A chroniosuchid from the Triassic of Kyrgyzstan and analysis of chroniosuchian relationships

A nearly complete skull and associated osteoderms from the Middle/Upper Triassic Madygen Formation of Kyrgyzstan are referred to a new chroniosuchid genus and species. The new taxon is characterized by a parabolic skull outline, pustular ornamentation, tabular‐squamosal contact, marked postparietal embayments, and the lack of an antorbital fontanelle. The palate is only preserved in part, showing broad palatines and ectopterygoids. Presence of a preorbital fenestra and characteristic osteoderm morphology are synapomorphies shared with all other chroniosuchids. According to the phylogenetic analysis performed, the new chroniosuchid nests with Chroniosaurus, with which it shares the wide, transversely extended osteoderms and pustular ornamentation. The chroniosuchians are robustly supported as a natural group, but their position within the reptiliomorph (stem‐amniote) clade is not adequately understood. Whereas the parasphenoid is similar to that of anthracosaurs, most other characters support a higher nesting of chroniosuchians within the stem‐amniotes. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 515–530.     

The sutural pattern of skull-roof bones in Lower Permian Discosauriscus austriacus from Moravia

Sutures between ornamented bones of Discosauriscus austriacus are mostly simple, but there are also more complicated, rarely serrated, sutures between some bones. In small individuals, the sutures are simple, but the same sutures also occur in the largest specimens. The character of the sutures and the incomplete ossification of bones around the pineal foramen indicate the larva type of organization of Discosauriscus The fenestra between premaxillaries and nasals appears to be absent. In the majority of specimens, a squamosal-intertemporal sutural contact is present, althought it is sometimes reduced and in a few cases interrupted by a postorbital and supratemporal contact Therefore the character 'intertemporal-squamosal suture present or absent' cannot be used in this rigorous sense for testing the relationships of early tetrapods. The configuration of the suture between both parietals in osteolepiforms, Discosauriscus , and various early amphibians and reptiles indicates that the bones enclosing the pineal foramen in osteolepifonns are frontals. *** D iscosauriscus . Seymouriamorpha, Lower Permian tetrapod, skull exoskeleton, sutures.     

Development and evolution of the tetrapod skull–neck boundary

The origin and evolution of the vertebrate skull have been topics of intense study for more than two centuries. Whereas early theories of skull origin, such as the influential vertebral theory, have been largely refuted with respect to the anterior (pre‐otic) region of the skull, the posterior (post‐otic) region is known to be derived from the anteriormost paraxial segments, i.e. the somites. Here we review the morphology and development of the occiput in both living and extinct tetrapods, taking into account revised knowledge of skull development by augmenting historical accounts with recent data. When occipital composition is evaluated relative to its position along the neural axis, and specifically to the hypoglossal nerve complex, much of the apparent interspecific variation in the location of the skull–neck boundary stabilizes in a phylogenetically informative way. Based on this criterion, three distinct conditions are identified in (i) frogs, (ii) salamanders and caecilians, and (iii) amniotes. The position of the posteriormost occipital segment relative to the hypoglossal nerve is key to understanding the evolution of the posterior limit of the skull. By using cranial foramina as osteological proxies of the hypoglossal nerve, a survey of fossil taxa reveals the amniote condition to be present at the base of Tetrapoda. This result challenges traditional theories of cranial evolution, which posit translocation of the occiput to a more posterior location in amniotes relative to lissamphibians (frogs, salamanders, caecilians), and instead supports the largely overlooked hypothesis that the reduced occiput in lissamphibians is secondarily derived. Recent advances in our understanding of the genetic basis of axial patterning and its regulation in amniotes support the hypothesis that the lissamphibian occipital form may have arisen as the product of a homeotic shift in segment fate from an amniote‐like condition.     

The cranial morphology of Greererpeton burkemorani Romer (Amphibia: Temnospondyli)

The skull of Greererpeton burkemorani Romer, a temnospondyl amphibian from the Upper Mississippian at Greer, West Virginia is described. A detailed account of the stapes of a Mississippian amphibian is given for the first time and its function is discussed. It is suggested that the stapes formed the principal element of support for the back of the braincase and resisted potential dislocation of the otico-occipital region from the skull roof during contraction of the hypaxial musculature.
Greererpeton is included in the Colosteidae and an amended diagnosis of the family is given. Erpetosaurus differs from Colosteus, Greererpeton and Pholidogaster in the pattern of bones in the skull roof and palate, the dentition and the otic region and, consequently, it is removed from the Colosteidae. The Temnospondyli are considered to be a monophyletic group characterized by the development of a connection between the dorsal portion of the occipital arch, the exoccipital bones, and the skull roof. The loxommatids are removed from the Temnospondyli as they retain the plesiomorphic condition of braincase attachment which relies exclusively on derivatives of the auditory capsules.
On the basis of similarities in the structure of the braincase, palate and manus it is suggested that microsaurs are the collateral descendants (sister group) of temnospondyls. This relationship may account for the large number of similarities in the three living groups of Amphibia: Anura are generally believed to have descended from temnospondyls, while the Urodela and Apoda are often considered to have descended from microsaurs. These systematic conclusions endorse the recent suggestions that neither the Lepospondyli nor the Labyrinthodontia are natural groups, and both terms should be abandoned.     

The subdivisions and fusions of the exoskeletal skull bones ofDiscosauriscus austriacus (Makowsky 1876) and their possible homologues in rhipidistians

Two localities in the Boskovice Furrow region of Moravia (Czecho-Slovakia) have produced new, well preserved material of the Lower Permian tetrapodDiscosauriscus austriacus (Makowsky 1876). A relatively large number of specimens have been found with some dermal skull roof bones partly or fully subdivided, and/or fused. These are the first records of such subdivisions and fusions in this tetrapod and the following bones are discussed: frontal, postfrontal, parietal, intertemporal, supratemporal, tabular and postparietal. The subdivided bones within the skull roof ofDiscosauriscus (and some dissorophoids) are situated at the same places as those found in Devonian rhipidistians, and may relate to the homology of the bones of the cranial exoskeletal roof inDiscosauriscus and in osteolepiforms. It shows that the “orthodox” terminology of the skull roof bones used in osteolepiforms is correct.     

The Evolution of the Tetrapod Middle Ear in the Rhipidistian-Amphibian Transition

From a survey ot the structure of the skull in rhipidistianfishes and early labylinthodont Amphibia and of the mechanismof hearing in these two groups, an account of the evolutionof the tetrapod middle ear is presented. The overall modificationof the otic region of the skull during the rhipidistian-amphibiantransition is analyzed in terms of changes in different organsystems in response to different selective pressures (affecting,for example, the feeding, respiratory, and locomotory mechanisms).These changes are seen to occur in a completely integrated pattern.Considerations of the different requirements for sound receptionunder water and in air, in connection with this correlated progressionof evolutionary change in the otic region of the head, revealthe manner in which the hyomandibular, spiracular diverticulum,and operculum of rhipidistian fishes became modified to formthe stapes, the tympanic cavity, and the outer portion of thetympanum, respectively, of tetrapods.     

Comparative morphology and functional interpretation of the sutures in the dermal skull roof of temnospondyl amphibians

Sutures in the dermal skull roof of several Palaeozoic temnospondyl amphibians were studied, including Archegosaurus decheni, Sclerocephalus haeuseri, Cheliderpeton latirostre (Archegosauridae), Acanthostomatops vorax (Zatrachydidae), Onchiodon labyrinthicus (Eryopidae), Micromelerpeton credneri and Branchierpeton amblystomum (Dissorophoidea). Lamellae, flat bevels, butt joints, steep walls, and grooves are the sutural types occurring. Morphological sutural differences cannot be used as taxonomically relevant characters, as they mainly differ in their functional manifestations. Similarities of sutural patterns in taxa not closely related are most probably based on convergence, therefore sutural morphology allows functional conclusions rather than revealing phylogenetic relationships. The basic sutural pattern is described in Micromelerpeton credneri. Different textures and obliquity patterns of the sutural surfaces are characteristic of each region of the skull roof. In all temnospondyls studied, the median plane and the margin of the skull roof show a nearly uniform sutural morphology; the circumorbital region is most complex. Although the skulls studied are akinetic, mesokinetic movements of dermal elements along their sutures must have been permitted to guarantee the dynamic stability of the skull construction.     

The identification of the dermal bones of the head

The discovery of the ichthyostegid Amphibia in Upper Devonian rocks by Säve-Söderbergh (1932) introduced further difficulties into the already complex problems of the dermal bones of the skull roof. For some years previously ideas about the origin of the tetrapods had been dominated by Watson's (1926) Croonian Lecture in which he had demonstrated beyond reasonable doubt that the crossopterygian fishes and not the Dipnoi were their ancestors, and had attempted to show that many of the features of the Carboniferous labyrinthodonts were a direct inheritance from these fishes. It was to be expected, therefore, that any Amphibia from the Upper Devonian would be intermediate in their structures between the Middle Devonian osteolepids and the Carboniferous labyrinthodonts, but when discovered the ichthyostegids did not conform at all well to this expectation. While their skulls showed some very primitive features which might have been expected, the pattern of the dermal bones did not conform to plan, for these new animals had lost, it seemed, the intertemporals, bones found in both the osteolepids and nearly all early labyrinthodonts, and had a single postparietal bone in place of the paired bones of all other early Amphibia. The osteolepid skull had many more bones than these earliest Amphibia.     

Creating morphological diversity in reptilian temporal skull region: A review of potential developmental mechanisms

Reptilian skull morphology is highly diverse and broadly categorized into three categories based on the number and position of the temporal fenestrations: anapsid, synapsid, and diapsid. According to recent phylogenetic analysis, temporal fenestrations evolved twice independently in amniotes, once in Synapsida and once in Diapsida. Although functional aspects underlying the evolution of tetrapod temporal fenestrations have been well investigated, few studies have investigated the developmental mechanisms responsible for differences in the pattern of temporal skull region. To determine what these mechanisms might be, we first examined how the five temporal bones develop by comparing embryonic cranial osteogenesis between representative extant reptilian species. The pattern of temporal skull region may depend on differences in temporal bone growth rate and growth direction during ontogeny. Next, we compared the histogenesis patterns and the expression of two key osteogenic genes, Runx2 and Msx2, in the temporal region of the representative reptilian embryos. Our comparative analyses suggest that the embryonic histological condition of the domain where temporal fenestrations would form predicts temporal skull morphology in adults and regulatory modifications of Runx2 and Msx2 expression in osteogenic mesenchymal precursor cells are likely involved in generating morphological diversity in the temporal skull region of reptiles.     

Adaptive problems and possibilities in the temporal fenestration of tetrapod skulls

Adaptive explanations for the temporal fenestration in reptiles are briefly reviewed. With few possible exceptions, fenestrate appeared first in the reptiles, and have seemingly evolved independently in several different phyletic lines. The several explanations for fenestration offered by previous authors include speculations that open spaces in the skull permitted bulging of the jaw-closing muscles, and that fenestrae formed in areas of reduced stress where the presence of bone would be functionally useless. The first of these does not readily apply to initial evolutionary stages; the second is more satisfactory. Certain features of muscular attachments to bones are dealt with, and their implications applied to the fenestration problem to add another possible explanation (which need not contradict previously published suggestions). Considerations of cranial strength in tetrapod skulls led to speculations on the lack of fenestration in temnospondyls, anthracosaurs, microsaurs and cotylosaurs. Emargination of the skull roof in turtles is also discussed.     

Osteocranial development in the viviparous surfperch Amphistichus argenteus (pisces: Embiotocidae)

The development of the cranial and branchial skeleton of the surfperch Amphistichus argenteus, a member of the family Embiotocidae, is described, and phylogenetic and functional aspects of the skull development of this species are discussed. The earliest bones to appear are those dermal elements of the branchial skeleton involved with feeding, and the bones, both dermal and endochondral, located in the basicranial region of the neurocranium. These are followed by dermal bones associated with the lateral line system and finally by the remainder of the bones of the branchial skeleton and the cartilaginous bones of the otic capsules. The last bone to develop is the ethmoid.     

The Parietal Problem: How to Cut This Gordian Knot?

Although no one has disputed that the piscine progenitors of the tetrapods have a homologue of the human parietal bone, opinions differ as to where in the skull roof this homologue is located. One view holds it to be either of two interorbital bones that together surround the foramen neuroepiphysium (and the so-called pineal plates); another, that it is each of the two mesial bones which comes next in order, behind the orbital cavities. Both of these views are untenable because neither of the proposed bones has proved to be amenable to conversion to the parietal bone of man. In seeking a solution to this issue thoughts turn to the tentorium cerebelli, whose topographic relationships and comparative morphology place it in the key position as a plausible derivative of the posterior half of the cranium of the tetrapod forerunners. Following this line of reasoning, it can be suggested that the tentorial exoskeleton, here called the pluteal bones, originally was situated in the dermis superjacent to the synotic tectum. In early therian phylogeny, these skull bones were covered by the backwardly expanding cerebral hemispheres which, concomitantly, became overgrown and thus protected by epiotic exoskeleton. It follows that the likely homologues of the human parietal bones are those parts of the skull roof of the piscine progenitors of the tetrapods that lie dorsolateral to the otic capsules.     

四川自贡大山铺蜀龙动物群——简报Ⅴ 两栖类

本文对扁头中国短头鲵(新属新种) (Sinobrachyops placenticephalus gen. et sp. nov.) 的形态特征和分类位置进行简述.标本得自著名的恐龙化石产地——自贡大山铺,产出时代为中侏罗世.中国短头鲵是目前迷齿类中在地史上最年轻的一个属.它的发现使迷齿类在地球上生存时代的上限推移到中侏罗世.     

Sutural bone frequency in synostotic rabbit crania

This study tests the hypothesis that crania with synostosed sutures will have a significantly higher incidence of calvarial sutural bones than normal crania. Sutural bones were counted in seven calvarial sutures and compared among four groups of adult New Zealand white rabbit skulls: normal in-colony (NI) controls (N = 14), normal out-colony (NO) controls (N = 12), skulls with familial delayed onset (DO) coronal synostosis (N = 25), and skulls with experimentally immobilized coronal sutures (EI) (N = 20). Comparisons among groups were made with a Kruskal-Wallis one-way ANOVA and between groups with a Mann-Whitney U-test, using a Bonferroni correction for multiple comparisons. Significant differences (P > 0.05) were noted only in the coronal and sagittal sutures, with EI crania having the greatest number of coronal sutural bones; between group differences were undetectable for sagittal sutural bones. A post hoc two-sample binomial test for equal proportions showed that the distribution of coronal sutural bones among individuals across groups was even, while the distribution of sagittal sutural bones was significantly higher in EI crania. These results suggest that altered sutural forces of the calvaria contribute to an increased occurrence of sutural bones. However, the influence of inheritance on increased occurrence of sutural bones cannot be discounted, as reflected in the equivalent number of individuals across groups that possessed coronal sutural bones. Am J Phys Anthropol 102:555–563, 1997. © 1997 Wiley-Liss, Inc.     

Laidleria uncovered: a redescription of Laidleria gracilis Kitching (1975), a temnospondyl from the Cynognathus Zone of South Africa

The holotype skull of Laidleria gracilis Kitching has been prepared from the dorsal surface, and the skull disarticulated from the vertebral column to expose the occiput. Unexpected features of the dorsal skull roof are the absence of an otic notch and tabular horn. The occiput has reduced paroccipital processes with extremely small or absent posttemporal fenestrae as a consequence of the lack of the tabular horns. Both stapes are in place with their distal ends anterior to an exceptionally deep occipital flange formed by ventral processes from the postparietals, tabulars, squamosals and quadratojugals. They thus abut the ventral surface of the skull roof rather than a tympanum. The vertebrae have well ossified, almost spool-shaped, intercentra associated with small, paired, pleurocentra. It is concluded that L. gracilis should be left in its own family, the Laidleriidae, which may prove the sister group to the Rhytidosteidae.