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.     

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.     

Cranial morphology of the Early Permian mesosaurid Mesosaurus tenuidens and the evolution of the lower temporal fenestration reassessed

The Early Permian mesosaurids are the oldest known aquatic amniotes with an exclusively Gondwanan distribution. Although several hundred of complete skeletons have been discovered and intensively studied, the anatomy and taxonomic composition of the group, as well as its phylogenetic relationships remain controversial. Several well-preserved mesosaurid specimens found in Uruguay justify a new anatomical reconstruction of the skull of Mesosaurus tenuidens, differing from earlier ones especially in the presence of a lower temporal fenestra. The significance of this structure for the evolution of temporal fenestration in amniotes is evaluated according to the two most recent phylogenetic hypotheses, in which mesosaurids are basalmost sauropsids or basalmost parareptiles. A synapsid-like fenestration may be the primitive condition for Amniota, and it may be also a basal condition for parareptiles, because recent phylogenies suggest a basal position for mesosaurids and lanthanosuchoids within that group, and both possess a lower temporal fenestra. Our results also give a moderately strengthened support for diapsid affinities of turtles.     

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.     

The temporal ligaments and their bearing on cranial kinesis in lizards

In the lateral temporal opening of the lacertilian skull there are three main types of ligaments: quadratojugal, jugomandibular, and quadratopostfrontal. The quadratojugal and ventral jugomandibular ligaments transmit the retraction force from the lower jaw to the rhinal unit of the maxillary segment. The dorsal jugomandibular ligament causes the coupled kinesis, that results in the protraction of the whole maxillary segment (in the amphikinetic skull with simple streptostyly) or in the protraction of the quadrate-mandibular complex (in the hyperstreptostylic skull), when opening the mouth.     

云南东部中三叠世始鳍龙类—新属种(英文)

根据一保存完整的骨骼(包括头骨、下颌和全部头后骨骼)建立了鳍龙类的一新属新种——利齿滇东龙(Diandongosaurus acutidentatus gen.et sp.nov.)。标本采自云南省罗平县中三叠世安尼期关岭组上段。新属种既具有肿肋龙类(包括Dactylosaurus,Anarosaurus,Serpianosaurus和Neusticosaurus)吻部两侧不收缩、眶前区域长于眶后区域、眼眶大于上颞孔等典型特征,同时又具有幻龙类(包括Simosaurus和nothosaurians)前颌和下颌前部犬齿型齿发育、上颌具1或2个犬齿型齿等典型特征。新种的额骨和顶骨均愈合,额骨两后外侧支的末端后延超过上颞孔前缘,轭骨和鳞骨相交将眶后骨排除于下颞孔之外,方轭骨发育,锁骨前外侧缘具一突起,3对荐肋以及最前部尾肋的外侧端均未见明显收缩。这些特征也多表现出肿肋龙类和幻龙类的混合特征。此外,新种还具有一些较为独特的特征,包括前额骨和后额骨沿眼眶背缘相交,第3至第8对尾肋的长度超过荐肋,后肢末端发育异常膨大的爪趾骨。系统关系分析表明,滇东龙既不是肿肋龙类也不是幻龙类;它可能与由乌蒙龙、幻龙类和传统的肿肋龙类所构成的分支亲缘关系最近,为始鳍龙类基干类群。     

云南早侏罗世一新具嵴冠兽脚类恐龙

报道了具嵴冠的兽脚类恐龙一新属种,安龙堡双柏龙(Shuangbaisaurus anlongbaoensis gen.et sp.nov.).双柏龙发现于云南省楚雄彝族自治州双柏县下侏罗统冯家河组中,保存了部分头骨带下颌.双柏龙沿两侧眼眶背缘向上有嵴冠发育,这在其他兽脚类中未曾报道过.与其他早侏罗世体型较大且具一对矢状嵴冠的兽脚类(双嵴龙属和中国龙属)相比,双柏龙还独具一些特征组合,如相对较高的前颌骨体、抬高的前颌骨腹缘、后腹向延伸的轭骨后突及较小的上颞孔.比较研究表明,尽管中国“双嵴龙”(“Dilophosaurus”sinensis)可能应归入中国龙属(Sinosaurus),但未必属于模式种.双柏龙的发现将有助于研究基干兽脚类的演化,尤其是各类头骨骨饰在其中的作用.     

Cranial morphology of Dvinia prima amalitzky (Cynodontia, Theromorpha)

The study of the skull of the Late Permian cynodont Dvinia prima Amalitzky, 1922 shows a combination of the general primitive skull design (many incisors, preservation of the precanine and large interpterygoid fenestra, etc) with the development of a number of “advanced” features (expansion of the temporal fenestra, development of the parietal crest, and closed pineal foramen, unusual structure of the premaxilla, complicated postcanines, and reduction of the angular wing). Dvinia prima is treated as a specialized omnivore and assigned to the family Dviniidae Sushkin, 1928 of the superfamily Thrinaxodontoidea Seeley, 1894.     

记裂头鳄属(Dibothrosuchus)一新种

本文记述的裂头鳄属-新种 (Dibothrosuchus xingsuensis sp. nov.)的标本采自云南禄丰盆地下禄丰组深红层.通过描述,对裂头鳄属属级特征作了补充.新种的头骨中鳞骨缺失降突;方骨极度向前背方伸展,形成大的耳凹,乌喙骨具后腹突,以及具有鳄类式的腕骨等,表明裂头鳄属应改属为楔形鳄科 (Sphenosuchidae).根据该科各属的头骨中方骨与脑颅侧壁的连接关系,可以清楚地看到楔形鳄科在这局部解剖学上存在一连续的发展过程.在形态上,裂头鳄属和南非的 Sphenosuehus (典型属)最为相近.     

CRANIAL CREST DEVELOPMENT IN THE AZHDARCHOID PTEROSAUR TUPUXUARA, WITH A REVIEW OF THE GENUS AND TAPEJARID MONOPHYLY

Abstract: A portion of pterosaur skull from the Romualdo Member of the Santana Formation (?Albian–?Turonian, Cretaceous) of north‐east Brazil provides new data on the morphology and ontogeny of azhdarchoid pterosaur cranial crests. The specimen consists of parts of the cranial bones posterodorsal to the nasoantorbital fenestra, including partial nasals, lacrimals, frontals and possibly the parietals. A posterodorsally directed premaxillary crest with a concave posterior border is located dorsal to the posterior border of the nasoantorbital fenestra. A well‐defined suture indicates overlapping, posterodorsally directed growth of the premaxilla over the skull roof, suggesting that the generation of the premaxillary crest is a late ontogenetic feature and thus probably related to sexual display. The systematics of Tupuxuara and its relationship to other azhdarchoids is reviewed and a cladistic analysis of the group is presented. Tupuxuara is found to be the sister‐taxon to Azhdarchidae. Tupuxuara longicristatus Kellner and Campos, 1988 is argued to be the only valid named species in this genus and Thalassodromeus Kellner and Campos, 2002 is considered a junior subjective synonym of this taxon. As originally conceived, Tapejaridae is paraphyletic: a new, more restrictive version of Tapejaridae (including Tapejara and Sinopterus dongi) might exist, but its monophyly is weakly supported. Furthermore, Tapejara was found to be paraphyletic in all trees.     

Morphospace occupation in thalattosuchian crocodylomorphs: skull shape variation, species delineation and temporal patterns

Abstract: Skull shape variation in thalattosuchians is examined using geometric morphometric techniques in order to delineate species, especially with respect to the classification of Callovian species, and to explore patterns of disparity during their evolutionary history. The pattern of morphological diversity in thalattosuchian skulls was found to be very similar to modern crocodilians: the main sources of variation are the length and the width of the snout, but these broad changes are correlated with size of supratemporal fenestra and frontal bone, length of the nasal bone, size of the orbit and premaxilla and position of the narial opening. Patterns of shape variation, in combination with discrete‐state morphology and stratigraphic and geographic range data were used to distinguish nine species of teleosaurid and 14 species of metriorhynchid, with the four currently recognized Callovian species being split into eight. Metriorhynchids were found to be more disparate from the average shape of morphospace than teleosaurids. However, short‐snouted metriorhynchids and long‐snouted teleosaurids showed the greatest amount of disparity with respect to snout morphotypes, indicating that each group tended to explore opposite areas of morphospace. Phylogeny was found to have a moderate influence on the pattern of morphospace occupation in metriorhynchids, but little effect in teleosaurids suggesting that other factors or constraints control the pattern of skull shape variation in thalattosuchians. A comparison of thalattosuchians with dyrosaur/pholidosaurids shows that thalattosuchians have a unique skull morphology, implying that there are multiple ways to construct a ‘long snout’. Moreover, the skull geometry of the problematic species Pelagosaurus typus was found to converge on the teleosaurid area of morphospace. Finally, the temporal distribution of thalattosuchian species and morphotypes demonstrate a clear and highly correlated relationship with sea level curves and mass extinction events through the Jurassic and the Early Cretaceous.     

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.     

三列齿类爬行动物化石在新疆的发现

本文报道的三列齿类化石是该类动物在我国西北地区之首次出现,与四川自贡所产大山铺恐龙动物群中的自贡似卞氏兽(Bienotheroides zigongensis)归同属同种,故认为五彩湾组可与下沙溪庙组相对比。新标本的发现在若干形态特征上补充了以往记述之不足。     

四川自贡肉食龙一新种

和平永川龙(Yongchuanosaurus hepingensis sp. nov.)为四川盆地发现的又一相当完整肉食龙新材料,本文对它进行了记述.     

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

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

Paedomorphosis and skull structure in Malagasy chamaeleons (Reptilia: Chamaeleoninae)

The phylogenetic analysis of skull characteristics, along with data derived from lung and hemipenial morphology, does not support monophyly of the predominantly terrestrial chamaeleons, Brookesiinae ( Brookesia plus Rhampholeon ), as sister-taxon of a monophyletic clade of arboreal chamaeleons, the Chamaeleoninae. Instead, the phylogenetic interrelationships of chamaeleonine lizards are ( Brookesia ("Rhampholeon" (Bradypodion, Chamaeleo) ). The monophyly of the genus Rhampholeon is not demonstrated. The arboreal chamaeleons form a monophyletic group comprising two genera, Bradypodion and Chamaeleo. Bradypodion may be placed as sister-group of the genus Chamaeleo , but its phyletic position and monophyly cannot at present be corroborated.
This study documents that there is a high degree of intraspecific variability in skull characters which may be due to paedomorphic reduction in the skull, especially of small species. Characters affected by a variable degree of ossification include the separation of a prefrontal fontanelle, the contact of the squamosal with the parietal, the size and position of the fenestra vestibuli, the degree of ossification in the basicranial rim of the fenestra, the extent of ossification of the pterygoid wing, and the shape of the skull roof as indicated by the bones forming the dorsal margin of the orbit. The closure of the lateral aperture of the occipital recess might indicate that miniaturization, and concomitant paedomorphosis, may have played a role in the initial phases of chamaeleonine phylogeny.