The last polar dinosaurs: high diversity of latest Cretaceous arctic dinosaurs in Russia

A latest Cretaceous (68 to 65 million years ago) vertebrate microfossil assemblage discovered at Kakanaut in northeastern Russia reveals that dinosaurs were still highly diversified in Arctic regions just before the Cretaceous–Tertiary mass extinction event. Dinosaur eggshell fragments, belonging to hadrosaurids and non-avian theropods, indicate that at least several latest Cretaceous dinosaur taxa could reproduce in polar region and were probably year-round residents of high latitudes. Palaeobotanical data suggest that these polar dinosaurs lived in a temperate climate (mean annual temperature about 10°C), but the climate was apparently too cold for amphibians and ectothermic reptiles. The high diversity of Late Maastrichtian dinosaurs in high latitudes, where ectotherms are absent, strongly questions hypotheses according to which dinosaur extinction was a result of temperature decline, caused or not by the Chicxulub impact.     

New stem-sauropodomorph (Dinosauria,Saurischia) from the Triassic of Brazil

Post-Triassic theropod, sauropodomorph, and ornithischian dinosaurs are readily recognized based on the set of traits that typically characterize each of these groups. On the contrary, most of the early members of those lineages lack such specializations, but share a range of generalized traits also seen in more basal dinosauromorphs. Here, we report on a new Late Triassic dinosaur from the Santa Maria Formation of Rio Grande do Sul, southern Brazil. The specimen comprises the disarticulated partial skeleton of a single individual, including most of the skull bones. Based on four phylogenetic analyses, the new dinosaur fits consistently on the sauropodomorph stem, but lacks several typical features of sauropodomorphs, showing dinosaur plesiomorphies together with some neotheropod traits. This is not an exception among basal dinosaurs, the early radiation of which is characterized by a mosaic pattern of character acquisition, resulting in the uncertain phylogenetic placement of various early members of the group.     

Bristle-like integumentary structures at the tail of the horned dinosaur Psittacosaurus

A specimen of the horned dinosaur Psittacosaurus from the early Cretaceous of China is described in which the integument is extraordinarily well-preserved. Most unusual is the presence of long bristle-like structures on the proximal part of tail. We interpret these structures as cylindrical and possibly tubular epidermal structures that were anchored deeply in the skin. They might have been used in display behavior and especially if one assumes that they were colored, they may have had a signal function. At present, there is no convincing evidence which shows these structures to be homologous to the structurally different integumentary filaments of theropod dinosaurs. Independent of their homology, however, the discovery of bristle-like structures in Psittacosaurus is of great evolutionary significance since it shows that the integumentary covering of at least some dinosaurs was much more complex than has ever been previously imagined.     

The first record of a sauropod dinosaur from Antarctica

Sauropoda is one of the most diverse and geographically widespread clades of herbivorous dinosaurs, and until now, their remains have now been recovered from all continental landmasses except Antarctica. We report the first record of a sauropod dinosaur from Antarctica, represented by an incomplete caudal vertebra from the Late Cretaceous of James Ross Island. The size and morphology of the specimen allows its identification as a lithostrotian titanosaur. Our finding indicates that advanced titanosaurs achieved a global distribution at least by the Late Cretaceous.     

Digital preparation of a probable neoceratopsian preserved within an egg, with comments on microstructural anatomy of ornithischian eggshells

We describe the first known embryo of a neoceratopsian dinosaur, perhaps the most ubiquitous Laurasian group of Cretaceous dinosaurs, which is preserved completely enclosed within an egg. This specimen was collected from Late Cretaceous beds of southern Mongolia, which commonly preserve fossils of the neoceratopsian, Yamaceratops dorngobiensis. The small egg was scanned using high-resolution X-ray computed tomography and digitally prepared from the matrix. The preserved and imaged elements support a diagnosis of the embryo to Neoceratopsia and allow preliminary observations of ontogenetic transformations within this group. The addition of an embryo also adds another important data point to the already impressive postnatal ontogenetic series that are available for this clade.     

The integument of Psittacosaurus from Liaoning Province, China: taphonomy, epidermal patterns and color of a ceratopsian dinosaur

Preserved skin of small dinosaurs is rare. Here, a specimen of the ceratopsian dinosaur, Psittacosaurus, presents some of the best preserved epidermal scales observed to date in a relatively small dinosaur, over wide areas extending from the head to the tail. We study the preserved epidermis of SMF R 4970, the different types of scales, color, and patterns, and their respective locations in the body. We use modern application of high-power digital imaging for close-up analysis of the tubercles and fragments of preserved color. Three types of scales are preserved, large plate-like scales, smaller polygonal scales or tubercles, and round pebble-like scales. The sizes of the plate-like scales vary in different parts of the body and vanish altogether posteriorly. Light and dark cryptic patterns are created by the associations of the tubercle and plate-like scales, and there is also evidence of countershading in the proximal caudal region, the body darker dorsally and lighter ventrally. Perhaps most impressive are the distinctive pigmented impressions of scales over most of the skeletal elements. The pigmentation follows the curvature of the bones implying that when it was deposited, the skin was still pliable and able to wrap around the visible parts of the elements. The present record of color is the first in a non-theropod dinosaur and only the second record in a non-avian dinosaur. Because of its resistance to degradation and ability to produce various color tones from yellows to blacks, we suggest that melanin was the dominant chemical involved in the coloration of Psittacosaurus. The data here enable us to reconstruct the colors of Psittacosaurus as predominantly black and amber/brown, in cryptic patterns, somewhat dull, but useful to a prey animal. Indeed, skin pigment within a partially degraded bone indicates that Psittacosaurus was scavenged shortly after death. The theropod dinosaur Sinosauropteryx has recently been reported to have naturally pigmented integumental structures, which the authors interpret as proof that they are protofeathers and not support fibers of collagen. Our findings in Psittacosaurus, on the other hand, indicate a more parsimonious and less profound alternative explanation, i.e., decomposition of the skin releases pigments that readily permeate underlying structures.     

Heterodonty and double occlusion in <Emphasis Type="Italic">Manidens condorensis</Emphasis>: a unique adaptation in an Early Jurassic ornithischian improving masticatory efficiency

New materials of the ornithischian dinosaur Manidens condorensis highlight a strong heterodonty between the upper and lower dentitions and reveal a novel occlusion type previously unreported in herbivorous dinosaurs. The diamond-shaped maxillary teeth have prominent cingular entolophs in a V- to Z-shaped configuration that are absent in dentary teeth. These cingular entolophs bear denticles and serrations with vertical wear that is coplanar with the apical wear facets, supporting their involvement in chewing. The separated apical and basal wear in dentary teeth is consistent with the apical and cingular wear in maxillary teeth, indicating an alternate occlusion, an orthal jaw motion, and shearing interactions between marginal and cingular edges in a double occlusion. Measurements of the length and wear area along the marginal and cingular edges indicate that the latter are functionally equivalent to adding eight teeth to a maxillary toothrow of ten, almost doubling the lengths of cutting edges and the degree of intraoral processing, while maintaining a plesiomorphic skull anatomy, an adaptation to herbivory unique in Ornithischia.     

Evolution of birds: ichthyosaur integumental fibers conform to dromaeosaur protofeathers

Filamentous integumentary structures have been reported as protofeathers in dromaeosaurs (non-avian dinosaurs). This hypothesis is considered against data on the complex architecture of dermal and subdermal collagenous fibers widely prevalent in living and extinct animals. Ichthyosaur integumental fibers, as dromaeosaur "protofeathers", are the most external structures preserved. Marked similarities are shown in branching patterns of these fibers compared with those of the dromaeosaur Sinornithosaurus; hence distinguishing between aberrant and primary features is difficult. Analysis of a pterosaur specimen shows that bent and straight fibers on the wings have functional implications. The fibers conform to the twofold shape of collagen and contradict the notion that bent integumental structures in the dinosaur Sinosauropteryx indicate softness and pliability. A suggestion also concerning Sinosauropteryx is that integumental structures with darker edges, compared with the middle, imply that they were hollow. Investigation of a similar condition in an ichthyosaur shows that it is more likely a consequence of mineralization. Dermal collagen fibers in, for example, sharks, dolphins, snakes, and turtles are shown to be grouped in bundles of varying sizes. Degradation of the dermis results in the breakdown of the fiber bundles and formation of myriad patterns of the disrupted fibers, as noted in decomposed dolphin skin. The overall findings of the study are that the thesis of dinosaur "protofeathers" requires more substantial support than exists at present.     

A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic

Non-avian theropod dinosaurs attained large body sizes, monopolising terrestrial apex predator niches in the Jurassic–Cretaceous. From the Middle Jurassic onwards, Allosauroidea and Megalosauroidea comprised almost all large-bodied predators for 85 million years. Despite their enormous success, however, they are usually considered absent from terminal Cretaceous ecosystems, replaced by tyrannosaurids and abelisaurids. We demonstrate that the problematic allosauroids Aerosteon, Australovenator, Fukuiraptor and Neovenator form a previously unrecognised but ecologically diverse and globally distributed clade (Neovenatoridae, new clade) with the hitherto enigmatic theropods Chilantaisaurus, Megaraptor and the Maastrichtian Orkoraptor. This refutes the notion that allosauroid extinction pre-dated the end of the Mesozoic. Neovenatoridae includes a derived group (Megaraptora, new clade) that developed long, raptorial forelimbs, cursorial hind limbs, appendicular pneumaticity and small size, features acquired convergently in bird-line theropods. Neovenatorids thus occupied a 14-fold adult size range from 175 kg (Fukuiraptor) to approximately 2,500 kg (Chilantaisaurus). Recognition of this major allosauroid radiation has implications for Gondwanan paleobiogeography: The distribution of early Cretaceous allosauroids does not strongly support the vicariant hypothesis of southern dinosaur evolution or any particular continental breakup sequence or dispersal scenario. Instead, clades were nearly cosmopolitan in their early history, and later distributions are explained by sampling failure or local extinction.     

A Middle Jurassic heterodontosaurid dinosaur from Patagonia and the evolution of heterodontosaurids

Heterodontosauridae is a morphologically divergent group of dinosaurs that has recently been interpreted as one of the most basal clades of Ornithischia. Heterodontosaurid remains were previously known from the Early Jurassic of southern Africa, but recent discoveries and studies have significantly increased the geographical and temporal range for this clade. Here, we report a new ornithischian dinosaur from the Middle Jurassic Ca?adón Asfalto Formation in central Patagonia, Argentina. This new taxon, Manidens condorensis gen. et sp. nov., includes well-preserved craniomandibular and postcranial remains and represents the only diagnostic ornithischian specimen yet discovered in the Jurassic of South America so far. Derived features of its anatomy indicate that Manidens belongs to Heterodontosauridae, as the sister taxon of Heterodontosaurus and other South African heterodontosaurids. The presence of posterior dentary teeth with high crowns but lacking extensive wear facets in Manidens suggests that this form represents an intermediate stage in the development of the remarkable adaptations to herbivory described for Heterodontosaurus. The dentition of Manidens condorensis also has autapomorphies, such as asymmetrically arranged denticles in posterior teeth and a mesially projected denticle in the posteriormost teeth. At an estimated total length of 60?C75?cm, Manidens furthermore confirms the small size of basal heterodontosaurids.     

The dinosaurian origin of feathers: perspectives from dolphin (Cetacea) collagen fibers

The early origin of birds is a hotly disputed debate and may be broadly framed as a conflict between paleontologists and ornithologists. The paleontological emphasis has shifted from Archaeopteryx and its origins to recent finds of Cretaceous birds and feathered dinosaurs from China. The identification of alleged feathers has, however, relied principally on the visual image. Some workers have interpreted these integumentary structures as collagen fibers. To test the latter hypothesis, using light microscopy, collagen from the hypodermis (blubber) and subdermal connective tissue sheath was examined from a dolphin that had been buried for a year as part of an experiment. Within the blubber, toward the central thicker parts of the material, the collagen fibers had compacted and the three-dimensional latticework of normal blubber had more or less collapsed. Chromatographic analysis of the blubber revealed pronounced oxidation of the unsaturated lipids, probably accounting for the collapse of the latticework. Fibers normally bound together in bundles became separated into individual fibers or smaller bundles by degradation of the glue-like substance binding them together. These degraded collagen fibers show, in many instances, feather-like patterns, strikingly reminiscent of many of those identified as either protofeathers or modern feathers in dromaeosaurid dinosaurs. The findings throw serious doubt on the virtually complete reliance on visual image by supporters of the feathered dinosaur thesis and emphasize the need for more rigorous methods of identification using modern feathers as a frame of reference. Since collagen is the main fiber type found in most supporting tissues, the results have wide implications regarding the degradation and fossilization of vertebrate integument, such as that of the ichthyosaurs, dinosaurs and birds.     

The evolutionary continuum of limb function from early theropods to birds

The bipedal stance and gait of theropod dinosaurs evolved gradually along the lineage leading to birds and at some point(s), flight evolved. How and when did these changes occur? We review the evidence from neontology and palaeontology, including pectoral and pelvic limb functional morphology, fossil footprints/trackways and biomechanical models and simulations. We emphasise that many false dichotomies or categories have been applied to theropod form and function, and sometimes, these impede research progress. For example, dichotomisation of locomotor function into ‘non-avian’ and ‘avian’ modes is only a conceptual crutch; the evidence supports a continuous transition. Simplification of pelvic limb function into cursorial/non-cursorial morphologies or flexed/columnar poses has outlived its utility. For the pectoral limbs, even the classic predatory strike vs. flight wing-stroke distinction and separation of theropods into non-flying and flying—or terrestrial and arboreal—categories may be missing important subtleties. Distinguishing locomotor function between taxa, even with quantitative approaches, will always be fraught with ambiguity, making it difficult to find real differences if that ambiguity is properly acknowledged. There must be an ‘interpretive asymptote’ for reconstructing dinosaur limb function that available methods and evidence cannot overcome. We may be close to that limit, but how far can it be stretched with improved methods and evidence, if at all? The way forward is a combination of techniques that emphasises integration of neontological and palaeontological evidence and quantitative assessment of limb function cautiously applied with validated techniques and sensitivity analysis of unknown variables.     

The First Detection of Human Bocavirus Species 2 and 3 in Raw Sewage and Mussels in South Africa

Human bocavirus (HBoV) has a global distribution and is associated with respiratory and enteric infections, particularly in the paediatric population. In this study, raw sewage and mussel samples were analysed for the presence of HBoV using nested PCR with primers targeting the VP1/VP2 junction. Amplification and sequencing of the 382 bp region followed by phylogenetic analysis indicated the presence of HBoV 2 in mussel samples and HBoV 3 in sewage samples. This is the first report describing the presence of enteric-associated HBoV in environmental samples from South Africa and in mussel samples from the African continent. The results signify the need for further studies examining the potential risk of foodborne transmission of HBoV and highlight the importance of continued screening to determine the prevalence and epidemiology of HBoV in South Africa.     

First complete sauropod dinosaur skull from the Cretaceous of the Americas and the evolution of sauropod dentition

Sauropod dinosaur bones are common in Mesozoic terrestrial sediments, but sauropod skulls are exceedingly rare—cranial materials are known for less than one third of sauropod genera and even fewer are known from complete skulls. Here we describe the first complete sauropod skull from the Cretaceous of the Americas, Abydosaurus mcintoshi, n. gen., n. sp., known from 104.46 ± 0.95 Ma (megannum) sediments from Dinosaur National Monument, USA. Abydosaurus shares close ancestry with Brachiosaurus, which appeared in the fossil record ca. 45 million years earlier and had substantially broader teeth. A survey of tooth shape in sauropodomorphs demonstrates that sauropods evolved broad crowns during the Early Jurassic but did not evolve narrow crowns until the Late Jurassic, when they occupied their greatest range of crown breadths. During the Cretaceous, brachiosaurids and other lineages independently underwent a marked diminution in tooth breadth, and before the latest Cretaceous broad-crowned sauropods were extinct on all continental landmasses. Differential survival and diversification of narrow-crowned sauropods in the Late Cretaceous appears to be a directed trend that was not correlated with changes in plant diversity or abundance, but may signal a shift towards elevated tooth replacement rates and high-wear dentition. Sauropods lacked many of the complex herbivorous adaptations present within contemporaneous ornithischian herbivores, such as beaks, cheeks, kinesis, and heterodonty. The spartan design of sauropod skulls may be related to their remarkably small size—sauropod skulls account for only 1/200th of total body volume compared to 1/30th body volume in ornithopod dinosaurs.     

恐龙骨骼化石与其它脊椎动物骨骼的微量元素的对比研究

采用中子活化分析 (INAA)技术 ,对比分析恐龙骨骼化石及其它脊椎动物 (乌龟、鱼和大量哺乳动物 )骨骼中微量元素的含量组合 ,结果表明稀土、砷、钡、铀元素在恐龙骨骼化石中的含量远高于其它脊椎动物骨骼 ,而锌元素的含量情况却与之相反 ;同时进一步分析推测得出 :这种砷、钡元素含量的高异常和锌元素的低异常 ,极可能是导致四川盆地恐龙动物群集群死亡的原因之一。     

Closely associated theropod trackways from the Jurassic of Zimbabwe

Eighty-eight tracks of large theropod dinosaurs were found in the mid-Jurassic of Zimbabwe. Among the tracks, at least five adjacent trackways are recorded. The adjacent tracks were probably made by animals traveling as a group, given that they are in relatively close succession; that there are three overlapping tracks (among just 23) suggesting reasonably close associations of the animals; that all the tracks are apparently of the same ichnotaxon; that the preservational types of the tracks are similar; and that the tracks are all of animals traveling in one general direction closely associated in time (there are no returning tracks of the same animals or of those of other species; presence of such tracks would be highly probable if the tracks were made over a period of time of even several hours). Nearby, recently discovered giant sauropod tracks, the first in sub-Saharan Africa, indicate a realistic potential of predator/prey interactions between the two groups of dinosaurs.     

First nonavian dinosaur from Lebanon: a brachiosaurid sauropod from the Lower Cretaceous of the Jezzine District

Two sauropod teeth from an Early Cretaceous (Neocomian) fluviodeltaic sandstone near Jezzine (Southern Lebanon) are the first nonavian dinosaur remains to be reported from Lebanon. Their distinctive character places them within Brachiosauridae. The sauropod teeth from Lebanon are a significant addition to the very scanty dinosaur record from the Levant, which hitherto consisted mainly of very poorly preserved and not easily identifiable specimens. The Basal Cretaceous Sandstone of Lebanon, thus, appears to be a potentially important source of fossil vertebrate material.     

粤北南雄盆地末代恐龙遗迹及其他

粤北南雄盆地是中生代末世和新生代初世的陆相沉积盆地。上白垩统富含末代恐龙、恐龙蛋和恐龙脚印化石。下第三系古新统则出现哺乳动物先驱化石。对研究恐龙的绝灭有重要科学意义。     

First ceratosaurian dinosaur from Australia

The basal theropod dinosaur clade Ceratosauria, and its subclade Abelisauroidea, is characteristic of late Mesozoic terrestrial vertebrate faunas in western Gondwana (South America, Africa, Madagascar, and India) and Europe. Yet unambiguous records of ceratosaurs have hitherto been absent from Australia, where the theropod assemblage appears to include several typically Laurasian clades. Here, we report the first evidence of ceratosaurs (and potentially abelisauroids) from eastern Gondwana––a diagnostic astragalocalcaneum from the Aptian (121–125?Ma) of Victoria, Australia. Ceratosauria thus occurred in both western and eastern Gondwana during the Early Cretaceous. This fossil adds to the poorly known dinosaur fauna of Australia, a major clade of basal theropods, emphasising that its mid-Cretaceous theropod diversity was surprisingly cosmopolitan despite relative geographic isolation, including clades that have been thought to be typical of both Gondwana and Laurasia––Ceratosauria, Spinosauridae, Carcharodontosauria, Tyrannosauroidea, and Deinonychosauria. Such a contemporaneous association of theropod clades is unknown from other Gondwanan continents and questions the views that the late Mesozoic dinosaur fauna of Australia was dominated by Gondwanan or Laurasian elements, extreme isolation, relictualism, and/or novelty as a ‘centre of origin’. The cosmopolitan theropod fauna of Australia probably reflects the global distribution of these clades early in their history, prior to significant continental breakup.     

Palaeoenvironmental controls on the distribution of Cretaceous herbivorous dinosaurs

Previous attempts to determine palaeoenvironmental preferences in dinosaurs have generally been qualitative assessments based upon data from restricted geographical areas. Here, we use a global database of Cretaceous herbivorous dinosaurs to identify significant associations between clades and broad palaeoenvironmental categories (‘terrestrial’, ‘coastal’, ‘marine’). Nodosaurid ankylosaurs and hadrosaurids show significant positive associations with marine sediments, while marginocephalians (Ceratopsia, Pachycephalosauria), saurischians (herbivorous theropods, Sauropoda) and ankylosaurid ankylosaurs are significantly positively associated with terrestrial sediments. These results provide quantitative support for the hypothesis that some clades (Nodosauridae, Hadrosauridae) were more abundant in coastal and/or fluvial environments, while others (e.g. Marginocephalia, Ankylosauridae) preferentially inhabited more distal environments. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. An erratum to this article can be found at