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.     

Minisauripus―the track of a diminutive dinosaur from the Cretaceous of China and South Korea: implications for stratigraphic correlation and theropod foot morphodynamics

The diminutive (2.5–3.0 cm long), Cretaceous dinosaur track ichnogenus Minisauripus, previously known only from the type ichnospecies, M. chuanzhuensis, from a single locality in Sichuan Province China, is here reported from two new localities in South Korea and one in China. Material from the new Chinese locality is assigned to the new ichnospecies M. zhenshuonani on the basis of its distinctive morphology. Most of the new material is well-preserved, revealing narrow asymmetric tracks with claw traces, long step and phalangeal formula (2-3-4 for digits II, III and IV, respectively), suggesting a theropod track maker rather than an ornithischian, as originally inferred for the Chinese type material.The South Korean samples (eight tracks), from two localities in the Haman Formation, are considered Early Cretaceous (Aptian-Albian) in age, whereas the Chinese type material (21 specimens) has been assigned both an Early and Late Cretaceous age. The former age is probably correct as suggested by a new Minisauripus locality (5 specimens) from the Early Cretaceous (Barremian-Albian) of Shandong Province, China.Other diminutive tracks from the Sichuan fauna include Aquatilavipes sinensis (2.5 cm long, a possible junior synonym of Koreanoris hamanensis), Grallator emeiensis (2 cm long) and Velociraptorichnus sichuanensis (10–11 cm), which occur, in various combinations, with Minisauripus at both the new Korean and Chinese localities.In Minisauripus, digit III is very short in comparison with other theropods and provides a striking contrast to G. emeinsis. This difference has significant implications for standard assumptions about theropod track allometry. Based on the classic Early Jurassic forms Grallator and Eubrontes, it has long been inferred that relative digit III length shrank with increasing size (up to foot lengths of 30–40 cm). The reiteration of reduction in relative length of digit III in specimens in the size range of 2–3 cm indicates that the allometric or morphodynamic ‘program’ that influenced development in large theropod clades reiterated fractally in theropod clades a full order of magnitude smaller. This shows that a given allometry can be size-dependent in one clade and size-independent in another. Thus, the developmental program appears ‘contracted’ or morphologically miniaturized by heterochrony to manifest paedomorphically in some clades and peramorphically in others. This strongly suggests that ‘formal’ developmental ‘programs operated’ along similar morphodynamic lines in quite different clades.     

Orkoraptor burkei nov. gen. et sp., a large theropod from the Maastrichtian Pari Aike Formation, Southern Patagonia, Argentina

A new large theropod, Orkoraptor burkei nov. gen. et nov. sp., from the Upper Cretaceous (Maastrichtian) Pari Aike Formation of southern Patagonia is based on a postorbital, quadratojugal, coronoid?, several teeth, an atlantal intercentrum and neurapophysis, two caudal vertebrae, and the proximal half of tibia. This new theropod exhibits characteristics of maniraptorans (i.e., coelurosaurians more derived than tyrannosaurids) including an upturned rostral process of the postorbital, the caudoventral corner of which is widely concave. Furthermore, teeth without denticles and carina in the mesial margin resemble the condition of the maniraptoran clades Compsognathidae and Deinonychosauria. The new Patagonian theropod differs from other coelurosaurians (Ornithomimosauria, Compsognathidae, Alvarezsauridae, Dromaeosauridae, Aves) in having caudal vertebrae with a single pair of small pleurocoels on each side, and in having a median depression flanked by two longitudinal and narrow furrows on each tooth. Orkoraptor represents one of the southernmost carnivorous dinosaurs yet found in South America and adds valuable information about the diversification of tetanuran theropods in Gondwana.     

First elaphrosaurine theropod dinosaur (Ceratosauria: Noasauridae) from Australia — A cervical vertebra from the Early Cretaceous of Victoria

Elaphrosaurinae is an enigmatic clade of gracile ceratosaurian theropod dinosaurs known from the Late Jurassic of Africa (Elaphrosaurus bambergi) and Asia (e.g., Limusaurus inextricabilis), and the early Late Cretaceous of Argentina (Huinculsaurus montesi). Elaphrosaurinae is often placed within Noasauridae as the sister taxon to Noasaurinae, a clade of small-bodied theropods that lived in South America, Africa, Madagascar and India throughout much of the Cretaceous. Herein, we report the first evidence of Elaphrosaurinae from Australia: a nearly complete middle cervical vertebra from the upper Lower Cretaceous (lower Albian) Eumeralla Formation of Cape Otway, Victoria, Australia. The fact that this site would have been situated at ~76°S towards the end of the Early Cretaceous (~110–107 Ma) implies that elaphrosaurines were capable of tolerating near-polar palaeoenvironments, whereas its age indicates that elaphrosaurines persisted in Australia until at least the late Early Cretaceous. The new Australian elaphrosaurine, in tandem with the recently described Huinculsaurus montesi from the Cenomanian–Turonian of Argentina, implies that the spatiotemporal distribution of Elaphrosaurinae has heretofore been greatly underestimated. Historic confusion of elaphrosaurines with coelurosaurs, especially ornithomimosaurs, coupled with our generally poor understanding of noasaurid evolution, might explain the apparent dearth of fossils of this theropod clade worldwide.     

黑龙江嘉荫地区晚白垩世兽脚类恐龙牙齿的发现及其意义

黑龙江嘉荫地区是中国最早发现恐龙化石的产地,在我国晚白垩世恐龙的研究史上具有极其重要的意义。该地区曾发掘出土大量鸭嘴龙类化石,是研究大型鸟脚类恐龙尤其是鸭嘴龙类系统分类和演化的理想地区。但相对应于鸭嘴龙类化石来说,兽脚类恐龙化石在该地区的发现却较为鲜见,主要以牙齿的形式出现。本文主要对最近在该地发现的一批兽脚类恐龙的牙齿化石进行详细描述对比,初步研究显示含有霸王龙类、驰龙类、白鲨齿龙、福井盗龙以及可能为新的兽脚类恐龙的牙齿。     

A theropod track assemblage including large deinonychosaur tracks from the Lower Cretaceous of Asia

A total of more than 40 tridactyl and didactyl tracks were preserved as natural casts on four fallen blocks of sandstone representing the Lower Cretaceous Jiaguan Formation of Gulin County in southeastern Sichuan Province, China. While several trackways can be distinctly followed, others are isolated imprints only. All have been flattened by overburden pressures. Tridactyl tracks are present with three size-classes being <10 cm, 10–20 cm and >20 cm in length. Morphologically they are similar to the ichnogenus Eubrontes, considering the relatively weak mesaxony. Eight of the tracks on one of the blocks are clearly didactyl and are here interpreted as representing large and medium sized dromaeosaurids. The largest track is about ∼30 cm long and comparable in size to the type of Dromaeopodus (∼28 cm), from the Lower Cretaceous of Shandong Province, which was the largest dromaeosaurid track previously reported. This report adds new data to the growing number of dromaeosaurid tracksites reports from China, and from the Jiaguan Formation, suggesting that this theropod group had a preference for fluvial paleoenvironments.     

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。     

Zurupleuropholis gen. nov. (Teleostei,Albian, Argentina), first pleuropholids from the Cretaceous of South America

Although the Cretaceous is characterized by a rich fish diversity, Cretaceous continental fishes from Gondwana are poorly known and comparatively scarce. Among these fishes, the family Pleuropholidae is only known by a few species relatively poorly preserved, from the Middle Jurassic to Lower Cretaceous of Europe, Africa, North America, and South America. In this paper, two new species of the pleuropholid new genus Zurupleuropholis are described, Z. quijadensis gen. et sp. nov. and Z. decollavi gen. et sp. nov. The new fishes were recovered in the Lower Cretaceous lacustrine Lagarcito Formation of central-west Argentina. This taxon constitutes a relevant finding considering that the representation of the family Pleuropholidae is rare worldwide. Zurupleuropholis gen. nov. appears to be the youngest known member of Pleuropholidae, and it represents the second record of the family in South America and the first record in the Cretaceous of the continent.     

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.     

Small-sized abelisaurid (Theropoda: Ceratosauria) remains from the Upper Cretaceous of northwest Patagonia,Argentina

A new abelisaurid theropod from the Cenomanian of Candeleros Formation (Neuquén Province, NW Patagonia, Argentina) is described. It includes an isolated frontal, partial pelvis, two fused sacral centra, femoral shaft, rib fragments, partial metatarsal II and shaft of metatarsal III. A paleohistological analysis shows that MMCh-PV 69 was somatically mature at time of death. The prevalence of relatively ordered intrinsic fibers in the primary bone indicates a relatively lower growth rate of MMCh-PV 69 compared with other abelisauroids, such as Aucasaurus garridoi. Phylogenetic analysis found MMCh-PV 69 as a basal abelisaurid, sister group of the node formed by Majungasaurinae and Brachyrostra. We estimated the body mass of MMCh-PV 69 in 240 kg, which makes it one of the smallest abelisaurids recorded. This new form adds to the extensive theropod fauna of the Candeleros Formation, probably the most diverse association of meat-eating dinosaurs recorded in a Cretaceous Formation from South America.     

First occurrence of tyrannosaurid theropods from the Corral de Enmedio Formation (Upper Cretaceous) Sonora,México

The Cabullona Basin in the state of Sonora, Mexico is becoming recognized due to its diversity of southern Laramidian continental vertebrates, especially dinosaurs. In this study we describe and analyze three theropod teeth (ERNO specimens) that were found isolated and surface collected in the Corral de Enmedio Formation (Cabullona Group, Upper Cretaceous). The three specimens possess similar morphological characteristics that match the ones present in Late Cretaceous Laramidian tyrannosaurids, so they were referred to the Tyrannosauridae, probably belonging to a new unknown taxon. The implementation of statistical and cladistic analyses corroborated their taxonomical assignment. ERNO specimens correspond to the first record of tyrannosaurid dinosaurs in the basal Corral de Enmedio Formation, extending the stratigraphic distribution of these dinosaurs in the Cabullona Basin. Although tyrannosaurids have been previously described in the Cabullona Basin, the ERNO specimens of the Corral de Enmedio Formation seem to be different, because they possess more labiolingually compressed teeth. This new evidence could indicate a higher taxonomic diversity of the tyrannosaurid theropods that were present in the Cabullona Basin, adding more information to the Tyrannosauridae diversification on one of the most southern Laramidian regions during the Late Cretaceous.     

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.     

A tyrannosaur trackway at Glenrock,Lance Formation (Maastrichtian), Wyoming

During the Campanian and Maastrichtian ages (86–66 million years ago), tyrannosaurids were the predominant large carnivorous dinosaurs throughout the Northern hemisphere. Despite the abundance of skeletal material, the fossil-footprint record of tyrannosaurids has been limited. Here we report a tyrannosaurid trackway in the Lance Formation, Wyoming. The trackway consists of three sequential tracks on a sandstone surface. Based on the age and size of the footprints, the trackmaker can be identified as either a sub-adult Tyrannosaurus rex or a Nanotyrannus lancensis. The trackway offers a record of a tyrannosaurid pace length, which permits the speed of the trackmaker to be calculated at 4.5–8.0 km/h. This result discounts previous speculation that tyrannosaurid walking speeds were notably slower than those of other large theropods.     

白垩纪四足动物足印的生物地层学、生物年代学与遗迹相

从全球范围来看,白垩纪四足动物的足印多数是非鸟恐龙与鸟类留下的痕迹;少量足印来自翼龙、鳄鱼、龟、哺乳动物和其他四足动物。白垩纪的足迹化石以东亚(尤其是中国和朝鲜)和北美西部的最为人所知。南美(主要是阿根廷和巴西)也有一定数量广泛分布的足迹化石,欧洲、非洲与澳大利亚的白垩纪足迹组合则鲜为人知。以白垩纪四足动物的足印记录为基础,我们对两个全球足印生物年代重新进行了检查。早白垩世生物年代以蜥脚类与鸟脚类的足迹为特征。晚白垩世生物年代中的蜥脚类足迹较少,但是鸭嘴龙、暴龙和角龙的足迹增多了。另外,白垩纪足印化石的记录中记载了许多重要的生物地层学信息,如北美白垩纪中期蜥脚类恐龙的消失,以及白垩纪末恐龙的绝灭。越来越多来自东亚的白垩纪足印记录使我们对更精细的地方性白垩纪足印生物年代学有了初步印象。因此,以地方性四足恐龙(包括鸟类)遗迹属的地层分布为基础,可以识别出三个或四个足印生物年代。种类丰富并具有地方性特色的东亚的白垩纪鸟类动物的遗迹群,可能指示白垩纪时东亚存在着一个独特而繁盛的鸟类动物群。以足印化石为基础的这一假说有待进一步的验证。     

Contribution to knowledge of the last dinosaur footprints in Europe. Persistence of ornithopods in the upper Maastrichtian of SE Spain

In the locality of Rambla de los Gavilanes (Betic Chain Murcia, Spain), six new outcrops with dinosaur footprints are reported from an uppermost Cretaceous succession of coastal marine deposits. Four of these sites are found in the upper part of the lower Maastrichtian and the other two in the upper Maastrichtian, these latter located only a few meters below the Cretaceous-Paleocene boundary (K/Pg). Traces appear as impressions or as natural casts, on tabular limestone beds consisting of micritic carbonate facies, these deposited in coastal marine areas that include a variety of environments (coastal lakes, tidal flats, salt marshes, and shallow coastal lakes). Two main types of traces have been recognized, respectively attributed to sauropod footprints and large ornithopod footprints with remarkable pointed hoof marks. The Maastrichtian sites with large ornithopod footprints in Europe are reported from Spain, Poland and Romania. Sauropod (titanosaurian), theropod (Irenesauripus and other undetermined theropods), and ornithopod footprints have been found at sites located up to 20 m below the K/Pg boundary. The Spanish dinosaur ichnites known closest to the K/Pg boundary are in the Pyrenean upper Maastrichtian, and are tridactyl, theropod and ornithopod (Hadrosauropodus and Amblydactylus) footprints. In this work, the assignation of the Rambla de los Gavilanes ornithopod footprints has been made following the recommended synthesis in several recent papers which reduce the large ornithopod ichnogenus to four.     

Diversity of theropod ootaxa and its implications for the latest Cretaceous dinosaur turnover in southwestern Europe

The scarcity of diagnostic skeletal elements in the latest Cretaceous theropod record of the Ibero-Armorican domain (southwestern Europe) prevents to perform accurate phylogenetic, paleobiogeographic, and diversity studies. In contrast, eggs and eggshells of theropod dinosaurs are relatively abundant and well known in this region from which several ootaxa have been described. Here, we describe the first Late Maastrichtian theropod ootaxon (Prismatoolithus trempii oosp. nov.) from SW Europe and demonstrate that oological record can be used as a proxy for assessing diversity of egg-producers and may help to complement their scarce bone record. The performed analyses indicate that the theropod taxa and ootaxa reach their diversity maxima during the Late Campanian and start to decrease near the Campanian–Maastrichtian boundary at both global and regional scales. The oological diversity of theropods in the Ibero-Armorican domain is consistent with the theropod diversity identified at high taxonomic level. Two distinct assemblages of theropod ootaxa can be recognized in the latest Cretaceous of the Ibero-Armorican domain. Their temporal transition can be correlated with other dinosaur faunal changes recorded in the region. This faunal turnover took place around the Early–Late Maastrichtian boundary, involving ornithopods, sauropods, ankylosaurs and, according to the present results, theropods as well.     

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.     

A large-clawed theropod (Dinosauria: Tetanurae) from the Lower Cretaceous of Australia and the Gondwanan origin of megaraptorid theropods

Megaraptoridae comprises a clade of enigmatic Gondwanan theropods with characteristic hypertrophied claws on the first and second manual digits. The majority of megaraptorids are known from South America, although a single genus (Australovenator) plus additional indeterminate material is also known from Australia. This clade has a controversial placement among theropods, and recently has been interpreted alternatively as a carcharodontosaurian or a tyrannosauroid lineage. We describe new fragmentary but associated postcranial remains from the opal fields of Lightning Ridge (middle-Albian, Griman Creek Formation) in north-central New South Wales. The new unnamed taxon exhibits a number of unusual features that suggest the presence of a hitherto unrecognised Australian megaraptorid. From an Australian perspective, the Lightning Ridge taxon predates Australovenator by ca. 10 Ma and is minimally coeval with megaraptoran material reported from the Eumeralla Formation of Victoria (but potentially 6.1–9.5 Ma younger). It is also notable as the largest predatory dinosaur yet identified from Australia and is only the second theropod known from more than a single element. A Bayesian phylogenetic approach integrating morphological, stratigraphic and palaeogeographic information tested both the carcharodontosaurian and tyrannosauroid placements for Megaraptora. Regardless of the preferred placement among Tetanurae, rigorous palaeobiogeographic analyses support an Asian origin of Megaraptora in the latest Jurassic (about 150–135 Ma), an Early Cretaceous (about 130–121 Ma) divergence of the Gondwanan lineage leading to Megaraptoridae, and an Australian root for megaraptorid radiation. These results indicate that Australia's Cretaceous dinosaur fauna did not comprise simply of immigrant taxa but was a source for complex two-way interchange between Australia–Antarctica–South America leading to the evolution of at least one group of apex predatory dinosaurs in Gondwana.     

Paleoecological implications of two closely associated egg types from the Upper Cretaceous St. Mary River Formation,Montana

Two closely associated egg types occur at the same locality in the Upper Cretaceous (Maastrichtian) St. Mary River Formation in north central Montana. These specimens represent the first fossil eggs described from this formation. At least fifteen small ovoid eggs or egg portions are scattered through a 25 cm interval of rock. Five significantly larger, round eggs overlie these smaller eggs and are in close proximity to one another on a single bedding plane. The best preserved egg of the smaller size measures 36 mm × 62 mm and exhibits the prismatic, two-layered eggshell structure of a theropod egg. The dispersed distribution and inconsistent angles of these small eggs likely resulted from disturbance by subsequent nesting activity and/or possibly nest predation. At least twelve additional small prismatic eggs also occur at this site. We assign the small eggs as a new oogenus and oospecies, Tetonoolithus nelsoni, within the Prismatoolithidae. The large round eggs measure 130 mm in diameter and the eggshell displays substantial diagenetic alteration. These eggs likely belonged to a hadrosaur due to their similarity in egg size, shape, and eggshell thickness to Maiasaura eggs from the stratigraphically lower Two Medicine Formation. Eggs at different stratigraphic levels at this site indicate that conditions favorable to both dinosaur species persisted for an extended period of time. However, determining whether these dinosaurs occupied the nesting site at the same or different years remains beyond the resolution of the rock record.