Molecular diversification of salamanders of the tropical American genus Bolitoglossa (Caudata: Plethodontidae) and its evolutionary and biogeographical implications

The largest genus of salamanders, Bolitoglossa (Plethodontidae), is widespread in tropical America, where it occurs in diverse habitats and elevations, from high elevation grasslands to lowland rain forest . It has the most extensive geographical range of any salamander genus. While most species occur in Middle America, it ranges throughout most of tropical South America as well. Phylogenetic analysis of 1196 bp of two mitochondrial genes (cytochrome b , 16S RNA) from 55 species offers strong support for the monophyly of the genus and sorts the species into a number of clades. Taking into account morphology, distribution, general ecology, and prior systematic and taxonomic studies, we recognize seven subgenera, four of them new: Bolitoglossa Duméril, Bibron et Duméril, 1854, Eladinea Miranda Ribeiro, 1937, Magnadigita Taylor, 1944, Mayamandra , Nanotriton , Oaxakia and Pachymandra . All South American and some lower Middle American species are included in a single well -supported clade, Eladinea . At the species level our analyses uncover the existence of large genetic diversity within morphologically homogeneous taxa. We propose the new combination: B. (Eladinea) paraensis (Unterstein, 1930) stat. nov. , for Brazilian salamanders previously included under B. altamazonica . We evaluate evidence for the multiple colonization of the tropical lowlands by morphologically derived species groups. South America was invaded by members of one clade, Eladinea , which we infer to have dispersed to South America prior to closure of the Panamanian Portal. Despite the relatively long history of salamanders in South America, that continent now accounts for a relatively small proportion of the lineages and species of neotropical salamanders.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 325–346.     

Cranial anatomy of the stem salamander Kokartus honorarius (Amphibia: Caudata) from the Middle Jurassic of Kyrgyzstan

The cranial anatomy of the stem salamander Kokartus honorarius, from the Middle Jurassic of Kyrgyzstan, is redescribed in detail on the basis of all available material using conventional methods and high‐resolution computed microtomography. In contrast to previous interpretations, the skull is characterized by posteriorly positioned external nostrils, the presence of an internarial fenestra between the premaxillae, the maxillae are not shortened, the overlapping of the frontal by the nasal and the parietal by the frontal, a dentate vomer, a dentate pterygoid with a long, strongly arcuate and anteromedially orientated anterior process, the absence of a fontanelle between frontals and pineal pit on the ventral surface of the frontals, and an inferred unique attachment of the adductor mandibulae internus to the lateral border and ventral ridge of the frontals and parietals. Kokartus is closely related to Karaurus and both these taxa constitute the monophyletic taxon Karauridae, which forms the sister group of crown‐group salamanders (Urodela). Karaurids were neotenic forms and neoteny was probably the ancestral life history strategy of salamanders. The morphological analysis of Kokartus supports the current phylogenetic hypothesis that Permian amphibamids (including branchiosaurids) are the closest relatives of salamanders amongst known non‐lissamphibian temnospondyls. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 816–838.     

The first find of chrysomelids (Insecta: Coleopetra: Chrysomeloidea) from Callovian-Oxfordian Daohugou biota of China

Tarsomegamerus mesozoicus nov. gen., nov. sp., an almost complete specimen of leaf beetle from the Callovian-Oxfordian of Daohugou Formation in Inner Mongolia, China is described. Tarsomegamerus mesozoicus is assigned to the Protoscelinae (Chrysomelidae, Coleoptera). This new leaf beetle represents the first record of chrysomeloids from Mesozoic of China and it extends the geographical distribution of Protoscelinae from central Asia into eastern Asia. This find provides new fossil material for studying the origin and evolution of Chrysomeloidea and by making a stratigraphic correlation of the correlative non-marine sedimentary rocks. Although the age of the Daohugou Formation is debatable it is most likely Callovian-Oxfordian (latest Middle Jurassic-earliest Late Jurassic) rather than early Middle Jurassic (late Aalenian-early Bajocian) or Early Cretaceous.     

First record of fossil Mesozoic Ctenopoda (Crustacea, Cladocera)

Pre-Pleistocene representatives of the crustacean order Ctenopoda Sars, 1865 are unknown. Here we describe Mesozoic fossil specimens of ctenopods from two localities in Mongolia: Khotont (Jurassic/Cretaceous boundary, about 145 Mya) and Khutel Khara (Lower Cretaceous, about 129 Mya). All specimens apparently belonged to the tribe Latonini Korovchinsky, 1986 of the subfamily Sidinae Baird, 1850. At the Khotont site, ctenopods were the most numerous microscopic animal fossils. We assigned these cladocerans to a new genus and species Archelatona zherikhini gen. nov., sp. nov. Our findings support a pre-Cretaceous origin for cladocerans.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 146 , 269–274.     

Taxonomy, evolutionary history and biogeography of the broad-toothed field mouse (Apodemus mystacinus) in the eastern Mediterranean area based on mitochondrial and nuclear genes

The broad-toothed field mouse ( Apodemus mystacinus ) is distributed throughout the Balkan Peninsula, Asia Minor and the Middle East. It is generally split into two different specific entities: Apodemus epimelas occurs on the Balkan Peninsula and A. mystacinus inhabits Asia Minor and the Middle East. This analysis, based on two mitochondrial regions (cytochrome b and the D-loop) and the interstitial retinol binding protein (IRBP) nuclear gene, confirms an important level of genetic divergence between the animals from these regions and their separation from each other at least 4.2–5.1 Mya, which is in favour of a distinct specific status. Finally, the broad-toothed field mice from south-western Turkey appear to be closely related to the animals from Crete but highly distinct from the populations of the other Oriental regions. This supports a distinct subspecific level ( A. m. rhodius ) for the insular animals and also for those from south-western Turkey. From a biogeographical point of view, it can be assumed that either late Pliocene or early Pleistocene cooling led to the isolation of two main groups of A. mystacinus , one in the Balkan region and the other one in Turkey and the Near East (Syria and Israel). In this region, it is suggested that a more recent event appeared during the Quaternary period, isolating broad-toothed field mice in Crete and leading to the appearance of two well-differentiated genetic groups: one in Crete and south-western Turkey, and the other widespread in northern and eastern Turkey as well as in the Near East.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 53–63.     

Historical biogeography of Hexabathynella , a cosmopolitan genus of groundwater Syncarida (Crustacea, Bathynellacea, Parabathynellidae)

Hexabathynella is the only cosmopolitan genus of the order Bathynellacea (Crustacea). The known species number 18, found in Europe (9), Africa (1), South America (2), North America (3) and Australia and New Zealand (3). Phylogenetic analysis suggests that the least derived species are those from South America and the most derived those from the Iberian Peninsula, North America and Australia. The five species with the most plesiomorphic characters occur in salt or brackish water, which supports a marine origin for the genus. Phylogenetic and biogeographical analyses suggest that the distribution of the genus can be explained by dispersion and a double vicariant biogeographical model based on plate tectonics and the evolution of the Tethys during the Mesozoic and Cenozoic.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 457–466.     

The earliest known Salamanders (Amphibia,Caudata):A record from the Middle Jurassic of England

Marmorerpeton gen. nov. represented by M. kermackisp. nov. and M. freemani sp. nov., is the earliest known genus of fossil salamander from the Upper Bathonian, Middle Jurassic of Kirtlington, Oxfordshire, England. Marmorerpeton is more primitive than any other known salamander in the absence of intravertebral spinal nerve foramina in the atlantal centrum, but in other features it resembles members of the family Scapherpetontidae, neotenous salamanders otherwise known from the Upper Cretaceous and Palaeocene. The Kirtlington herpetofauna is a unique freshwater assemblage of Middle Jurassic small amphibians and reptiles, several of which represent the earliest known occurrences of their respective groups.     

The tenth skeletal specimen of Archaeopteryx

We describe the tenth skeletal specimen of the Upper Jurassic Archaeopterygidae. The almost complete and well-preserved skeleton is assigned to Archaeopteryx siemensii Dames, 1897 and provides significant new information on the osteology of the Archaeopterygidae. As is evident from the new specimen, the palatine of Archaeopteryx was tetraradiate as in non-avian theropods, and not triradiate as in other avians. Also with respect to the position of the ectopterygoid, the data obtained from the new specimen lead to a revision of a previous reconstruction of the palate of Archaeopteryx . The morphology of the coracoid and that of the proximal tarsals is, for the first time, clearly visible in the new specimen. The new specimen demonstrates the presence of a hyperextendible second toe in Archaeopteryx . This feature is otherwise known only from the basal avian Rahonavis and deinonychosaurs (Dromaeosauridae and Troodontidae), and its presence in Archaeopteryx provides additional evidence for a close relationship between deinonychosaurs and avians. The new specimen also shows that the first toe of Archaeopteryx was not fully reversed but spread medially, supporting previous assumptions that Archaeopteryx was only facultatively arboreal. Finally, we comment on the taxonomic composition of the Archaeopterygidae and conclude that Archaeopteryx bavarica Wellnhofer, 1993 is likely to be a junior synonym of A. siemensii , and Wellnhoferia grandis Elżanowski, 2001 a junior synonym of A. lithographica von Meyer, 1861.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 97–116.     

Seasonality, microhabitat and cryptic variation in tropical salamander reproductive cycles

In many salamanders, most aspects of reproduction coincide with seasonal changes in abiotic factors such as temperature and humidity. However, while these reproductive patterns have been well documented for temperate salamanders, detailed data for tropical species are relatively sparse. I used histological techniques to examine temporal variation in the spermatogenetic cycles of four species of Guatemalan bolitoglossine salamanders ( Bolitoglossa occidentalis , B. rostrata , Dendrotriton bromeliacia and Pseudoeurycea goebeli ) from different microhabitats along an elevational gradient. All four species have mature sperm present in the testis throughout the year, irrespective of patterns of courtship and egg-laying. Additionally, the two species from more seasonal habitats ( B. occidentalis and B. rostrata ) exhibit significant but cryptic levels of variation in the amount of spermatozoa present in the testis not detectable by external appearance or the presence of secondary sexual characteristics. For the two species from less seasonal cloud forests ( D. bromeliacia and P. goebeli ), there were no detectable patterns to variation within the testis. Regional variation in climate undoubtedly influences reproductive cycles, however, microhabitat and the immediate environment are also important determinants of reproductive strategy.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 489–496.     

Sunosuchus (Archosauria, Crocodyliformes) from the Toutunhe Formation (Middle Jurassic) of the Southern Junggar Basin (Xinjiang, NW-China)

A new specimen of the widespread Middle Jurassic to Lower Cretaceous Asian gonipholidid crocodilian genus Sunosuchus is described on the basis of a partial skeleton from the Upper Toutunhe Formation (Middle Jurassic, ?Bathonian-Callovian) of Liuhonggou, SW of Urumqi, Xinjiang Uygur Autonomous Region, People’s Republic of China. The specimen is represented by a partial mandible, teeth, vertebrae, limb and girdle bones and osteoderms. It can be distinguished from other nominal species of the genus by a unique combination of characters: slightly heterodontous dentition, strongly sculptured posteroventral part of the mandible, short fenestra mandibularis, convex dorsal surface of the retroarticular process, keeled cervical vertebral centra and ventral osteoderms with a distinctive sculpture of wide pits and narrow ridges. The heterodontous dentition is a potential autapomorphy of this form. The new specimen is closest in morphology to material described recently from the Callovian of Kirghisia as Sunosuchus sp. It represents the second Middle Jurassic record of the genus, the first crocodile from the Toutunhe Formation, the first substantial crocodile find from the Mesozoic of the Southern Junggar Basin, and the first Middle Jurassic record of Sunosuchus from China. This extends both the paleobiogeographical distribution of the genus in Asia and its stratigraphic distribution in China considerably.     

Metamorphosis and evolution of feeding behaviour in salamanders of the family Plethodontidae

Plethodontid salamanders capture prey with enhanced tongue protraction relative to other salamander taxa, yet metamorphosing plethodontids are hypothesized to be constrained relative to direct-developing plethodontids in their degree of tongue evolution (protraction length and velocity) by the presence of a larval stage in development. In this biphasic life history the hyobranchial apparatus serves the conflicting functions of larval suction feeding and adult tongue protraction. The deletion of the larval stage removes one of the conflicting functions and has thus permitted direct-developing plethodontids to circumvent this constraint and evolve extremely long tongues, which in some species can be projected to 80% of body length. To evaluate this constraint hypothesis and explore taxonomic diversity of feeding behaviours, we studied feeding in larvae, adults and metamorphosing individuals of seven species of metamorphosing plethodontids from the basal taxa Desmognathinae and Hemidactyliini using direct observations, high-speed videography and kinematic analysis. We found that larval plethodontids suction feed, but feeding is suspended entirely during metamorphosis, and aquatic adults do not suction feed. Adults have exapted the terrestrial modes of tongue and jaw prehension for aquatic prey capture. These findings substantiate the premise that suction feeding and tongue protraction are conflicting functions, and thus our results support the constraint hypothesis. Plethodontid adults have evolved their extreme tongue protraction ability at the expense of adult suction feeding. The rapid metamorphosis that characterizes plethodontids may be an adaptation that minimizes the non-feeding period imposed by the evolution of derived tongue protraction in adults. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 134 , 375–400.     

Mitochondrial DNA variation in the western house mouse (Mus musculus domesticus) close to its site of origin: studies in Turkey

In contrast to a substantial inventory of mitochondrial d -loop sequences from the West European part of the range of the western house mouse Mus musculus domesticus , there are few such data from close to the Middle Eastern origin of the subspecies. This paper attempts to rectify this situation, with details of d -loop sequences of 92 mice from 45 localities in Turkey, adding to the six mice from three localities previously studied from that country. The molecular data were found to be consistent with Turkey being close to the site of origin of M. m. domesticus . The 54 Turkish haplotypes were distributed throughout a phylogenetic tree incorporating representative sequences described previously, and there was high nucleotide diversity among the Turkish mice. There was weak genetic structure in Turkey, although haplotypes of the 'Main Turkish Clade', while common over much of Turkey, were absent from Southern Anatolia. Haplotypes of the Main Turkish Clade were also found in western Europe and it is believed that they derived from Turkey, which thus represents one of the source areas for the westwards expansion of M. m. domesticus .  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 473–485.     

Comparative morphology of seeds of four closely related species of Aristolochia subgenus Siphisia (Aristolochiaceae, Piperales)

The primary purpose of this study was to determine whether morphological divergence has occurred in seeds of species in a subclade of Aristolochia subgenus Siphisia . Thus, we compared seed mass, surface area, and wings (presence or absence), embryo shape and state, and embryo length to seed length (E:S) ratio in A. californica Torr., endemic to California, A. macrophylla Lamk. and A. tomentosa Sims of eastern USA, and A. manshuriensis Komarov of eastern Asia. We also compared capsule length and number of seeds per capsule in the three US species. All four species have linear, underdeveloped embryos, but for the other morphological features statistically significant differences occurred among the taxa. However, none of the seed morphological features differed between the eastern North American species A. macrophylla and its eastern Asian sister species A. manshuriensis.   © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 433–436.     

Postcranial anatomy of Haldanodon exspectatus (Mammalia, Docodonta) from the Late Jurassic (Kimmeridgian) of Portugal and its bearing for mammalian evolution

The postcranium of the Late Jurassic docodont Haldanodon exspectatus Kühne & Krusat, 1972 is represented by a partial skeleton and isolated bones of other individuals from the Late Jurassic (Kimmeridgian) of the Guimarota coal mine in Portugal. Haldanodon exhibits adaptations for a fossorial lifestyle such as stout and short limb bones and humeri with greatly expanded distal joints and strong deltopectoral crests. Short first and second phalanges and moderately curved and laterally compressed terminal phalanges with lateral grooves suggest that Haldanodon was a scratch-digger. The dorso-ventrally elongated, triangular scapula has a convex transverse profile with strongly laterally reflected anterior and posterior scapula margins, enclosing a deep trough-like 'infraspinous fossa'. A supraspinous fossa is not developed. The saddle-shaped glenoid facet is mainly formed by the coracoid and orientated antero-ventrally indicating a sprawling gait. No epiphyses were detected and the wide size range of humerus and femur possibly indicate a lifelong growth. Haldanodon is more derived than Morganucodon by complete reduction of the procoracoid, absence of the procoracoid foramen, and a peg-like coracoid. It shares with monotremes a postscapular fossa that is absent in Morganucodon . A PAUP analysis based on 280 cranio-dental and postcranial characters corroborated the position of Haldanodon above morganucodontids and below Hadrocodium .  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 219–248.     

On the origin and development of Artemisia (Asteraceae) in the geological past

Fossil records combined with phytogeographical data reveal that Artemisia L. possibly originated from the existing arid or subarid area of temperate Asia in the mid-Tertiary. The earliest recorded Asteraceae pollen grains are mostly ornamented with medium to long spines, whereas Artemisia L. with short spinules is recorded slightly later. It may be deduced that the latter evolved from the former. Pollen morphological data confirm the coexistence of the two pollen patterns in the subtribe Artemisiinae containing Artemisia L. The development and evolution of the genus suggest three major stages in its history. Artemisia L. did not become abundant until the Middle to Late Miocene in spite of its earlier presence in the Late Palaeogene. Its representation remained low even though there are many records of the genus in the Late Miocene around the world. The Pliocene was an important time for the development of angiosperm herbs including Artemisia L., and the Quaternary has been a major period for the diversification and further worldwide expansion of the genus. The origin and subsequent development of Artemisia L. are considered to be closely related with increased environmental variation, including global climate changes and some important tectonic movements.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 331–336.