A comparative analysis of the post‐cranial skeleton of fossorial and non‐fossorial gymnophthalmid lizards

Squamates are found in a wide range of habitats and show a corresponding diversity of morphologies that can often be correlated with locomotor mode. The evolution of a snake‐like body form, frequently associated with fossoriality, from a typical lacertiform morphology involves changes in the morphology of vertebrae, girdles, and limbs; the changes are mainly manifested by the reduction or loss of limbs and body elongation. In this study, we describe the axial and appendicular skeletons of six closely related gymnophthalmid species. Three of them show a lizard‐like morphology, with a four‐digit forelimb and a five‐digit hindlimb, and the other three show a snake‐like morphology associated with a burrowing habit, with reduced limbs and a longer body in comparison to the former three species. We show that vertebral morphology is similar among the six species, with the differences being accounted for by an increase in the number of vertebrae and by the structural reduction of girdles and limbs in the snake‐like species. Skeletal morphology provides valuable information on locomotion type, physiology, diet, and other biological features. The burrowing morphology usually involves accentuated reduction of girdle and limb elements, reflecting an undulating type of locomotion in which the limbs play little or no role in propelling the body; in contrast, well‐developed limbs and girdles indicate a greater reliance on the limbs for body propulsion. Limb reduction is frequent among vertebrates, but many different phenotypes are found in species exhibiting some kind of reduction, indicating that different mechanisms and evolutionary pressures may be involved in generating the diverse morphologies. J. Morphol. 274:845–858, 2013. © 2013 Wiley Periodicals, Inc.     

A molecular perspective on the evolution of microteiid lizards (Squamata, Gymnophthalmidae), and a new classification for the family

A molecular phylogeny was reconstructed for 26 recognized genera of the Gymnophthalmidae using a total of 2379 bp of mitochondrial (12S, 16S and ND4) and nuclear (18S and c-mos) DNA sequences. We performed maximum parsimony (MP) and maximum likelihood (ML) analyses, and data partitions were analysed separately and in combination under MP. ML analyses were carried out only on the combined sequences for computational simplicity. Robustness for the recovered nodes was assessed with bootstrap and partitioned Bremer support (PBS) analyses. The total molecular evidence provided a better-resolved hypothesis than did separate analysis of individual partitions, and the PBS analysis indicates congruence among independent partitions for support of some internal nodes. Based on this hypothesis, a new classification for the family is proposed. Alopoglossus , the sister group of all the other Gymnophthalmidae was allocated to a new subfamily Alopoglossinae, and Rhachisaurus (a new genus for Anotosaura brachylepis) to the new Rhachisaurinae. Two tribes are recognized within the subfamily Gymnophthalminae: Heterodactylini and Gymnophthalmini, and two others within Cercosaurinae (Ecpleopini and Cercosaurini). Some ecological and evolutionary implications of the phylogenetic hypothesis are considered, including the independent occurrence of limb reduction, body elongation, and other characters associated with fossoriality.     

Comparative study of sperm ultrastructure of five species of teiid lizards (Teiidae, Squamata), and Cercosaura ocellata (Gymnophthalmidae, Squamata)

Sperm ultrastructure of five teiid lizards (Callopistes flavipunctatus, Crocodilurus amazonicus, Dicrodon guttulatum, Dracaena guianensis, and Teius oculatus), and the gymnophthalmid Cercosaura ocellata is described for the first time. Comparisons of sperm ultrastructure among these species and with those of previously examined teiids and gymnophthalmids revealed that the two groups of Teiioidea (Gymnophthalmidae and Teiidae), and the two subfamilies of Teiidae (Teiinae and Tupinambinae) could be distinguished on the basis of sperm ultrastructure data. Significant differences in sperm dimensions between Cnemidophorus and Aspidoscelis support the recent splitting of these two lineages into different genera. Our results revealed high levels of inter-generic variability in sperm ultrastructure within Teiidae, which produces a data set useful in analyzing relationships between genera and families. In phylogenetic analyses, however, sampling multiple species within teiid genera is essential and recording sperm measurements may profitably complement qualitative ultrastructural characters, maximizing the information content of these structures.     

Appendicular skeleton in Bachia bicolor (Squamata: Gymnophthalmidae): osteology,limb reduction and postnatal skeletal ontogeny

The osteology of the appendicular skeleton and its postnatal development are described in Bachia bicolor, a serpentiform lizard with reduced limbs. The pectoral girdle is well developed and the forelimb consists of a humerus, ulna, radius, five carpal elements (ulnare, radiale, distal carpals 4–3, centrale), four metacarpals (II, III, IV, V) and phalanges (phalangeal formula X‐2‐2‐2‐2). In the hindlimb, the femur is small and slender, and articulates distally with a series of ossified amorphous and extremely reduced elements that correspond to a fibula, tibia and proximal and distal tarsals 4 and 3. The pelvic girdle consists of ischium, pubis and ilium, but its two halves are widely separated; the ilium is the least reduced element. We describe the ossification and development during postnatal skeletal ontogeny, especially of epiphyseal secondary centres, ossifications of carpal elements, apophyseal ossifications and sesamoids. Compared to other squamates, B. bicolor shows an overall reduction in limb size, an absence of skeletal elements, a fusion of carpal elements, an early differentiation of apophyseal centres, and a low number of sesamoids and apophyseal centres. These observations suggest that the reductions are produced by heterochronic changes during postnatal development and probably during embryonic development; therefore the appendicular skeleton exhibits a pattern of paedomorphic features.     

The cranial anatomy,ontogeny, and relationships of Karpinskiosaurus secundus (Amalitzky) (Seymouriamorpha,Karpinskiosauridae) from the Upper Permian of European Russia

The Upper Permian seymouriamorph tetrapod Karpinskiosaurus from European Russia includes two species: Karpinskiosaurus secundus and Karpinskiosaurus ultimus. Karpinskiosaurus secundus is represented by two specimens with skull lengths of about 75 mm. All specimens of K. ultimus are smaller than those of K. secundus. Revision of the cranial anatomy of all previously known and several new specimens of Karpinskiosaurus shows that the specimens of K. secundus and most of the specimens of K. ultimus represent the ontogenetic series of one species: K. secundus. The holotype specimen of K. ultimus requires revision, with the aim to find out whether it represents a second species of Karpinskiosaurus or not. The available material permits new reconstructions of the largest, holotype skull, and one smaller skull with a length of about 36 mm. Karpinskiosaurus secundus is included in a cladistic analysis for the first time here. The analysis shows it to form a sister taxon to Discosauriscidae. The clade comprising Karpinskiosaurus secundus plus Discosauriscidae forms a sister group to Seymouriidae. Karpinskiosaurus secundus has a large postorbital and a short preorbital region, and the orbits are placed in the posterior portion of the anterior half of the skull length. Among all seymouriamorphs, such cranial proportions are exhibited only by the largest known specimens of Discosauriscus austriacus. None of the specimens of K. secundus described here exhibits the presence of sensory grooves; thus, all specimens composing the ontogenetic sequence of K. secundus are considered to be terrestrial. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010.     

Roles for modularity and constraint in the evolution of cranial diversity among Anolis lizards

Complex organismal structures are organized into modules, suites of traits that develop, function, and vary in a coordinated fashion. By limiting or directing covariation among component traits, modules are expected to represent evolutionary building blocks and to play an important role in morphological diversification. But how stable are patterns of modularity over macroevolutionary timescales? Comparative analyses are needed to address the macroevolutionary effect of modularity, but to date few have been conducted. We describe patterns of skull diversity and modularity in Caribbean Anolis lizards. We first diagnose the primary axes of variation in skull shape and then examine whether diversification of skull shape is concentrated to changes within modules or whether changes arose across the structure as a whole. We find no support for the hypothesis that cranial modules are conserved as species diversify in overall skull shape. Instead we find that anole skull shape and modularity patterns independently converge. In anoles, skull modularity is evolutionarily labile and may reflect the functional demands of unique skull shapes. Our results suggest that constraints have played little role in limiting or directing the diversification of head shape in Anolis lizards.     

Comparative skull osteology of Karsenia koreana (Amphibia,Caudata, Plethodontidae)

The recent discovery of a plethodontid salamander, Karsenia koreana, in Korea challenged our understanding of the biogeographic history of the family Plethodontidae, by far the largest family of salamanders, which otherwise is distributed in the New World with a few European species. Molecular studies suggest that Karsenia forms a clade with Hydromantes (sensu lato), which includes among its species the only other Old World plethodontids. We studied the skull of K. koreana and compared it with that of other plethodontid genera, especially members of the subfamily Plethodontinae, which it resembles most closely in general anatomy. The anatomy of its skull corresponds to the most generalized and apparently ancestral condition for plethodontids. No clearly autapomorphic states were detected, and no synapomorphies can be found that would link it to other genera. The Karsenia skull is cylindrical and well ossified, giving an impression of strength. In contrast, the skull of Hydromantes is highly derived; the skull is flattened and the bones are weakly ossified and articulated. Hydromantes and Karsenia share no unique anatomical features; differences between them are especially evident in the hyobranchial skeleton, which is generalized in Karsenia but highly modified in Hydromantes, which is well known for its highly projectile tongue. Plethodon and Plethodon‐like species, including Karsenia and to a lesser degree Ensatina, represent the more generalized and apparently ancestral plethodontid morphology. Specialized morphologies have evolved along only a few morphological axes within the Plethodontidae, resulting in a pattern of rampant homoplasy. Our analysis of the anatomy of the new Asiatic lineage illuminates some potential mechanisms underlying adaptive morphological evolution within the Plethodontidae. J. Morphol. 2010. © 2009 Wiley‐Liss, Inc.     

Comparative morphology of Western Australian varanid lizards (Squamata: Varanidae)

Varanid lizards, which vary considerably in body mass both interspecifically and intraspecifically, are generally considered to be morphologically similar. However, significant and non-isometric variation in the relative appendage dimensions for 17 species of Western Australian goannas suggest that these lizards are not morphologically conservative. The first and second canonical variates clearly distinguish the two subgeneral Odatria and Varanus, and species are generally sexually dimorphic. The morphological variation observed among these 17 species of goanna is associated with foraging mode and ecology. However, no single or small group of morphological dimensions discriminates phylogenetic groups, sexes, or ecological groups, and body size is an important component in these analyses. J. Morphol. 233:127–152, 1997. © 1997 Wiley-Liss, Inc.     

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.     

Chromosomal evolution in the Brazilian lizards of genus Leposoma (Squamata, Gymnophthalmidae) from Amazon and Atlantic rain forests: banding patterns and FISH of telomeric sequences

An extensive karyotype differentiation was found among three species of gymnophthalmid lizard genus Leposoma which occur in the tropical forest areas of Brazil. We examined the chromosomes of the Amazonic species L. guianense (LOU) and L. oswaldoi (LOS) and the Atlantic forest species L. scincoides (LSC) after conventional and differential staining, and FISH of telomeric sequences. Both Amazonic species shared very similar 2n = 44 karyotypes, including 20 biarmed macrochromosomes and 24 microchromosomes (20 M + 24 m). However, the location of Ag-NORs and the amount of constitutive heterochromatin differed in these karyotypes. The Atlantic forest species L. scincoides has a very distinct karyotype with 52 acrocentric and subtelocentric chromosomes of decreasing size. Comparative R-banding analysis revealed complete homeology of the macrochromosomes of LGU and LOS and correspondence of banding patterns between LSC acrocentrics and subtelocentrics and some arms of biarmed LGU and LOS chromosomes. Pair 1 had similar banding patterns in the three species, implying the occurrence of a pericentric inversion. Interstitial telomeric bands (ITBs) detected by FISH at the pericentromeric region of some biarmed LGU and LOS chromosomes could be remnants of chromosomal rearrangements occurred during the differentiation of the karyotypes. Robertsonian rearrangements as well as pericentric inversions events probable were involved in the karyotype evolution of these Amazon and Atlantic forests species of Leposoma.     

New species of Schulzia (Nematoda: Molineidae) in Ptychoglossus festae (Squamata: Gymnophthalmidae) from Panama

Schulzia ptychoglossi n. sp. (Strongylida: Molineidae) from the intestines of Ptychoglossus festae (Squamata: Gymnophthalmidae) is described and illustrated. Schulzia ptychoglossi n. sp. represents the fourth species assigned to the genus and is most similar to the Venezuelan species S. usu by possessing a cervical inflation that begins a short distance from the anterior end of the body. Schulzia ptychoglossi differs from S. usu in that ray 8 separates midway between the root and tip of the dorsal ray in S. ptychoglossi, but separates close to the root of the dorsal ray in S. usu.     

Skull osteology of Vipera walser (Squamata,Viperidae): Description,variability, ontogeny,and diagnostic characters in comparison to other Italian vipers

Vipera walser is the most recently recognized European viper. This rare species is endemic to a small area in the Piedmont Alps of Italy, but its closest relatives are found among the Caucasian viper species. In order to provide a starting point for a phylogenetic and biogeographic investigation based on osteology, and including fossils remains, we analyzed four specimens of V. walser and compared them with specimens of the four other Italian viper species. Based on these specimens, we improved the diagnosis of V. walser and provided a first evaluation of intraspecific variability and ontogenetic variation. The skull of V. walser is subject to significant variation, most likely related to ontogeny in some cases (i.e., development of the parietal crest, development of the basioccipital process, shape of the posterior margin of the parabasisphenoid, shape of the quadrate). Based on the studied material, it is possible to distinguish V. walser from the other Italian vipers by the shape of the occipital crest of the supraoccipital, which is posteriorly directed, whereas it is laterally directed in the other species. The osteological diagnosibility provides further support for the validity of V. walser as a distinct species from Vipera berus.     

Comparative finite element analysis of the cranial performance of four herbivorous marsupials

Marsupial herbivores exhibit a wide variety of skull shapes and sizes to exploit different ecological niches. Several studies on teeth, dentaries, and jaw adductor muscles indicate that marsupial herbivores exhibit different specializations for grazing and browsing. No studies, however, have examined the skulls of marsupial herbivores to determine the relationship between stress and strain, and the evolution of skull shape. The relationship between skull morphology, biomechanical performance, and diet was tested by applying the finite element method to the skulls of four marsupial herbivores: the common wombat (Vombatus ursinus), koala (Phascolarctos cinereus), swamp wallaby (Wallabia bicolor), and red kangaroo (Macropus rufus). It was hypothesized that grazers, requiring stronger skulls to process tougher food, would have higher biomechanical performance than browsers. This was true when comparing the koala and wallaby (browsers) to the wombat (a grazer). The cranial model of the wombat resulted in low stress and high mechanical efficiency in relation to a robust skull capable of generating high bite forces. However, the kangaroo, also a grazer, has evolved a very different strategy to process tough food. The cranium is much more gracile and has higher stress and lower mechanical efficiency, but they adopt a different method of processing food by having a curved tooth row to concentrate force in a smaller area and molar progression to remove worn teeth from the tooth row. Therefore, the position of the bite is crucial for the structural performance of the kangaroo skull, while it is not for the wombat which process food along the entire tooth row. In accordance with previous studies, the results from this study show the mammalian skull is optimized to resist forces generated during feeding. However, other factors, including the lifestyle of the animal and its environment, also affect selection for skull morphology to meet multiple functional demands. J. Morphol. 276:1230–1243, 2015. © 2015 Wiley Periodicals, Inc.     

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     

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.     

Functional cranial analysis of large animalivorous bats (Microchiroptera)

Large animalivorous bats include carnivorous, piscivorous and insectivorous microchiropterans. Skull proportions and tooth morphology are examined and interpreted functionally. Four wide- faced bats from four families are convergent in having wide skulls, large masseter muscle volumes and stout jaws, indicating a powerful bite. Three of the four also have long canine teeth relative to their maxillary toothrows. Carnivorous bats have more elongate skulls, larger brain volumes and larger pinnae. The wide-faced bats are all dral emitters and have heads positively tilted relative to the basicranial axis. The carnivorous species are nasal-emitting bats and have negatively tilted heads. The orientation of the head relative to the basicranial axis affects several characters of the skull and jaws and is not correlated with size. The speculation that the type of echolocation may be more of a determinant of evolutionary change than the feeding mechanism is addressed. Wide-faced bats are thought to be capable of eating hard prey items (durophagus) and are probably non- discriminating, aurally less sophisticated insect generalists while the carnivorous and non- durophagus insectivorous bats may be more discriminating and aurally more sophisticated in what they eat.