Theropod Lineages Research Articles

The evolution of femoral morphology in giant non-avian theropod dinosaurs.

Theropods are obligate bipedal dinosaurs that appeared 230 million years ago and are still extant as birds. Their history is characterized by extreme variations in body mass, with gigantism evolving convergently between many lineages. However, no quantification of hindlimb functional morphology has shown if these body mass increases led to similar specializations between distinct lineages. Here we studied femoral shape variation across 41 species of theropods (n= 68 specimens) using a high-density 3D geometric morphometric approach. We demonstrated that the heaviest theropods evolved wider epiphyses and a more distally located fourth trochanter, as previously demonstrated in early archosaurs, along with an upturned femoral head and a mediodistal crest that extended proximally along the shaft. Phylogenetically informed analyses highlighted that these traits evolved convergently within six major theropod lineages, regardless of their maximum body mass. Conversely, the most gracile femora were distinct from the rest of the dataset, which we interpret as a femoral specialization to "miniaturization" evolving close to Avialae (bird lineage). Our results support a gradual evolution of known "avian" features, such as the fusion between lesser and greater trochanters and a reduction of the epiphyses' offset, independently from body mass variations, which may relate to a more "avian" type of locomotion (more knee-than hip-driven). The distinction between body mass variations and a more "avian" locomotion is represented by a decoupling in the mediodistal crest morphology, whose biomechanical nature should be studied to better understand the importance of its functional role in gigantism, miniaturization and higher parasagittal abilities.

Quantifying the gastral mass in Early Cretaceous ornithuromorphs (Aves, Ornithothoraces) from the Jehol avifauna

AbstractSome birds intentionally ingest stones to facilitate digestion of hard foodstuffs, a behaviour inherited from non‐avian dinosaurs and present in some of the earliest birds, as evidenced by clusters of gastroliths preserved within the abdominal cavity of a wide range of dinosaurs and Cretaceous birds. For the first time, high‐resolution computed laminographic and computed tomographic scans were used to reconstruct the gastral mass in two species of non‐neornithine ornithuromorph birds from the Lower Cretaceous Jehol Group. Four specimens of each taxon were analysed. Preservation of the gastral mass in most of these specimens is in situ and regarded as complete or nearly so. The number of gastroliths, their total volume, and their total mass relative to the estimated body mass were calculated for each specimen. The resultant gastral mass to body mass ratios fall within the range observed in extant birds, supporting previous inferences that the digestive system in non‐neornithine ornithuromorphs was comparable to that of extant taxa. Compared to available data for non‐volant non‐avian theropods, the gastral mass is proportionately smaller in birds suggesting that the evolution of flight constrained gastral mass size in the theropod lineage. Currently available data on gastral mass characteristics suggests that Iteravis ate larger food particles compared to Archaeorhynchus but cannot be used to determine diet more precisely. Better understanding of the relationship between gastral mass characteristics and food items across a broader range of extant taxa may provide an indirect but important method through which to infer diet and digestive function in archosaurs.

Evolutionary process toward avian-like cephalic thermoregulation system in Theropoda elucidated based on nasal structures.

It has long been discussed whether non-avian dinosaurs were physiologically closer to ectotherms or endotherms, with the internal nasal structure called the respiratory turbinate present in extant endotherms having been regarded as an important clue for this conundrum. However, the physiological function and relevance of this structure for dinosaur physiology are still controversial. Here, we found that the size of the nasal cavity relative to the head size of extant endotherms is larger than those of extant ectotherms, with that of the dromaeosaurid Velociraptor being below the extant endotherms level. The result suggests that a large nasal cavity accommodating a well-developed respiratory turbinate is primarily important as a thermoregulation apparatus for large brains characteristic of endothermic birds and mammals, and the nasal cavity of Velociraptor was apparently not large enough to carry out this role required for an endothermic-sized brain. In addition, a hypothesis that the enlargement of the nasal cavity for brain cooling has been associated with the skull modification in the theropod lineage toward modern birds is proposed herein. In particular, the reduction of the maxilla in derived avialans may have coincided with acquisition of the avian-like cephalic thermoregulation system.

A Jurassic Tibetan theropod tooth reveals dental convergency and its implication for identifying fragmentary fossils

<p><b>The Tibetan Plateau is among the least explored areas in terms of dinosaur paleontology in the world. Here, we report a dromaeosaurid-like tooth from the Middle Jurassic Dongdaqiao Formation at the eastern part of the plateau. The tooth exhibits dromaeosaurid dental features such as the absence of constriction at the cervix, a relatively small crown-to-base ratio, a strongly recurved apex, and it lacks large transverse and marginal undulations. Parsimonious phylogenetic analyses using both dentition-based and complete morphological matrices support a dromaeosaurid affinity for this tooth and place this Tibetan theropod as the sister taxon to</b> <i><b>Velociraptor</b></i><b>. However, Bayesian tip dating analyses based on the velociraptorine affinity of the Tibetan tooth place the majority of the diversifications of early-diverging Paravian within �C or earlier than �C the Middle Jurassic, which conflicts with the paravian fossil record. Furthermore, our discriminant analysis based on morphometric data places the Tibetan theropod within Metriacanthosauridae. These results may suggest that the broad resemblance between the Tibetan specimen and velociraptorine teeth is due to convergent evolution. Based on our results, other dromaeosaurid-like teeth reported from the Middle and Late Jurassic deposits throughout Eurasia could potentially be from non-dromaeosaurids. This study suggests the presence of an early diverged theropod lineage with a wide geographical distribution and potentially a dromaeosaurid-like dietary niche. This study also highlights the importance of using multiple lines of data to identify fragmentary fossils.</b></p>

Macroevolutionary trends in theropod dinosaur feeding mechanics

Theropod dinosaurs underwent some of the most remarkable dietary changes in vertebrate evolutionary history, shifting from ancestral carnivory1-3 to hypercarnivory4,5 and omnivory/herbivory,6-9 with some taxa eventually reverting to carnivory.10-12 The mandible is an important tool for food acquisition in vertebrates and reflects adaptations to feeding modes and diets.13,14 The morphofunctional modifications accompanying the dietary changes in theropod dinosaurs are not well understood because most of the previous studies focused solely on the cranium and/or were phylogenetically limited in scope,12,15-21 while studies that include multiple clades are usually based on linear measurements and/or discrete osteological characters.8,22 Given the potential relationship between macroevolutionary change and ontogenetic pattern,23 we explore whether functional morphological patterns observed in theropod mandibular evolution show similarities to the ontogenetic trajectory. Here, we use finite element analysis to study the mandibles of non-avialan coelurosaurian theropods and demonstrate how feeding mechanics vary between dietary groups and major clades. We reveal an overall reduction in feeding-induced stresses along all theropod lineages through time. This is facilitated by a post-dentary expansion and the development of a downturned dentary in herbivores and an upturned dentary in carnivores likely via the "curved bone effect." We also observed the same reduction in feeding-induced stress in an ontogenetic series of jaws of the tyrannosaurids Tarbosaurus and Tyrannosaurus, which is best attributed to bone functional adaptation. This suggests that this common tendency for structural strengthening of the theropod mandible through time, irrespective of diet, is linked to "functional peramorphosis" of bone functional adaptations acquired during ontogeny.

Aerodynamics Show Membranous-Winged Theropods Were a Poor Gliding Dead-End

The bizarre Scansoriopterygidae theropods Yi and Ambopteryx are the only dinosaurs thought to have had skin stretched between elongate fingers that form a potential membranous wing. While these structures have been suggested to have been used in aerial locomotion, this has never been shown to be the case. Demonstrating this function is of great importance because if these wings permitted flight it would support the growing view that theropod dinosaurs evolved flight multiple times in distinctive ways before the origin of modern birds. Here using Laser-Stimulated Fluorescence imaging we re-evaluate the anatomy of Yi and integrate this new information into several different equation-based aerodynamic calculations covering flight potential, methods of take-off (terrestrial-based running, arboreal launch), and powered versus gliding flight capabilities. For the first time we test whether Yi and Ambopteryx had flight capabilities and if so how efficient and effective it was. We also evaluate wing-based behaviours including wing-assisted incline running (WAIR). We find that Yi and Ambopteryx were highly unlikely to have any form of powered flight and had limited gliding abilities compared to similar-sized living and fossil gliders. We find that both taxa could not take-off from a terrestrial setting, supporting the idea that they were likely arboreal. Compared to similar-sized early-diverging fossil paravians, like Archaeopteryx and Microraptor, Scansoriopterygidae show significant deficiencies and differences in flapping-based locomotion models that indicate a much less competent flight stroke with limited aerial abilities. Combining these findings, our results show that the Scansoriopterygidae are not models for the early evolution of bird flight and that their structurally distinct and inefficient wings differed greatly from contemporaneous paravians including the earliest birds. Instead, we propose that Scansoriopterygidae represent a unique but failed flight architecture of non-avialan theropods. These findings conform with emerging data supporting multiple independent origins of flight in modern lineages. They show that the battle to capture the vertebrate aerial morphospace in the Middle to Late Jurassic was a dynamic and complex one not restricted to pterosaurs and only one theropod lineage.

The fast and the frugal: Divergent locomotory strategies drive limb lengthening in theropod dinosaurs

Limb length, cursoriality and speed have long been areas of significant interest in theropod paleobiology, since locomotory capacity, especially running ability, is critical in the pursuit of prey and to avoid becoming prey. The impact of allometry on running ability, and the limiting effect of large body size, are aspects that are traditionally overlooked. Since several different non-avian theropod lineages have each independently evolved body sizes greater than any known terrestrial carnivorous mammal, ~1000kg or more, the effect that such large mass has on movement ability and energetics is an area with significant implications for Mesozoic paleoecology. Here, using expansive datasets that incorporate several different metrics to estimate body size, limb length and running speed, we calculate the effects of allometry on running ability. We test traditional metrics used to evaluate cursoriality in non-avian theropods such as distal limb length, relative hindlimb length, and compare the energetic cost savings of relative hindlimb elongation between members of the Tyrannosauridae and more basal megacarnivores such as Allosauroidea or Ceratosauridae. We find that once the limiting effects of body size increase is incorporated there is no significant correlation to top speed between any of the commonly used metrics, including the newly suggested distal limb index (Tibia + Metatarsus/ Femur length). The data also shows a significant split between large and small bodied theropods in terms of maximizing running potential suggesting two distinct strategies for promoting limb elongation based on the organisms' size. For small and medium sized theropods increased leg length seems to correlate with a desire to increase top speed while amongst larger taxa it corresponds more closely to energetic efficiency and reducing foraging costs. We also find, using 3D volumetric mass estimates, that the Tyrannosauridae show significant cost of transport savings compared to more basal clades, indicating reduced energy expenditures during foraging and likely reduced need for hunting forays. This suggests that amongst theropods, hindlimb evolution was not dictated by one particular strategy. Amongst smaller bodied taxa the competing pressures of being both a predator and a prey item dominant while larger ones, freed from predation pressure, seek to maximize foraging ability. We also discuss the implications both for interactions amongst specific clades and Mesozoic paleobiology and paleoecological reconstructions as a whole.

Late Jurassic theropod dinosaur bones from the Langenberg Quarry (Lower Saxony, Germany) provide evidence for several theropod lineages in the central European archipelago.

Marine limestones and marls in the Langenberg Quarry provide unique insights into a Late Jurassic island ecosystem in central Europe. The beds yield a varied assemblage of terrestrial vertebrates including extremely rare bones of theropod from theropod dinosaurs, which we describe here for the first time. All of the theropod bones belong to relatively small individuals but represent a wide taxonomic range. The material comprises an allosauroid small pedal ungual and pedal phalanx, a ceratosaurian anterior chevron, a left fibula of a megalosauroid, and a distal caudal vertebra of a tetanuran. Additionally, a small pedal phalanx III-1 and the proximal part of a small right fibula can be assigned to indeterminate theropods. The ontogenetic stages of the material are currently unknown, although the assignment of some of the bones to juvenile individuals is plausible. The finds confirm the presence of several taxa of theropod dinosaurs in the archipelago and add to our growing understanding of theropod diversity and evolution during the Late Jurassic of Europe.

Two Early Cretaceous Fossils Document Transitional Stages in Alvarezsaurian Dinosaur Evolution

Highly specialized animals are often difficult to place phylogenetically. The Late Cretaceous members of Alvarezsauria represent such an example, having been posited as members of various theropod lineages, including birds [1-11]. A 70-million-year ghost lineage exists between them and the Late Jurassic putative alvarezsaurian Haplocheirus [12], which preserves so few derived features that its membership in Alvarezsauria has recently been questioned [13]. If Haplocheirus is indeed an alvarezsaurian, then the 70-million-year gap between Haplocheirus and other alvarezsaurians represents the longest temporal hiatus within the fossil record of any theropod subgroup [14]. Here we report two new alvarezsaurians from the Early Cretaceous of Western China that document successive, transitional stages in alvarezsaurian evolution. They provide further support for Haplocheirus as an alvarezsaurian and for alvarezsaurians as basal maniraptorans. Furthermore, they suggest that the early biogeographic history of the Alvarezsauria involved dispersals from Asia to other continents. The new specimens are temporally, morphologically, and functionally intermediate between Haplocheirus and other known alvarezsaurians and provide a striking example of the evolutionary transition from a typical theropod forelimb configuration (i.e., the relatively long arm and three-digit grasping hand of typical tetanuran form in early-branching alvarezsaurians) to a highly specialized one (i.e., the highly modified and shortened arm and one-digit digging hand of Late Cretaceous parvicursorines such as Linhenykus [1, 15]). Comprehensive analyses incorporating data from these new finds show that the specialized alvarezsaurian forelimb morphology evolved slowly and in a mosaic fashion during the Cretaceous.

The trophic habits of early birds

Although direct associations that reveal diet are extremely rare in the fossil record, the rich Lower Cretaceous Jehol Lagerstätte has produced dozens of specimens preserving ingested items, which together reveal important information regarding the early evolution of the avian alimentary canal. Direct evidence indicates Jeholornis and Sapeornis ate seeds and like living granivores utilized a gastric mill although only the more derived Sapeornis possessed a crop for food storage. Despite their smaller numbers in the Early Cretaceous, most direct evidence pertains to the Ornithuromorpha, indicating a structurally and functionally modern alimentary canal was present in even the earliest members of this clade. Similar evidence is altogether lacking in the Enantiornithes suggesting this clade was characterized by a primitive alimentary canal, potentially factoring into the ultimate extinction of this successful Cretaceous lineage. The role of the gizzard in the loss of teeth in Aves is more complex than in non-avian theropod lineages. As in neornithines, the presence of gastroliths in basal ornithuromorphs may not be entirely indicative of herbivory although this diet does appear to correlate with complete tooth loss in this clade. Positive selection for specialized tooth morphologies persists throughout the evolution of Aves and in the Ornithuromorpha appears to be linked to piscivory. Although represented by the greatest numbers, no direct indicator of diet preference is preserved in any confuciusornithiform or enantiornithine suggesting these clades utilized different nutritional strategies. The absence of teeth and gastroliths in all confuciusornithiforms suggest this lineage may have secondarily switched to a non-herbivorous, soft diet. Similarly, the absence of gizzard stones in enantiornithines also suggests a soft diet while their small body size suggests Early Cretaceous species would have fed on invertebrates. The diversity of recognized dental patterns including enamel specializations observed in Jehol enantiornithines suggests effective resource partitioning of available invertebrate faunas and that teeth had an active participation in enantiornithine feeding strategies.

Evolution of the hallux in non-avian theropod dinosaurs

ABSTRACTThe emergence of a reversed hallux (first pedal digit) in the theropod lineage is regarded as an important indicator of increasing grasping ability and possibly arboreality. However, functions of the pes with a plesiomorphic, non-reversed hallux in non-avian theropods, other than the major role in supporting and propelling the body during the stance phase, have not been fully addressed. To clarify hallucal function, 21 specimens of non-avian theropods were examined in this study. Ancestrally in Theropoda, the proximal end of the first metatarsal reached the ankle joint as in extant crocodilians. The size of the first metatarsal was then reduced, lost direct contact with the ankle joint, and shifted more plantarly in basal neotheropods. In articulated coelophysoid specimens, the proximal end of the first metatarsal attaches to the medioplantar margin of the second metatarsal, whereas the distal part of the first metatarsal and phalanges of the hallux lies on the plantar aspect of the pes. In four articulated specimens of the dromaeosaurid Velociraptor, preserved positions of the first metatarsal on the second metatarsal vary from the medial to plantar aspects, and correspondingly the rotational axis of the extension/flexion movement at its distal articulation ranges from mediolateral to dorsoplantar. Similar conditions are also observed in troodontid specimens, suggesting the presence of intermetatarsal mobility between the first and second metatarsals enabled by a convex proximal articular surface of the first metatarsal in deinonychosaurians similar to the one present in extant birds.Citation for this article: Hattori, S. 2016. Evolution of the hallux in non-avian theropod dinosaurs. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2016.1116995.

Dinosaur evolution. Sustained miniaturization and anatomical innovation in the dinosaurian ancestors of birds.

Recent discoveries have highlighted the dramatic evolutionary transformation of massive, ground-dwelling theropod dinosaurs into light, volant birds. Here, we apply Bayesian approaches (originally developed for inferring geographic spread and rates of molecular evolution in viruses) in a different context: to infer size changes and rates of anatomical innovation (across up to 1549 skeletal characters) in fossils. These approaches identify two drivers underlying the dinosaur-bird transition. The theropod lineage directly ancestral to birds undergoes sustained miniaturization across 50 million years and at least 12 consecutive branches (internodes) and evolves skeletal adaptations four times faster than other dinosaurs. The distinct, prolonged phase of miniaturization along the bird stem would have facilitated the evolution of many novelties associated with small body size, such as reorientation of body mass, increased aerial ability, and paedomorphic skulls with reduced snouts but enlarged eyes and brains.

The origins of Dinosauria: much ado about nothing

AbstractResearch this century has greatly improved our knowledge of the origin and early radiation of dinosaurs. The unearthing of several new dinosaurs and close outgroups from Triassic rocks from various parts of the world, coupled with improved phylogenetic analyses, has set a basic framework in terms of timing of events and macroevolutionary patterns. However, important parts of the early dinosauromorph evolutionary history are still poorly understood, rendering uncertain the phylogenetic position of silesaurids as either non‐dinosaur Dinosauriformes or ornithischians, as well as that of various early saurischians, such as Eoraptor lunensis and herrerasaurs, as either noneusaurischians or members of the sauropodomorph or theropod lineages. This lack of agreement in part derives from a patchy distribution of traits among early members of the main dinosauromorph lineages and requires a more meticulous assessment of characters and homologies than those recently conducted. Presently, the oldest uncontroversial dinosaur records come from Late Triassic (Carnian) rocks of South America, southern Africa and India, hinting at a south‐western Pangaea origin of the group. Besides, macroevolutionary approaches suggest that the rise of dinosaurs was a more gradual process than previously understood. Obviously, these tentative scenarios need to be tested by new fossil finds, which should also help close the major gaps recognized in the fossil record of Triassic dinosauromorphs.

Shake a tail feather: the evolution of the theropod tail into a stiff aerodynamic surface.

Theropod dinosaurs show striking morphological and functional tail variation; e.g., a long, robust, basal theropod tail used for counterbalance, or a short, modern avian tail used as an aerodynamic surface. We used a quantitative morphological and functional analysis to reconstruct intervertebral joint stiffness in the tail along the theropod lineage to extant birds. This provides new details of the tail’s morphological transformation, and for the first time quantitatively evaluates its biomechanical consequences. We observe that both dorsoventral and lateral joint stiffness decreased along the non-avian theropod lineage (between nodes Theropoda and Paraves). Our results show how the tail structure of non-avian theropods was mechanically appropriate for holding itself up against gravity and maintaining passive balance. However, as dorsoventral and lateral joint stiffness decreased, the tail may have become more effective for dynamically maintaining balance. This supports our hypothesis of a reduction of dorsoventral and lateral joint stiffness in shorter tails. Along the avian theropod lineage (Avialae to crown group birds), dorsoventral and lateral joint stiffness increased overall, which appears to contradict our null expectation. We infer that this departure in joint stiffness is specific to the tail’s aerodynamic role and the functional constraints imposed by it. Increased dorsoventral and lateral joint stiffness may have facilitated a gradually improved capacity to lift, depress, and swing the tail. The associated morphological changes should have resulted in a tail capable of producing larger muscular forces to utilise larger lift forces in flight. Improved joint mobility in neornithine birds potentially permitted an increase in the range of lift force vector orientations, which might have improved flight proficiency and manoeuvrability. The tail morphology of modern birds with tail fanning capabilities originated in early ornithuromorph birds. Hence, these capabilities should have been present in the early Cretaceous, with incipient tail-fanning capacity in the earliest pygostylian birds.

Regulation of bone-renal mineral and energy metabolism: the PHEX, FGF23, DMP1, MEPE ASARM pathway.

More than 300 million years ago, vertebrates emerged from the vast oceans to conquer gravity and the dry land. With this transition, new adaptations occurred that included ingenious changes in reproduction, waste secretion, and bone physiology. One new innovation, the egg shell, contained an ancestral protein (ovocleidin-116) that likely first appeared with the dinosaurs and was preserved through the theropod lineage in modern birds and reptiles. Ovocleidin-116 is an avian homolog of matrix extracellular phosphoglycoprotein (MEPE) and belongs to a group of proteins called short integrin-binding ligand-interacting glycoproteins (SIBLINGs). These proteins are all localized to a defined region on chromosome 5q in mice and chromosome 4q in humans. A unifying feature of SIBLING proteins is an acidic serine aspartate-rich MEPE-associated motif (ASARM). Recent research has shown that the ASARM motif and the released ASARM peptide have regulatory roles in mineralization (bone and teeth), phosphate regulation, vascularization, soft-tissue calcification, osteoclastogenesis, mechanotransduction, and fat energy metabolism. The MEPE ASARM motif and peptide are physiological substrates for PHEX, a zinc metalloendopeptidase. Defects in PHEX are responsible for X-linked hypophosphatemic rickets (HYP). There is evidence that PHEX interacts with another ASARM motif containing SIBLING protein, dentin matrix protein-1 (DMP1). DMP1 mutations cause bone and renal defects that are identical with the defects caused by a loss of PHEX function. This results in autosomal recessive hypophosphatemic rickets (ARHR). In both HYP and ARHR, increased FGF23 expression plays a major role in the disease and in autosomal dominant hypophosphatemic rickets (ADHR), FGF23 half-life is increased by activating mutations. ASARM peptide administration in vitro and in vivo also induces increased FGF23 expression. FGF23 is a member of the fibroblast growth factor (FGF) family of cytokines, which surfaced 500 million years ago with the boney fish (i.e., teleosts) that do not contain SIBLING proteins. In terrestrial vertebrates, FGF23, like SIBLING proteins, is expressed in the osteocyte. The boney fish, however, are an-osteocytic, so a physiological bone-renal link with FGF23 and the SIBLINGs was cemented when life ventured from the oceans to the land during the Triassic period, approximately 300 million years ago. This link has been revealed by recent research that indicates a competitive displacement of a PHEX-DMP1 interaction by an ASARM peptide that leads to increased FGF23 expression. This review discusses the new discoveries that reveal a novel PHEX, DMP1, MEPE, ASARM peptide, and FGF23 bone-renal pathway. This pathway impacts not only bone formation, bone-renal mineralization, and renal phosphate homeostasis but also energy metabolism. The study of this new pathway is relevant for developing therapies for several diseases: bone-teeth mineral loss disorders, renal osteodystrophy, chronic kidney disease and bone mineralization disorders (CKD-MBD), end-stage renal diseases, ectopic arterial-calcification, cardiovascular disease renal calcification, diabetes, and obesity.

Air‐filled postcranial bones in theropod dinosaurs: physiological implications and the ‘reptile’–bird transition

Pneumatic (air-filled) postcranial bones are unique to birds among extant tetrapods. Unambiguous skeletal correlates of postcranial pneumaticity first appeared in the Late Triassic (approximately 210 million years ago), when they evolved independently in several groups of bird-line archosaurs (ornithodirans). These include the theropod dinosaurs (of which birds are extant representatives), the pterosaurs, and sauropodomorph dinosaurs. Postulated functions of skeletal pneumatisation include weight reduction in large-bodied or flying taxa, and density reduction resulting in energetic savings during foraging and locomotion. However, the influence of these hypotheses on the early evolution of pneumaticity has not been studied in detail previously. We review recent work on the significance of pneumaticity for understanding the biology of extinct ornithodirans, and present detailed new data on the proportion of the skeleton that was pneumatised in 131 non-avian theropods and Archaeopteryx. This includes all taxa known from significant postcranial remains. Pneumaticity of the cervical and anterior dorsal vertebrae occurred early in theropod evolution. This 'common pattern' was conserved on the line leading to birds, and is likely present in Archaeopteryx. Increases in skeletal pneumaticity occurred independently in as many as 12 lineages, highlighting a remarkably high number of parallel acquisitions of a bird-like feature among non-avian theropods. Using a quantitative comparative framework, we show that evolutionary increases in skeletal pneumaticity are significantly concentrated in lineages with large body size, suggesting that mass reduction in response to gravitational constraints at large body sizes influenced the early evolution of pneumaticity. However, the body size threshold for extensive pneumatisation is lower in theropod lineages more closely related to birds (maniraptorans). Thus, relaxation of the relationship between body size and pneumatisation preceded the origin of birds and cannot be explained as an adaptation for flight. We hypothesise that skeletal density modulation in small, non-volant, maniraptorans resulted in energetic savings as part of a multi-system response to increased metabolic demands. Acquisition of extensive postcranial pneumaticity in small-bodied maniraptorans may indicate avian-like high-performance endothermy.

Bird-Like Anatomy, Posture, and Behavior Revealed by an Early Jurassic Theropod Dinosaur Resting Trace

BackgroundFossil tracks made by non-avian theropod dinosaurs commonly reflect the habitual bipedal stance retained in living birds. Only rarely-captured behaviors, such as crouching, might create impressions made by the hands. Such tracks provide valuable information concerning the often poorly understood functional morphology of the early theropod forelimb.Methodology/Principal FindingsHere we describe a well-preserved theropod trackway in a Lower Jurassic (∼198 million-year-old) lacustrine beach sandstone in the Whitmore Point Member of the Moenave Formation in southwestern Utah. The trackway consists of prints of typical morphology, intermittent tail drags and, unusually, traces made by the animal resting on the substrate in a posture very similar to modern birds. The resting trace includes symmetrical pes impressions and well-defined impressions made by both hands, the tail, and the ischial callosity.Conclusions/SignificanceThe manus impressions corroborate that early theropods, like later birds, held their palms facing medially, in contrast to manus prints previously attributed to theropods that have forward-pointing digits. Both the symmetrical resting posture and the medially-facing palms therefore evolved by the Early Jurassic, much earlier in the theropod lineage than previously recognized, and may characterize all theropods.

An integrative phylogenetic and extrapolatory approach to the reconstruction of dromaeosaur (Theropoda: Eumaniraptora) shoulder musculature

Dromaeosauridae is the sister taxon of the Avialae; thus, an investigation of dromaeosaur shoulder girdle musculature and forelimb function provides substantial information regarding changes in the size and performance of the theropod shoulder girdle musculature leading to avian powered flight. Twenty-two shoulder girdle muscles were reconstructed for the dromaeosaurid shoulder apparatus, based on phylogenetic inference, which involves the comparison of lepidosaurian, crocodilian and avian musculature, and extrapolatory inference, which involves a secondary comparison with functional analogues of theropods. In addition to these comparative methodologies, osteological correlates of shoulder musculature preserved in eumaniraptorans are identified, and comparisons with those of extant archosaurs allow these muscles to be definitively inferred in dromaeosaurids. This muscle reconstruction provides a foundation for subsequent investigation of differences in muscular attachment and function, based on scapulocoracoid morphology, across the theropod lineage leading to birds. © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 146, 301–344.

Theropod pedal unguals from the Late Cretaceous (Cenomanian) of Morocco, Africa

Two different theropod pedal unguals from the Upper Cretaceous (Cenomanian) of Tafilalt, Morocco are described. One of them belongs to a slender kind of abelisauroid theropod. Comparison with other members of this group reveals that abelisauroid unguals show high morphological diversity in congruence with what is known on other parts of the skeleton. Another large-sized ungual resembles that described as “Spinosaurus B” by Stromer (1934), and is here referred with caution to Theropoda, although ignoring to which theropod lineage it belongs. Comparisons between northern (e.g., Sahara, Africa and Brazil) and southern (e.g., Patagonia) Gondwanan Cretaceous dinosaur faunas reveal some differences leading to the conclusion that Gondwanan faunas were not uniform across this supercontinent.

Mid-Cretaceous turnover of saurischian dinosaur communities: evidence from the Neuquén Basin

Abstract Successional changes are recorded among saurischian dinosaur faunas throughout the Cretaceous of the Neuquén Basin. Both sauropod and theropod lineages present in NW Patagonia show comparable transformations and indicate that some clades did not survive the Late Turonian boundary. Similar patterns of changes are observed in other Patagonian sedimentary basins. Diplodocoid sauropods and carcharodontosaurid theropods are not yet recorded in levels younger than Turonian, while titanosaurians and abelisauroids are present before and after this period. Important floral changes during the mid-Cretaceous, mainly related to the proliferation of angiosperms, could be one element that influenced some dinosaurian associations.