. fasciata Meek, and the other formed by P. presidionis Jordan and P. turneri Miller) and the clade of Poecilia butleri, with northern and southern individuals within each clade grouping into separate lineages. There is also evidence for slower substitution rates in Poecilia compared with Poeciliopsis. After taking into account these substitution rate discrepancies with Bayesian relaxed molecular clock analyses, north–south divergences in Poecilia butleri were equivalent to those reported for Poeciliopsis latidens‐fasciata and P. presidionis‐turneri. Main conclusions The same Plio‐Pleistocene vicariant event associated with geological activity of the Trans‐Mexican Volcanic Belt (TMVB) appears to have caused divergence in three different freshwater fish lineages. This study is an example of how comparative phylogeography can strengthen inferences about vicariant events in regions of high biological diversity and complex geological history such as the TMVB.     

Phylogeny and speciation of the eastern Asian cyprinid genus Sarcocheilichthys

The genus Sarcocheilichthys is a group of small cyprinid fishes comprising 10 species/sub‐species widely distributed in East Asia, which represents a valuable model for understanding the speciation of freshwater fishes in East Asia. In the present study, the molecular phylogenetic relationship of the genus Sarcocheilichthys was investigated using a 1140 bp section of the mitochondrial cytochrome b gene. Two different tree‐building methods, maximum parsimony (MP) and Bayesian methods, yielded trees with almost the same topology, yielding high bootstrap values or posterior probabilities. The results showed that the genus Sarcocheilichthys consists of two large clades, clades I and II. Clade I contains Sarcocheilichthys lacustris, Sarcocheilichthys sinensis and Sarcocheilichthys parvus, with S. parvus at a basal position. In clade II, Sarcocheilichthys variegatus microoculus is at a basal position; samples of the widespread species, Sarcocheilichthys nigripinnis, form a large subclade containing another valid species Sarcocheilichthys czerskii. Sarcocheilichthys kiangsiensis is retained at an intermediate position. Since S. czerskii is a valid species in the S. nigripinnis clade, remaining samples of S. nigripinnis form a paraphyly. This speciation process is attributed to geographical isolation and special environmental conditions experienced by S. czerskii and stable environments experienced by the other S. nigripinnis populations. This type of speciation process was suggested to be very common. Samples of Sarcocheilichthys sinensis sinensis and Sarcocheilichthys sinensis fukiensis that did not form their own monophyletic groups suggest an early stage of speciation and support their sub‐species status. Molecular clock analysis indicates that the two major lineages of the genus Sarcocheilichthys, clades I and II diverged c. 8·89 million years ago (mya). Sarcocheilichthys v. microoculus from Japan probably diverged 4·78 mya from the Chinese group. The northern–southern clades of S. nigripinnis began to diverge c. 2·12 mya, while one lineage of S. nigripinnis evolved into a new species, S. czerski, c. 0·34 mya.     

Phylogeny and the fossil record of the Helophoridae reveal Jurassic origin of extant hydrophiloid lineages (Coleoptera: Polyphaga)

We performed a phylogenetic analysis focused on the hydrophiloid family Helophoridae (Coleoptera: Polyphaga) in order to test the phylogenetic position of selected Mesozoic fossils assigned to the Hydrophiloidea. The analysis is based on 92 characters of larvae and adults, and includes all extant subgenera of Helophorus and representatives of all other extant hydrophiloid families. Based on this analysis, we provide additional evidence for the monophyly of the helophorid lineage containing the families Helophoridae, Georissidae and Epimetopidae, as well as the first hypothesis on the phylogenetic relationships within Helophorus, revealing three main clades: Lihelophorus, Rhopalohelophorus and the clade of sculptured small subgenera; the subgenera Helophorus s.str., Gephelophorus, Trichohelophorus and Transithelophorus are recognized as paraphyletic or polyphyletic. Inclusion of fossil species in the analysis reveals the Mesozoic genera Hydrophilopsia Ponomarenko, Laetopsia Fiká?ek et al. (adult forms) and Cretotaenia Ponomarenko (larval form) as basal extinct clades of the helophorid lineage, the former genus Mesosperchus Ponomarenko as containing probable stem taxa of Helophorus and the former genus Mesohelophorus Ponomarenko as a member of the Helophorus clade containing extant sculptured subgenera. The extant subgenus Thaumhelophorus syn.nov. is synonymized with Rhopalohelophorus. Our results show that the family Helophoridae may be dated back to the late Jurassic (c. 150 Ma) and the extant clades of Helophorus back to the Early Cretaceous (c. 136 Ma). The basal groups of Helophorus and the supposed basal extinct lineages of the helophorid lineages are shown to be aquatic as adults. A single origin of trichobothria and ventral hydrophobic pubescence in the common ancestor of the Hydrophiloidea is hypothesized, indicating ancestral aquatic habits in the adult stage for the whole Hydrophiloidea.     

Geographically widespread mitochondrial lineages of the African saw-wings inconsistent with species boundaries

The African saw-wings (genus Psalidoprocne) are a group of swallows endemic to sub-Saharan Africa. Although currently described as five species with several subspecies, the taxonomy of the saw-wings is unclear. We sequenced two mitochondrial genes (ND2 and cytb; 1 717 sites) from 20 individuals in order to investigate phylogenetic relationships across the range of Psalidoprocne. We obtained similar results from maximum likelihood (RAxML) and Bayesian (BEAST) phylogenetic analyses, which reveal multiple well-supported mitochondrial clades in the genus. The geographic overlap of distinct lineages suggests reproductive isolation, although these clades do not correspond entirely to current species designations.     

Unmasking alpha diversity,cladogenesis and biogeographical patterning in an ancient panarthropod lineage (Onychophora: Peripatopsidae: Opisthopatus cinctipes) with the description of five novel species

Speciation and biogeographical patterning in the velvet worm Opisthopatus cinctipes was examined under a null hypothesis that numerous discrete lineages are nested within the species. A total of 184 O. cinctipes specimens, together with a single specimen of each of the two congeneric point endemic sister species (O. roseus and O. herbertorum), were collected throughout the forest archipelago in the Eastern Cape, KwaZulu‐Natal and Mpumalanga provinces of South Africa. All specimens were sequenced for two partial mitochondrial DNA loci (COI and 12S rRNA), while a single specimen from each locality was sequenced for the nuclear 18S rRNA locus. Evolutionary relationships were assessed using maximum‐likelihood and Bayesian inferences, while divergence time estimations were conducted using BEAST. A Bayesian species delimitation approach was undertaken to explore the number of possible novel lineages nested within Opisthopatus, while population genetic structure was examined for the COI locus using ARLEQUIN. Phylogenetic results revealed that O. cinctipes is a species complex comprising seven geographically discrete and statistically well‐supported clades. An independent statistical approach to species delimitations circumscribed ca. 67 species. Results from divergence time estimation and rate constancy tests revealed near constant net diversification occurring throughout the Eocene and Oligocene with subdivision of ranges during the Miocene. Gross morphological characters such as leg pair number within O. cinctipes were invariant, while dorsal and ventral integument colour was highly polymorphic. However, scanning electron microscopy revealed considerable differences both between and within clades. The caveats associated with both morphological and algorithmic delineation of species boundaries are discussed. The five novel Opisthopatus species are described.     

A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods

Megalosaurus bucklandii (Dinosauria: Theropoda), the oldest named dinosaur taxon, from the Bathonian (Middle Jurassic) of England, is a valid taxon diagnosed by a unique character combination of the lectotype dentary. Abundant referred material is described and several autapomorphies are identified: ventral surfaces of first and third to fifth sacral centra evenly rounded, ventral surface of second sacral centrum bearing longitudinal, angular ridge; dorsally directed flange around midheight on the scapular blade; an array of posterodorsally inclined grooves on the lateral surface of the median iliac ridge; anteroposteriorly thick ischial apron with an almost flat medial surface; and complementary groove and ridge structures on the articular surfaces between metatarsals II and III. A new phylogenetic analysis focuses on basal tetanurans and includes 41 taxa, six of which have never been included in a cladistic analysis, and 213 characters, 29 of which are new. This is the first phylogenetic analysis to focus on basal tetanuran relationships, and it reveals several new results. Megalosauroidea (= Spinosauroidea) includes two clades, basal to the traditional content of Megalosauridae + Spinosauridae. These comprise Xuanhanosaurus, Marshosaurus, Condorraptor + Piatnitzkysaurus and Chuandongocoelurus + Monolophosaurus. Almost all large‐bodied Middle Jurassic theropods are megalosauroids, but Poekilopleuron is an allosauroid. Megalosauroids show geographical differentiation among clades, indicating the development of endemic theropod faunas across Pangaea during the Middle Jurassic. Notably, megalosaurids are not known from outside of Europe during this epoch. Megalosauroids are less diverse and abundant during the Late Jurassic, when most theropods are neotetanurans and allosauroids dominate the large‐bodied predator niche. This indicates faunal turnover between the Middle and Late Jurassic. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 882–935.     

Origins and diversification of Indo-West Pacific marine fauna: evolutionary history and biogeography of turban shells (Gastropoda, Turbinidae)

The present study aimed to assess the consequences of tectonic events and temperature regime on the diversification of Indo‐West Pacific (IWP) turban shell species. Bayesian and parsimony methods were used to construct a phylogenetic hypothesis using sequence data from three genes for the turban shell genus Turbo and for a larger data set including representatives of all known genera in the subfamily Turbininae. Phylogenetic analyses indicate that all IWP Turbo species form a single clade approximately 68 Myr in age, predating the closure of the Tethys Sea and therefore predating the physical separation of the IWP from other biogeographical regions. All but one of the IWP subgenera tested in Turbo also predate the closure of the Tethys Sea. Fossil evidence for Marmarostoma, the largest subgenus, confirms that at least some Turbo lineages currently restricted to the IWP were previously more widespread. The combination of the phylogeny with the fossil evidence suggests that present day diversity in IWP Turbo is the result of the evolutionary persistence within the IWP of several, morphologically distinct lineages, some of which were more widespread in the Oligocene. Some IWP lineages show significant increases in diversification in the early Miocene, probably as a result of the increased availability of both shallow‐water habitats due to tectonic plate movements and increased carbonate platforms in the central IWP resulting from coincident diversification of zooxanthellate corals. The IWP is therefore behaving as both a cradle of diversity (with new species originating in situ) and a museum of diversity (with lineages that predate its isolation also being maintained). Bayesian and parsimony analyses of the subfamily recovered five clades and mapping the temperature regime (tropical or temperate) of each species onto the molecular tree using parsimony suggested that temperate habitat is an ancestral character in at least four of the clades. This result was also supported by Bayesian reconstruction of ancestral states. Astralium (the fifth clade) may also prove to have a temperate origin, but this could not be determined with certainty given the available data. The origin of diversity in tropical regions is of particular interest because it has been suggested that the tropics are the source of many evolutionary novelties and have provided a species pool, from which temperate regions were populated. The present study suggests that Turbininae may be an exception to this rule. The tree shape also suggests that temperature has had an effect on speciation rates; temperate Turbininae are apparently evolving more slowly or suffering more extinction than their tropical sister clades, which show greater diversity. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 573–592.     

Combining mitochondrial DNA sequences and morphological data to infer species boundaries: phylogeography of lanceheaded pitvipers in the Brazilian Atlantic forest,and the status of Bothrops pradoi (Squamata: Serpentes: Viperidae)

Phylogeographic studies using mitochondrial DNA sequence information are frequently used as the principal source of evidence to infer species boundaries. However, a critical analysis of further evidence is essential to test whether different haplotype clades identify different species. We demonstrate a hypothesis‐testing approach, using a combination of phylogeographic methods, multivariate morphometrics and matrix association tests, to investigate species boundaries in eastern Brazilian pitvipers conventionally assigned to the species Bothrops leucurus and B. pradoi. Two basal haplotype clades with partly overlapping geographical distributions are identified, which could either represent two partly sympatric species, or multiple haplotypes within one organismal lineage. We use partial Mantel matrix association tests to verify whether generalized morphology, or any of four supposedly diagnostic characters for the two species, show any association with mtDNA variation. Negative results lead to the conclusion that the haplotype clades do not denote independently evolving organismal lineages, and do not constitute separate species under any criterion.     

A large‐scale phylogeny of the lycophyte genus Selaginella (Selaginellaceae: Lycopodiopsida) based on plastid and nuclear loci

The lycophyte genus Selaginella alone constitutes the family Selaginellaceae, the largest of the lycophyte families. The genus is estimated to contain 700–800 species distributed on all continents except Antarctica, with highest species diversity in tropical and subtropical regions. The monophyly of Selaginella in this broad sense has rarely been doubted, whereas its intrageneric classification has been notoriously contentious. Previous molecular studies were based on very sparse sampling of Selaginella (up to 62 species) and often used DNA sequence data from one genome. In the present study, DNA sequences of one plastid (rbcL) and one nuclear (ITS) locus from 394 accessions representing approximately 200 species of Selaginella worldwide were used to infer a phylogeny using maximum likelihood, Bayesian inference and maximum parsimony methods. The study identifies strongly supported major clades and well resolves relationships among them. Major results include: (i) six deep‐level clades are discovered representing the deep splits of Selaginella; and (ii) 20 major clades representing 20 major evolutionary lineages are identified, which differ from one another in molecular, macro‐morphological, ecological and spore features, and/or geographical distribution.     

Phylogeny and Evolutionary History of the Ground Squirrels (Rodentia: Marmotinae)

Although ground squirrels (Spermophilus) and prairie dogs (Cynomys) are among the most intensively studied groups of mammals with respect to their ecology and behavior, a well-resolved phylogeny has not been available to provide a framework for comparative and historical analyses. We used complete mitochondrial cytochrome b sequences to construct a phylogeny that includes all 43 currently recognized species in the two genera, as well as representatives of two closely related genera (Marmota and Ammospermophilus). In addition, divergence times for ground squirrel lineages were estimated using Bayesian techniques that do not assume a molecular clock. All methods of phylogenetic analysis recovered the same major clades, and showed the genus Spermophilus to be paraphyletic with respect to both Marmota and Cynomys. Not only is the phylogeny at odds with previous hypotheses of ground squirrel relationships, but it suggests that convergence in morphology has been a common theme in ground squirrel evolution. A well-supported basal clade, including Ammospermophilus and two species in the subgenus Otospermophilus, diverged from all other ground squirrels an estimated 17.5 million years ago. Between 10 and 14 million years ago, a relatively rapid diversification gave rise to lineages leading to marmots and to several distinct groups of ground squirrels. The Eurasian ground squirrels diverged from their North American relatives during this period, far earlier than previously hypothesized. This period of diversification corresponded to warming climate and spread of grasslands in western North America and Eurasia. Close geographic proximity of related forms suggests that most species evolved in or near their current ranges.     

Morphological characters add support for some members of the basal grade of Asteraceae

Recent molecular studies in Asteraceae have divided tribe Mutisieae (sensu Cabrera) into 13 tribes and eight subfamilies. Each of the major clades is well supported but the relationships among them are not always clear. Some of the new taxa are easily characterized by morphological data but others are not, chief among the latter being three subfamilies (Stifftioideae, Wunderlichioideae and Gochnatioideae) and the tribe Hyalideae. To understand evolution in the family it is critical to investigate potential morphological characters that can help to evaluate the basal lineages of the Asteraceae. The data for this study were taken from 52 species in 24 genera representing the basal groups in the family. Many characters were examined but most of the useful ones were from reproductive structures. Several apomorphies supported a few of the clades. For instance, members of subfamily Wunderlichioideae (Hyalideae and Wunderlichieae) share predominantly ten‐ribbed achenes and members of Wunderlichioideae + Stifftioideae share two synapomorphies: 100–150 (200) pappus elements, arranged in (three) four or five series. These apomorphies can be viewed as an indication of a sister‐group relationship between the two subfamilies as the placement of Stifftieae was not well resolved by the molecular data. Members of Wunderlichieae are characterized by having a paleaceous receptacle, style branches that are strongly papillose above and below the bifurcation, and a pappus of scales. Hyalis and Ianthopappus (Hyalideae) share venation type and an apiculate anther appendage but these are also found in Gochnatieae. Other clades have fewer supporting characters. These characters are just a beginning. Cladograms with morphology characters plotted, illustrations and a key to the basal grade of Asteraceae are provided. © 2013 The Linnean Society of London