Epidemiologic study of tumors in dinosaurs

Occasional reports in isolated fragments of dinosaur bones have suggested that tumors might represent a population phenomenon. Previous study of humans has demonstrated that vertebral radiology is a powerful diagnostic tool for population screening. The epidemiology of tumors in dinosaurs was here investigated by fluoroscopically screening dinosaur vertebrae for evidence of tumors. Computerized tomography (CT) and cross-sections were obtained where appropriate. Among more than 10,000 specimens x-rayed, tumors were only found in Cretaceous hadrosaurs (duck-billed dinosaurs). These included hemangiomas and metastatic cancer (previously identified in dinosaurs), desmoplastic fibroma, and osteoblastoma. The epidemiology of tumors in dinosaurs seems to reflect a familial pattern. A genetic propensity or environmental mutagens are suspected.     

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.     

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     

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.     

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.     

A large Cretaceous theropod from Patagonia, Argentina, and the evolution of carcharodontosaurids

The Cretaceous Carcharodontosauridae is the latest clade of carnosaurs, including the largest predatory dinosaurs yet recorded. Albeit spectacular for their size, the skeletal anatomy of these theropods remains poorly-known, and their diversity was until recently restricted to two Cenomanian species: the highly derived Giganotosaurus carolinii, from southern South America, and the incompletely known Carcharodontosaurus saharicus, from northern Africa. Here we describe an older and basal member of the group, Tyrannotitan chubutensis gen. et sp. nov., from Aptian strata of Patagonia, Argentina. The new taxon gives new insights into the systematics and evolution of carcharodontosaurids and offers a better understanding of the evolution of Southern theropod faunas. We suggest that carcharodontosaurids radiated in Gondwana sharing with spinosaurids the role of top-predators until their extinction in Cenomanian–Turonian times. During this interval, the diplodocoid sauropods and giant titanosaurians went extinct (probably as part of a global-scale crisis), and the smaller abelisaurid theropods took dominance, reigning until the end of the Cretaceous. Electronic Supplementary Material is available.Electronic Supplementary Information  Supplementary material is available for this article at     

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.     

Chicken-sized oviraptorid dinosaurs from central China and their ontogenetic implications

Oviraptorids are a group of specialized non-avian theropod dinosaurs that were generally one to 8 m in body length. New specimens of baby oviraptorids from the Late Cretaceous of Henan Province are some of the smallest individuals known. They include diagnostic characters such as the relative position of the antorbital fenestra and the external naris, distinct opening in the premaxilla anteroventral to the external naris, antorbital fossa partly bordered by premaxilla posterodorsally, lacrimal process of premaxilla does not contact the anterodorsal process of the lacrimal, parietal almost as long as frontal; in dorsal view, posterior margin forms a straight line between the postzygapophyses in each of the fourth and fifth cervicals; femur longer than ilium. They also elucidate the ontogenetic processes of oviraptorids, including fusion of cranial elements and changes in relative body proportions. Hind limb proportions are constant in oviraptorids, regardless of absolute body size or ontogenetic stage. This suggests a sedentary lifestyle that did not involve the pursuit of similar-sized prey. The functional implications for bite force and therefore dietary preferences are better understood through the study of such small animals. The comparison of the measurements of 115 skeletons indicates that oviraptorids maintain their hind limb proportions regardless of ontogenetic stage or absolute size, which is a pattern seen more commonly in herbivores than in carnivores. This may weakly support the hypothesis that oviraptorids are herbivores rather than active carnivores.     

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.     

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.     

The first definitive carcharodontosaurid (Dinosauria: Theropoda) from Asia and the delayed ascent of tyrannosaurids

Little is known about the evolution of large-bodied theropod dinosaurs during the Early to mid Cretaceous in Asia. Prior to this time, Asia was home to an endemic fauna of basal tetanurans, whereas terminal Cretaceous ecosystems were dominated by tyrannosaurids, but the intervening 60 million years left a sparse fossil record. Here, we redescribe the enigmatic large-bodied Chilantaisaurus maortuensis from the Turonian of Inner Mongolia, China. We refer this species to a new genus, Shaochilong, and analyze its systematic affinities. Although Shaochilong has previously been allied with several disparate theropod groups (Megalosauridae, Allosauridae, Tyrannosauroidea, Maniraptora), we find strong support for a derived carcharodontosaurid placement. As such, Shaochilong is the first unequivocal Asian member of Carcharodontosauridae, which was once thought to be restricted to Gondwana. The discovery of an Asian carcharodontosaurid indicates that this clade was cosmopolitan in the Early to mid Cretaceous and that Asian large-bodied theropod faunas were no longer endemic at this time. It may also suggest that the ascent of tyrannosaurids into the large-bodied dinosaurian predator niche was a late event that occurred towards the end of the Cretaceous, between the Turonian and the Campanian. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

<Emphasis Type="Italic">Bonitasaura salgadoi</Emphasis> gen. et sp. nov.: a beaked sauropod from the Late Cretaceous of Patagonia

Ornithischian and theropod dinosaurs were morphologically diverse during the Cretaceous. In contrast, sauropods were relatively more conservative. The anatomy of Bonitasaura salgadoi, a new 9-m titanosaurian sauropod from Upper Cretaceous beds of Patagonia, suggests that sauropod anatomical diversity would have included unexpected items. Its unusual, rectangular lower jaw possesses narrow, anteriorly restricted teeth and shows evidence of a sharp keratinous sheath over the non-dentigerous region that probably worked to guillotine plant material. This discovery definitely demonstrates that titanosaurs acquired a mandibular configuration similar to that of some basal diplodocoids, as had already been suggested by the lower jaw of the controversial genus Antarctosaurus. This oral configuration, plus the beak-like structure and the skull shape, resemble some traits more commonly seen in Laurasian ornithischians, mostly unexpressed in southern continents. A high sauropod morphological diversity seems to be in agreement with the poorly represented ornithischian clades of the southern hemisphere.Communicated by G. Mayr     

Largest bird from the Early Cretaceous and its implications for the earliest avian ecological diversification

With only one known exception, early Cretaceous birds were smaller than their closest theropod dinosaur relatives. Here we report on a new bird from the Early Cretaceous feathered-dinosaur-bearing continental deposits of Liaoning, northeast China, which is not only larger than Archaeopteryx but is nearly twice as large as the basal dromaeosaur Microraptor. The new taxon, Sapeornis chaoyangensis gen. et sp. nov., has a more basal phylogenetic position than all other birds except for Archaeopteryx. Its exceptionally long forelimbs, well-developed deltoid crest of the humerus, proximally fused metacarpals, relatively short hindlimbs and short pygostyle indicate powerful soaring capability and further suggest that by the Early Cretaceous ecological diversification of early birds was greater than previously assumed. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00114-001-0276-9.     

Postnatal nutrition influences male attractiveness and promotes plasticity in male mating preferences

The fossil record of Late Cretaceous–Paleogene modern birds in the Southern Hemisphere includes the Maastrichtian Neogaeornis wetzeli from Chile, Polarornis gregorii and Vegavis iaai from Antarctica, and Australornis lovei from the Paleogene of New Zealand. The recent finding of a new and nearly complete Vegavis skeleton constitutes the most informative source for anatomical comparisons among Australornis, Polarornis, and Vegavis. The present contribution includes, for the first time, Vegavis, Polarornis, and Australornis in a comprehensive phylogenetic analysis. This analysis resulted in the recognition of these taxa as a clade of basal Anseriformes that we call Vegaviidae. Vegaviids share a combination of characters related to diving adaptations, including compact and thickened cortex of hindlimb bones, femur with anteroposteriorly compressed and bowed shaft, deep and wide popliteal fossa delimited by a medial ridge, tibiotarsus showing notably proximally expanded cnemial crests, expanded fibular crest, anteroposterior compression of the tibial shaft, and a tarsometatarsus with a strong transverse compression of the shaft. Isolated bones coming from the Cretaceous and Paleogene of South America, Antarctica, and New Zealand are also referred to here to Vegaviidae and support the view that these basal anseriforms were abundant and diverse at high southern latitudes. Moreover, vegaviids represent the first avian lineage to have definitely crossed the K–Pg boundary, supporting the idea that some avian clades were not affected by the end Mesozoic mass extinction event, countering previous interpretations. Recognition of Vegaviidae indicates that modern birds were diversified in southern continents by the Cretaceous and reinforces the hypothesis indicating the important role of Gondwana for the evolutionary history of Anseriformes and Neornithes as a whole.

     

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.     

Vegaviidae,a new clade of southern diving birds that survived the K/T boundary

The fossil record of Late Cretaceous–Paleogene modern birds in the Southern Hemisphere includes the Maastrichtian Neogaeornis wetzeli from Chile, Polarornis gregorii and Vegavis iaai from Antarctica, and Australornis lovei from the Paleogene of New Zealand. The recent finding of a new and nearly complete Vegavis skeleton constitutes the most informative source for anatomical comparisons among Australornis, Polarornis, and Vegavis. The present contribution includes, for the first time, Vegavis, Polarornis, and Australornis in a comprehensive phylogenetic analysis. This analysis resulted in the recognition of these taxa as a clade of basal Anseriformes that we call Vegaviidae. Vegaviids share a combination of characters related to diving adaptations, including compact and thickened cortex of hindlimb bones, femur with anteroposteriorly compressed and bowed shaft, deep and wide popliteal fossa delimited by a medial ridge, tibiotarsus showing notably proximally expanded cnemial crests, expanded fibular crest, anteroposterior compression of the tibial shaft, and a tarsometatarsus with a strong transverse compression of the shaft. Isolated bones coming from the Cretaceous and Paleogene of South America, Antarctica, and New Zealand are also referred to here to Vegaviidae and support the view that these basal anseriforms were abundant and diverse at high southern latitudes. Moreover, vegaviids represent the first avian lineage to have definitely crossed the K–Pg boundary, supporting the idea that some avian clades were not affected by the end Mesozoic mass extinction event, countering previous interpretations. Recognition of Vegaviidae indicates that modern birds were diversified in southern continents by the Cretaceous and reinforces the hypothesis indicating the important role of Gondwana for the evolutionary history of Anseriformes and Neornithes as a whole.     

松属植物地质历史及生物地理演变

松属（Pinus L.）约113种,是松科现代属中最原始的类群。松属植物种类丰富且研究领域广泛,对其已经积累的资料数据进行系统梳理总结十分必要。本文通过总结国内外松属大化石资料,结合分子系统发育、地质背景和地理隔离事件讨论了其地史分布及植物地理学意义;该属化石在早白垩世至全新世地层中均有记录。化石证据表明松属很可能在早白垩世（之前）起源于西欧地区,从这一起源地通过北大西洋陆桥扩散到北美洲东部,而东亚的类群可能是从北美洲西部经过白令陆桥散布的。在晚白垩世分化出双维管束松亚属Subgenus Pinus L.及单维管束松亚属Subgenus Strobus (D. Don) Lemmon,前者更接近祖先类群。古新世由于全球显著增温以及白垩纪末期大灭绝等地质事件的影响使松属数量急剧减少,在晚始新世至中新世时期随着气温转凉转冷再次分化扩散,中新世达到其发展高峰且分布面貌与现代类群近似。松属多样性时空历史可能和新生代气候变迁及晚新生代构造运动塑造的山地隆升等环境变化紧密相关。     

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.     

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。