A new basal snake from the mid-Cretaceous of Morocco

Fossil snakes are relatively well represented in the Upper Cretaceous of northern Africa, with material known from Morocco, Sudan, Egypt, Libya, Algeria, and Niger. The Moroccan Kem Kem beds have yielded a particularly diverse snake assemblage, with Simoliophiidae, Madtsoiidae, ?Nigerophiidae and several unnamed taxa co-occurring. These fossils are important for our understanding of the early evolutionary history of snakes, and may shed light on the ecology and initial diversification of basal snakes. We describe a new taxon, Norisophis begaa gen. et sp. nov., from the Kem Kem beds of Begaa, in southeast Morocco. It is characterised by a marked interzygapophyseal constriction, parazygantral foramina, an incipient prezygapophyseal process, and an anterio-posteriorly short centrum. Several characteristics shared with Najash, Seismophis, Madtsoiidae, and Coniophis suggest that Norisophis is a stem ophidian. N. begaa further increases the diversity and disparity of snakes within the Kem Kem beds, supporting the hypothesis that Africa was a mid-Cretaceous hotspot for snake diversity.     

An unusual modification of the jaws in cf. Alanqa,a mid-Cretaceous azhdarchid pterosaur from the Kem Kem beds of Morocco

A jaw bone attributed to the mid-Cretaceous azhdarchid pterosaur Alanqa saharica Ibrahim et al., 2010 is described. The new specimen from the Kem Kem beds of south eastern Morocco is unusual for the presence of expanded rostral bones that protrude above the occlusal surface of the jaws. The function of this unusual osteological modification is uncertain, but it is considered likely that it served a role in food processing, or perhaps jaw stability.     

Kazaklambia convincens comb. nov., a primitive juvenile lambeosaurine from the Santonian of Kazakhstan

The holotype of ‘Procheneosaurus’ convincens, a juvenile lambeosaurine, is the most complete hadrosaurid known from Kazakhstan. North American species of Procheneosaurus are considered juveniles of Corythosaurus, Hypacrosaurus, and Lambeosaurus, rendering the generic name unusable. A replacement name, Kazaklambia convincens comb. nov., is herein proposed as this specimen is morphologically distinct from other Eurasian taxa and known juvenile lambeosaurines at a similar ontogenetic stage in having a prefrontal process of the postorbital with a dorsal thickening forming a dome lateral to the frontal dome, doming of the nasal anterodorsal to the orbit, and a frontal length/width ratio <1. The juvenile status of Kazaklambia makes phylogenetic placement difficult; however, morphometric and morphological information (particularly in relation to the hollow cranial crest and the length of the frontal) suggest a close affiliation with the basal lambeosaurines Amurosaurus and Tsintaosaurus, and support the hypothesis for an Asian origin for Lambeosaurinae.     

Distinctive azhdarchoid pterosaur jaws from the mid-Cretaceous Cambridge Greensand of eastern England and the Kem Kem Group of Morocco

An isolated jaw fragment from the Late Cretaceous (Cenomanian) Cambridge Greensand Member of the West Melbury Marly Chalk Formation previously identified as a cestraciontid shark fin spine is referred to the pterosaur clade Azhdarchoidea on account of its lateral and occlusal foramina and edentuly. The specimen differs from the azhdarchoid Ornithostoma sedgwicki from the same deposit in having flat lateral surfaces and an acute dorsal/ventral apex. The specimen is similar in overall morphology to CAMSM B40085 from the same horizon and probably represents the corresponding jaw but from a different individual. Likely these specimens represent a new taxon but are considered too fragmentary to diagnose at present. A remarkably similar and distinctive morphology is found in unnamed pterosaur jaws from the Kem Kem Group (?Albian-Cenomanian) of Morocco, supporting the idea of faunal similarity between these two distant localities.     

Overabundance of piscivorous dinosaurs (Theropoda: Spinosauridae) in the mid-Cretaceous of North Africa: The Algerian dilemma

The informally called ‘Continental intercalaire’ is a series of continental and brackish deposits that outcrops in several regions of North Africa. The age of the series is not well-constrained, but its upper part, visible in the ‘Kem Kem beds’ in Morocco and in Bahariya in Egypt, is regarded as early Cenomanian in age. Spinosaurid remains are an important component of this series, but records of this dinosaur are surprisingly rare in Algerian localities of the ‘Continental intercalaire’. Here, we describe a vertebrate assemblage from two localities, Kénadsa and Menaguir, situated in the Guir basin, Western Algeria. The assemblage comprises hybodont sharks, sarcopterygian fishes, ray-finned fishes, turtles, crocodiles and dinosaurs. Among the latter, only teeth of theropods have been recovered and 94% belong to Spinosaurus. The assemblage is taxonomically very similar to the Moroccan and Egyptian assemblages mentioned above. This study: 1) suggests a likely early Cenomanian age for the Guir basin deposits containing the assemblage; 2) provides a new evidence of the homogeneity of the early Cenomanian vertebrate fauna throughout North Africa; and 3) confirms the overabundance of theropod dinosaurs, especially spinosaurs, in the assemblage showing a possible shortcut in the vertebrate food chain. The northern most locality, Menaguir, shows sedimentological and ichnological evidence of marine influences indicating that the palaeoenvironment shows spatial heterogeneities.     

Additions to the sauropod dinosaur fauna of the Cenomanian (early Late Cretaceous) Kem Kem beds of Morocco: Palaeobiogeographical implications of the mid-Cretaceous African sauropod fossil record

Determining the relationships of mid-Late Cretaceous African taxa is central to understanding the timing and resultant palaeobiogeographical patterns of Gondwanan fragmentation. The early Late Cretaceous (Cenomanian) Kem Kem beds of southeastern Morocco preserve a diverse vertebrate fauna, including sauropod dinosaurs. Sauropod material includes the holotype of the rebbachisaurid diplodocoid Rebbachisaurus garasbae and fragmentary remains representing indeterminate titanosauriforms and rebbachisaurids. Here, we describe two new specimens from the Kem Kem beds. A dorsal neural arch with complex internal pneumaticity is tentatively attributed to a somphospondylan titanosauriform. A caudal vertebra possessing several rebbachisaurid synapomorphies is excavated by a large and pervasive lateral pneumatic foramen, a feature undocumented in other rebbachisaurids. However, caudal vertebrae are currently unknown for the sympatric R. garasbae, so this element could be referable to that taxon or a second, previously unknown, rebbachisaurid species. Interestingly, this new caudal vertebra displays a mosaic of features otherwise restricted to limaysaurine or nigersaurine rebbachisaurids, suggesting a placement basal to these clades, which is the position usually recovered for R. garasbae in phylogenetic analyses. A review of the mid-Cretaceous African sauropod fossil record removes the Cretaceous record of dicraeosaurids from Africa, restricting this clade to a single post-Jurassic occurrence in Argentina. All diagnostic sauropod remains can be attributed to titanosauriforms or rebbachisaurids. Whereas rebbachisaurids were seemingly restricted to northwestern Africa and disappeared post-Cenomanian, titanosauriforms were widespread across the African continent and survived until the latest Cretaceous. The development of the mid-Cretaceous Trans-Saharan Seaway might have acted as a dispersal barrier for rebbachisaurids and other vertebrate groups. In contrast, titanosauriforms might have been able to cross this barrier, but it is possible that they were also unable to disperse, and that northwestern African titanosauriforms were not closely related to taxa from the rest of the African continent. New materials and a better understanding of titanosaur interrelationships will be crucial in teasing these scenarios apart.     

中国早白垩世热河生物群中大型霸王龙类化石的发现

一件包括部分头骨、下颌、肠骨等的不完整兽脚类恐龙骨架发现于辽宁喀左早白垩世九佛堂组中,估计身体全长可达9~10m。其前颌骨主体部分高、肠骨外侧面具显著的直立的嵴等特征表明该化石应归霸王龙类。它以外鼻孔大、上颌骨背缘前部略内凹、上颌骨孔向前达眶前窝前缘、腹缘与眶前窝腹缘存在较大距离等特征不同于晚白垩世的霸王龙科分子,故被命名为一新属种——喀左中国暴龙(Sinotyrannus kazuoensis gen. et sp. nov.),或许代表了最早的霸王龙科类型。该属种是已知个体最大的前晚白垩世霸王龙类,也是辽西及周边地区热河生物群中个体最大的兽脚类恐龙。它的发现不仅表明东亚是霸王龙类最主要的演化地区之一,而且为探讨霸王龙科的起源、热河生物群的组成与生态系统等提供了重要依据。     

A new tyrannosaurine specimen (Theropoda: Tyrannosauroidea) with insect borings from the Upper Cretaceous Honglishan Formation of Northwestern China

A large theropod ilium was recently collected from the Upper Cretaceous Honglishan Formation in the Sangequan area of the northern Junggar Basin, Xinjiang, China, which represents the first officially reported dinosaur fossil from this formation. Several morphological features, including robust supracetabular ridge, reduced supracetabular crest, concave anterior margin of the pubic peduncle, ventral flange on the pubic peduncle, converging dorsal surface of the iliac blades, laterally visible cuppedicus shelf, and ventral flange on the posterior surface of pubic peduncle, suggest that this specimen can be referred to Tyrannosaurinae, and furthermore, a few differences between this specimen and other tyrannosaurines in particular the contemporary Asian tyrannosaurine Tarbosaurus suggest that IVPP V22757 may represent a new tyrannosaurine species. However, in the absence of extensive data that would make it possible to properly evaluate these differences, we refrain from naming a new taxon based on this specimen. Some insect borings are also identified in this specimen, and are referable to the ichnogenus Cubiculum, which is interpreted as the insect pupichnia. This new fossil documents the presence of a gigantic theropod in the Upper Cretaceous of Junggar Basin, adding new information on its poorly studied ecosystems.     

New Cenomanian florule and a leaf mine from southeastern Morocco: Palaeoecological and climatological inferences

The recently found Gara Sbaa florule of southeastern Morocco comes from distinctive carbonate lamellites with abundant fish fauna above the terrestrial/paralic Kem Kem sequence and correlated with the basal horizon of the open shelf carbonate Akrabou Formation. It is a “mixed” assemblage of ferns and gymnosperms of “Wealden aspect” associated with relatively advanced angiosperms, suggesting floristic exchanges between the insular land masses of northern Africa and southern Europe. The climatic conditions are inferred to have been similar to the mildly dry subtropical climate of the western Canary Islands. New taxa of aquatic angiosperms Garasbahia fexuosa Krassilov et Bacchia gen. et sp nov. and a leaf mine Troponoma constricta Krassilov, sp. nov. are described.     

A New Basal Sauropod Dinosaur from the Lufeng Basin,Yunnan Province, Southwestern China

<正>A new dinosaur Chuxiongosaurus lufengensis gen.et sp.nov.is erected based on a nearly complete skull.The taxon is characterized by the lacrimal perpendicular to the ventral margin of the upper jaw,which is similar to that of Thecodontosaurus;a depression present on the dorsal profile of the snout behind the naris;the rostral profile of the maxilla slopes continuously towards the rostral tip;and the presence of 25 dentary teeth.It also displays prosauropod characters such as a relatively long skull,the slope of the maxillary rostral profile,and teeth that do not have basically constricted crowns.The new specimen is more basal than Anchisaurus and represents the first basal sauropod dinosaur from the Early Jurassic of China.     

Distribution of the dentary groove of theropod dinosaurs: Implications for theropod phylogeny and the validity of the genus Nanotyrannus Bakker et al., 1988

This study examines the phylogenetic distribution of a morphologic character, described as a groove containing pores, on the lateral surface of the dentary bone in theropod dinosaurs. The nature of this groove is a feature unique to theropods. Of the 92 theropod taxa examined for the presence and absence of this feature, 48 possessed and 44 lacked this feature. Distribution of this character was compared to published phylogenetic analyses of theropods, in order to evaluate the utility of the dentary groove as a diagnostic feature. 80% of pre-Tyrannoraptoran theropods possessed the dentary groove, with only 6 reversals in basal theropod clades. Theropods with beaks or edentulous jaws all lacked a dentary groove. Tyrannosauroidea is marked by mosaic distribution of this character. Among tyrannosauroids, the dentary groove occurs only in Dryptosaurus and the Albertosaurinae (Albertosaurus + Gorgosaurus). Nanotyrannus lancensis, sometimes described as representing juvenile Tyrannosaurus rex, also possesses this groove, unlike the remainder of the Tyrannosaurinae. Nanotyrannus lancensis was included in a phylogenetic analysis of Tyrannosauroidea and was recovered within Albertosaurinae. We recommend that Nanotyrannus stand as a valid taxon nested within the Albertosaurinae, based on the presence of this groove, as well as other features of the skull.     

A new, large tyrannosaurine theropod from the Upper Cretaceous of China

Tyrannosaurids are primarily gigantic, predatory theropod dinosaurs of the Cretaceous. Here we report a new member of the tyrannosaurid clade Tyrannosaurinae from the Upper Cretaceous Wangshi Group of Zhucheng, Shandong Province, China, based on a maxilla and associated dentary. The discovery of this animal, here named Zhuchengtyrannus magnus gen. et sp. nov., adds to the known diversity of tyrannosaurids in Asia. Z. magnus can be identified by a horizontal shelf on the lateral surface of the base of the ascending process, and a rounded notch in the anterior margin of the maxillary fenestra. Several additional features contribute to a unique combination of character states that serves to further distinguish Z. magnus from other taxa. Comparisons with other tyrannosaurids suggest that Zhuchengtyrannus was a very large theropod, comparable in size to both Tarbosaurus and Tyrannosaurus.     

CT scanning, rapid prototyping and re-examination of a partial skull of a basal crocodylomorph from the Late Triassic of Germany

An incomplete skull (SMNS 12352) from the Norian (Late Triassic), found at the same locality (Pfaffenhofen, Germany) as Procompsognathus triassicus, has previously been attributed to either a theropod or a crocodylomorph. The specimen is partially embedded in matrix and, therefore, was CT scanned so that the concealed portion could be visualized and a 3D model could be printed by means of rapid prototyping. Mainly based on the separation between the nasal and the antorbital fossa, the skull likely pertains to an indeterminate basal crocodylomorph (non-Crocodyliformes), which is, however, distinct from Saltoposuchus connectens.     

A New Carcharodontosauria (Theropoda) from the Early Cretaceous of Guangxi, Southern China

A new large-bodied theropod dinosaur,Datanglong guangxiensis gen.et sp.nov.,was recovered from the Lower Cretaceous Xinlong Formation of the Datang Basin,Guangxi.It is unique in several features including:posteriormost dorsal vertebra with teardrop-shaped pneumatic foramen confined by enlarged pcdl,acpl and the centrum; posteriormost dorsal with well-developed,horizontal prpl; posteriormost dorsal with a parapophysis projecting more laterally than the diapophysis; brevis fossa shallow with short,ridge-like medial blade; and iliac pubic peduncle with posteroventrally expanded margin.Cladistic analysis supports the idea that this new taxon Datanglong guangxiensis is a primitive member of the Carcharodontosauria in possessing two unambiguous synapomorphies:large external pneumatic foramina and internal spaces present in the lateral surface of ilium,and a peg-and-socket ischiac articulation with the ilium.The presence of the new taxon from Guangxi further confirms that Carcharodontosauria were cosmopolitan large-bodied predators during the Early–mid Cretaceous.     

The longest theropod trackway from East Asia,and a diverse sauropod-, theropod-, and ornithopod-track assemblage from the Lower Cretaceous Jiaguan Formation,southwest China

Here we report a large dinosaur tracksite from an extensive fluvial sandstone surface in the Lower Cretaceous Jiaguan Formation of Sichuan Province, China. The site contains over 250 individual tracks comprising at least 18 recognizable trackways, including the longest theropod trackway (cf. Eubrontes) known from China. This exceptional theropod trackway consists of 81 successive footprints covering a distance of 69 m. The tracks are well-preserved and are expressed both as true tracks on the main “upper” surface and as transmitted undertracks on a locally exposed “lower” bed. Also recorded are six other theropod trackways, including small Grallator-like ichnites, eight sauropod trackways (cf. Brontopodus), and three small ornithopod (cf. Ornithopodichnus) trackways with a parallel orientation, which may indicate gregarious behavior. Several trackways of a larger theropod trackmaker show pes imprints with elongated traces of the metatarsals, suggesting extramorphological (substrate-controlled) variation and/or plantigrade posture, which is here interpreted as indicating a change in gait assumed in response to deep and soft sediment. The assemblage indicates a diverse dinosaur fauna in the Lower Cretaceous Sichuan Basin with variously sized theropods, sauropods, and ornithopods. The late occurrence of footprints of the Grallator-Eubrontes plexus in Lower Cretaceous strata is further evidence of the extended stratigraphic range of this morphotype and the distinct palaeobiogeographic distribution of these trackmakers in East Asia.     

A New Triassic Primitive Ichthyosaur from Yuanan, South China

Based on the systematic study of two fossil skeleton specimens collected from the top of the third member of the marine Lower Triassic Jialingjiang Formation of Yuanan, Hubei Province, South China, a new Early Triassic primitive ichthyosaur Chaohusaurus zhangjiawanensis sp. nov. is reported and described. The beds yielding the type material are correlated with the Neospathodus homeri-N. triangularis Conodont zone. The new taxon is most similar to Chaohusaurus geishanensis Young and Dong, 1972 in the shape and configuration of the scapula, forefin, pelvic girdle and hindfin, presacral vertebral count and well-developed caudal peak, but distinguished from the latter by its larger size, the position of the pineal foramen in the centre of the parietal, the occurrence of a larger calcaneum in hindfin and the first sacral rib with distal expansion. The new species exhibits common features of primitive ichthyosaurs such as: (1) the prefrontal has an antero-dorsal shelf projecting towards the orbit; (2) the upper temporal fenestra is small; (3) the postorbital and the squamosal meet laterally to the upper temporal fenestra; and (4) cylindrical centra. However, more derived ichthyosaur characters are seen with the frontal separated from the orbital dorsal margin by the contact of the prefrontal and postfrontal, which offer new clues to the early radiation of ichthyosaurs.     

A diversified vertebrate ichnite fauna from the Feitianshan Formation (Lower Cretaceous) of southwestern Sichuan,China

The Bajiu tracksite exposed on a fine sandstone surface at the top of the Feitianshan Formation, close the contact with the overlying Xiaoba Formation, reveals a saurischian dominated ichnofauna consisting of four sauropod and at least eight theropod trackways. The sauropod trackways display a medium-wide gauge pattern characteristic of titanosauriforms, while the theropod trackways can be subdivided into two distinct morphotypes. One is similar to the ichnogenera Eubrontes and Megalosauripus, the other (cf. Dromaeopodus) appears to represent a functionally didactyl dromaeosaur where pedal digit II is represented only by an oval basal pad. A single swim track is possibly attributable to a turtle or crocodylian. Such sauropod-theropod dominated ichnofaunas are consistently typical of the red-bed tracksites in the region and useful for characterizing a fauna that is otherwise poorly known from body fossils. The stratigraphic position and late occurrence of the (typical Jurassic) Eubrontes-Megalosauripus morphotype is well known from other Cretaceous localities in China and seems to reflect a peculiarity in the theropod faunas of East Asia.     

Troodontidae (Dinosauria: Theropoda) from the Upper Cretaceous of Uzbekistan

To date three taxa of troodontid theropod dinosaurs have been recognized from Upper Cretaceous strata in two regions of the Kyzylkum Desert in Uzbekistan. The Cenomanian Khodzhakul Formation in the southwestern Kyzylkum Desert has yielded isolated serrated teeth and some postcranial bones of an indeterminate troodontid. In the central Kyzylkum Desert troodontids are known from the Cenomanian Dzharakuduk Formation (Urbacodon itemirensis) and the Turonian Bissekty Formation (Urbacodon sp.). Urbacodon itemirensis is known from a single dentary whereas Urbacodon sp. is represented by isolated teeth, maxilla and dentary fragments, a partial braincase, and some postcranial bones. The troodontid affinities of Urbacodon are supported by several synapomorphies: presence of a subotic recess; reduced basal tubera placed directly under the occipital condyle; maxilla participating in the margin of the external naris; nutrient foramina on dentary situated within a deep lateral groove; dentary without distinct interdental plates; large number of small dentary and maxillary teeth; teeth constricted between root and crown; anterior dentary teeth smaller, more numerous, more closely spaced than those in the middle of the tooth row, and implanted in a groove; posterior dorsal vertebrae with tall and posterodorsally tapering neural spines; and presence of a midline sulcus on the neural arches of distal caudals. Among Troodontidae, Urbacodon resembles Byronosaurus, Gobivenator, and Xixiasaurus in the absence of serrations on the tooth crowns and having premaxillary teeth that are D-shaped in cross-section. However, phylogenetic analysis did not recover a clade of Asiatic troodontids with unserrated teeth.     

Evolution of the carnivorous dinosaurs during the Cretaceous: The evidence from Patagonia

Patagonia has yielded the most comprehensive fossil record of Cretaceous theropods from Gondwana, consisting of 31 nominal species belonging to singleton taxa and six families: Abelisauridae, Noasauridae, Carcharodontosauridae, Megaraptoridae nov. fam., Alvarezsauridae, and Unenlagiidae. They provide anatomical information that allows improved interpretation of theropods discovered in other regions of Gondwana. Abelisauroids are the best represented theropods in Patagonia. They underwent an evolutionary radiation documented from the Early Cretaceous through to the latest Cretaceous, and are represented by the clades Abelisauridae and Noasauridae. Patagonian carcharodontosaurids are known from three taxa (Tyrannotitan, Giganotosaurus and Mapusaurus), as well as from isolated teeth, collected from Aptian to Cenomanian beds. These allosauroids constituted the top predators during the mid-Cretaceous, during which gigantic titanosaur sauropods were the largest herbivores. Megaraptorans have become better documented in recent years with the discovery of more complete remains. Megaraptor, Aerosteon and Orkoraptor have been described from Cretaceous beds from Argentina, and these taxa exhibit close relationships with the Aptian genera Australovenator, from Australia, and Fukuiraptor, from Japan. The Gondwanan megaraptorans are gathered into the new family Megaraptoridae, and the Asiatic Fukuiraptor is recovered as the immediate sister taxon of this clade. Although megaraptorans have been recently interpreted as members of Allosauroidea, we present evidence that they are deeply nested within Coelurosauria. Moreover, anatomical information supports Megaraptora as more closely related to the Asiamerican Tyrannosauridae than thought. Megaraptorans improve our knowledge about the scarcely documented basal radiation of Gondwanan coelurosaurs and tyrannosauroids as a whole. Information at hand indicates that South America was a cradle for the evolutionary radiation for different coelurosaurian lineages, including some basal forms (e.g., Bicentenaria, Aniksosaurus), megaraptorans, alvarezsaurids less derived than those of Laurasia, and unenlagiids, revealing that Gondwanan coelurosaurs played sharply differing ecological roles, and that they were taxonomically as diverse as in the northern continents. The unenlagiids represent an endemic South American clade that has been recently found to be more closely related to birds than to dromaeosaurid theropods. Analysis of the theropod fossil record from Gondwana shows the highest peak of origination index occurred during the Aptian–Albian and a less intense one in the Campanian time spans. Additionally, peaks of extinction index are recognized for the Cenomanian and Turonian–Coniacian time spans. In comparison, the Laurasian pattern differs from that of Gondwana in the presence of an older extinction event during the Aptian–Albian time-span and a high origination rate during the Cenomanian time-bin. Both Laurasian and Gondwanan theropod records show a peak of origination rates during the Campanian.