Cranial osteology of Exostinus serratus (Squamata: Anguimorpha), fossil sister taxon to the enigmatic clade Xenosaurus

Within the squamate clade Anguimorpha, Xenosaurus is an enigmatic taxon combining several apparently primitive features with a highly specialized set of autapomorphies. This combination makes the fossil record along the Xenosaurus stem particularly important for resolving the relationships of Xenosaurus with other anguimorphs, and between species of Xenosaurus.Exostinus serratus Cope, 1873 from the Oligocene of the western United States is currently hypothesized to be the immediate sister taxon to Xenosaurus. The cranial osteology of this pivotal taxon is described here for the first time from all known material, using high‐resolution X‐ray computerized tomography (CT) scanning to visualize individual elements. Exostinus serratus displays a mosaic of ancestral anguimorph features that are transformed in Xenosaurus, as well as unique synapomorphies shared with Xenosaurus. The region of the external nares is less transformed than in Xenosaurus, as are the general proportions of the bones surrounding the cartilaginous nasal capsule. However, the forms of the teeth and of the osteodermal sculpture, as well as several details of maxillary and mandibular morphology, are distinctly Xenosaurus‐like. Several autapomorphies are also present, including an abbreviated tooth row and an unusually wide palatal shelf of the maxilla. An understanding of the ways in which the non‐ossified tissues of the head influenced the development of the bones is crucial to interpreting their morphology. Furthermore, subtle anatomical features often provide important comparative information, and are emphasized herein. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 921–953.     

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 anatomy of the spade-headed amphisbaenian Diplometopon zarudnyi (Squamata, Amphisbaenia) based on high-resolution X-ray computed tomography

The skull of the trogonophid amphisbaenian Diplometopon zarudnyi is described from high-resolution X-ray computed tomographic (HRXCT) imagery of a whole museum specimen preserved in ETOH. The skull was digitally resliced and disarticulated into individual elements, producing novel visualizations that allow detailed morphological analysis of its three-dimensionally complex structure. The prefrontal and jugal are absent in Diplometopon. The septomaxilla is present but hidden entirely from superficial view. In contrast to previous studies, we recognize a splenial fused to the compound bone of the mandible and a squamosal fused to the otic-occipital complex. Comparison of Diplometopon to the two other amphisbaenians previously described in comparable detail, Rhineura hatcherii and Amphisbaena alba, reveals a mosaic of cranial similarities and differences. Both Diplometopon and Rhineura exhibit a craniofacial angulation and expanded rostral blade related to use of the head as a digging tool, but the detailed architecture of these features is quite different. Additionally, whereas the snout of Rhineura exhibits a high degree of sculpturing and sensory innervation, this is not the case in Diplometopon. Unlike in Rhineura and Amphisbaena, the cranial elements of Diplometopon do not exhibit an extensive degree of overlap or complex interlocking sutures; instead, most of the cranial elements lie in loose apposition to each other. The degree to which this mosaic of features reflects functional demands, shared ancestry, and/or convergence is unclear in the absence of a stable hypothesis of amphisbaenian phylogeny.     

Cranial shape variation and molecular phylogenetic structure of crested newts (Triturus cristatus superspecies: Caudata,Salamandridae) in the Balkans

In the present study, we investigated the degree of congruence between phylogeny, as inferred from mitochondrial (mt)DNA sequences, and cranium shape variation of crested newts (Triturus cristatus superspecies) in the Balkans. These newts belong to four phylogenetic clades defined by mtDNA analysis, and significantly differed in cranial shape. Allometry explained a high percentage of shape variation in crested newts. The clade‐specific allometric slopes significantly diverged for both the ventral cranium and dorsal cranium, indicating that differences in shape between clades could not be a simple consequence of their difference in size. The analysis of hierarchical and spatial variation showed similarity in the patterns of global and spatially localized hierarchical variation of cranial shape. We also found significant congruence between the pattern of cranial shape variation and molecular phylogeny. The differences in morphology of Triturus dobrogicus in comparison to other crested newt clades, including marked differences in cranium shape, is discussed in the context of the evolution and ecology of crested newts. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 348–360.     

Cranial anatomy of the extinct amphisbaenian Rhineura hatcherii (Squamata, Amphisbaenia) based on high-resolution X-ray computed tomography

The fossilized skull of a small extinct amphisbaenian referable to Rhineura hatcherii Baur is described from high-resolution X-ray computed tomographic (HRXCT) imagery of a well-preserved mature specimen from the Brule Formation of Badlands National Park, South Dakota. Marked density contrast between bones and surrounding matrix and at bone-to-bone sutures enabled the digital disarticulation of individual skull elements. These novel visualizations provide insight into the otherwise inaccessible three-dimensionally complex structure of the bones of the skull and their relationships to one another, and to the internal cavities and passageways that they enclose. This study corrects several previous misidentifications of elements in the rhineurid skull and sheds light on skull construction generally in "shovel-headed" amphisbaenians. The orbitosphenoids in R. hatcherii are paired and entirely enclosed within the braincase by the frontals; this is in contrast to the condition in many extant amphisbaenians, in which a large azygous orbitosphenoid occupies a topologically distinct area of the skull, closing the anterolateral braincase wall. Rhineura hatcherii retains a vestigial jugal and a partially fused squamosal, both of which are absent in many extant species. Sculpturing on the snout of R. hatcherii represents perforating canals conveying sensory innervation; thus, the face of R. hatcherii receives cutaneous innervation to an unprecedented degree. The HRXCT data (available at www.digimorph.org) corroborate and extend previous hypotheses that the mechanical organization of the head in Rhineura is organized to a large degree around its burrowing lifestyle.     

Cranial osteology and phylogenetic position of the theropod dinosaur Proceratosaurus bradleyi (Woodward, 1910) from the Middle Jurassic of England

The cranial osteology of the small theropod dinosaur Proceratosaurus from the Bathonian of Minchinhampton, England, is described in detail, based on new preparation and computed tomography (CT) scan images of the type, and only known, specimen. Proceratosaurus is an unusual theropod with markedly enlarged external nares and a cranial crest starting at the premaxillary–nasal junction. The skull is highly pneumatic, with pneumatized nasals, jugals, and maxillae, as well as a highly pneumatic braincase, featuring basisphenoid, anterior tympanic, basipterygoid, and carotid recesses. The dentition is unusual, with small premaxillary teeth and much larger lateral teeth, with a pronounced size difference of the serrations between the mesial and distal carina. The first dentary tooth is somewhat procumbent and flexed anteriorly. Phylogenetic analysis places Proceratosaurus in the Tyrannosauroidea, in a monophyletic clade Proceratosauridae, together with the Oxfordian Chinese taxon Guanlong. The Bathonian age of Proceratosaurus extends the origin of all clades of basal coelurosaurs back into the Middle Jurassic, and provides evidence for an early, Laurasia‐wide, dispersal of the Tyrannosauroidea during the late Middle to Late Jurassic. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009.     

An Unusual,Fangless Short-tailed Snake(Squamata,Serpentes, Homalopsidae) from Sumatra,Indonesia

Two genera of fangless homalopsid snakes Brachyorrhos and Calamophis from eastern Indonesia have been described as basal members of the clade. A third genus belonging to this group from Sumatra, Indonesia is described here based upon morphology. Fangless homalopsid snakes share 19 dorsal scale rows at mid-body, fused dorsal scales above the cloaca, 5 to 7 upper labials, divided anal plate, divided subcaudals, 15–21 teeth on the dentary, frontal bone about 23% of skull length and other morphological characters that suggest they are related. All the three genera are known only from Indonesia. The new genus and species have a distinctive skull morphology with an absent premaxilla, a more depressed skull, and a larger eye than are present in Brachyorrhos.     

Development of the Osteocranium in Natrix natrix (Serpentes,Colubridae) Embryogenesis II: Development of the jaws,palatal complex and associated bones

Snakes differ from the other vertebrates with their hyperkinetic skull. To establish the developmental features of the skull bones, involved in prey capture and ingestion, the Grass snake Natrix natrix (Serpentes, Colubridae) embryos are studied at all the successive stages of embryogenesis. Thirty-five N. natrix embryos are examined. Twenty embryos are studied with histological methods; fifteen embryos are cleared and double-stained with alizarin red and alcian blue. The sequence of appearance and formation of the upper and lower jaw bones, palatal complex and associated bones is described in accordance with the table of developmental stages. New features in the ossification mode of some bones are revealed: each bone, namely, the vomer, septomaxilla and maxilla, is formed from three separate ossification centres. Three ossification centres in the maxilla, two ossification centres in the bodies of the septomaxilla and vomer, as well as the unknown additional ossification centre in the vomer had not been previously described in snake embryos. The new data can be used in further comparative research on the reptile skull development and vertebrate phylogeny.     

Skull osteology and osteological phylogeny of the Western whip snake Hierophis viridiflavus (Squamata,Colubridae)

The skull osteology of Hierophis viridiflavus is here described and figured in detail on the basis of 18 specimens. The sample includes specimens from the ranges of both H. viridiflavus viridiflavus and H. viridiflavus carbonarius as well as specimens not identified at sub-specific level. The main characters that define H. viridiflavus in comparison to the parapatric congeneric species Hierophis gemonensis are wide maxillary diastema, basioccipital crest well distinct in three lobes and basioccipital process well marked. The foramina of the otoccipital and prootic, and the basioccipital process of the basioccipital are among the most ontogenetically variable characters, as indicated by two juvenile specimens included in the sample. A specimen-level phylogenetic analysis including H. gemonensis and other outgroups (overall 6 species, 26 specimens, 64 skull characters) recovered all H. viridiflavus specimens in one clade, indicating the presence of a clear phylogenetic signal in the applied characters. However, the resolution within the H. viridiflavus clade is poor the monophyly of H. viridiflavus carbonarius was retrieved, but not that of Hierophis v. viridiflavus. Probably due to the relatively high variability, the skull morphology does not support the recently proposed specific status of the two subspecies.     

Phylogenetic relationships and molecular evolution in uropeltid snakes (Serpentes: Uropeltidae): allozymes and albumin immunology

Multilocus electrophoretic methods and microcomplement fixation comparisons of serum albumin are used to assess phylogenetic relationships among species of uropeltid snakes, to infer aspects of their population biology and biogeography, and to evaluate their relationships to other primitive snakes (Henophidia). There is very good agreement between phylogenetic inferences derived from the electrophoretic data and those derived from the albumin immunological data. Protein variation detected by electrophoresis is relatively high among 17 operational taxonomic units (OTUs) examined. The mean number of alleles per locus (5.1 across all OTUs), levels of polymorphism (25% of loci), and heterozygosity (4–6%), are typical of, or greater than, values reported for other snakes. Species of uropeltids are genetically highly differentiated, as measured by genetic distances (lowest interspecific Nei's unbiased genetic distances, 0.22-0.27 among several Sri Lankan species; 2.3 between Teretrurus of India and other uropeltines). The phylogenetic tree most consistent with both the immunological and electrophoretic data shows uropeltines from Sri Lanka to be monophyletic, but the Indian species are paraphyletic with respect to those from Sri Lanka. Rhinophis travancoricus of India is inferred to be the sister taxon to the Sri Lankan radiation. As the genera are presently understood, neither Rhinophis nor Uropeltis appears to be monophyletic. A biogeographic scenario derived from the phylogenetic hypothesis suggests an early diversification of uropeltids in India, followed by a single invasion into the lowlands of Sri Lanka. Subsequent evolution on Sri Lanka resulted in occupation of montane biotopes. Cylindrophis is the sister group to uropeltines and is considered a member of the Uropeltidae. The immunological data indicate no phylogenetic association between uropeltids and other ‘anilioid’ taxa, specifically Anilius, Loxocemus or Xenopeltis, although we cannot rule out a very remote relationship. We specifically reject the hypothesis that uropeltines and scolecophidians form a clade relative to henophidians. High levels of genetic variation and a trend toward negative F_IS values for polymorphic loci in three populations suggest generally large effective population sizes and outbreeding in these species. The niche-width variation hypothesis for allozyme loci is not supported by the uropeltid data. In comparison to other vertebrates, the relationship between Nei's genetic distance and albumin immunological distance in uropeltids suggests either conservative albumin evolution or strong differentiation at electrophoretic loci.     

Wood anatomy of Penaeaceae (Myrtales): comparative, phylogenetic, and ecological implications

Wood samples of stems, lignotubers, and roots of the majority of species of Penaeaceae were analyzed with respect to qualitative and quantitative features. Virtually no data have hitherto been presented on xylem features of this family, restricted to Cape Province, South Africa. Presence of vestured pits in vessels, septate crystalliferous parenchyma in wood, intraxylary phloem, predominantly erect ray cells in the typically narrow, multiseriate rays and in the uniseriate rays, and amorphous deposits in ray cells place Penaeaceae securely in Myrtales and help to define that order. By comparison of ecological preferences of the species, as observed during field work, with quantitative analysis of conductive tissue, close correspondence of the wood structure to habit and habitat is demonstrated.     

A combined morphological and molecular phylogeny of the genus Chironius Fitzinger, 1826 (Serpentes: Colubridae)

Chironius is one of the most speciose genera of the South American colubrid snakes. Although the genus represents a well‐known radiation of diurnal racers, its monophyly, affinities with other Neotropical colubrid genera, and intrageneric relationships are open questions. Here, we present a phylogenetic analysis of Chironius based on a data matrix that combines one nuclear (c‐mos) and two mitochondrial (12S and 16S rRNA) genes with 37 morphological characters derived from scutellation, skull, and hemipenial features. Phylogenetic relationships were inferred using maximum parsimony (MP) and maximum likelihood (ML). Our combined morphological and molecular analyses strongly support the monophyly of the genus Chironius and its sister‐group relationship with a clade formed by the genera Dendrophidion and Drymobius. Phylogenetic relationships within the genus Chironius is still controversial, although five clades are retrieved with medium to strong support. © 2014 The Linnean Society of London     

Image assessment of MSCT and CBCT scans for rapid maxillary expansion: a pilot study

Objectives: The accuracy of cone-beam technique, cone-beam computed tomography (CBCT), compared with that of the multislice spiral CT (MSCT), for image-based linear measurements of midpalatal suture was assessed.

Material and Methods: Two measurements were performed by one investigator on the dry skull by using one digital caliper and in the axial cuts by using software. A 2D object-based image registration process was applied to determine the best affine transformation that maps a 2D input image (CBCT) in a reference image (MSCT).

Results: The value of the intraclass correlation coefficient was approximately 0.9%. The results suggest that differences between two scanners did not exist (observation 1, p = 0.964 and observation 2, p = 0.795). With regard to the dry skull and the image, the significance probabilities equaled zero (observation 1, p = 0.002 and observation 2, p = 0.004), therefore, indicating significant differences.

Conclusion: Measurements acquired in the images were similar and these findings contribute to stimulate the use of CBCT for evaluation of the maxillary expansion procedure.     

The embryonic development of the Egyptian cobra Naja h. haje (Squamata: Serpentes: Elapidae)

The Egyptian cobra, Naja h. haje, is the largest of the African cobras and is a member of a successful and medically important species complex found throughout Africa, north and south of the Sahara, as well as across the Arabian Peninsula to Oman. Although its phylogenetic position and venom characteristics have been well studied, its development has not. Here, we present a normal staging table for N. h. haje, based on external features. Comparison with firstly the Asian monocled cobra, Naja kaouthia, and then with the small number of other oviparous snake species, allowed us to examine whether differences between two species in the same genus were of the same type and magnitude as those between unrelated genera. In fact, at least with respect to external features, we found a similar level of disparity. N. h. haje embryos lagged behind those of N. kaouthia in body and head scale development, size in ovo and hatchling length, despite having a slightly shorter incubation period and a somewhat larger adult size. Some of these differences may have been the result of differing incubation temperatures. Nonetheless, there does appear to be a broadly conserved pattern of in ovo development in at least macrostomatan snakes.     

Composition and phylogenetic affinities of vangas (Vangidae, Oscines, Passeriformes) based on morphological characters

The first cladistic analysis of the relationship of Malagasy vangas (Vangidae) based on morphological characters is presented here. The comparative morphological analysis provides new arguments for the exclusion of several taxa traditionally or recently assigned to Vangidae such as Tylas 'Vanga' ( Tylas eduardi ), Crossley's Babbler ( Mystacornis crossleyi ) and newtonias ( Newtonia spp.), but also for the exclusion of the Red-tailed 'Vanga' ( Calicalicus madagascariensis ) from the Vangidae. It leads also to the hypothesis that vangas sensu stricto , the Australasian woodswallows (Artamidae) and the Australo-Papuan butcherbirds (Cracticidae) form a monophyletic group, with the New Guinean peltops ( Peltops spp.) as its probable sister taxon. Such a relationship implies that the last common ancestor of vangas reached Madagascar from the east by means of a dispersal event across the Indian Ocean. This hypothesis is partly in accordance with that of William Pycraft (1907) published a century ago, but in conflict with contemporary hypotheses based on DNA sequence analysis which favour helmet-shrikes, bush shrikes and relatives as the closest relatives of vangas and therefore an African origin of this group.     

The osteology of the golden grey mullet Liza aurata (Teleostei: Mugiliformes: Mugilidae) including interactive three-dimensional reconstructions

Grey mullets are remarkably characterized by their overall uniform external morphology. Identifying species as well as positioning the Mugiliformes in a phylogenetic context is rather difficult. Most recently they were placed in the newly erected Ovalentaria, but more detailed relationships to potential sister taxa were not resolved. Studying the internal morphology, especially the osteology, might provide new insights into the evolution of the Mugiliformes as well as help clarify their phylogenetic position within the Ovalentaria. A detailed osteology of the golden grey mullet Liza aurata is presented. The use of cleared and stained specimens allowed for a complete examination of bony and cartilaginous structures, and a 3D reconstruction from a μCT data set provided additional information on the positional relationships of the bones. Following this, the data obtained were compared with different mugilid species, particularly with the flathead grey mullet Mugil cephalus. Several differences between these species could be identified, such as the position of the basisphenoid, the shape of the hyomandibular and the composition of the branchial arches. These characters might help in understanding the evolutionary changes happening within the mugiliforms and will provide the basis to study this taxon in detail, finally allowing the reconstruction of the body plan of grey mullets.     

Reappraisal of patterns of nonmarine cryptodiran turtle carotid circulation: evidence from osteological correlates and soft tissues

The turtle cranial circulation has been employed as an important source of phylogenetic information, but recent conflicting hypotheses of relationship within Testudinata suggest reevaluation of the utility of characters drawn from this complex. As a component of a comprehensive character analysis, the osteological correlates of the nonmarine cryptodiran turtle carotid circulation are herein subjected to high-resolution X-ray computed tomography, reassessed, and statistically investigated. Three different patterns of osteological correlates, indicating three disparate cranial circulatory patterns, are described, and this finding is corroborated by evidence from circulatory soft tissues. Members of the Trionychia and Kinosternoidea exhibit patterns that differ from the more widespread condition found in testudinoid taxa. This result differs from previous work, which has indicated the presence of only two major cranial circulatory patterns, and suggests that while cranial circulatory features may be phylogenetically informative, the information contained within them indicates patterns of relationship different from those previously hypothesized.