A new avialan theropod from an emerging Jurassic terrestrial fauna.

Birds are descended from non-avialan theropod dinosaurs of the Late Jurassic period, but the earliest phase of this evolutionary process remains unclear owing to the exceedingly sparse and spatio-temporally restricted fossil record1-5. Information about the early-diverging species along the avialan line is crucial to understand the evolution of the characteristic bird bauplan, and to reconcile phylogenetic controversies over the origin of birds3,4. Here we describe one of the stratigraphically youngest and geographically southernmost Jurassic avialans, Fujianvenator prodigiosus gen. et sp. nov., from the Tithonian age of China. This specimen exhibits an unusual set of morphological features that are shared with other stem avialans, troodontids and dromaeosaurids, showing the effects of evolutionary mosaicism in deep avialan phylogeny. F. prodigiosus is distinct from all other Mesozoic avialan and non-avialan theropods in having a particularly elongated hindlimb, suggestive of a terrestrial or wading lifestyle-in contrast with other early avialans, which exhibit morphological adaptations to arboreal or aerial environments. During our fieldwork in Zhenghe where F. prodigiosus was found, we discovered a diverse assemblage of vertebrates dominated by aquatic and semi-aquatic species, including teleosts, testudines and choristoderes. Using in situ radioisotopic dating and stratigraphic surveys, we were able to date the fossil-containing horizons in this locality-which we name the Zhenghe Fauna-to 148-150 million years ago. The diversity of the Zhenghe Fauna and its precise chronological framework will provide key insights into terrestrial ecosystems of the Late Jurassic.

Dental microwear of a basal ankylosaurine dinosaur, Jinyunpelta and its implication on evolution of chewing mechanism in ankylosaurs.

Jinyunpelta sinensis is a basal ankylosaurine dinosaur excavated from the mid Cretaceous Liangtoutang Formation of Jinyun County, Zhejiang Province, China. In the present study, its dental microwear was observed using a confocal laser microscope. Jinyunpelta had steep wear facets that covered most of buccal surfaces of posterior dentary teeth. Observation of dental microwear on the wear facet revealed that scratch orientation varied according to its location within the wear facet: vertically (i.e. apicobasally) oriented scratches were dominant in the upper half of the wear facet, and horizontally (i.e. mesiolaterally) oriented ones were in the bottom of the facet. These findings indicated that Jinyunpelta adopted precise tooth occlusion and biphasal jaw movement (orthal closure and palinal lower jaw movement). The biphasal jaw movement was widely observed among nodosaurids, among ankylosaurids, it was previously only known from the Late Cretaceous North American taxa, and not known among Asian ankylosaurids. The finding of biphasal jaw movement in Jinyunpelta showed sophisticate feeding adaptations emerged among ankylosaurids much earlier (during Albian or Cenomanian) than previously thought (during Campanian). The Evolution of the biphasal jaw mechanism that contemporaneously occurred among two lineages of ankylosaurs, ankylosaurids and nodosaurids, showed high evolutionary plasticity of ankylosaur jaw mechanics.

A new species of early-diverging Sauropodiformes from the Lower Jurassic Fengjiahe Formation of Yunnan Province, China.

Sauropodomorpha were herbivorous saurischian dinosaurs that incorporate Sauropoda and early-diverging sauropodomorphs. The oldest sauropodomorph remains are known from Late Triassic deposits, most of them Gondwanan. The Laurasian record comprises some Triassic forms, but the bulk is Jurassic in age. Among the 14 Jurassic non-sauropodan sauropodomorphs from Laurasia described in the past, 8 are from China. Here we describe a new non-sauropodan sauropodomorph, Irisosaurus yimenensis gen. et sp. nov., from the Early Jurassic Fengjiahe Formation of China. Nearly all of the non-sauropodan sauropodomorph genera currently known from China were first reported from the Lufeng Formation. The Fengjiahe Formation is its Southern equivalent, bringing a fauna similar to that of the Lufeng Formation to light. The new genus is defined based on an incomplete but unique maxilla, with a premaxillary ramus higher than long prior to the nasal process, a large and deep neurovascular foramen within the perinarial fossa, and a deep perinarial fossa defined by a sharp rim. Phylogenetic analysis places Irisosaurus at the very base of Sauropodiformes, as the sister-taxon of the Argentinean genus Mussaurus. This specimen adds to a growing assemblage of Chinese Jurassic non-sauropodan sauropodomorphs that offers new insight into the Laurasian evolution of this clade.

Novel track morphotypes from new tracksites indicate increased Middle Jurassic dinosaur diversity on the Isle of Skye, Scotland.

Dinosaur fossils from the Middle Jurassic are rare globally, but the Isle of Skye (Scotland, UK) preserves a varied dinosaur record of abundant trace fossils and rare body fossils from this time. Here we describe two new tracksites from Rubha nam Brathairean (Brothers' Point) near where the first dinosaur footprint in Scotland was found in the 1980s. These sites were formed in subaerially exposed mudstones of the Lealt Shale Formation of the Great Estuarine Group and record a dynamic, subtropical, coastal margin. These tracksites preserve a wide variety of dinosaur track types, including a novel morphotype for Skye: Deltapodus which has a probable stegosaur trackmaker. Additionally, a wide variety of tridactyl tracks shows evidence of multiple theropods of different sizes and possibly hints at the presence of large-bodied ornithopods. Overall, the new tracksites show the dinosaur fauna of Skye is more diverse than previously recognized and give insight into the early evolution of major dinosaur groups whose Middle Jurassic body fossil records are currently sparse.

Inter-amphibian predation in the Early Cretaceous of China.

For most fossil taxa, dietary inference relies primarily on indirect evidence from jaw morphology and the dentition. In rare cases, however, preserved gut contents provide direct evidence of feeding strategy and species interaction. This is important in the reconstruction of food webs and energy flow through ancient ecosystems. The Early Cretaceous Chinese Jehol Biota has yielded several such examples, with lizards, birds, small dinosaurs, and mammals as both predator and prey. Here we describe an Early Cretaceous fossil frog specimen, genus Genibatrachus, that contains an adult salamander within its body cavity. The salamander is attributed to the hynobiid-like genus Nuominerpeton. The salamander skeleton is complete and articulated, suggesting it was caught and swallowed shortly before the frog itself died and was buried.

Possible bite-induced abscess and osteomyelitis in Lufengosaurus (Dinosauria: sauropodomorph) from the Lower Jurassic of the Yimen Basin, China.

We report an osseous abnormality on a specimen of the sauropod dinosaur Lufengosaurus huenei from the Fengjiahe Formation in Yuxi Basin, China. A gross pathological defect occurs on the right third rib, which was subjected to micro-computed tomographic imaging as an aid in diagnosis. The analysis of pathological characteristics and the shape of the abnormality is incompatible with impact or healed trauma, such as a common rib fracture, and instead suggests focal penetration of the rib, possibly due to a failed predator attack. The identification of characteristics based on gross morphology and internal micro-morphology presented by the specimen, suggests an abscess with osteomyelitis as the most parsimonious explanation. Osteomyelitis is a severe infection originating in the bone marrow, usually resulting from the introduction of pyogenic (pus-producing) bacteria into the bone. Micro-tomographic imaging of the lesion suggests a degree of healing and bone remodelling following post-traumatic wound infection with evidence of sclerotic bone formation at the site of pathological focus, indicating that L. huenei survived the initial trauma. However, as osteomyelitis can express through widespread systemic effects, including a lowering of immune response and overall condition, this disease may have been a contributing factor to the eventual death of the individual.

The bone microstructure of polar "hypsilophodontid" dinosaurs from Victoria, Australia.

High-latitude (i.e., "polar") Mesozoic fauna endured months of twilight and relatively low mean annual temperatures. Yet non-avian dinosaurs flourished in this taxing environment. Fossils of basal ornithopod dinosaurs ("hypsilophodontids") are common in the Early Cretaceous high-latitude sediments of Victoria, Australia, and four taxa have been described; although their ontogenetic histories are largely unexplored. In the present study, eighteen tibiae and femora were utilized in the first multi-specimen ontogenetic histological analysis of Australian polar hypsilophodontids. The sample consists of eleven individuals from the Flat Rocks locality (Late Valanginian or Barremian), and five from the Dinosaur Cove locality (Albian). In both groups, growth was most rapid during the first three years, and skeletal maturity occurred between five and seven years. There is a weak asymptotic trend in a plot of growth mark count versus femur length, with considerable individual variation. Histology suggests two genera are present within the Dinosaur Cove sample, but bone microstructure alone could not distinguish genera within the Flat Rocks sample, or across the two geologically separate (~ 26 Ma) localities. Additional histologic sampling, combined with morphological analyses, may facilitate further differentiation between ontogenetic, individual, and species variation.

Mosaic evolution in an asymmetrically feathered troodontid dinosaur with transitional features.

Asymmetrical feathers have been associated with flight capability but are also found in species that do not fly, and their appearance was a major event in feather evolution. Among non-avialan theropods, they are only known in microraptorine dromaeosaurids. Here we report a new troodontid, Jianianhualong tengi gen. et sp. nov., from the Lower Cretaceous Jehol Group of China, that has anatomical features that are transitional between long-armed basal troodontids and derived short-armed ones, shedding new light on troodontid character evolution. It indicates that troodontid feathering is similar to Archaeopteryx in having large arm and leg feathers as well as frond-like tail feathering, confirming that these feathering characteristics were widely present among basal paravians. Most significantly, the taxon has the earliest known asymmetrical troodontid feathers, suggesting that feather asymmetry was ancestral to Paraves. This taxon also displays a mosaic distribution of characters like Sinusonasus, another troodontid with transitional anatomical features.

Diverse transitional giant fleas from the Mesozoic era of China.

Fleas are one of the major lineages of ectoparasitic insects and are now highly specialized for feeding on the blood of birds or mammals. This has isolated them among holometabolan insect orders, although they derive from the Antliophora (scorpionflies and true flies). Like most ectoparasitic lineages, their fossil record is meagre and confined to Cenozoic-era representatives of modern families, so that we lack evidence of the origins of fleas in the Mesozoic era. The origins of the first recognized Cretaceous stem-group flea, Tarwinia, remains highly controversial. Here we report fossils of the oldest definitive fleas--giant forms from the Middle Jurassic and Early Cretaceous periods of China. They exhibit many defining features of fleas but retain primitive traits such as non-jumping hindlegs. More importantly, all have stout and elongate sucking siphons for piercing the hides of their hosts, implying that these fleas may be rooted among the pollinating 'long siphonate' scorpionflies of the Mesozoic. Their special morphology suggests that their earliest hosts were hairy or feathered 'reptilians', and that they radiated to mammalian and bird hosts later in the Cenozoic.

Mineralized soft-tissue structure and chemistry in a mummified hadrosaur from the Hell Creek Formation, North Dakota (USA).

An extremely well-preserved dinosaur (Cf. Edmontosaurus sp.) found in the Hell Creek Formation (Upper Cretaceous, North Dakota) retains soft-tissue replacement structures and associated organic compounds. Mineral cements precipitated in the skin apparently follow original cell boundaries, partially preserving epidermis microstructure. Infrared and electron microprobe images of ossified tendon clearly show preserved mineral zonation, with silica and trapped carbon dioxide forming thin linings on Haversian canals within apatite. Furthermore, Fourier transform infrared spectroscopy (FTIR) of materials recovered from the skin and terminal ungual phalanx suggests the presence of compounds containing amide groups. Amino acid composition analyses of the mineralized skin envelope clearly differ from the surrounding matrix; however, intact proteins could not be obtained using protein mass spectrometry. The presence of endogenously derived organics from the skin was further demonstrated by pyrolysis gas chromatography mass spectrometry (Py-GCMS), indicating survival and presence of macromolecules that were in part aliphatic (see the electronic supplementary material).

An Early Cretaceous heterodontosaurid dinosaur with filamentous integumentary structures.

Ornithischia is one of the two major groups of dinosaurs, with heterodontosauridae as one of its major clades. Heterodontosauridae is characterized by small, gracile bodies and a problematic phylogenetic position. Recent phylogenetic work indicates that it represents the most basal group of all well-known ornithischians. Previous heterodontosaurid records are mainly from the Early Jurassic period (205-190 million years ago) of Africa. Here we report a new heterodontosaurid, Tianyulong confuciusi gen. et sp. nov., from the Early Cretaceous period (144-99 million years ago) of western Liaoning Province, China. Tianyulong extends the geographical distribution of heterodontosaurids to Asia and confirms the clade's previously questionable temporal range extension into the Early Cretaceous period. More surprisingly, Tianyulong bears long, singular and unbranched filamentous integumentary (outer skin) structures. This represents the first confirmed report, to our knowledge, of filamentous integumentary structures in an ornithischian dinosaur.

A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers.

Recent coelurosaurian discoveries have greatly enriched our knowledge of the transition from dinosaurs to birds, but all reported taxa close to this transition are from relatively well known coelurosaurian groups. Here we report a new basal avialan, Epidexipteryx hui gen. et sp. nov., from the Middle to Late Jurassic of Inner Mongolia, China. This new species is characterized by an unexpected combination of characters seen in several different theropod groups, particularly the Oviraptorosauria. Phylogenetic analysis shows it to be the sister taxon to Epidendrosaurus, forming a new clade at the base of Avialae. Epidexipteryx also possesses two pairs of elongate ribbon-like tail feathers, and its limbs lack contour feathers for flight. This finding shows that a member of the avialan lineage experimented with integumentary ornamentation as early as the Middle to Late Jurassic, and provides further evidence relating to this aspect of the transition from non-avian theropods to birds.

[Mapping metal elements of Shuangbai dinosaur fossil by synchrotron X-ray fluorescence microprobe].

The metal elements mapping of Shuangbai dinosaur fossil, was obtained by synchrotron x-ray fluorescence (SXRF). Eight elements, Ca, Mn, Fe, Cu, Zn, As, Y and Sr were determined. Elements As and Y were detected for the first time in the dinosaur fossil. The data indicated that metal elements are asymmetrical on fossil section. This is different from common minerals. Mapping metals showed that metal element As is few. The dinosaur most likely belongs to natural death. This is different from Zigong dinosaurs which were found dead from poisoning. This method has been used to find that metals Fe and Mn are accrete, and the same is ture for Sr and Y. This study indicated that colloid granule Fe and Mn, as well as Sr and Y had opposite electric charges in lithification process of fossils. By this analysis, compound forms can be ascertained. Synchrotron light source x-ray fluorescence is a complementary method that shows mapping of metal elements at the dinosaur fossil, and is rapid, exact and intuitionist. This study shows that dinosaur fossil mineral imaging has a potential in reconstructing the paleoenvironment and ancient geology.

A gigantic bird-like dinosaur from the Late Cretaceous of China.

An evolutionary trend of decreasing size is present along the line to birds in coelurosaurian theropod evolution, but size increases are seen in many coelurosaurian subgroups, in which large forms are less bird-like. Here we report on a new non-avian dinosaur, Gigantoraptor erlianensis, gen. et sp. nov., from the Late Cretaceous Iren Dabasu Formation of Nei Mongol, China. Although it has a body mass of about 1,400 kg, a phylogenetic analysis positions this new taxon within the Oviraptorosauria, a group of small, feathered theropods rarely exceeding 40 kg in body mass. A histological analysis suggests that Gigantoraptor gained this size by a growth rate considerably faster than large North American tyrannosaurs such as Albertosaurus and Gorgosaurus. Gigantoraptor possesses several salient features previously unknown in any other dinosaur and its hind limb bone scaling and proportions are significantly different from those of other coelurosaurs, thus increasing the morphological diversity among dinosaurs. Most significantly, the gigantic Gigantoraptor shows many bird-like features absent in its smaller oviraptorosaurian relatives, unlike the evolutionary trend seen in many other coelurosaurian subgroups.

A new Chinese specimen indicates that 'protofeathers' in the Early Cretaceous theropod dinosaur Sinosauropteryx are degraded collagen fibres.

Alleged primitive feathers or protofeathers in the theropod dinosaur Sinosauropteryx have potentially profound implications concerning feather morphogenesis, evolution offlight, dinosaur physiology and perhaps even the origin of birds, yet their existence has never been adequately documented. We report on a new specimen of Sinosauropteryx which shows that the integumental structures proposed as protofeathers are the remains of structural fibres that provide toughness. The preservation in the proximal tail area reveals an architecture of closely associated bands offibres parallel to the tail's long axis, which originate from the skin. In adjacent more exposed areas, the fibres are short, fragmented and disorganized. Fibres preserved dorsal to the neck and back and in the distal part of the tail are the remains of a stiffening system of a frill, peripheral to the body and extending from the head to the tip of the tail. These findings are confirmed in the holotype Sinosauropteryx and NIGP 127587. The fibres show a striking similarity to the structure and levels of organization of dermal collagen. The proposal that these fibres are protofeathers is dismissed.