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.     

An unusual new archosauriform from the Middle–Late Triassic of southern Brazil and the monophyly of Doswelliidae

Until now the Doswelliidae was considered a monospecific family including Doswellia kaltenbachi from the Late Triassic of North America. The phylogenetic position of this taxon remained enigmatic until recently, when a sister‐group relationship with the Proterochampsidae was suggested. In the present contribution we describe the new doswelliid species Archeopelta arborensis gen. et sp. nov. from the Middle–Late Triassic of Brazil. A cladistic analysis recovered Archeopelta, Doswellia, and Tarjadia within a monophyletic group of basal archosauriforms, the Doswelliidae. The monophyly of this family is supported by the presence of osteoderm ornamentation that is coarse, incised, and composed of regular pits and the presence of an unornamented anterior articular lamina. Archeopelta is more closely related to Doswellia than to other archosauriforms by the presence of basipterygoid processes anterolaterally orientated, dorsal centra with a convex surface, width of the neural arch plus ribs of the first primordial sacral that are three times the length of the neural arch, and iliac blade laterally deflected, with strongly convex dorsal margin, and a length less than three times its height. The phylogenetic analysis indicates that Doswellidae is the closest large monophyletic entity to Archosauria, which achieved a wide palaeolatitudinal distribution during the late Middle and Late Triassic time span. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 839–871.     

董氏中华盗龙(兽脚类:异特龙超科)脑颅新知:筛区、颅腔解剖和具气腔(英文)

董氏中华盗龙(Sinraptor dongi) 是产自中国西北部新疆侏罗纪石树沟组上部的一种大型兽脚类。其正型标本包括了保存完好的脑颅,这是在最近的化石修理中额外发现的。脑颅中骨化筛骨的横切面为 U 形。在脑颅的解剖方位,蝶筛骨和眶蝶骨之间有一未骨化空间,表明该恐龙存活时具有软骨质的中隔蝶骨。利用乳胶颅腔模型和 CT 扫描技术对颅腔和含气隐窝的形态进行了研究,发现了一些此前未描述过的特点。其中尤为引人注目的是,发育良好的尾鼓室隐窝[一般认为是虚骨龙类(coelurosaurs) 的典型特征]以及基蝶骨隐窝和相关气腔的内部形态。侧鼓室隐窝和基蝶骨隐窝有纵向通道连接,很可能同时具有气腔和血管功能。模型也表明,颅腔没有髓质隆起,小脑绒球隐窝的开孔为沙漏状。若不包含嗅束和嗅球,颅腔容积为 95 ml,脑容商在其他基干坚尾龙类( tetanurans) 的范围内。虽然这些类群的第Ⅱ-Ⅳ对脑神经相对位置各有差异,但前脑、中脑和后脑之间的角度与鲨齿龙类( carcharodontosaurids) [鲨齿龙(Carcharodontosaurus) 及南方巨兽龙(Giganotosaurus) ]相似。第Ⅸ,Ⅹ和Ⅺ对脑神经穿过一个半月形的孔。在异特龙超科( Allosauroidea) 中,中华盗龙(Sinraptor) 在颅腔形态上与鲨齿龙、南方巨兽龙和异特龙( Allosaurus) 的相似程度比它与高棘龙(Acrocanthosaurus) 的相似程度更高。     

Ontogeny and homology of the basipterygoid articulation in Pantodon buchholzi (Teleostei: Osteoglossomorpha)

The basipterygoid articulation, an articular connection between the base of the braincase and the palatoquadrate, is widespread among gnathostome vertebrates. However, among living teleosts it is present only in the osteoglossomorphs Arapaima , Heterotis , Scleropages , Osteoglossum and Pantodon . Study of the development of the hyopalatine arch and the basipterygoid articulation in Pantodon buchholzi based on an ontogenetic series of cleared and double stained specimens yielded the following results: the symplectic process of the hyosymplectic cartilage never develops and the symplectic is absent; the pars hyomandibularis fuses with the palatoquadrate; the dermopalatine and ectopterygoid originate as separate bones, but fuse subsequently; the basal process is first visible on the pars metapterygoidea of the palatoquadrate at 7.0 mm standard length (SL); at 11.5 mm SL the basipterygoid process appears on the parasphenoid and contacts the basal process, establishing the basipterygoid articulation; the endopterygoid is initially not involved in the articulation, but during subsequent development enlarges and eventually forms an articular groove for the reception of the basipterygoid process of the parasphenoid; the distal tip of the basal process of the metapterygoid, however, still forms the caudolateral part of the articular groove in the adult. We discuss previous hypotheses about the homology of the basipterygoid articulation of osteoglossoids in light of these findings. Based on the numerous shared similarities and its occurrence in all major actinopterygian lineages, we argue that the basipterygoid articulation in osteoglossoids is homologous to that in non teleostean actinopterygians and represents a plesiomorphic character state at the level of Osteoglossomorpha.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 1−13.     

On the origin of high growth rates in archosaurs and their ancient relatives: Complementary histological studies on Triassic archosauriforms and the problem of a “phylogenetic signal” in bone histology

Three possible hypotheses could explain the polarity of the histological features of basal archosauriform and archosauromorph reptiles: either, the fibrolamellar complex is basal; or, the lamellar-zonal complex is basal or finally, the condition varied, and each complex evolved more than once in these early groups. The answer to this question would have broad implications for our understanding of the physiological, ecological, and behavioral features of the first archosaurs. To this end, we sampled the bone histology of various archosauriforms and basal archosaurs from the Triassic and Lower Jurassic: erythrosuchids, proterochampsids, euparkeriids, and basal ornithischian dinosaurs, including forms close to the origin of archosaurs but poorly assessed phylogenetically. The new data suggest that the possibility of reaching and maintaining very high growth rates through ontogeny could have been a basal characteristic of archosauriforms. This was partly retained (at least during early ontogeny) in most lineages of Triassic pseudosuchians, which nevertheless generally relied on lower growth rates to reach large body sizes. This trend to slower growth seems to have been further emphasized among Crocodylomorpha, which may thus have secondarily reverted toward more generalized reptilian growth strategies. Accordingly, their “typical ectothermic reptilian condition” may be a derived condition within archosauriforms, homoplastic to the generalized physiological condition of basal amniotes. On the other hand, ornithosuchians apparently retained and even enhanced the high growth rates of many basal archosauriforms during most of their ontogenetic trajectories. The Triassic may have been a time of “experimentation” in growth strategies for several archosauriform lineages, only one of which (ornithodirans) eventually stayed with the higher investment strategy successfully.     

Cranial anatomy and palaeobiology of the Miocene marsupial Hondalagus altiplanensis and a phylogeny of argyrolagids

New cranial material of Hondalagus altiplanensis, from the middle Miocene of southern Bolivia, allows a rediagnosis of the genus and an assessment of its palaeobiology and phylogenetic relationships with other argyrolagid marsupials. The new specimens demonstrate several derived (synapomorphic) cranial features shared by HondalagusArgyrolagus: a globular braincase, ventrally directed occipital condyles, a broad zygomatic arch, and a short, deep dentary with a flat and long coronoid notch. Hondalagus had powerful masticatory muscles and its cementum-encased hypselodont cheek teeth suggests it had a very abrasive diet. The deep fossae on the lateral aspect of the skull of argyrolagids, interpreted by Simpson as large, laterally-facing orbits, are actually sharply margined temporal fossae. Hondalagus has a very large carotid foramen medially situated within the suture of the basisphenoid and basioccipital. A phylogenetic analysis of five argyrolagid genera was conducted using 32 characters (16 cranial, 16 dental) and a didelphid and a caenolestid as outgroups. Hondalagus-Argyrolagus-Microtragulus form a monophyletic group with an undescribed gen. et sp. nov. (MACN-Ch-1305) from the lower Miocene (Colhuehuapian) of Argentina as its sister taxon. Proargyrolagus appears as sister group to the other taxa of argyrolagids.     

A new fossil from the Jurassic of Patagonia reveals the early basicranial evolution and the origins of Crocodyliformes

Extant crocodylians have a limited taxonomic and ecological diversity but they belong to a lineage (Crocodylomorpha) that includes basal and rather generalized species and a highly diverse clade, Crocodyliformes. The latter was among the most successful groups of Mesozoic tetrapods, both in terms of taxonomic and ecological diversity. Crocodyliforms thrived in terrestrial, semiaquatic, and marine environments, and their fossil diversity includes carnivorous, piscivorous, insectivorous, and herbivorous species. This remarkable ecological and trophic diversity is thought only to occur in forms with a completely akinetic skull, characterized by a functionally integrated and tightly sutured braincase‐quadrate‐palate complex. However, the patterns of evolutionary change that led to the highly modified skull of crocodyliforms and that likely enabled their diversification remain poorly understood. Herein, a new basal crocodylomorph from the Late Jurassic of Patagonia is described, Almadasuchus figarii gen. et sp. nov. The new taxon is known from a well‐preserved posterior region of the skull as well as other craniomandibular and postcranial remains. Almadasuchus figarii differs from all other crocodylomorphs in the presence of six autapomorphic features, including the presence of a large lateral notch on the upper temporal bar, an otic shelf of the squamosal that is wider than long, a deep subtriangular concavity on the posterolateral surface of the squamosal, and an elongated pneumatopore on the ventral surface of the quadrate. Phylogenetic analysis focused on the origin of Crocodyliformes places Almadasuchus as the sister group of Crocodyliformes, supported by synapomorphic features of the skull (e.g. subtriangular basisphenoid, absence of basipterygoid process, absence of a sagittal ridge on the frontal, and a flat anterior skull roof with an ornamented dorsal surface). New braincase information provided by Almadasuchus and other crocodylomorphs indicates that most of the modifications on the posterior region of the skull of crocodyliforms, including the strongly sutured braincase, quadrate, and the extensive secondary palate appeared in a stepwise manner, and pre‐dated the evolutionary changes in the snout, jaws, and dentition. This indicates that the progressively increased rigidity of the skull provided the structural framework that allowed the great ecological diversification of crocodyliforms during the course of the Mesozoic. The phylogenetic pattern of character acquisition inferred for the strongly sutured (akinetic) skull and the appearance of more diverse feeding behaviours that create high mechanical loads on the skull provides another interesting parallel between the evolution of Mesozoic crocodyliforms and the evolutionary origins of mammals.     

The evolution of the basicranium in the Henophidia (Reptilia: Serpentes)

In lizards, a short Vidian canal pierces the base of the basipterygoid process. In snakes, the anterior opening of the primitively short Vidian canal lies on the dorsal surface of the basisphenoid, trapped in an intracranial position by downgrowths of the frontal and parietal which meet the lateral edges of the basisphenoid-parasphenoid complex. This condition is observed in anilioid snakes which retain other primitive features in the braincase: the paroccipital process and the spheno-occipital tubercle (Aniliidae only) and participation of the basioccipital in the apertura lateralis of the recessus scalae tympani.
Subsequent evolution within booid snakes shows a shift of the anterior opening of the Vidian canal around the anterior edge of the basisphenoid, thus acquiring a secondary extracranial position. This occurs in parallel within boine and pythonine snakes. Dinilysia shows a parallel development of die condition observed in advanced booid snakes. Pseudoboa , with its short Vidian canal opening in-tracranially, demonstrates that caenophidians originated from a basal henophidian or pre-henophidian stock. The Acrochordidae show a basicranium that can be interpreted as either primitive henophidian or primitive caenophidian.     

Validity of <Emphasis Type="Italic">Scorpaena jacksoniensis</Emphasis> and a redescription of <Emphasis Type="Italic">S. cardinalis</Emphasis>, a senior synonym of <Emphasis Type="Italic">S. cookii</Emphasis> (Scorpaeniformes: Scorpaenidae)

The Scorpaena cardinalis complex, including S. cardinalis, S. jacksoniensis and S. orgila, is defined. The genus Ruboralga (type species: S. jacksoniensis) is regarded as a junior synonym of Scorpaena. Scorpaena jacksoniensis Steindachner 1866, previously treated as a junior synonym of Scorpaena cardinalis Solander and Richardson 1842, is regarded here as a valid species. Scorpaena cookii Günther 1874, previously treated as a valid species, is regarded here as a junior synonym of S. cardinalis. Thus, recent recognition of the two Australasian scorpionfishes, i.e., S. cardinalis and S. cookii, are re-identified in this study as S. jacksoniensis and S. cardinalis, respectively. Scorpaena plebeia Solander 1842 is regarded as a junior synonym of S. cardinalis. Scorpaena jacksoniensis is distributed along the east coast of Australia from southern Queensland to eastern Victoria, whereas S. cardinalis occurs around northern New Zealand, the Kermadec Islands and offshore islands of the Tasman Sea. A neotype is designated for S. cardinalis. Morphological changes with growth in the two species are described in detail.     

Braincase evolution in suchian archosaurs (Reptilia: Diapsida): evidence from the rauisuchian Batrachotomus kupferzellensis

The osteology of an almost complete braincase of the rauisuchian archosaur Batrachotomus kupferzellensis Gower from the Middle Triassic of Germany is described. There is a possibly discrete epiotic ossification, the metotic fissure is undivided by bone (i.e. there is a metotic foramen), the medial wall of the otic capsule is mostly ossified, the cerebral branch of the internal carotid artery entered the lateral surface of the parabasisphenoid, the ventral ramus of the opisthotic is more prominent laterally than a strong subvertical ridge on the exoccipital and basioccipital that lies posterior to the external foramen for the hypoglossal nerve, and the perilymphatic foramen faces away from the otic capsule in a posterior direction. Braincase morphology in the rauisuchians Saurosuchus galilei , Postosuchus kirkpatricki, and Tikisuchus romeri is reviewed. A matrix of 27 braincase characters for 12 archosaurian taxa is analysed. The most parsimonious hypothesis is consistent with the currently orthodox view of archosaurian phylogeny, except in that aetosaurians are more closely related to crocodylomorphs than is any rauisuchian. This phylogeny is used in a brief interpretation of the evolution of derived braincase features present in extant crocodilians. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 136 , 49–76.     

Skeletal development of the direct-developing caecilian <Emphasis Type="Italic">Gegeneophis ramaswamii</Emphasis> (Amphibia: Gymnophiona: Caeciliidae)

A few previous studies of skeletal and especially skull development in Gymnophiona often provided contradictory results. We studied the development of the skull and vertebral column of Gegeneophis ramaswamii, a direct-developing Indian caeciliid, based on 13 specimens. The chondrocranium forms at (Brauer in Zool Jahrb Anat 12:477-508,1899) stage 38. First dermal and perichondral ossifications occur at stage 40. The first dermal bones to form are the mentomeckelian, dentary, angular, vomer, and premaxillary. These are followed by the coronoid, palatine, pterygoid, maxillary, and the skull-roofing bones. The last occurring dermal ossifications are the parasphenoid and the squamosal. We present evidence for the occurrence of a lacrimal bone. No ectopterygoid, basioccipital, supraoccipital, pleurosphenoid, postorbital, or supratemporal elements were found. We assess the homology of the bones constituting the caecilian skull and discuss the above-mentioned terminologies. The phylogenetic implications of our findings are briefly discussed and we conclude that the evidence from developmental morphology is at present consistent with a monophyletic Lissamphibia of temnospondyl origin.     

The basicranium of dicynodonts (Synapsida) and its use in phylogenetic analysis

Current phylogenetic hypotheses for the dicynodonts conflict, probably because the characters used, especially those of the jaws and facial region, show considerable convergence. Characters of the braincase and basipterygoid articulation of the Late Permian–Middle Triassic dicynodonts Diictodon , Dicynodon , Kingoria, Lystrosaurus , Rechnisaurus , and 14 other genera, may have phylogenetic value. Parsimony analysis and the character compatability permutation test suggest, at the highest possible confidence level, that the data set contains significant hierarchical structure, interpreted as a result of phylogeny. The most parsimonious tree broadly agrees with all recent hypotheses on the relationships among dicynodonts. However, it conflicts with the recent suggestion that Lystrosaurus is part of a clade of Middle–Late Triassic dicynodonts, but supports the basal position of Kingoria . The use of Eodicynodon as an outgroup does not perturb the parsimonious relationship of the included taxa. Topological constraints reveal that phylogenetic hypotheses based only on basicranial characters are not robust. Characters of the basipterygoid articulation and inner braincase have high consistency and retention indices, which suggests that the main evolutionary transformations in the dicynodont basicranium occurred within these structures.     

Synonymy of two species of the genus <Emphasis Type="Italic">Platycephalus</Emphasis> and validity of <Emphasis Type="Italic">Platycephalus westraliae</Emphasis> (Teleostei: Platycephalidae)

The type specimens of platycephalid Platycephalus endrachtensis Quoy and Gaimard 1825 are regarded as being conspecific with Platycephalus arenarius Ramsay and Ogilby 1886, so the latter becomes a junior synonym. This species is characterized as having a caudal fin with four or more longitudinal dark bands and lacking a yellow blotch. It is also found that Platycephalus westraliae (Whitley 1938), which had been considered to be a junior synonym of Platycephalus bassensis Cuvier 1829, is a valid species. Specimens that recently had been mistakenly identified as “P. endrachtensis,” having the caudal fin with three or four longitudinal dark bands and a yellow blotch on the upper lobe, should be referred to P. westraliae.     

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.