Non-avian Theropods Research Articles

Intelligent Objective Osteon Segmentation Based on Deep Learning

Histology is key to understand physiology, development, growth and even reproduction of extinct animals. However, the identification and interpretation of certain structures, such as osteons, medullary bone (MB), and Lines of Arrested Growth (LAGs), are not only based on personal judgments, but also require considerable labor for subsequent analysis. Due to the dearth of available specimens, only a few quantitative histological studies have been proceeded for limited dinosaur taxa, most of which focus primarily on their growth, namely, LAGs and other growth lines without much attention to other histological structures. Here we develop a deep convolutional neural network-based method for automated osteohistological segmentation. Raw images are firstly divided into sub-images and the borders are expanded to guarantee the osteon regions integrity. ResNet-50 is employed as feature extractor and atrous spatial pyramid pooling (ASPP) is used to capture multi-scale information. A dual-resolution segmentation strategy is designed to observe the primary and secondary osteon regions from the matrix background. Finally, a segmented map with different osteon regions is obtained. This deep convolutional neural network-based model is tested on a histological dataset derived from various taxa in Alvarezsauria, a highly specialized group of non-avian theropod dinosaurs. The results show that large-scale quantitative histological analysis can be achieved by neural network-based methods, and previously hidden information by traditional methods can be revealed. Phylogenetic mapping of osteon segmentation results suggests a developmental pathway towards miniaturized body sizes in the evolution of Alvarezsauria, which may resemble the transition from non-avian dinosaurs to birds.

A survey of the uncinate bone and other poorly known ossicles associated with the lacrimal/ectethmoid complex of the avian skull.

In several taxa of Neornithes (crown group birds), the lacrimal/ectethmoid complex exhibits small bones, the comparative osteology of which is poorly studied. Some of these ossicles-which are commonly known as uncinate bones (ossa uncinata or ossa lacrimopalatina)-were already described two centuries ago, but knowledge of their distribution and morphological variability in higher level clades is incomplete. In the present study, a detailed survey of the occurrence of uncinate bones is given, and these ossicles are for the first time reported in the gruiform Psophiidae, some Rallidae, and in the Otidiformes. Their presence in the latter taxon is of particular interest, because in current molecular analyses, the Otidiformes result as close relatives of the Musophagiformes, in which the uncinate bone is particularly large. The uncinate bones of most other neornithine clades, however, appear to have evolved multiple times independently through parallel evolution from the same ligamentous structures. A few earlier authors assumed that the uncinate bone is homologous to the ectopterygoid of non-avian theropods. Although this remains a viable hypothesis, more data on the occurrence of the ectopterygoid in Mesozoic birds are needed for well-supported conclusions. Here, it is noted that the ontogenetic development of the uncinate bone appears to be correlated with that of the ectethmoid, which is another bone in the skull of neornithine birds that is of unknown origin.

A newly recognized theropod assemblage from the Lewisville Formation (Woodbine Group; Cenomanian) and its implications for understanding Late Cretaceous Appalachian terrestrial ecosystems.

While the terrestrial fossil record of the mid-Cretaceous interval (Aptian to Cenomanian) in North America has been poorly studied, the recent focus on fossil localities from the western United States has offered a more detailed picture of vertebrate diversity, ecosystem dynamics and faunal turnover that took place on the western landmass of Laramidia. This is in stark contrast to the terrestrial record from the eastern landmass of Appalachia, where vertebrate fossils are rare and consist mostly of isolated and fragmentary remains. However, a detailed understanding of these fossil communities during this interval is necessary for comparison of the faunal patterns that developed during the opening of the Western Interior Seaway (WIS). The Woodbine Group of Texas is a Cenomanian age (95–100 mya) deposit consisting of shallow marine, deltaic, and terrestrial communities, which were only recently separated from their western counterparts. These deposits have yielded a wealth of vertebrate remains, yet non-avian theropods are still largely unknown. Recently, multiple localities in the Lewisville Formation of the Woodbine Group have yielded new non-avian theropod material, including numerous isolated teeth and postcranial remains. While largely fragmentary, this material is sufficiently diagnostic to identify the following taxa: a large-bodied carcharodontosaur, a mid-sized tyrannosauroid, a large ornithomimosaur, a large dromaeosaurine, a small dromaeosaurid, a small troodontid, and a small coelurosaur. Some of these groups represent the first occurrence for Appalachia and provide a broader understanding of a newly expanded faunal diversity for the Eastern landmass. The Lewisville Formation theropod fauna is similar in taxonomic composition to contemporaneous deposits in Laramidia, confirming that these groups were widespread across the continent prior to extension of the WIS. The Lewisville Formation documents the transitional nature of Cenomanian coastal ecosystems in Texas while providing additional details on the evolution of Appalachian communities shortly after WIS extension.

3D model related to the publication: Morphology and distribution of scales, dermal ossifications, and other non-feather integumentary structures in non-avialan theropod dinosaurs

3D model related to the publication: Morphology and distribution of scales, dermal ossifications, and other non-feather integumentary structures in non-avialan theropod dinosaurs

Morphology and distribution of scales, dermal ossifications, and other non-feather integumentary structures in non-avialan theropod dinosaurs.

Modern birds are typified by the presence of feathers, complex evolutionary innovations that were already widespread in the group of theropod dinosaurs (Maniraptoriformes) that include crown Aves. Squamous or scaly reptilian-like skin is, however, considered the plesiomorphic condition for theropods and dinosaurs more broadly. Here, we review the morphology and distribution of non-feathered integumentary structures in non-avialan theropods, covering squamous skin and naked skin as well as dermal ossifications. The integumentary record of non-averostran theropods is limited to tracks, which ubiquitously show a covering of tiny reticulate scales on the plantar surface of the pes. This is consistent also with younger averostran body fossils, which confirm an arthral arrangement of the digital pads. Among averostrans, squamous skin is confirmed in Ceratosauria (Carnotaurus), Allosauroidea (Allosaurus, Concavenator, Lourinhanosaurus), Compsognathidae (Juravenator), and Tyrannosauroidea (Santanaraptor, Albertosaurus, Daspletosaurus, Gorgosaurus, Tarbosaurus, Tyrannosaurus), whereas dermal ossifications consisting of sagittate and mosaic osteoderms are restricted to Ceratosaurus. Naked, non-scale bearing skin is found in the contentious tetanuran Sciurumimus, ornithomimosaurians (Ornithomimus) and possibly tyrannosauroids (Santanaraptor), and also on the patagia of scansoriopterygids (Ambopteryx, Yi). Scales are surprisingly conservative among non-avialan theropods compared to some dinosaurian groups (e.g. hadrosaurids); however, the limited preservation of tegument on most specimens hinders further interrogation. Scale patterns vary among and/or within body regions in Carnotaurus, Concavenator and Juravenator, and include polarised, snake-like ventral scales on the tail of the latter two genera. Unusual but more uniformly distributed patterning also occurs in Tyrannosaurus, whereas feature scales are present only in Albertosaurus and Carnotaurus. Few theropods currently show compelling evidence for the co-occurrence of scales and feathers (e.g. Juravenator, Sinornithosaurus), although reticulate scales were probably retained on the mani and pedes of many theropods with a heavy plumage. Feathers and filamentous structures appear to have replaced widespread scaly integuments in maniraptorans. Theropod skin, and that of dinosaurs more broadly, remains a virtually untapped area of study and the appropriation of commonly used techniques in other palaeontological fields to the study of skin holds great promise for future insights into the biology, taphonomy and relationships of these extinct animals.

A high-density Grallator assemblage from the Haman Formation (Cretaceous), Korea: implications for Cretaceous distribution of grallatorids in east Asia

ABSTRACT Monospecific assemblages of non-avian theropod tracks are relatively rare in the Cretaceous of Korea, where assemblages dominated by the tracks of diverse tetrapods (birds, sauropods, ornithopods, and pterosaurs) are well known and widely distributed in the Haman and Jindong formations (Hayang Group). To date, few high-density assemblages of theropod tracks have been documented. A Haman Formation site on Adu Island reveals a small high-density assemblage of Grallator tracks (~29 tracks/m2). The tracks, which indicate registration on a sandy substrate infused with microbial mat, are indistinguishable from Grallator tracks known from a few other Lower Cretaceous sites in China and east Asia. Early and Middle Jurassic Grallator-dominated assemblages are common in east Asia, and persist into the Late Jurassic- Early Cretaceous (Neocomian) transition in China, where they are typically only associated with sauropodomorph tracks. Post-Neocomian (Aptian-Albian) Cretaceous assemblages from Korea are much more diverse, and characterised by tracks of sauropod and ornithopod dinosaurs, non-avian and avian theropods (birds), pterosaurs and other tetrapods. Post-Neocomian assemblages dominated by theropod tracks, are rare and no large monospecific grallatorid assemblages are currently known.

New prospects on the cranial evolution of non-avialan paravian theropods based on geometric morphometrics

Abstract The cranial morphology of theropod dinosaurs has been used to examine the phylogeny, ontogeny, ecology and biomechanics of the clade. Previous studies have recognized that paedomorphosis and peramorphosis occurred multiple times throughout theropod evolution, with skull paedomorphism being one of the major changes during the transition from non-avialan theropods to birds. This study supplemented previous works with more detailed sampling of the morphological data of non-avialan paravians. Principal component analyses based on the cranial geometry confirm that the small-bodied non-avialan paravians have paedomorphic skulls compared with the early-diverging theropods, but independent peramorphosis is also observed in various groups. The secondary elongation of the preorbital portion of the cranium was present in both the late-diverging troodontids and the late-diverging dromaeosaurids, but it was achieved through different morphological configurations in these two lineages.

An exquisitely preserved in-ovo theropod dinosaur embryo sheds light on avian-like prehatching postures

SummaryDespite the discovery of many dinosaur eggs and nests over the past 100 years, articulated in-ovo embryos are remarkably rare. Here we report an exceptionally preserved, articulated oviraptorid embryo inside an elongatoolithid egg, from the Late Cretaceous Hekou Formation of southern China. The head lies ventral to the body, with the feet on either side, and the back curled along the blunt pole of the egg, in a posture previously unrecognized in a non-avian dinosaur, but reminiscent of a late-stage modern bird embryo. Comparison to other late-stage oviraptorid embryos suggests that prehatch oviraptorids developed avian-like postures late in incubation, which in modern birds are related to coordinated embryonic movements associated with tucking — a behavior controlled by the central nervous system, critical for hatching success. We propose that such pre-hatching behavior, previously considered unique to birds, may have originated among non-avian theropods, which can be further investigated with additional discoveries of embryo fossils.

A new dinosaur tracksite from the Lower Cretaceous (Aptian–Albian) Eumeralla Formation of Wattle Hill, Victoria, Australia: a preliminary investigation

ABSTRACT Dinosaur fossils of the Lower Cretaceous (Aptian–Albian) Eumeralla Formation (Otway Basin) of Victoria provide insights into the faunal and ichnofaunal diversity of southern hemisphere high latitude dinosaurs during the initial separation of Australia from Antarctica. To date, at least eight dinosaur fossil locations have been reported from this formation. In this paper, we describe a new tracksite locality along the coastline of Wattle Hill that represents the most westerly occurrence of dinosaur fossils in the state. Some of the Wattle Hill prints resemble tracks attributed to ornithopodan trackmakers from neighbouring tracksites of the Eumeralla Formation (Knowledge Creek, Browns Creek), while others resemble bird prints. The site also preserves a large non-avian theropod footprint that resembles Megalosauropus prints from the Lower Cretaceous (Valanginian–Barremian) Broome Sandstone, Western Australia that if correct, extends the temporospatial range of this ichnogenus. The Wattle Hill tracks vary in their erosion between and within tracksites, from a natural mould (concave epirelief), transmitted tracks, possibly penetrative tracks and infilled reliefs that resemble (but differ from) natural casts. With the constant weathering of the Eumeralla Formation exposures, it seems likely that additional dinosaur tracks fossils will be discovered with continued Victorian coastal explorations.

Large, unwebbed bird and bird‐like footprints from the Mesozoic and Cenozoic: a review of ichnotaxonomy and trackmaker affinity

The Mesozoic and Cenozoic track record of large birds (avian theropods) with footprint lengths (FL) > 10.0 cm is quite limited, whereas small tracks (FL < 10.0 cm) are abundant from the Early Cretaceous onwards. This large versus small threshold value is consistent with the track record of extant birds among which only ˜10% are large, and so is scaled appropriately to Class Aves. The proportion of large pachydactylous (thick-toed) or robust non-avian theropod ichnotaxa reported from the Mesozoic considerably exceeds the few large leptodactylous (thin-toed) or gracile forms such as Archaeornithipus and Magnoavipes, named, albeit controversially, to suggest avian affinity. The Cenozoic record of large avian tracks is quite different, with large pachydactylous tracks limited to a few Eocene-Oligocene (Palaeogene) ichnogenera (Ornithoformipes and Rivavipes), and rare unnamed Neogene and Holocene dinornithiform tracks. Thus, large flightless species are less well represented by tracks than body fossils. This suggests probable preservational bias in favour of waterbird or shorebird-like tracks in shoreline facies. However, the Cenozoic record of large leptodactylous tracks attributable to cranes, herons and related trackmakers is quite extensive, mostly confined to the Neogene and formally named to suggest representatives of extant avian clades capable of flight. These distribution patterns apparently reflect the dominance of large pachydactylous non-avian and avian theropods in the Mesozoic and early Palaeogene, and contrast with the rise of large, as well ongoing diversification of smaller leptodactylous Aequornithes in the Neogene. These distribution patterns in space and time likely reflect changing selection pressures as today's modern avifauna evolved.

Becoming-Dinosaur: Towards a Morphology of Creative Involution

Palaeontologist Jack Horner, along with molecular biologists and other experts, has been working on building an animal that he calls “the Chickenosaurus.” Horner’s purpose is to reverse-evolve a bird into a non-avian theropod by intervening on the genome expression of a chick’s embryo in ways that stimulate the formation of ancestral characteristics (activation of atavism). After a discussion of the project in its scientific detail, I offer a theoretical perspective on the topic that relies on concepts developed by Gilles Deleuze. In particular, I argue that the Chickenosaurus can be regarded as an eligible example of Deleuzian “becoming” and that it deserves philosophical attention in that it displays the “creative involution” through which novelty is produced.

The first edentulous ceratosaur from South America

The recognition of ontogenetic edentulism in the Jurassic noasaurid Limusaurus inextricabilis shed new light on the dietary diversity within Ceratosauria, a stem lineage of non-avian theropod dinosaurs known for peculiar craniomandibular adaptations. Until now, edentulism in Ceratosauria was exclusive to adult individuals of Limusaurus. Here, an exceptionally complete skeleton of a new toothless ceratosaur, Berthasaura leopoldinae gen. et sp. nov., is described from the Cretaceous aeolian sandstones of the Bauru Basin, Southern Brazil. The specimen resembles adult individuals of Limusaurus by the absence of teeth but based on the unfused condition of several elements (e.g., skull, vertebral column) it clearly represents an ontogenetically immature individual, indicating that it might never have had teeth. The phylogenetic analysis performed here has nested Berthasaura leopoldinae as an early-divergent Noasauridae, not closely related to Limusaurus. It represents the most complete non-avian theropod from the Brazilian Cretaceous and preserves the most complete noasaurid axial series known so far. Moreover, the new taxon exhibits many novel osteological features, uncommon in non-avian theropods, and unprecedented even among South American ceratosaurs. These include not only toothless jaws but also a premaxilla with cutting occlusal edge, and a slightly downturned rostral tip. This indicate that B. leopoldinae unlikely had the same diet as other ceratosaurs, most being regarded as carnivorous. As the ontogenetically more mature specimens of Limusaurus, Berthasaura might have been herbivorous or at least omnivorous, corroborating with an early evolutionary divergence of noasaurids from the ceratosaurian bauplan by disparate feeding modes.

Intraspecific facial bite marks in tyrannosaurids provide insight into sexual maturity and evolution of bird-like intersexual display

AbstractIntraspecific aggression, or agonism, is a widespread intrasexual selective behavior important to understanding animal behavioral ecology and reproductive systems. Such behavior can be studied either by direct observation or inferred from wound/scar frequency in extant species but is difficult to document in extinct taxa, limiting understanding of its evolution. Among extant archosaurs, crocodylians display extensive intrasexual aggression, whereas birds show extreme visual/vocal intersexual display. The evolutionary origin of this behavioral divergence, and pattern in non-avian dinosaurs, is unknown. Here we document the morphology, frequency, and ontogeny of intraspecific facial bite lesions (324 lesions) in a large sample of tyrannosaurids (202 specimens, 528 elements) to infer patterns of intraspecific aggression in non-avian theropods. Facial scars are consistent in position and orientation across tyrannosaurid species, suggesting bites were inflicted due to repeated/postured behavior. Facial scars are absent in young tyrannosaurids, first appear in immature animals (~50% adult skull length), are present in ~60% of the adult-sized specimens, and show aggressor:victim size isometry. The ontogenetic distribution of bite scars suggests agonistic behavior is associated with the onset of sexual maturity, and scar presence in approximately half the specimens may relate to a sexual pattern. Considered in a phylogenetic context, intraspecific bite marks are consistent and widely distributed in fossil and extant crocodyliforms and non-maniraptoriform theropods, suggesting a potential plesiomorphic behavior in archosaurs. Their absence in maniraptoriform theropods, including birds, may reflect a transition from boney cranial ornamentation and crocodylian-like intrasexual aggression to avian-like intersexual display with the evolution of pennaceous feathers.

The diet of early birds based on modern and fossil evidence and a new framework for its reconstruction.

ABSTRACTBirds are some of the most diverse organisms on Earth, with species inhabiting a wide variety of niches across every major biome. As such, birds are vital to our understanding of modern ecosystems. Unfortunately, our understanding of the evolutionary history of modern ecosystems is hampered by knowledge gaps in the origin of modern bird diversity and ecosystem ecology. A crucial part of addressing these shortcomings is improving our understanding of the earliest birds, the non‐avian avialans (i.e. non‐crown birds), particularly of their diet. The diet of non‐avian avialans has been a matter of debate, in large part because of the ambiguous qualitative approaches that have been used to reconstruct it. Here we review methods for determining diet in modern and fossil avians (i.e. crown birds) as well as non‐avian theropods, and comment on their usefulness when applied to non‐avian avialans. We use this to propose a set of comparable, quantitative approaches to ascertain fossil bird diet and on this basis provide a consensus of what we currently know about fossil bird diet. While no single approach can precisely predict diet in birds, each can exclude some diets and narrow the dietary possibilities. We recommend combining (i) dental microwear, (ii) landmark‐based muscular reconstruction, (iii) stable isotope geochemistry, (iv) body mass estimations, (v) traditional and/or geometric morphometric analysis, (vi) lever modelling, and (vii) finite element analysis to reconstruct fossil bird diet accurately. Our review provides specific methodologies to implement each approach and discusses complications future researchers should keep in mind. We note that current forms of assessment of dental mesowear, skull traditional morphometrics, geometric morphometrics, and certain stable isotope systems have yet to be proven effective at discerning fossil bird diet. On this basis we report the current state of knowledge of non‐avian avialan diet which remains very incomplete. The ancestral dietary condition in non‐avian avialans remains unclear due to scarce data and contradictory evidence in Archaeopteryx. Among early non‐avian pygostylians, Confuciusornis has finite element analysis and mechanical advantage evidence pointing to herbivory, whilst Sapeornis only has mechanical advantage evidence indicating granivory, agreeing with fossilised ingested material known for this taxon. The enantiornithine ornithothoracine Shenqiornis has mechanical advantage and pedal morphometric evidence pointing to carnivory. In the hongshanornithid ornithuromorph Hongshanornis only mechanical advantage evidence indicates granivory, but this agrees with evidence of gastrolith ingestion in this taxon. Mechanical advantage and ingested fish support carnivory in the songlingornithid ornithuromorph Yanornis. Due to the sparsity of robust dietary assignments, no clear trends in non‐avian avialan dietary evolution have yet emerged. Dietary diversity seems to increase through time, but this is a preservational bias associated with a predominance of data from the Early Cretaceous Jehol Lagerstätte. With this new framework and our synthesis of the current knowledge of non‐avian avialan diet, we expect dietary knowledge and evolutionary trends to become much clearer in the coming years, especially as fossils from other locations and climates are found. This will allow for a deeper and more robust understanding of the role birds played in Mesozoic ecosystems and how this developed into their pivotal role in modern ecosystems.

Taxonomic identification bias does not drive patterns of abundance and diversity in theropod dinosaurs.

The ability of palaeontologists to correctly diagnose and classify new fossil species from incomplete morphological data is fundamental to our understanding of evolution. Different parts of the vertebrate skeleton have different likelihoods of fossil preservation and varying amounts of taxonomic information, which could bias our interpretations of fossil material. Substantial previous research has focused on the diversity and macroevolution of non-avian theropod dinosaurs. Theropods provide a rich dataset for analysis of the interactions between taxonomic diagnosability and fossil preservation. We use specimen data and formal taxonomic diagnoses to create a new metric, the Likelihood of Diagnosis, which quantifies the diagnostic likelihood of fossil species in relation to bone preservation potential. We use this to assess whether a taxonomic identification bias impacts the non-avian theropod fossil record. We find that the patterns of differential species abundance and clade diversity are not a consequence of their relative diagnosability. Although there are other factors that bias the theropod fossil record that are not investigated here, our results suggest that patterns of relative abundance and diversity for theropods might be more representative of Mesozoic ecology than often considered.

Rare evidence for ‘gnawing-like’ behavior in a small-bodied theropod dinosaur

Mammalian carnivores show a higher degree of prey bone utilization relative to non-avian theropod dinosaurs, with this major ecological difference reflected in the frequency and morphology of tooth marks in modern and Cenozoic assemblages relative to Mesozoic ones. As such, prey bone utilization (i.e., gnawing, bone-breaking, osteophagy) may represent a key ecological strategy repeatedly exploited by mammalian carnivores but rarely in theropod dinosaurs. Here we describe an isolated adult-sized hadrosaurid pedal ungual (III-4) from the Dinosaur Park Formation (Campanian) of southern Alberta which shows a unique pattern of bite marks from a small- to medium-sized theropod dinosaur. Thirteen distinct tooth marks occur in a restricted area of the ungual, and the pattern suggests up to six repeated, high-power bites delivered to the bone. The tracemaker cannot be definitively identified, but was likely a dromaeosaurid or very young tyrannosaurid. Tooth marks on at least four other Dinosaur Park Formation hadrosaur pedal unguals are reported, but the overall frequency of occurrence in unguals (< 1%) is much lower than that reported for other bones. The pattern of tooth marks on this specimen deviates from most described theropods tooth marks, and given the low volume of meat associated with the ungual, may represent theropod prey bone utilization as part of late-stage carcass consumption, and a behavior similar to mammalian gnawing.

The smallest non-avian dinosaur track in China (Lower Jurassic, Sichuan Province)

ABSTRACT The smallest non-avian theropods have often attracted attention of palaeobiological and ichnological research. Here, we describe a diminutive tridactyl track (10.2 mm in length) from the Lower Jurassic Ma’anshan Member of the Ziliujing Formation, Wuli site, Sichuan Province, China. The Wuli specimen may be the smallest theropod track in the world. However, as with other diminutive tracks, it remains unresolved if it was made by a juvenile or by a small species. The body length of the trackmaker is estimated at 12 cm, equivalent to the size of a modern sparrow. Possibly the track belongs to a juvenile suggesting precociality or superprecociality.

Vertebral pneumaticity of the paravian theropod Unenlagia comahuensis, from the Upper Cretaceous of Patagonia, Argentina

Postcranial skeletal pneumaticity (PSP) characterizes extant birds. This feature is related to a series of air sacs connected to the lungs and prolonged in diverticula that invade bones internally. Previous works revealed that PSP was present along the line to birds, being distinctive of pterosaurs and saurischian dinosaurs. PSP is profuse in the vertebral column of sauropods and theropods and was very studied in sauropods, although scarcely in non-avian theropods. Here we analyze the vertebral pneumaticity of the unenlagiine theropod Unenlagia comahuensis, including the observation through CT scans. Unenlagiinae is a clade of southern dromaeosaurid theropods that is closely related to birds. The vertebral centra have lateral pneumatic foramina (lpf) within fossae (commonly termed ‘pleurocoels’) in middle and posterior dorsals, an unusual feature among extant birds and many non-avian theropods. Another possibly pneumatic fossa stands out at both sides of the neural spine base, which is not present in dorsals of other non-avian theropods, except the unenlagiine Unenlagia paynemili. CT scans revealed camellate tissue in the centra, consisting of small chambers separated by thin trabeculae. Camellae are also observed in the unenlagiines U. paynemili and Austroraptor cabazai, other dromaeosaurids, other coelurosaurs, and some non-coelurosaurian tetanurans. Instead, more primitive groups generally have camerae (larger chambers separated by scarce thick septa). Thus, a possible trend of the vertebral inner pneumaticity types is observed throughout non-avian theropod evolution, as indicated by previous authors. This study provides valuable information that helps to clarify this trend, not only in dromaeosaurids but also throughout theropod evolution.

Lower cretaceous avian-dominated, theropod, thyreophoran, pterosaur and turtle track assemblages from the Tugulu Group, Xinjiang, China: ichnotaxonomy and palaeoecology.

Rich tetrapod ichnofaunas, known for more than a decade, from the Huangyangquan Reservoir (Wuerhe District, Karamay City, Xinjiang) have been an abundant source of some of the largest Lower Cretaceous track collections from China. They originate from inland lacustrine clastic exposures of the 581–877 m thick Tugulu Group, variously divided into four formations and subgroups in the northwestern margin of the Junggar Basin. The large Huangyangquan track assemblages occur in the Lower layer/Subgroup II. Similarly-composed track assemblages also occur at the smaller Asphaltite site in the Upper Layer/Subgroup III. The Huangyangquan assemblages have yielded more than 1,500 identified tracks including abundant tracks of avian and non-avian theropods, pterosaurs and turtles and less abundant tracks of stegosaurs. Previous avian track identifications have been reassessed to conclude that Moguiornipes robustus is a taphotaxon and Koreanaornis dodsoni might be better accommodated in the ichnogenus Aquatilavipes which appears to be the dominant avian ichnotaxon. The avian track Ignotornis is also recognized and represents the first occurrence of this ichnogenus in China. Although the Huangyangquan assemblages lack some of the larger components (e.g., sauropodan and ornithopodan tracks) known from other Lower Cretaceous localities, the association of abundant tracks of smaller tetrapods (avian and non-avian theropods, pterosaurs and turtles) appears to be representative of lacustrine basin faunas of this region, and are an excellent example of the shorebird ichnocoenosis/ichnofacies concept. This is the first comprehensive review and re-analysis of an important Lower Cretaceous ecosystem.

Shifts in eggshell thickness are related to changes in locomotor ecology in dinosaurs.

Birds share an array of unique characteristics among extant land vertebrates. Among these, external and microstructural characteristics of extant bird eggs have been linked to changes in reproductive strategy that arose among non-avian theropod dinosaurs. More recently, differences in egg proportions recovered in crown birds relative to other dinosaurs were suggested as possibly linked to avian flight, but dense sampling close to its proposed origin was lacking. Here we assess the evolution of eggshell thickness in a targeted sample of 114 dinosaurs including birds, and test the relationship of eggshell thickness with potential life history correlates and locomotor mode using phylogenetic comparative methods. Only egg mass and flight are identified as significant predictors of eggshell thickness. While a high correlation between egg mass and eggshell thickness is expected, that relationship is much stronger in flying taxa, which show a significantly higher slope and lower residual variance than flightless species. This suggests stabilizing selection of eggshell thickness among theropods, as recovered for other traits in extant birds (e.g. genome size, metabolic rate). Within living birds, Eufalconimorphae present an apomorphic increase in relative eggshell thickness which remains unexplained, as few morphological synapomorphies of this clade have been identified.