The geometry of taking flight: Limb morphometrics in Mesozoic theropods

Theropoda was one of the most successful dinosaurian clades during the Mesozoic and has remained a dominant component of faunas throughout the Cenozoic, with nearly 10,000 extant representatives. The discovery of Archaeopteryx provides evidence that avian theropods evolved at least 155 million years ago and that more than half of the tenure of avian theropods on Earth was during the Mesozoic. Considering the major changes in niche occupation for theropods resulting from the evolution of arboreal and flight capabilities, we have analyzed forelimb and hindlimb proportions among nonmaniraptoriform theropods, nonavian maniraptoriforms, and basal avialans using reduced major axis regressions, principal components analysis, canonical variates analysis, and discriminant function analysis. Our study is the first analysis on theropod limb proportions to apply phylogenetic independent contrasts and size corrections to the data to ensure that all the data are statistically independent and amenable to statistical analyses. The three ordination analyses we performed did not show any significant groupings or deviations between nonavian theropods and Mesozoic avian forms when including all limb elements. However, the bivariate regression analyses did show some significant trends between individual elements that suggested evolutionary trends of increased forelimb length relative to hindlimb length from nonmaniraptoriform theropods to nonavian maniraptoriforms to basal avialans. The increase in disparity and divergence away from the nonavian theropod body plan is well documented within Cenozoic forms. The lack of significant groupings among Mesozoic forms when examining the entire theropod body plan concurrently suggests that nonavian theropods and avian theropods did not substantially diverge in limb proportions until the Cenozoic. J. Morphol. 276:152–166, 2015. © 2014 Wiley Periodicals, Inc.     

Oldest known avian footprints from Australia: Eumeralla Formation (Albian), Dinosaur Cove,Victoria

Two thin‐toed tridactyl tracks in a fluvial sandstone bed of the Eumeralla Formation (Albian) at Dinosaur Cove (Victoria, Australia) were likely made by avian trackmakers, making these the oldest known fossil bird tracks in Australia and the only Early Cretaceous ones from Gondwana. These tracks, which co‐occur on the same surface with a slightly larger nonavian theropod track, are distinguishable by their anisodactyl form, hallux impressions and wide digit II–IV divarication angles. A lengthy hallux impression and other deformational structures associated with one track indicate foot movement consistent with an abrupt stop, suggesting its tracemaker landed after either flight or a hop. The single nonavian theropod track is similar to other tracks described from the Eumeralla Formation at another locality. The avian footprints are larger than most Early Cretaceous avian tracks recorded worldwide, indicating sizeable enantiornithine or ornithurine species in formerly polar environments of Australia. The avian tracks further supplement scant body fossil evidence of Early Cretaceous birds in southern Australia, which includes a furcula from the Wonthaggi Formation. Because of this discovery, Dinosaur Cove, previously known for its vertebrate body fossils, is added to a growing list of Early Cretaceous vertebrate tracksites in southern Australia.     

Parallel evolution of theropod dinosaurs and birds

The hypothesis of the direct origin of birds from theropod dinosaurs has recently become widespread. Direct sisterly relationships between theropods and birds were assumed in the basis of random and formal synapomorphies, such as the number of caudal vertebrae, relative length of the humerus, and flattening of the dorsal margin of the pubis. In essence, this hypothesis is supported by the characters of theropods and birds, such as the presence of feathering, furcula, uncinate processes of ribs, pygostyle, double-condyled dorsal joint of the quadrate, and posteriorly turned pubis, which are recognized as homologies. Until recently, these characters have been regarded as avian apomorphies; however, they are presently known in various coelurosaurian groups. At the same time, they occur in various combinations in the Dromaeosauridae, Troodontidae, Oviraptoridae, Therizinosauridae, and Tyrannosauridae. None of the theropod groups possesses the entire set of these characters. This suggests that theropods and birds acquired them in parallel. Theropod dinosaurs and Sauriurae (Archaeornithes and Enantiornithes) show a number of important system synapomorphies, which indicate that they are closely related. Ornithurine birds lack such synapomorphies; however, their monophyly is supported by a large number of diagnostic characters. The hypothesis of independent origin of Sauriurae and Ornithurae is substantiated; the former are considered to have evolved from theropods in the Jurassic, while the latter deviated from a basal archosauromorph group in the Late Triassic. The hypothesis that birds existed in the Early Mesozoic is supported by the findings of small avian footprints in the Upper Triassic and Lower Jurassic of different continents.     

Morphological and functional diversity in therizinosaur claws and the implications for theropod claw evolution

Therizinosaurs are a group of herbivorous theropod dinosaurs from the Cretaceous of North America and Asia, best known for their iconically large and elongate manual claws. However, among Therizinosauria, ungual morphology is highly variable, reflecting a general trend found in derived theropod dinosaurs (Maniraptoriformes). A combined approach of shape analysis to characterize changes in manual ungual morphology across theropods and finite-element analysis to assess the biomechanical properties of different ungual shapes in therizinosaurs reveals a functional diversity related to ungual morphology. While some therizinosaur taxa used their claws in a generalist fashion, other taxa were functionally adapted to use the claws as grasping hooks during foraging. Results further indicate that maniraptoriform dinosaurs deviated from the plesiomorphic theropod ungual morphology resulting in increased functional diversity. This trend parallels modifications of the cranial skeleton in derived theropods in response to dietary adaptation, suggesting that dietary diversification was a major driver for morphological and functional disparity in theropod evolution.     

Feeding behaviour and bone utilization by theropod dinosaurs

Hone, D.W.E. & Rauhut, O.W.M. 2009: Feeding behaviour and bone utilization by theropod dinosaurs. Lethaia, Vol. 43, pp. 232–244. Examples of bone exploitation by carnivorous theropod dinosaurs are relatively rare, representing an apparent waste of both mineral and energetic resources. A review of the known incidences and possible ecological implications of theropod bone use concludes that there is currently no definitive evidence supporting the regular deliberate ingestion of bone by these predators. However, further investigation is required as the small bones of juvenile dinosaurs missing from the fossil record may be absent as a result of theropods preferentially hunting and consuming juveniles. We discuss implications for both hunting and feeding in theropods based on the existing data. We conclude that, like modern predators, theropods preferentially hunted and ate juvenile animals leading to the absence of small, and especially young, dinosaurs in the fossil record. The traditional view of large theropods hunting the adults of large or giant dinosaur species is therefore considered unlikely and such events rare. □Behaviour, carnivory, palaeoecology, predation, resource utilization.     

Biomechanical modeling and sensitivity analysis of bipedal running ability. II. Extinct taxa

Using an inverse dynamics biomechanical analysis that was previously validated for extant bipeds, I calculated the minimum amount of actively contracting hindlimb extensor muscle that would have been needed for rapid bipedal running in several extinct dinosaur taxa. I analyzed models of nine theropod dinosaurs (including birds) covering over five orders of magnitude in size. My results uphold previous findings that large theropods such as Tyrannosaurus could not run very quickly, whereas smaller theropods (including some extinct birds) were adept runners. Furthermore, my results strengthen the contention that many nonavian theropods, especially larger individuals, used fairly upright limb orientations, which would have reduced required muscular force, and hence muscle mass. Additional sensitivity analysis of muscle fascicle lengths, moment arms, and limb orientation supports these conclusions and points out directions for future research on the musculoskeletal limits on running ability. Although ankle extensor muscle support is shown to have been important for all taxa, the ability of hip extensor muscles to support the body appears to be a crucial limit for running capacity in larger taxa. I discuss what speeds were possible for different theropod dinosaurs, and how running ability evolved in an inverse relationship to body size in archosaurs.     

A new peculiar dinosaur egg,Sankofa pyrenaica oogen. nov. oosp. nov. from the Upper Cretaceous coastal deposits of the Aren Formation,south‐central Pyrenees,Lleida, Catalonia,Spain

Abstract: A new type of small, ovoid dinosaur egg, Sankofa pyrenaica oogen. nov. oosp. nov., with a prismatic type eggshell is described from upper Cretaceous (upper Campanian–Maastrichtian) deposits of the Montsec area, South Pyrenean Central Unit, Lleida, Catalonia, Spain. This egg type was sub‐vertically laid in only two rich monospecific sites of a single stratigraphic layer from coastal deposits of the Aren Formation, interpreted as an emerged beach ridge of a barrier island – lagoon depositional system. The size and shape of these eggs with their asymmetric poles are roughly similar to modern hen eggs, which is unusual in the Cretaceous fossil egg record. Its phylogenetic position clusters with bird and Troodontid eggs. A morphospace analysis of egg shapes shows the similarity of the new egg to a Campanian fossil bird egg from Argentina, both being intermediate between modern‐bird eggs and extinct nonavian theropod eggs. However, the eggshell microstructure of Sankofa pyrenaica differs from that of bird eggs in its incipient squamatic texture. It has a peculiar pattern of interlocking small crystals in the middle of the palisade layer, instead of the thick squamatic structure commonly present in modern avian eggshells. This new egg type is attributed to a small theropod, probably with a single oviduct like birds and whose mosaic distribution of features is a combination between that of birds and nonavian theropods. This enhances the arguments supporting the close phylogenetic relationships between both groups.     

Ventilatory mechanics from maniraptoran theropods to extant birds

Shared behavioural, morphological and physiological characteristics are indicative of the evolution of extant birds from nonavian maniraptoran dinosaurs. One such shared character is the presence of uncinate processes and respiratory structures in extant birds. Recent research has suggested a respiratory role for these processes found in oviraptorid and dromaeosaurid dinosaurs. By measuring the geometry of fossil rib cage morphology, we demonstrate that the mechanical advantage, conferred by uncinate processes, for movements of the ribs in the oviraptorid theropod dinosaur, Citipati osmolskae, basal avialan species Zhongjianornis yangi, Confuciusornis sanctus and the more derived ornithurine Yixianornis grabaui, is of the same magnitude as found in extant birds. These skeletal characteristics provide further evidence of a flow-through respiratory system in nonavian theropod dinosaurs and basal avialans, and indicate that uncinate processes are a key adaptation facilitating the ventilation of a lung air sac system that diverged earlier than extant birds.     

The evolutionary origin and diversification of feathers

Progress on the evolutionary origin and diversification of feathers has been hampered by conceptual problems and by the lack of plesiomorphic feather fossils. Recently, both of these limitations have been overcome by the proposal of the developmental theory of the origin of feathers, and the discovery of primitive feather fossils on nonavian theropod dinosaurs. The conceptual problems of previous theories of the origin of feathers are reviewed, and the alternative developmental theory is presented and discussed. The developmental theory proposes that feathers evolved through a series of evolutionary novelties in developmental mechanisms of the follicle and feather germ. The discovery of primitive and derived fossil feathers on a diversity of coelurosaurian theropod dinosaurs documents that feathers evolved and diversified in nonavian theropods before the origin of birds and before the origin of flight. The morphologies of these primitive feathers are congruent with the predictions of the developmental theory. Alternatives to the theropod origin of feathers are critique and rejected. Hypotheses for the initial function of feathers are reviewed. The aerodynamic theory of feather origins is falsified, but many other functions remain developmentally and phylogenetically plausible. Whatever their function, feathers evolved by selection for a follicle that would grow an emergent tubular appendage. Feathers are inherently tubular structures. The homology of feathers and scales is weakly supported. Feathers are composed of a suite of evolutionary novelties that evolved by the duplication, hierarchical organization, interaction, dissociation, and differentiation of morphological modules. The unique capacity for modular subdivision of the tubular feather follicle and germ has fostered the evolution of numerous innovations that characterize feathers. The evolution of feather keratin and the molecular basis of feather development are also discussed.     

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

Background

Fossil 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 Findings

Here 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/Significance

The 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.