The osteology and relationships of the Millerettidae (Reptilia: Cotylosauria)

The morphology of the Millerettidae is described in detail, based on careful preparation using the acetic acid technique. This shows that the Millerettidae constitute a closely knit natural group and results in a more satisfactory classification. Common to the group is the possession of anthracosaur-like vomerine fangs. A second feature of major importance is that these animals do not have a solid lower temporal bar. It is suggested that the reason for this is to be sought in an understanding of cranial kinesis. The reason for the lack of a temporal bar can be traced back to Anthracosaurian amphibia and is due to expansion and contraction across the cheek region during jaw movements. (These movements were necessarily minimized during the transition from amphibian to reptile.) Following from this it is suggested that the Millerettidae may have given rise directly to the Squamata. The Archo-sauria and Rhynchocephalia probably had an entirely independent origin. Only three genera are considered valid, viz. Miileretta (Broom, 1948) in which the temporal opening first appears, Millerosaurus (Broom, 1948) in which contact between jugal and quadratojugal is lost, though the jugal retains a crenellated posterior edge, and a new genus Milleropsis based on Millerosaurus pricei (Watson, 1957) in which there is still greater separation between jugal and quadratojugal and in which the posterior border of the jugal is quite smooth.     

Proterochampsa barrionuevoi (Archosauriformes: Proterochampsia) from the Late Triassic (Carnian) of Argentina and a phylogenetic analysis of Proterochampsia

Abstract: Restudy of skulls and available postcrania of the proterochampsian archosauriform Proterochampsa barrionuevoi from the Ischigualasto Formation (Upper Triassic, Carnian) in the San Juan Province, Argentina, confirms that the genus is diagnosed by autapomorphies that include dermal sculpturing consisting of prominent ridges and nodular protuberances, a large hook‐like lateral projection on the quadratojugal, an antorbital fossa restricted to a depression along the maxilla, lateral expansion of the premaxilla anterior to the premaxilla–maxilla contact, absence of a supratemporal fossa, exclusion of jugal from suborbital fenestra, basal tubera of parabasisphenoid facing ventrally and reaching laterally beyond the basipterygoid process, and a ventral lamina on the angular. Proterochampsa nodosa is a valid species distinguished from P. barrionuevoi by fewer cranial ridges with larger protuberances, relatively smaller supratemporal fenestrae and width of frontals between orbits less than that of the nasals. A phylogenetic analysis supports the monophyly of Proterochampsia consisting of Proterochampsa, Chanaresuchus bonapartei, Gualosuchus reigi, Tropidosuchus romeri and Cerritosaurus binsfeldi. A temporal separation between the two basal proterochampsians with earliest records in the Late Triassic (Proterochampsa and Cerritosaurus) and Chanaresuchus, Gualosuchus and Tropidosuchus in the Middle Triassic indicates hidden proterochampsian diversity in the Middle Triassic.     

A Late Triassic Invertebrate Ichnofauna from Ghost Ranch,New Mexico

Invertebrate trace fossils are reported from the Late Triassic Chinle Formation at Ghost Ranch, New Mexico, for the first time. They occur in beds higher in the section than the Coelophysis Quarry. Six ichnotaxa are recognized: Steinichnus milfordensis, Planolites montanus, Palaeophycus tubularis, Taenidium serpentium, ?Arenicolites sp. and Skolithos sp. This ichnofauna is consistent with the Scoyenia ichnofacies, considered typical of lake margins in semiarid to arid regions.     

新铺龙(爬行纲:海龙目)的分类(英文)

比较了已经描述的新铺龙(Xinpusaurus)的三个种,确定以往确立的种间鉴别特征是无效的。吻部比例以及齿骨形态都不能用来区分它们,戈氏种(X.kohi)的头后骨骼鉴别特征基础不牢,大多因标本保存欠佳不能明确其有效性。因此,建议废除巴毛林种(X.bamaolinensis)以及戈氏种,新铺龙属仅保留属型种——孙氏新铺龙(X.suni)——为有效种。文中还进一步阐明了本属种脑颅的形态。     

Osteoderm histology of Proterochampsia and Doswelliidae (Reptilia: Archosauriformes) and their evolutionary and paleobiological implications

Postcranial osteoderms are commonly developed in the major lineages of Archosauriformes, including forms such as proterochampsids and doswelliids. Here, we survey the histology of osteoderms of the doswelliids Archeopelta arborensis and Tarjadia ruthae, and the proterochampsids Chanaresuchus bonapartei and Pseudochampsa ischigualastensis to understand better the morphogenesis of these skeletal elements. Whereas, the Doswelliid osteoderms possess a trilaminar organization, in which two cortices (external and basal) can be differentiated from an internal core of cancellous bone, these elements are compact structures in proterochampsids. The osteoderms of P. ischigualastensis are avascular and they consist entirely of parallel‐fibered bone. Conversely, the osteoderms of C. bonapartei are well vascularized structures composed of zones of woven‐fibered bone and annuli of parallel‐fibered bone. The rather simple microstructure observed in P. ischigualastensis osteoderms suggests that these elements grew at a constant, low rate. Compared with proterochampsids, doswelliid osteoderms possess a more complex histology, which appears to be linked to variations in the growth rate during the osteoderm formation and also to the development of the external ornamentation. A comparison of our findings with the results of earlier studies on other archosauriforms (phytosaurs and pseudosuchians) reveals that the general osteoderm histology of doswelliids bears a closer resemblance to that of phytosaurs and pseudosuchians than the proterochampsid osteoderm microstructure. If all archosauriform osteoderms are homologous structures, the closer resemblance of doswellid osteoderm microstructures to that of phytosaurs and pseudosuchians is in agreement with the hypothesis that doswellids are more closely related to archosaurs than proterochampsids. J. Morphol. 276:385–402, 2015. © 2015 Wiley Periodicals, Inc.     

Hind foot drumming: Morphofunctional analysis of the hind limb osteology in three species of African mole-rats (Bathyergidae)

Hind foot drumming is a form of seismic signaling that plays a vital role in the communication of several Bathyergidae species. Hind foot drumming is initiated by the rapid movement of the whole hind limb by flexion and extension of the hip and knee. This study aimed to determine if morphological adaptations of the hind limb osteology were measurable using established morphometric analyses in two drumming (Bathyergus suillus and Georychus capensis) and one non-drumming (Cryptomys hottentotus natalensis) African mole-rat species. Forty-three linear measurements of the hind limb were taken in 48 limbs (n = 16 limbs per species) and 32 indices were calculated. Mixed model analysis of variance was used to compare the three species and sexes within a species. Thirteen indices had significant differences between species. Eleven indices had significant differences between sexes within a species. Significant differences between the drumming (B. suillus and G. capensis) and the non-drumming species were observed in three indices. The femoral greater trochanter was relatively shorter in the drumming species compared to the non-drumming species, which is proposed to allow for increased hip joint mobility, thereby permitting drummers to move their limbs at the rapid speed required to generate seismic signals. Furthermore, the small in-lever (shorter greater trochanter) may increase the velocity of limb motion. The robust tibias in the drumming species, as indicated by the tibial robustness index, are likely to counter the additional biomechanical load caused by the muscles involved in hind foot drumming. The relatively small hind feet seen in the drumming species allows for reduced limb weight needed for the rapid extension and flexion motion required during hind foot drumming. The significant differences reflected in the hind limb osteological indices between B. suillus and G. capensis and the non-drumming species are indicative of adaptations for hind foot drumming.     

Comparative osteology and relationships of the Umbridae (Pisces: Salmoniformes)

The phylogenetic relationships of the five living species of umbrids are examined through a comparative osteological study based on a series of cleared-and-stained specimens of each species. The Umbridae appear to be strictly monophyletic. In addition, for 45 characters, the outgroup Esox shares one character-state with at least one umbrid species, while two or more umbrid species share the other state. Assuming that the states shared with Esox are primitive for the Umbridae, the hypothesis that Dallia is more closely related to Umbra than to Novumbra is supported by compatibility and Wagner-tree analysis of the data. Thirteen derived characters are shared by Dallia and Umbra , 20 more are shared by the three species of Umbra , and four more by U. limi and U. pygmaea. Eight characters are considered likely to have undergone reversal or parallel evolution.     

The Late Triassic Schilderia Adamanica and Woodworthia Arizonica Trees of the Petrified Forest National Park, Arizona, Usa

Two fossil tree species, both with unusual characteristics, occur in the Upper Triassic of the Petrified Forest National Park, Arizona, USA and adjacent areas. The first, Schilderia adamanica , has a highly idiosyncratic secondary xylem structure which contains normal uniseriate and broad complex multiseriate 'herring-bone' rays. The trunk cross-section of the secondary xylem may be either of a normal pycnoxylic type with a central pith and rays radiating from that or it may consist of appressed 'xylem masses' with rays curving towards one another at their extremities. The second, Woodworthia arizonica , has narrow, horizontal vascular traces traversing the entire radial width of the secondary xylem. By analogy with extant tree species, these traces would have terminated on preventitious buds deeply embedded in the bark which, in the case of these fossil trunks, have failed to be silicified. Such buds have the capacity to develop into epicormic shoots when the crown foliage of the tree is damaged. A further specimen of W. arizonica is recorded for the first time from the Permian of southern Brazil. Reconstruction drawings of both trees are produced.     

中国肯氏兽-山西鳄组合带的新发现之三:山西临县的主龙型类

近几十年,华北的二马营组上部地层以产出中国肯氏兽-山西鳄四足动物化石组合而闻名.最近在山西临县白道峪于上覆的铜川组一段发现了中国肯氏兽.本文描述了同一地点同一层位产出的主龙型类化石,包括一具山西鳄的部分骨架以及一些可以归入suchian的主龙类.它们是铜川组一段首次记述的主龙型类.最有鉴定特征的suchian材料包括一个大的髂骨以及一个小的、形状很奇特的、可能是跟骨的骨骼.髂骨可能可以归入一个奇异的波波龙类(poposauroid).因为中国肯氏兽与山西鳄同时出现在白道峪,表明中国肯氏兽-山西鳄组合可以向上延伸到铜川组一段.髂骨与跟骨大小悬殊,可能代表两个从未在中国肯氏兽-山西鳄组合报道过的物种.白道峪发现了特化的波波龙类,支持了波波龙类在中三叠世大量分化的观点.     

Heterochrony and tooth evolution in hyperodapedontine rhynchosaurs (Reptilia, Diapsida)

The tooth arrangement of hyperodapedontine rhynchosaurs shows clear patterns of morphological derivation, which can be summarized as three main apomorphic trends: the increase in the number of tooth rows lateral to the main maxillary groove, the loss of dental structures (medial groove and lingual teeth) medial to the main maxillary groove, and the loss of dental structures (medial crest and lingual teeth) medial to the main dentary crest. The analysis of these trends from a heterochronic viewpoint reveals that acceleration was the most probable process involved in the increase in number of the lateral maxillary tooth rows, while the loss of the medial structures of the maxilla and dentary seem to be related respectively to neoteny and post-displacement. Both peramorphic and paedomorphic processes are, therefore, thought to have directed the main modifications seen in the tooth arrangement of the hyperodapedontine rhynchosaurs. Heterochrony plays an important role in the evolution of the Late Triassic rhynchosaurs, which are differentiated mainly on the basis of their dental morphology.     

The comparative osteology and phylogenetic relationships of African and South American lungfishes (Sarcopterygii: Dipnoi)

Lepidosirenidae is a clade of freshwater lungfishes that include the extant South American Lepidosiren paradoxa Fitzinger, 1837 and African species of the genus Protopterus. These genera have been geographically separated since the break‐up of Gondwana in the Early Cretaceous, but they display similar biology and morphology. Species were distinguished by a combination of features such as head‐to‐body ratios, the number of pairs of ribs, and the presence of external gills, but no discrete skeletal characters were identified, and no comparative studies including all extant species have been published. I used computed tomography (CT), X‐ray photography, and specimens from museum collections to describe the skeletal morphology of all species of lepidosirenid in a comparative context. I digitally disarticulated the bones in each specimen to compile a comparative atlas of the cranial and pectoral elements of all extant lungfishes, which has the potential to increase the correct identifications of specimens in museum collections. The morphology of the frontoparietal, parasphenoid, supraorbital, and suboperculum differ between species. I used those characters, along with molecular sequence data from the ribosomal RNA gene 16S, to run combined morphological and molecular phylogenetic analyses. Lepidosirenidae is monophyletic in all analyses, but the interrelationships of the species of Protopterus vary with the different sources of character data. © 2015 The Linnean Society of London     

Cranial morphology of Platypterygius longmani Wade, 1990 (Reptilia: Ichthyosauria) from the Lower Cretaceous of Australia

Exceptionally well-preserved remains belonging to the Australian Early Cretaceous (Albian) ichthyosaur Platypterygius longmani Wade, 1990 are described in detail. The material is used to reconstruct some of the cranial musculature and provide a brief functional analysis of the skull and mandible. Preparation of specimens using acetic acid and application of high-resolution X-ray computed tomographic analyses has revealed many previously unrecorded anatomical details, including the absence of a coronoid and squamosal, and the presence of well-developed epipterygoid facets on the pterygoid and parietal (possibly indicating retention of an unossified epipterygoid element). Reconstruction of the jaw musculature suggests a well-developed m. adductor mandibulae internus pterygoideus (serving to close the jaws against inertia and drag of the surrounding water), and possibly an m. intramandibularis (acting to accentuate static pressure along the elongate mandible when the jaws were closed). Despite its large size (maximum total body length of around 7 m), the long, narrow snout of P. longmani (together with preserved gut contents) indicates selective feeding on relatively small prey such as fish, small tetrapods and probably cephalopods. Large orbits are indicative of a visual hunter, but extensive external passages for nerves and blood vessels might suggest the presence of a dermal sensory system. The massive stapes appears to have lost all sound conductive function, serving primarily as structural support in the basicranial region.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 583–622.     

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.     

A new choristodere (Reptilia: Diapsida) from the Lower Cretaceous of western Liaoning Province, China, and phylogenetic relationships of Monjurosuchidae

The publication of the scientific name Monjurosuchus splendens in 1940 documented the first tetrapod fossil of the later world-renowned Jehol Biota. For more than half a century since this discovery, however, Monjurosuchus has remained as a monotypic genus of the family Monjurosuchidae, and the relationships of the family with choristoderes have not been correctly recognized until quite recently. In this paper, a new monjurosuchid is named and described based on a nearly complete skull and postcranial skeleton from the Early Cretaceous Chiufotang Formation exposed near Chaoyang, western Liaoning Province, China. This new material documents the first occurrence of monjurosuchid choristoderes outside the type Lingyuan area, and extends the geological range of the family from the Yixian Formation to the younger Chiufotang Formation. Cladistic analyses were conducted with inclusion of monjurosuchids, and the results support the placement of the family Monjurosuchidae as a primitive clade outside the Neochoristodera. A new classification scheme is proposed for choristoderes on the basis of the recovered phylogenetic framework of the group.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 427–444.     

Neurocranial osteology and systematic relationships of Varanus (Megalania) prisca Owen, 1859 (Squamata: Varanidae)

The neurocranial osteology of the giant monitor lizard Varanus (Megalania) prisca Owen, 1859 is described in detail for the first time. Optimization of neurocranial characters onto phylogenetic topologies for varanoids, including Lanthanotus, Heloderma, and Varanus species nests V. prisca within an Indo‐Australian clade of Varanus on the basis of characters of the otic capsule. A sister‐taxon relationship between V. prisca and Varanus komodoensis Ouwens, 1912 is proposed based on apomorphies of the crista prootica, fenestra vestibuli, occipital recess, and supraoccipital. These results support a monophyletic clade of giant monitors among Indo‐Australian species, and unambiguously synonymize Megalania with Varanus at both generic and subgeneric levels. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 445–457.