A BASAL MIXOSAURID ICHTHYOSAUR FROM THE MIDDLE TRIASSIC OF CHINA

Abstract:  New specimens of a mixosaurid ichthyosaur are described from the People's Republic of China. They consist of an almost complete skull, forefin and shoulder girdle of a single individual, and a second partial skull from the Anisian Guanling Formation of Guizhou Province that belongs to the same taxon. A phylogenetic analysis shows this species to be the sister-taxon of all other mixosaurids. It is distinguished from Mixosaurus , Contecotpalatus and Phalarodon by features of its dentition and cranial morphology. There is only one maxillary tooth row. The posterior maxillary teeth are blunt and rounded, and much wider than the premaxillary teeth, but they are not strongly elongated anteroposteriorly. The teeth are subthecodontously implanted. The postorbital skull segment is long. The jugal-quadratojugal notch is deep and these two elements remain separated externally. The postorbital separates supratemporal and postfrontal. There are no postaxial ossifications of the forefin. Autapomorphies include a long ventral process of the postorbital lamina of the postorbital as well as characteristic morphologies of the coracoid and humerus. The material is therefore clearly distinguished from other known mixosaurids. It can be referred, on the basis of the features of the humerus and coracoid, to Mixosaurus maotaiensis Young, 1965. As it is generically distinct from Mixosaurus , a new generic name, Barracudasaurus gen. nov. is proposed. The phylogenetic relevance of the external separation of jugal and quadratojugal in some Triassic ichthyosaurs is discussed and it is concluded that it does not demonstrate the initial presence of a lower temporal fenestra in ichthyosaurs. The new material suggests that mixosaurids could have also originated in the Eastern rather than the Western Tethys, as previously assumed.     

Skull of Grippia Longirostris: no contradiction with a diapsid affinity for the Ichthyopterygia

The skull of Grippia longirostris , one of the basal ichthyopterygians, is redescribed, based on a newly prepared referred specimen and the examination of others. Grippia had been considered to be the most basal ichthyopterygian until recently, and it has been suggested that the quadratojugal enters the upper temporal fenestra in this species. This and other unique features, which were considered plesiomorphic for ichthyopterygians, obscured the relationship of the group to other amniotes. It is shown here that the quadratojugal is much smaller than previously described, and does not enter the upper temporal fenestra. The skull configuration of the species is very similar to the condition in the recently-described skull of Utatsusaurus hataii , another ichthyopterygian belonging to the basal grade of the group. The close similarity between the two help clarify ichthyopterygian plesiomorphies. There is no feature in the skull of Grippia that particularly contradicts the emerging consensus that ichthyopterygians are diapsids.     

A new lizard-like reptile (Diapsida: Lepidosauromorpha) from the Middle Jurassic of England

The Middle Jurassic (Late Bathonian) Mammal Bed at Kirtlington, Oxfordshire, has yielded a rich assemblage of small vertebrates including mammals, frogs, salamanders and small reptiles. This paper describes the skull of a new diapsid reptile, Marmorelta oxoniensis, which was common within the fauna. The skull is gracile with large orbits and relatively long external nares. The frontals and parietals are single in the adult and almost separated on the roof of the skull by the postfrontals. The temporal region shows a mosaic of primitive and derived character states. The quadrate was firmly held to the skull by a large squamosal and a small, free, quadratojugal was retained. However, reduction of the anterior process of the quadratojugal and of the posterior process of the jugal left the lower temporal fenestra broadly open. Comparison with other diapsids leads to the conclusion that Marmoretta was a lepidosauromorph, most probably the sister taxon of Lepidosauria.     

The skull of a large Lower Triassic ichthyosaur from Spitzbergen and its implications for the origin of the Ichthyosauria

An incomplete ichthyosaur skull from the Lower Triassic (Spathian) Sticky Keep Formation of Spitzbergen is one of the earliest ichthyosaurs known and by far the largest of the most basal ichthyosaurs. The specimen shows an extremely plesiomorphic cheek region of the skull and allows for the first time a confident reconstruction of the grundplan condition of the lower cheek region of ichthyosaurs. It unequivocally demonstrates the presence of a deep emargination of the lower cheek, which is bordered dorsally by the quadratojugal, squamosal and jugal, and therefore a plesiomorphic lack of contact between jugal and quadratojugal. The specimen also allows an evaluation of recent skull reconstructions of several Lower Triassic taxa which probably contain severe misinterpretations. The structure of the temporal and cheek region of the specimen can be most parsimoniously explained by the initial presence of a lower temporal fenestra. This provides important new evidence that the ancestry of ichthyosaurs might have lain within the Diapsida or their stem-group.     

Cranial anatomy of Shunosaurus, a basal sauropod dinosaur from the Middle Jurassic of China

Shunosaurus, from the Middle Jurassic of China, is probably the best‐known basal sauropod and is represented by several complete skeletons. It is unique among sauropods in having a small, bony club at the end of its tail. New skull material provides critical information about its anatomy, brain morphology, tooth replacement pattern, feeding habits and phylogenetic relationships. The skull is akinetic and monimostylic. The brain is relatively small, narrow and primitively designed. The tooth replacement pattern exhibits back to front replacement waves in alternating tooth position. The teeth are spatulate, stout and show well‐developed wear facets indicative of coarser plant food. Upper and lower tooth rows interdigitate and shear past each other. Tooth morphology, skull architecture, and neck posture indicate that Shunosaurus was adapted to ground feeding or low browsing. Shunosaurus exhibits the following cranial autapomorphies: emargination of the ventral margin of the jugal/quadratojugal bar behind the tooth row; postorbital contains a lateral pit; vomers do not participate in the formation of the choanae; pterygoid is extremely slender and small with a dorsal fossa; quadrate ramus of the pterygoid is forked; quadratojugal participates in the jaw articulation; tooth morphology is a combination of cylindrical and spatulate form; basipterygoid process is not wrapped by the caudal process of the pterygoid; trochlear nerve has two exits; occlusal level of the maxillary tooth row is convex downward, whereas that of the dentary is concave upward, acting like a pair of garden shears; dentary tooth count is 25 or more; and the replacing teeth invade the labial side of the functional teeth. Cranial characters among the basal sauropods are reviewed. As Shunosaurus is the earliest sauropod for which cranial remains are known, it occupies an important position phylogenetically, showing the modification of skull morphology from the prosauropod condition. Although the skull synapomorphies of Sauropoda are unknown at present, 27 cranial synapomorphies are known for the clade Eusauropoda. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society, 2002, 136 , 145?169.     

早白垩世热河生物群一新的有尾两栖类

描述了内蒙古宁城下白垩统义县组下部一新的有尾两栖类：奇异热河螈（新属、新种）Ｊｅｈｏｌｏｔｒｉｔｏｎ ｐａｒａｄｏｘｕｓ ｇｅｎ． ｅｔ ｓｐ． ｎｏｖ．。标本保存了完好的相关节的骨架印痕,头骨中的翼 骨具有一个不与上颌骨相连,而与头骨中部相连的前内侧突,这使它区别于其他早期有尾类。     

In-vitro results of rapid maxillary expansion on adults compared with finite element simulations

This study was mainly performed to investigate the effects of high maxillary expansion forces on the skull with fresh and thiel-fixed human skulls. The maxillary suture was not weakened except in one experiment. This study compares the strain measured on the zygomatic process of the skull with the results of a finite element model generated for this purpose. An increasing transversal force was applied on the alveolar process (teeth) until rupture. Strain on the zygomatic process, maxilla displacement and the expanding forces were registered.The results of this study show linear material behaviour of the skull before rupture. The highest stress during the experiments and FE simulation was observed on the alveolar process.Conclusions of this study are the necessity of the existence of appropriate models and that female specimens seem to rupture at a lower force than male ones. Both male and female specimens show a similar linear behaviour in the force/strain curve within each gender group. The probability of maxillary suture opening in adults during ultra-rapid maxillary expansion with tooth anchorage is very low. Complications and unwanted rupture could occur.     

Feeding Motor Patterns in Anurans: Insights from Biomechanical Modeling

During feeding in anurans, the mouth opens while the tongue,which is attached to the mandible at the front of the mouth,rotates forward. Due to the relative simplicity of its anatomyand the complexity of its motion, tongue protraction in frogspresents an ideal system for exploring the neural control ofmultijoint movements. In this study, we used a forward dynamic,rigid body model with four segments and two muscles to investigateopen loop control of tongue protraction in the Australian white-lippedtree frog, Litoria caerulea. Model parameters include the massdistribution, initial position and initial angular velocityof each segment and the anatomy and physiology of each muscle.Model variables include the level of muscle activation at eachtime step and impulsive torques to open and close the mouth.The model gives X,Y coordinates of each segment and joint anglesat each time step as output. The model was tested using scaled,normalized EMG signals and impulsive joint torques to predictthe paths of the lower jaw tip and tongue tip. Predicted pathswere compared to experimentally observed paths using Pearsonproduct-moment correlation coefficients. Simulations demonstratethat the genioglossus muscles likely play a minor role, if any,in determining the trajectory of the tongue in most anurans.Most of the force for tongue protraction comes from angularmomentum transferred to the tongue by the opening jaws. In anurans,tongue protraction is dynamically stable and will occur as longas the musculoskeletal elements are in the correct initial position.     

The comparative feeding mechanism of gadidae and macrouridae. I. Functional morphology of the feeding apparatus

The comparative feeding apparatus of Gadidae and Macrouridae has been studied from a functional point of view. Some important modifications in macrourid species in comparison with the gadid ones were found: rostral end of the cranium, rostral ligaments and disposition of the jaw muscles. Anyways, all the modifications seem secondary, motivated by the habitat. That is true in the case of the beak, in relation with the swimming system which had been already observed by Marshall and Bourne (1964). The question of the rostral ligaments modification would be related with the protraction of the upper jaw, but a further analysis is required. Also discussed are two schemes of possible changes in the jaw musculature.     

A Basal Lithostrotian Titanosaur (Dinosauria: Sauropoda) with a Complete Skull: Implications for the Evolution and Paleobiology of Titanosauria

We describe Sarmientosaurus musacchioi gen. et sp. nov., a titanosaurian sauropod dinosaur from the Upper Cretaceous (Cenomanian—Turonian) Lower Member of the Bajo Barreal Formation of southern Chubut Province in central Patagonia, Argentina. The holotypic and only known specimen consists of an articulated, virtually complete skull and part of the cranial and middle cervical series. Sarmientosaurus exhibits the following distinctive features that we interpret as autapomorphies: (1) maximum diameter of orbit nearly 40% rostrocaudal length of cranium; (2) complex maxilla—lacrimal articulation, in which the lacrimal clasps the ascending ramus of the maxilla; (3) medial edge of caudal sector of maxillary ascending ramus bordering bony nasal aperture with low but distinct ridge; (4) ‘tongue-like’ ventral process of quadratojugal that overlaps quadrate caudally; (5) separate foramina for all three branches of the trigeminal nerve; (6) absence of median venous canal connecting infundibular region to ventral part of brainstem; (7) subvertical premaxillary, procumbent maxillary, and recumbent dentary teeth; (8) cervical vertebrae with ‘strut-like’ centroprezygapophyseal laminae; (9) extremely elongate and slender ossified tendon positioned ventrolateral to cervical vertebrae and ribs. The cranial endocast of Sarmientosaurus preserves some of the most complete information obtained to date regarding the brain and sensory systems of sauropods. Phylogenetic analysis recovers the new taxon as a basal member of Lithostrotia, as the most plesiomorphic titanosaurian to be preserved with a complete skull. Sarmientosaurus provides a wealth of new cranial evidence that reaffirms the close relationship of titanosaurs to Brachiosauridae. Moreover, the presence of the relatively derived lithostrotian Tapuiasaurus in Aptian deposits indicates that the new Patagonian genus represents a ‘ghost lineage’ with a comparatively plesiomorphic craniodental form, the evolutionary history of which is missing for at least 13 million years of the Cretaceous. The skull anatomy of Sarmientosaurus suggests that multiple titanosaurian species with dissimilar cranial structures coexisted in the early Late Cretaceous of southern South America. Furthermore, the new taxon possesses a number of distinctive morphologies—such as the ossified cervical tendon, extremely pneumatized cervical vertebrae, and a habitually downward-facing snout—that have rarely, if ever, been documented in other titanosaurs, thus broadening our understanding of the anatomical diversity of this remarkable sauropod clade. The latter two features were convergently acquired by at least one penecontemporaneous diplodocoid, and may represent mutual specializations for consuming low-growing vegetation.     

How do ants stick out their tongues?

The mouthparts are very important tools for almost any task performed by ants. In particular, the labiomaxillary complex is essential for food intake. In the present study we investigated the anatomical design of the labiomaxillary complex in various ant species, focusing on movement mechanisms. Six labial and six maxillary muscles with different functions control the several joints and ensure the proper performance of the labiomaxillary complex. According to our measurements of sarcomere lengths, muscle fiber lengths and diameters, and the relative muscle volumes, the labial and maxillary muscles feature rather slow than fast muscle characteristics and do not seem to be specialized for specific tasks. Since glossa protractor muscles are absent, the protraction of the glossa, the distal end of the labium, is a nonmuscular movement. By histological measurements of hemolymph volumes we could exclude a pressure-driven mechanism. Additional experiments showed that, upon relaxation of the glossa retractor muscles, the glossa protracts elastically. This elastic mechanism possibly sets an upper limit to licking frequency, thus influencing food intake rates and ultimately foraging behavior. In contrast to many other elastic mechanisms among arthropods, glossa protraction in ants is based on a mechanism where elasticity works as an actual antagonist to muscles. We compared the design of the labiomaxillary complex of ants with that of the honeybee and suggest an elastic mechanism for glossa protraction in honeybees as well.     

New Evidence for a Discrete Supratemporal Bone in the Jurassic Ichthyosaur Temnodontosaurus

The temporal region in Temnodontosaurus trigonodon and Temnodontosaurus platyodon consists of three discrete ossifications. These bones are identified as the supratemporal, bordering much of the temporal opening, and in mid-cheek, the squamosal, which is in contact with the quadratojugal along its ventral margin. Interpretation of the bone in mid-cheek position as a neomorphic "supernumerary" ossification is rejected. The supratemporal bone of ichthyosaurs is functionally convergent with the squamosal of diapsids. The size and configuration of the supratemporal casts doubt on the supposition that ichthyosaurs were diapsids.     

Revision of<Emphasis Type="Italic">Cheliderpeton vranyi</Emphasis><Emphasis Type="SmallCaps">Fritsch</Emphasis>, 1877 (Amphibia,Temnospondyli) from the Lower Permian of Bohemia (Czech Republic)

The temnospondylCheliderpeton vranyi Fritsch, 1877 from the Lower Permian Ruprechtice horizon (Rotliegend) of the Intrasudetic Basin (Bohemia, Czech Republic) is redescribed. Many features of the skeleton permit a new understanding of the type species and consequently of the genus. Diagnostic characters are the narrow and round-tipped snout, straight to convex outline of the skull roof, narrow and long otic notch, posteriorly expanded quadratojugal, and the relatively wide and short rhombic interclavicle. The ilium with a short, expanded dorsal branch and the missing contact of nasal/maxilla are features shared with the related Upper PermianIntasuchus from Russia and the Eryopidae.Actinodon germanicus is a junior synonym ofCheliderpeton vranyi.      

The cranial osteology and hyolaryngeal apparatus of Rhinophrynus dorsalis (Anura: Rhinophrynidae) with comparisons to recent pipid frogs

The cranial osteology (including the hyolaryngeal apparatus) of Rhinophrynus dorsalis (Anura: Rhinophrynidae) is described from whole skeletons and serial cross sections. Some unique features of the extensively ossified skull include the enlarged and protracted olfactory region, for which the nasals form part of the septum nasi; the relatively short maxillaries and broad premaxillaries, and the immense quadratojugal; the extreme forward position of the quadrate; the lack of a firm articulation of the pterygoid and quadrate with the neurocranium and crista parotica; the quadrate lacking the distinct processes typical of other frogs; a single foramen for Nn. II–VII; a large, distinct operculum; and a bipartite hyale. Rhinophrynus shares other unusual cranial characteristics with the other pipoid frogs, Xenopus, Pipa, Hemipipa, and Hymenochirus. Among these features are the presence of a single frontoparietal in the adult, and the absence of parasphenoid alae, palatines, and mentomeckelian bones. Rhinophrynus differs from the pipids in the lack of a columella and a palatine process on the premaxilla, and in the possession of a quadratojugal, parahyoid bone, paired prevomers, olfactory eminence, massive quadrate that lacks distinguishable processes, a modified squamosal, and a bipartite hyale. Although the cranium of Rhinophrynus is distinctive, the evolutionary significance of its unusual features will remain obscure until comparable data are gathered from other closely related groups, the Discoglossoidea and the Pelobatoidea.     

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.     

The Effects of Maxillary Protraction with or without Rapid Maxillary Expansion and Age Factors in Treating Class III Malocclusion: A Meta-Analysis

We conducted a comprehensive meta-analysis of 12 studies to examine whether maxillary protraction face mask associated with rapid maxillary expansion (FM/RME) could be an effective treatment for Class III malocclusion and to evaluate the effect of timing on treatment response. Patients with a maxillary deficiency who were treated with FM with or without RME were compared with those who had an untreated Class III malocclusion. In both treatment groups, forward displacement of the maxilla and skeletal changes were found to be statistically significant. In addition, posterior rotation of the mandible and increased facial height were more evident in the FM group compared with the control group. However, no significant differences were observed between the early treatment groups and late treatment groups. The results indicated that both FM/RME and FM therapy produced favorable skeletal changes for correcting anterior crossbite, and the curative time was not affected by the presence of deciduous teeth, early mixed dentition or late mixed dentition in the patient.     

Mechanical regulation of secondary chondrogenesis

The development of the skull is characterised by its dependence upon epigenetic influences. One of the most important of these is secondary chondrogenesis, which occurs following ossification within certain membrane bone periostea, as a result of biomechanical articulation. We have studied the genesis, character and function of the secondary chondrocytes of the quadratojugal of the chick between embryonic days 11 and 14. Analysis of gene expression revealed that secondary chondrocytes formed coincident with Sox9 upregulation from a precursor population expressing Cbfa1/Runx2: a reversal of the normal sequence. Such secondary chondrocytes rapidly acquired a phenotype that is a compound of prehypertrophic and hypertrophic chondrocytes, exited from the cell cycle and upregulated Ihh. Pulse and pulse/chase experiments with BrdU confirmed the germinal region as the highly proliferative source of the secondary chondrocytes, which formed by division of chondrocyte-committed precursors. By blocking Hh signalling in explant cultures we show that the enhanced proliferation of the germinal region surrounding the secondary chondrocytes derives from this Ihh source. Additionally, in vitro studies on membrane bone periosteal cells (nongerminal region) demonstrated that these cells can also respond to Ihh, and do so both by enhanced proliferation and precocious osteogenesis. Despite the pro-osteogenic effects of Ihh on periosteal cell differentiation, mechanical articulation of the quadratojugal/quadrate joint in explant culture revealed a negative role for articulation in the regulation of osteocalcin by germinal region descendants. Thus, the mechanical stimulus that is the spur to secondary chondrocyte formation appears able to override the osteogenic influence of Ihh on the periosteum, but does not interfere with the cell cycle-promoting component of Hh signalling.