A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods

Megalosaurus bucklandii (Dinosauria: Theropoda), the oldest named dinosaur taxon, from the Bathonian (Middle Jurassic) of England, is a valid taxon diagnosed by a unique character combination of the lectotype dentary. Abundant referred material is described and several autapomorphies are identified: ventral surfaces of first and third to fifth sacral centra evenly rounded, ventral surface of second sacral centrum bearing longitudinal, angular ridge; dorsally directed flange around midheight on the scapular blade; an array of posterodorsally inclined grooves on the lateral surface of the median iliac ridge; anteroposteriorly thick ischial apron with an almost flat medial surface; and complementary groove and ridge structures on the articular surfaces between metatarsals II and III. A new phylogenetic analysis focuses on basal tetanurans and includes 41 taxa, six of which have never been included in a cladistic analysis, and 213 characters, 29 of which are new. This is the first phylogenetic analysis to focus on basal tetanuran relationships, and it reveals several new results. Megalosauroidea (= Spinosauroidea) includes two clades, basal to the traditional content of Megalosauridae + Spinosauridae. These comprise Xuanhanosaurus, Marshosaurus, Condorraptor + Piatnitzkysaurus and Chuandongocoelurus + Monolophosaurus. Almost all large‐bodied Middle Jurassic theropods are megalosauroids, but Poekilopleuron is an allosauroid. Megalosauroids show geographical differentiation among clades, indicating the development of endemic theropod faunas across Pangaea during the Middle Jurassic. Notably, megalosaurids are not known from outside of Europe during this epoch. Megalosauroids are less diverse and abundant during the Late Jurassic, when most theropods are neotetanurans and allosauroids dominate the large‐bodied predator niche. This indicates faunal turnover between the Middle and Late Jurassic. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 882–935.     

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.     

Osteology and relationships of a new theropod dinosaur from the Middle Jurassic of Patagonia

Abstract:  A new taxon of theropod dinosaur is described as Condorraptor currumili gen. et sp. nov., from the Middle Jurassic (Callovian) Cañadón Asfalto Formation of Chubut Province, Argentinean Patagonia. The taxon is represented by a single fragmentary postcranial skeleton. Although incompletely known, Condorraptor is the second most complete theropod from the Middle Jurassic of Gondwana. The new taxon is characterized by the absence of a posterior incision between the fibular condyle and the medial side of the proximal articular end of the tibia, the pleurocoels in the anterior cervicals being situated posteroventral to the parapophyses, and the presence of a pronounced 'step' between the distal articular facet and shaft of Mt IV. Pneumatic features of the vertebral column show strong variation between the left and right side. Condorraptor gen. nov. can be referred to the Tetanurae and is a representative of a global radiation of basal tetanurans in the Early to Mid Jurassic.     

New information on the cranial anatomy of Acrocanthosaurus atokensis and its implications for the phylogeny of Allosauroidea (Dinosauria: Theropoda)

Background

Allosauroidea has a contentious taxonomic and systematic history. Within this group of theropod dinosaurs, considerable debate has surrounded the phylogenetic position of the large-bodied allosauroid Acrocanthosaurus atokensis from the Lower Cretaceous Antlers Formation of North America. Several prior analyses recover Acrocanthosaurus atokensis as sister taxon to the smaller-bodied Allosaurus fragilis known from North America and Europe, and others nest Acrocanthosaurus atokensis within Carcharodontosauridae, a large-bodied group of allosauroids that attained a cosmopolitan distribution during the Early Cretaceous.

Methodology/Principal Findings

Re-evaluation of a well-preserved skull of Acrocanthosaurus atokensis (NCSM 14345) provides new information regarding the palatal complex and inner surfaces of the skull and mandible. Previously inaccessible internal views and articular surfaces of nearly every element of the skull are described. Twenty-four new morphological characters are identified as variable in Allosauroidea, combined with 153 previously published characters, and evaluated for eighteen terminal taxa. Systematic analysis of this dataset recovers a single most parsimonious topology placing Acrocanthosaurus atokensis as a member of Allosauroidea, in agreement with several recent analyses that nest the taxon well within Carcharodontosauridae.

Conclusions/Significance

A revised diagnosis of Acrocanthosaurus atokensis finds that the species is distinguished by four primary characters, including: presence of a knob on the lateral surangular shelf; enlarged posterior surangular foramen; supraoccipital protruding as a double-boss posterior to the nuchal crest; and pneumatic recess within the medial surface of the quadrate. Furthermore, the recovered phylogeny more closely agrees with the stratigraphic record than hypotheses that place Acrocanthosaurus atokensis as more closely related to Allosaurus fragilis. Fitch optimization of body size is also more consistent with the placement of Acrocanthosaurus atokensis within a clade of larger carcharodontosaurid taxa than with smaller-bodied taxa near the base of Allosauroidea. This placement of Acrocanthosaurus atokensis supports previous hypotheses of a global carcharodontosaurid radiation during the Early Cretaceous.     

Limusaurus inextricabilis (Theropoda: Ceratosauria) gives a hand to evolutionary teratology: a complementary view on avian manual digits identities

It is widely accepted that birds are rooted within theropod dinosaurs. However, there is controversy between palaeontological and developmental data regarding manual digit identities of birds and their tetanuran ancestors (I, II and III vs. II, III and IV). To resolve this conflict, the principle of a frame‐shift has been considered. Identities of digits I–III would develop on condensations 2–4. Nevertheless, the discovery of the basal Ceratosauria Limusaurus inextricabilis has been used as a reference to define the digital identity of Tetanurae as II–IV. The new concept of evolutionary teratology states that certain anatomical structures identified in evolutionary lineages are viable developmental anomalies (‘adaptive’ or not), becoming part of the considered groups. The features of Limusaurus' forelimb match teratological characterization. This diagnosis, associated with the variations previously identified in derived Ceratosauria taxa (Carnotaurinae), underline an anatomical and developmental independence (regarding evolutionary conserved mechanisms) compared with Tetanurae and therefore birds. Consequently, Limusaurus should not be used as a reference concerning the identity of avian manuals digits. Evolutionary teratology supports identities I, II and III of the tetanuran manus via a frame‐shift that did not occur in the Ceratosauria lineage. © 2016 The Linnean Society of London     

Large sauropod and theropod tracks from the Middle Jurassic Chuanjie Formation of Lufeng County,Yunnan Province and palaeobiogeography of the Middle Jurassic sauropod tracks from southwestern China

Tracks of large theropods and a single sauropod footprint are reported from red beds at Beikeshan locality in the Middle Jurassic Chuanjie Formation, of Lufeng County, near the large World Dinosaur Valley Park complex. The Chuanjie theropod tracks are assigned to the ichnogenus Eubrontes and the large sauropod track is given the provisional label Brontopodus. All occur as isolated tracks, i.e., trackways are not preserved. Saurischian dominated ichnofaunas are relatively common in the Jurassic of China. The producers of the Chuanjie tracks may have been similar to the basal tetanuran theropod Shidaisaurus and to mamenchisaurid sauropods, which were widely distributed throughout China, during the Jurassic, and are known from skeletal remains found in the same unit. Other potential sauropod trackmakers include titanosauriforms or as-yet-unknown basal eusauropods. The ichno- and skeletal records from the Jurassic of the Lufeng Basin are largely consistent, and both document the presence of middle-large sized theropods and sauropods.     

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.     

Cranial anatomy and systematics of Hypacrosaurus altispinus,and a comparative analysis of skull growth in lambeosaurine hadrosaurids (Dinosauria: Ornithischia)

The cranial anatomy of the helmet‐crested lambeosaurine Hypacrosaurus altispinus (Ornithischia: Hadrosauridae) is described, with a focus on ontogenetic and individual variation in phylogenetically significant characters of the cranial crest, braincase, and facial skeleton. Cranial material of H. altispinus represents a relatively complete growth series that includes crestless juveniles of less than half the size of large individuals with fully developed crests. Cranial ontogeny is compared with other lambeosaurines using bivariate morphometrics and through qualitative comparison of a size‐standardized cranial growth series. Bivariate analyses reveal that the relative growth of the skull and cranial crest of H. altispinus and H. stebingeri are similar, and that Hypacrosaurus more closely resembles Corythosaurus than Lambeosaurus. Hypacrosaurus altispinus is systematically revised. The taxon is characterized by five autapomorphies, most of which are concentrated in the skull, as well as an enlarged terminal ischial foot. Maximum parsimony and Bayesian likelihood (Mk+gamma) phylogenetic analyses were conducted to test the monophyly of the genus. Hypacrosaurus monophyly is corroborated in light of new anatomical data. Although H. stebingeri and H. altispinus share few derived characters of the skull, the hypothesis that H. stebingeri is a metaspecies that represents the ancestor of H. altispinus cannot be rejected. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 398–434.     

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。     

The early evolution of titanosauriform sauropod dinosaurs

Titanosauriformes was a globally distributed, long‐lived clade of dinosaurs that contains both the largest and smallest known sauropods. These common and diverse megaherbivores evolved a suite of cranial and locomotory specializations perhaps related to their near‐ubiquity in Mesozoic ecosystems. In an effort to understand the phylogenetic relationships of their early (Late Jurassic–Early Cretaceous) members, this paper presents a lower‐level cladistic analysis of basal titanosauriforms in which 25 ingroup and three outgroup taxa were scored for 119 characters. Analysis of these characters resulted in the recovery of three main clades: Brachiosauridae, a cosmopolitan mix of Late Jurassic and Early Cretaceous sauropods, Euhelopodidae, a clade of mid‐Cretaceous East Asian sauropods, and Titanosauria, a large Cretaceous clade made up of mostly Gondwanan genera. Several putative brachiosaurids were instead found to represent non‐titanosauriforms or more derived taxa, and no support for a Laurasia‐wide clade of titanosauriforms was found. This analysis establishes robust synapomorphies for many titanosauriform subclades. A re‐evaluation of the phylogenetic affinities of fragmentary taxa based on these synapomorphies found no body fossil evidence for titanosaurs before the middle Cretaceous (Aptian), in contrast to previous reports of Middle and Late Jurassic forms. Purported titanosaur track‐ways from the Middle Jurassic either indicate a substantial ghost lineage for the group or – more likely – represent non‐titanosaurs. Titanosauriform palaeobiogeographical history is the result of several factors including differential extinction and dispersal. This study provides a foundation for future study of basal titanosauriform phylogeny and the origins of Titanosauria. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 624–671.     

Pterosaur and dinosaur remains from the Middle Jurassic Balabansai Svita in the northern Fergana Depression, Kyrgyzstan (Central Asia)

Abstract: Isolated pterosaur and dinosaur teeth and a sauropod metatarsal I and manual phalanx V-1 from the Middle Jurassic (Callovian) Balabansai Svita in the northern Fergana Valley, Kyrgyzstan, are described and attributed to the pterosaur taxon Rhamphorhynchinae indet., a theropod Tetanurae indet., a sauropod Neosauropoda indet., and a new pachycephalosaurid Ferganocephale adenticulatum gen. et sp. nov. The Balabansai theropod is possibly a stem-lineage representative of Dromaeosauridae. The new pachycephalosaurid is the oldest representative of the group and extends its known history by 10–20 myr. The Balabansai vertebrate assemblage is most similar to the Callovian assemblages from the Qigu and Upper Shaximiao formations in China, and intermediate in the evolutionary level of the taxa present between the Bathonian assemblages from Wucaiwan and the Lower Shaximiao formations (China) and the Late Jurassic Shar Teg fauna from Mongolia.     

The plagiosaurid temnospondyl Plagiosuchus pustuliferus (Amphibia: Temnospondyli) from the Middle Triassic of Germany: anatomy and functional morphology of the skull

The cranial anatomy of the plagiosaurid temnospondyl Plagiosuchus pustuliferus, from the Middle Triassic of Germany, is described in detail on the basis of a newly discovered skull and mandibular material. The highly derived skull is characterized by huge orbitotemporal fenestrae, a reduction of the circumorbital bones – the prefrontal, postfrontal and (probably) postorbital are lost – and the expansion of the jugal to occupy most of the lateral skull margin. Ventrally the extremely long subtemporal vacuities correlate with the elongate adductor fossa of the mandible. The dentition is feebly developed on both skull and mandible. Ossified ?ceratobranchials and ‘branchial denticles’ indicate the presence of open gills clefts in life. The remarkably divergent cranial morphology of P. pustuliferus highlights the extraordinary cranial diversity within the Plagiosauridae, probably unsurpassed within the Temnospondyli. Specific structural aspects of the skull – including an extremely short marginal tooth row, feeble dentition and an elongated chamber for adductor musculature – together with evidence for a hyobranchial skeleton, suggests that P. pustuliferus utilized directed suction feeding for prey capture. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 348–373.     

Comparative ossification and development of the skull in palaeognathous birds (Aves: Palaeognathae)

Ratites and tinamous are a morphologically diverse group of flightless and weakly flighted birds. As one of the most basal clades of extant birds, they are frequently used as an outgroup for studies discussing character evolution within other avian orders. Their skeletal development is not well known in spite of their important phylogenetic position, and studies have historically been plagued with small sample sizes and limited anatomical and temporal scope. Here, I describe the ossification of the skull in the emu (Dromaius novaehollandiae), ostrich (Struthio camelus), greater rhea (Rhea americana), and elegant crested‐tinamou (Eudromia elegans). Skeletal development is remarkably consistent within palaeognaths, in spite of large differences in absolute size and incubation period. Adult morphology appears to play a role in interordinal differences in the sequence and timing of ossification of certain bones. Neither the timing of cranial ossification events relative to stage nor the sequence of ossification events provides any evidence in support of a paedomorphic origin of the palaeognathous palate. This study provides an important first look at the timing and sequence of skull development in palaeognathous birds, providing data that can be compared to better‐studied avian systems in order to polarize ontogenetic characters. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 184–200.     

A review of theropod dinosaurs from the Late Jurassic to mid-Cretaceous of Southeast Asia

Several non-avian theropod dinosaurs, as well as some Mesozoic birds, have been reported from Southeast Asia. The fossils are dominantly found in northeastern Thailand, however, one bizarre theropod has been described from Laos, one theropod has been reported from Malaysia, and some avian and non-avian theropods have been recently reported from Myanmar. The temporal distribution of Southeast Asian theropods ranges from the Late Jurassic to the mid-Cretaceous. All non-avian theropod faunas from Southeast Asia consist of non-maniraptoran tetanurans. They show similarity to Chinese plus Japanese theropods during the Early Cretaceous in broad systematic terms. During this time, megaraptorans can be found only in Japan, Australia, Brazil, and possibly Thailand, whereas tyrannosauroids can be found in China, Europe, possibly Brazil and Australia. Spinosaurids, carcharodontosaurians, and some coelurosaurs such as ornithomimosaurs were almost cosmopolitan. Metriacanthosaurids, on the other hand, were endemic to Europe and Asia including China and Thailand during the Middle to Late Jurassic.     

Theropod and possible ornithopod track assemblages from the Jurassic–Cretaceous boundary Houcheng Formation,Shangyi, northern Hebei,China

Dinosaur track assemblages from the Houcheng Formation in the small continental Shangyi Basin of northern Hebei Province, China bridge a gap in the record of vertebrates from this unit and enrich our knowledge of ichnofaunas from the Jurassic–Cretaceous boundary. Their stratigraphic position between the Middle Jurassic Yan-Liao Biota and the Lower Cretaceous Jehol Biota gives them a special importance. New discoveries allow a re-assessment of theropod and possible ornithopod tracks that are present with several trackways. Seventy-three footprints were examined and documented. Despite their smaller size, the tridactyl mesaxonic theropod tracks show morphological similarities with the ichnogenus Therangospodus known from the Upper Jurassic deposits of North America, Europe, and Central Asia. The possible ornithopod tracks lack an associated manus imprint, suggesting a bipedal trackmaker. These possible ornithopod tracks from the Houcheng Formation provide evidence for the presence of small basal ornithopods or basal Cerapoda in the Upper Jurassic–Lower Cretaceous in this region. The depositional environment was the margin of an extensive shallow lake with fluctuating water levels under seasonally dry climate.