New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers.

Discoveries of bird-like theropod dinosaurs and basal avialans in recent decades have helped to put the iconic 'Urvogel' Archaeopteryx into context and have yielded important new data on the origin and early evolution of feathers. However, the biological context under which pennaceous feathers evolved is still debated. Here we describe a new specimen of Archaeopteryx with extensive feather preservation, not only on the wings and tail, but also on the body and legs. The new specimen shows that the entire body was covered in pennaceous feathers, and that the hindlimbs had long, symmetrical feathers along the tibiotarsus but short feathers on the tarsometatarsus. Furthermore, the wing plumage demonstrates that several recent interpretations are problematic. An analysis of the phylogenetic distribution of pennaceous feathers on the tail, hindlimb and arms of advanced maniraptorans and basal avialans strongly indicates that these structures evolved in a functional context other than flight, most probably in relation to display, as suggested by some previous studies. Pennaceous feathers thus represented an exaptation and were later, in several lineages and following different patterns, recruited for aerodynamic functions. This indicates that the origin of flight in avialans was more complex than previously thought and might have involved several convergent achievements of aerial abilities.

Heterodonty and double occlusion in Manidens condorensis: 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.

Re-evaluation of the Haarlem Archaeopteryx and the radiation of maniraptoran theropod dinosaurs.

BACKGROUND: Archaeopteryx is an iconic fossil that has long been pivotal for our understanding of the origin of birds. Remains of this important taxon have only been found in the Late Jurassic lithographic limestones of Bavaria, Germany. Twelve skeletal specimens are reported so far. Archaeopteryx was long the only pre-Cretaceous paravian theropod known, but recent discoveries from the Tiaojishan Formation, China, yielded a remarkable diversity of this clade, including the possibly oldest and most basal known clade of avialan, here named Anchiornithidae. However, Archaeopteryx remains the only Jurassic paravian theropod based on diagnostic material reported outside China. RESULTS: Re-examination of the incomplete Haarlem Archaeopteryx specimen did not find any diagnostic features of this genus. In contrast, the specimen markedly differs in proportions from other Archaeopteryx specimens and shares two distinct characters with anchiornithids. Phylogenetic analysis confirms it as the first anchiornithid recorded outside the Tiaojushan Formation of China, for which the new generic name Ostromia is proposed here. CONCLUSIONS: In combination with a biogeographic analysis of coelurosaurian theropods and palaeogeographic and stratigraphic data, our results indicate an explosive radiation of maniraptoran coelurosaurs probably in isolation in eastern Asia in the late Middle Jurassic and a rapid, at least Laurasian dispersal of the different subclades in the Late Jurassic. Small body size and, possibly, a multiple origin of flight capabilities enhanced dispersal capabilities of paravian theropods and might thus have been crucial for their evolutionary success.

New insights into the vertebral Hox code of archosaurs.

Variation in axial formulae (i.e., number and identity of vertebrae) is an important feature in the evolution of vertebrates. Vertebrae at different axial positions exhibit a region-specific morphology. Key determinants for the establishment of particular vertebral shapes are the highly conserved Hox genes. Here, we analyzed Hox gene expression in the presacral vertebral column in the Nile crocodile in order to complement and extend a previous examination in the alligator and thus establish a Hox code for the axial skeleton of crocodilians in general. The newly determined expression of HoxA-4, C-5, B-7, and B-8 all revealed a crocodilian-specific pattern. HoxA-4 and HoxC-5 characterize cervical morphologies and the latter furthermore is associated with the position of the forelimb relative to the axial skeleton. HoxB-7 and HoxB-8 map exclusively to the dorsal vertebral region. The resulting expression patterns of these two Hox genes is the first description of their exact expression in the archosaurian embryo. Our comparative analyses of the Hox code in several amniote taxa provide new evidence that evolutionary differences in the axial skeleton correspond to changes in Hox gene expression domains. We detect two general processes: (i) expansion of a Hox gene's expression domain as well as (ii) a shift of gene expression. We infer that the ancestral archosaur Hox code may have resembled that of the crocodile. In association with the evolution of morphological traits, it may have been modified to patterns that can be observed in birds.

Correlation between Hox code and vertebral morphology in archosaurs.

The relationship between developmental genes and phenotypic variation is of central interest in evolutionary biology. An excellent example is the role of Hox genes in the anteroposterior regionalization of the vertebral column in vertebrates. Archosaurs (crocodiles, dinosaurs including birds) are highly variable both in vertebral morphology and number. Nevertheless, functionally equivalent Hox genes are active in the axial skeleton during embryonic development, indicating that the morphological variation across taxa is likely owing to modifications in the pattern of Hox gene expression. By using geometric morphometrics, we demonstrate a correlation between vertebral Hox code and quantifiable vertebral morphology in modern archosaurs, in which the boundaries between morphological subgroups of vertebrae can be linked to anterior Hox gene expression boundaries. Our findings reveal homologous units of cervical vertebrae in modern archosaurs, each with their specific Hox gene pattern, enabling us to trace these homologies in the extinct sauropodomorph dinosaurs, a group with highly variable vertebral counts. Based on the quantifiable vertebral morphology, this allows us to infer the underlying genetic mechanisms in vertebral evolution in fossils, which represents not only an important case study, but will lead to a better understanding of the origin of morphological disparity in recent archosaur vertebral columns.

Exceptionally preserved juvenile megalosauroid theropod dinosaur with filamentous integument from the Late Jurassic of Germany.

Recent discoveries in Asia have greatly increased our understanding of the evolution of dinosaurs' integumentary structures, revealing a previously unexpected diversity of "protofeathers" and feathers. However, all theropod dinosaurs with preserved feathers reported so far are coelurosaurs. Evidence for filaments or feathers in noncoelurosaurian theropods is circumstantial and debated. Here we report an exceptionally preserved skeleton of a juvenile megalosauroid, Sciurumimus albersdoerferi n. gen., n. sp., from the Late Jurassic of Germany, which preserves a filamentous plumage at the tail base and on parts of the body. These structures are identical to the type 1 feathers that have been reported in some ornithischians, the basal tyrannosaur Dilong, the basal therizinosauroid Beipiaosaurus, and, probably, in the basal coelurosaur Sinosauropteryx. Sciurumimus albersdoerferi represents the phylogenetically most basal theropod that preserves direct evidence for feathers and helps close the gap between feathers reported in coelurosaurian theropods and filaments in ornithischian dinosaurs, further supporting the homology of these structures. The specimen of Sciurumimus is the most complete megalosauroid yet discovered and helps clarify significant anatomical details of this important basal theropod clade, such as the complete absence of the fourth digit of the manus. The dentition of this probably early-posthatchling individual is markedly similar to that of basal coelurosaurian theropods, indicating that coelurosaur occurrences based on isolated teeth should be used with caution.

Reassessment of the enigmatic "Prestosuchus" loricatus (Archosauria: Pseudosuchia) from the Middle-Late Triassic of southern Brazil.

Our knowledge of the diversity and evolution of South American Triassic pseudosuchians has greatly improved in the past 15 years, due to new discoveries, but also to the revision of several historically important specimens. One of the earliest descriptions of pseudosuchians from the Triassic of Brazil stems from the classic work of Huene from the first half of the 20th century, who described several species, including such influential taxa as Rauisuchus tiradentes or Prestosuchus chiniquensis, which have recently been reviewed. The more poorly known proposed second species of Prestosuchus, P. loricatus, is the focus of the present work. The original material included some elements of the axial skeleton (cervical and caudal vertebrae, ribs, osteoderms) and the hindlimb (ischia, calcaneum, metatarsus), collected from the Dinodontosaurus Assemblage Zone of the Chiniquá area, west of São Pedro do Sul. "Prestosuchus" loricatus shows numerous differences to P. chiniquensis, including the shape of cervical neural spines, presence of epipophyses on the cervical vertebrae, presence of a pit in the iliac articulation of the ischium, lack of longitudinal furrows in the dorsolateral surface of the ischial shafts, the more slender calcaneal tuber and a less pronounced ventral pit in the calcaneum, and is thus referred to a new genus, Schultzsuchus gen. nov. Phylogenetic analysis indicates an early branching position within Poposauroidea for Schultzsuchus, making it the oldest known member of this clade in South America.

A juvenile pleurosaurid (Lepidosauria: Rhynchocephalia) from the Tithonian of the Mörnsheim Formation, Germany.

Late Jurassic rhynchocephalians from the Solnhofen Archipelago have been known for almost two centuries. The number of specimens and taxa is constantly increasing, but little is known about the ontogeny of these animals. The well-documented marine taxon Pleurosaurus is one of such cases. With over 15 described (and many more undescribed) specimens, there were no unambiguous juveniles so far. Some authors have argued that Acrosaurus, another common component of the Solnhofen Archipelago herpetofauna, might represent an early ontogenetic stage of Pleurosaurus, but the lack of proper descriptions for this taxon makes this assignment tentative, at best. Here, we describe the first unambiguous post-hatchling juvenile of Pleurosaurus and tentatively attribute it to Pleurosaurus cf. P. ginsburgi. The new specimen comes from the Lower Tithonian of the Mörnsheim Formation, Germany. This specimen is small, disarticulated, and incomplete, but preserves several of its craniomandibular bones and presacral vertebrae. It shares with Pleurosaurus a set of diagnostic features, such as an elongated and triangular skull, a low anterior flange in its dentition, and an elongated axial skeleton. It can be identified as a juvenile due to the presence of an unworn dentition, well-spaced posteriormost dentary teeth, a large gap between the last teeth and the coronoid process of the dentary, and poorly ossified vertebrae with unfused neural arches. Acrosaurus shares many anatomical features with both this specimen and Pleurosaurus, which could indicate that the two genera are indeed synonyms. The early ontogenetic stage inferred for the new Pleurosaurus specimen argues for an even earlier ontogenetic placement for specimens referred to Acrosaurus, the latter possibly pertaining to hatchlings.

A Middle Jurassic abelisaurid from Patagonia and the early diversification of theropod dinosaurs.

Abelisaurids are a clade of large, bizarre predatory dinosaurs, most notable for their high, short skulls and extremely reduced forelimbs. They were common in Gondwana during the Cretaceous, but exceedingly rare in the Northern Hemisphere. The oldest definitive abelisaurids so far come from the late Early Cretaceous of South America and Africa, and the early evolutionary history of the clade is still poorly known. Here, we report a new abelisaurid from the Middle Jurassic of Patagonia, Eoabelisaurus mefi gen. et sp. nov., which predates the so far oldest known secure member of this lineage by more than 40 Myr. The almost complete skeleton reveals the earliest evolutionary stages of the distinctive features of abelisaurids, such as the modification of the forelimb, which started with a reduction of the distal elements. The find underlines the explosive radiation of theropod dinosaurs in the Middle Jurassic and indicates an unexpected diversity of ceratosaurs at that time. The apparent endemism of abelisauroids to southern Gondwana during Pangean times might be due to the presence of a large, central Gondwanan desert. This indicates that, apart from continent-scale geography, aspects such as regional geography and climate are important to reconstruct the biogeographical history of Mesozoic vertebrates.

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 Cañadón 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 60-75 cm, Manidens furthermore confirms the small size of basal heterodontosaurids.

Discovery of a short-necked sauropod dinosaur from the Late Jurassic period of Patagonia.

Sauropod dinosaurs are one of the most conspicuous groups of Mesozoic terrestrial vertebrates. They show general trends towards an overall increase in size and elongation of the neck, by means of considerable elongation of the length of individual vertebrae and a cervical vertebra count that, in some cases, increases to 19 (ref. 1). The long neck is a particular hallmark of sauropod dinosaurs and is usually regarded as a key feeding adaptation. Here we describe a new dicraeosaurid sauropod, from the latest Jurassic period of Patagonia, that has a particularly short neck. With a neck that is about 40% shorter than in other known dicraeosaurs, this taxon demonstrates a trend opposite to that seen in most sauropods and indicates that the ecology of dicraeosaurids might have differed considerably from that of other sauropods. The new taxon indicates that there was a rapid radiation and dispersal of dicraeosaurids in the Late Jurassic of the Southern Hemisphere, after the separation of Gondwana from the northern continents by the late Middle Jurassic.

Sphenofontis velserae gen. et sp. nov., a new rhynchocephalian from the Late Jurassic of Brunn (Solnhofen Archipelago, southern Germany).

The Solnhofen Archipelago is well known for its fossil vertebrates of Late Jurassic age, among which figure numerous rhynchocephalian specimens, representing at least six and up to nine genera. A new taxon, named Sphenofontis velserae gen. et sp. nov., increases rhynchocephalian diversity in the Solnhofen Archipelago and is herein described based on a single, well-preserved specimen originating from the Late Kimmeridgian of the Brunn quarry, near Regensburg. The exquisite preservation of the holotype allowed a detailed description of the animal, revealing a skeletal morphology that includes both plesiomorphic and derived features within rhynchocephalians. Sphenofontis is herein referred to Neosphenodontia and tentatively to sphenodontine sphenodontids. It notably differs from all other rhynchocephalians known from the Jurassic of Europe, showing instead closer resemblance with the Middle Jurassic Cynosphenodon from Mexico and especially the extant Sphenodon. This is evidence for a wide distribution of taxa related to the extant tuatara early in the Mesozoic, and also for the presence of less-specialized rhynchocephalians coexisting with more derived forms during the earliest time in the history of the Solnhofen Archipelago.

Neuroanatomy of the spinosaurid Irritator challengeri (Dinosauria: Theropoda) indicates potential adaptations for piscivory.

Spinosauridae, a theropod group characterized by elongated snouts, conical teeth, enlarged forelimbs, and often elongated neural spines, show evidence for semiaquatic adaptations and piscivory. It is currently debated if these animals represent terrestrial carnivores with adaptations for a piscivorous diet, or if they largely lived and foraged in aquatic habitats. The holotype of Irritator challengeri, a nearly complete skull from the late Early Cretaceous Santana Formation of northeastern Brazil, includes one of the few preserved spinosaurid braincases and can provide insights into neuroanatomical structures that might be expected to reflect ecological affinities. We generated digital models of the neuroanatomical cavities within the braincase, using computer tomography (CT) data. The cranial endocast of Irritator is generally similar to that of other non-maniraptoriform theropods, with weakly developed distinctions of hindbrain and midbrain features, relatively pronounced cranial flexures and relatively long olfactory tracts. The endosseous labyrinth has a long anterior semicircular canal, a posteriorly inclined common crus and a very large floccular recess fills the area between the semicircular canals. These features indicate that Irritator had the ability for fast and well-controlled pitch-down head movements. The skull table and lateral semicircular canal plane are strongly angled to one another, suggesting a downward angling of approximately 45° of the snout, which reduces interference of the snout with the field of vision of Irritator. These neuroanatomical features are consistent with fast, downward snatching movements in the act of predation, such as are needed for piscivory.

Notes on the cheek region of the Late Jurassic theropod dinosaur Allosaurus.

Allosaurus, from the Late Jurassic of North America and Europe, is a model taxon for Jurassic basal tetanuran theropod dinosaurs. It has achieved an almost iconic status due to its early discovery in the late, 19th century, and due to the abundance of material from the Morrison Formation of the western U.S.A., making Allosaurus one of the best-known theropod taxa. Despite this, various aspects of the cranial anatomy of Allosaurus are surprisingly poorly understood. Here, we discuss the osteology of the cheek region, comprised by the jugal, maxilla, and lacrimal. This region of the skull is of importance for Allosaurus taxonomy and phylogeny, particularly because Allosaurus has traditionally been reconstructed with an unusual cheek configuration, and because the European species Allosaurus europaeus has been said to be different from North American material in the configuration of these bones. Based on re-examination of articulated and disarticulated material from a number of repositories, we show that the jugal participates in the antorbital fenestra, contradicting the common interpretation. The jugal laterally overlies the lacrimal, and forms an extended antorbital fossa with this bone. Furthermore, we document previously unrecorded pneumatic features of the jugal of Allosaurus.

Probable basal allosauroid from the early Middle Jurassic Cañadón Asfalto Formation of Argentina highlights phylogenetic uncertainty in tetanuran theropod dinosaurs.

Tetanurae, the most successful clade of theropod dinosaurs, including modern birds, split into three major clades early in their evolutionary history: Megalosauroidea, Coelurosauria, and Allosauroidea. The oldest tetanurans occur in the earliest Middle Jurassic, but the early fossil record of the clade is still poor. Here we report one of the oldest known and most complete pre-Late Jurassic tetanuran, the probable allosauroid Asfaltovenator vialidadi gen. et sp. nov., which has an unusual character combination, uniting features currently considered to be apomorphic of different tetanuran lineages. A phylogenetic analysis resulted in a monophyletic Carnosauria (Allosauroidea + Megalosauroidea), and the inclusion of the new taxon significantly changes topology within carnosaurs. The analysis shows concentrated homoplasy in proximal nodes at the base of Tetanurae, and a temporal peak at the Pliensbachian-Toarcian extinction event, recently identified as a potential driver of tetanuran radiation. These results highlight the complex morphological evolution in the early radiation of tetanuran theropods, in which convergences and parallelisms were extremely common. This pattern seems to be a common feature in rapid radiation events of major clades of vertebrates and might explain the common difficulties to unravel phylogenetic relationships of important lineages at the base of major clades.

The oldest Archaeopteryx (Theropoda: Avialiae): a new specimen from the Kimmeridgian/Tithonian boundary of Schamhaupten, Bavaria.

The iconic primeval bird Archaeopteryx was so far mainly known from the Altmühltal Formation (early Tithonian) of Bavaria, southern Germany, with one specimen having been found in the overlying Mörnsheim Formation. A new specimen (the 12th skeletal specimen) from the earliest Tithonian Painten Formation of Schamhaupten (Bavaria) represents the so far oldest representative of the genus. The new specimen shows several interesting anatomical details, including the presence of a postorbital in contact with the jugal, the presence of a separate prefrontal and coronoid, and opisthocoelous mid-cervical vertebrae. Based on observations on the new specimen, we discuss several problematic issues concerning Archaeopteryx, including the monophyly and diagnosis of the genus, the absence/presence of the sternum, the position of the gastralia, and variation in morphometrics and dental morphology in that genus. Based on a new diagnosis for the genus Archaeopteryx, the Berlin, Eichstätt, Solnhofen, Munich, Daiting, Thermopolis, 11th, and 12th specimens can be referred to this genus with high certainty. The Maxberg specimen is very probably also an Archaeopteryx, based on overall similarity, although none of the diagnostic characters can be evaluated with certainty. The ninth specimen ('chicken wing') might be Archaeopteryx, but cannot be referred to the genus with any certainty. In comparison with other paravians, the presence of distally thickened anterior pectoral ribs indicates that a rather large cartilagenous sternum was present in this taxon. In contrast to non-opisthopubic theropods, opisthopubic taxa, such as Archaeopteryx and many other paravians, have the posterior end of the gastral basket preserved at about half-length of the pubis, which might reflect the post-mortem collapse of enlarged abdominal air sacs in these taxa. Specimens that can be referred to Archaeopteryx show a high amount of variation, both in the morphometrics of the limb bones as well as in the dentition. In respect to the latter aspect, variation is found in tooth number, spacing, orientation, and morphology, with no two specimens showing the exact same pattern. The significance of this variation is unclear, and possible explanations reach from high intraspecific (and possibly ontogenetic and/or sexual dimorphic) variation to the possibility that the known specimens represent a 'species flock' of Archaeopteryx, possibly due to island speciation after the initial dispersal of the genus into the Solnhofen Archipelago.

The largest European theropod dinosaurs: remains of a gigantic megalosaurid and giant theropod tracks from the Kimmeridgian of Asturias, Spain.

The Kimmeridgian Vega, Tereñes and Lastres formations of Asturias have yielded a rich vertebrate fauna, represented by both abundant tracks and osteological remains. However, skeletal remains of theropod dinosaurs are rare, and the diversity of theropod tracks has only partially been documented in the literature. Here we describe the only non-dental osteological theropod remain recovered so far, an isolated anterior caudal vertebra, as well as the largest theropod tracks found. The caudal vertebra can be shown to represent a megalosaurine megalosaurid and represents the largest theropod skeletal remain described from Europe so far. The tracks are also amongst the largest theropod footprints reported from any setting and can be assigned to two different morphotypes, one being characterized by its robustness and a weak mesaxony, and the other characterized by a strong mesaxony, representing a more gracile trackmaker. We discuss the recently proposed distinction between robust and gracile large to giant theropod tracks and their possible trackmakers during the Late Jurassic-Berriasian. In the absence of complete pedal skeletons of most basal tetanurans, the identity of the maker of Jurassic giant theropod tracks is difficult to establish. However, the notable robustness of megalosaurine megalosaurids fits well with the described robust morphotypes, whereas more slender large theropod tracks might have been made by a variety of basal tetanurans, including allosaurids, metriocanthosaurids or afrovenatorine megalosaurids, or even exceptionally large ceratosaurs. Concerning osteological remains of large theropods from the Late Jurassic of Europe, megalosaurids seem to be more abundant than previously recognized and occur in basically all Jurassic deposits where theropod remains have been found, whereas allosauroids seem to be represented by allosaurids in Western Europe and metriacanthosaurids in more eastern areas. Short-term fluctuations in sea level might have allowed exchange of large theropods between the islands that constituted Europe during the Late Jurassic.

Endocast of the Late Triassic (Carnian) dinosaur Saturnalia tupiniquim: implications for the evolution of brain tissue in Sauropodomorpha.

The evolutionary history of dinosaurs might date back to the first stages of the Triassic (c. 250-240 Ma), but the oldest unequivocal records of the group come from Late Triassic (Carnian - c. 230 Ma) rocks of South America. Here, we present the first braincase endocast of a Carnian dinosaur, the sauropodomorph Saturnalia tupiniquim, and provide new data regarding the evolution of the floccular and parafloccular lobe of the cerebellum (FFL), which has been extensively discussed in the field of palaeoneurology. Previous studies proposed that the development of a permanent quadrupedal stance was one of the factors leading to the volume reduction of the FFL of sauropods. However, based on the new data for S. tupiniquim we identified a first moment of FFL volume reduction in non-sauropodan Sauropodomorpha, preceding the acquisition of a fully quadrupedal stance. Analysing variations in FFL volume alongside other morphological changes in the group, we suggest that this reduction is potentially related to the adoption of a more restricted herbivore diet. In this context, the FFL of sauropods might represent a vestigial trait, retained in a reduced version from the bipedal and predatory early sauropodomorphs.