Dinosaur speed demon: the caudal musculature of Carnotaurus sastrei and implications for the evolution of South American abelisaurids

In the South American abelisaurids Carnotaurus sastrei, Aucasaurus garridoi, and, to a lesser extent Skorpiovenator bustingorryi, the anterior caudal ribs project at a high dorsolateral inclination and have interlocking lateral tips. This unique morphology facilitated the expansion of the caudal hypaxial musculature at the expense of the epaxial musculature. Distinct ridges on the ventrolateral surfaces of the caudal ribs of Aucasaurus garridoi are interpreted as attachment scars from the intra caudofemoralis/ilio-ischiocaudalis septa, and confirm that the M. caudofemoralis of advanced South American abelisaurids originated from a portion of the caudal ribs. Digital muscle models indicate that, relative to its overall body size, Carnotaurus sastrei had a substantially larger M. caudofemoralis than any other theropod yet studied. In most non-avian theropods, as in many extant sauropsids, the M. caudofemoralis served as the primary femoral retractor muscle during the locomotive power stroke. This large investment in the M. caudofemoralis suggests that Carnotaurus sastrei had the potential for great cursorial abilities, particularly short-burst sprinting. However, the tightly interlocking morphology of the anterior caudal vertebrae implies a reduced ability to make tight turns. Examination of these vertebral traits in evolutionary context reveals a progressive sequence of increasing caudofemoral mass and tail rigidity among the Abelisauridae of South America.     

Kinematic model of tyrannosaurid (Dinosauria: Theropoda) arctometatarsus function

We present a hypothesis of tyrannosaurid foot function termed the "tensile keystone model," in which the triangular central metatarsal and elastic ligaments dynamically strengthened the foot. The tyrannosaurid arctometatarsus, in which the central metatarsal is proximally constricted, displays osteological correlates of distal intermetatarsal ligaments. The distal wedge-like imbrication of tyrannosaurid metatarsals indicates that rebounding ligaments drew the outer elements towards the middle digit early in the stance phase, unifying the arctometatarsus under high loadings. This suggests increased stability and resistance to dissociation and implies, but does not demonstrate, greater agility than in large theropods without an arctometatarsus.     

THE TAXONOMIC STATUS OF MEGALOSAURUS BUCKLANDII (DINOSAURIA, THEROPODA) FROM THE MIDDLE JURASSIC OF OXFORDSHIRE, UK

Abstract:  The lectotype of the Middle Jurassic theropod dinosaur Megalosaurus bucklandii , a right dentary, can be diagnosed on the basis of two unique characters: a longitudinal groove on the ventral part of the lateral surface of the dentary and a slit-like anterior Meckelian foramen. This taxon, the first dinosaur to be scientifically described, is therefore valid. Currently, however, no further material can be referred to this species with any certainty. Megalosaurus bucklandii occupies an uncertain systematic position but is not an abelisaurid or coelophysoid. Additionally, it does not possess the diagnostic dentary characters that are present in all known spinosauroids. Owing to this uncertainty, use of the family Megalosauridae should be discontinued until such time as its systematic position becomes clearer.     

The skull evolution of oviraptorosaurian dinosaurs: the role of niche partitioning in diversification

Oviraptorosaurs are bird‐like theropod dinosaurs that thrived in the final pre‐extinction ecosystems during the latest Cretaceous, and the beaked, toothless skulls of derived species are regarded as some of the most peculiar among dinosaurs. Their aberrant morphologies are hypothesized to have been caused by rapid evolution triggered by an ecological/biological driver, but little is known about how their skull shapes and functional abilities diversified. Here, we use quantitative techniques to study oviraptorosaur skull form and mandibular function. We demonstrate that the snout is particularly variable, that mandibular form and upper/lower beak form are significantly correlated with phylogeny, and that there is a strong and significant correlation between mandibular function and mandible/lower beak shape, suggesting a form–function association. The form–function relationship and phylogenetic signals, along with a moderate allometric signal in lower beak form, indicate that similar mechanisms governed beak shape in oviraptorosaurs and extant birds. The two derived oviraptorosaur clades, oviraptorids and caenagnathids, are significantly separated in morphospace and functional space, indicating that they partitioned niches. Oviraptorids coexisting in the same ecosystem are also widely spread in morphological and functional space, suggesting that they finely partitioned feeding niches, whereas caenagnathids exhibit extreme disparity in beak size. The diversity of skull form and function was likely key to the diversification and evolutionary success of oviraptorosaurs in the latest Cretaceous.     

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.     

The trackmaker of Apatopus (Late Triassic, North America): implications for the evolution of archosaur stance and gait

Abstract:  For some decades, a major focus of research has been on how locomotor modes changed in some archosaurian reptiles from a more or less 'sprawling' to an 'erect' posture, whether there were discrete intermediate stages, and how many times 'erect' posture evolved. The classic paradigm for the evolution of stance and gait in archosaurs, a three-stage transition from sprawling to 'semi-erect' to erect posture, has been replaced by a subtler understanding of a continuum of changing limb joint angles. We suggest a further separation of terminology related to stance vs. gait so as not to entail different processes: 'sprawling' and 'erect' should refer to continua of stance; 'rotatory' and 'parasagittal' are more appropriate ends of a continuum that describes the motions of gait. We show that the Triassic trackway Apatopus best fits the anatomy and proportions of phytosaurs, based on a new reconstruction of their foot skeleton; it is less likely to have been made by another pseudosuchian or non-archosaurian archosauromorph. Moreover, the trackmaker was performing the high walk. A phytosaurian trackmaker would imply that the common ancestor of pseudosuchians, and therefore archosaurs could approximate the high walk (depending on phylogeny), and if so, erect stance and parasagittal gait did not evolve independently in pseudosuchians and ornithosuchians, although the kinematic mechanisms differed in the two groups. It remains to be seen how far outside Archosauria, if at all, more or less erect posture and parasagittal gait may have evolved.     

Poekilopleuron bucklandii, the theropod dinosaur from the Middle Jurassic (Bathonian) of Normandy

Poekilopleuron bucklandii , described by Eudes–Deslongchamps in 1838, is one of the earliest discovered dinosaurs. Although incomplete, it is one of the best preserved Middle Jurassic theropods known from Europe. Unfortunately, the only specimen of P. bucklandii , housed in the Musée de la Faculté des Sciences de Caen, was destroyed during World War II. However, casts of some parts of the type skeleton have been found in the collections of the Museum National d'Histoire Naturelle, Paris. These casts and Eudes–Deslongchamps' monograph are used to redescribe the specimen. Poekilopleuron shares one synapomorphy with the Spinosauroidea and we tentatively assign it to that clade. The possible synonymy between Poekilopleuron and Megalosaurus is examined and we conclude that Megalosaurus is a nomen dubium and that the name should be restricted to the type dentary.     

小盗龙亚科(兽脚亚目:驰龙科)——新标本及简评复合骨在兽脚亚目中的演化（英文）

小盗龙亚科是驰龙科兽脚类恐龙中新近发现的一个亚群,多数属种产自中国辽宁西部下白垩统热河群。本文描记一件产自辽宁省朝阳县大平房地区九佛堂组(热河群上段)的小盗龙亚科的新标本,该标本具有一些未曾报道过的有趣形态特征。尤其值得关注的是,一些特征呈现出介于中国鸟龙和小盗龙之间的过渡状态,其中部分来自牙齿和坐骨。这些特征突出显示了在更接近典型驰龙类的中国鸟龙和具有许多伤齿龙科特征的小盗龙之间存在着一系列的形态变异。然而,这些变异特征在分类上的意义尚未得到充分评估。特别值得一提的是该标本中相互愈合的耻骨和肠骨,这一特征有助于阐明复合骨在兽脚类恐龙中的演化模式。初步分析显示许多鸟类的复合骨都是通过在过型形成过程中各组分的顺序愈合形成的。     

Limusaurus inextricabilis (Theropoda: Ceratosauria) gives a hand to evolutionary teratology: a complementary view on avian manual digits identities

It is widely accepted that birds are rooted within theropod dinosaurs. However, there is controversy between palaeontological and developmental data regarding manual digit identities of birds and their tetanuran ancestors (I, II and III vs. II, III and IV). To resolve this conflict, the principle of a frame‐shift has been considered. Identities of digits I–III would develop on condensations 2–4. Nevertheless, the discovery of the basal Ceratosauria Limusaurus inextricabilis has been used as a reference to define the digital identity of Tetanurae as II–IV. The new concept of evolutionary teratology states that certain anatomical structures identified in evolutionary lineages are viable developmental anomalies (‘adaptive’ or not), becoming part of the considered groups. The features of Limusaurus' forelimb match teratological characterization. This diagnosis, associated with the variations previously identified in derived Ceratosauria taxa (Carnotaurinae), underline an anatomical and developmental independence (regarding evolutionary conserved mechanisms) compared with Tetanurae and therefore birds. Consequently, Limusaurus should not be used as a reference concerning the identity of avian manuals digits. Evolutionary teratology supports identities I, II and III of the tetanuran manus via a frame‐shift that did not occur in the Ceratosauria lineage. © 2016 The Linnean Society of London     

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.     

The skull of Monolophosaurus jiangi (Dinosauria: Theropoda) and its implications for early theropod phylogeny and evolution

The Middle Jurassic was a critical time in the evolution of theropod dinosaurs, highlighted by the origination and initial radiation of the large‐bodied and morphologically diverse Tetanurae. Middle Jurassic tetanurans are rare, but have been described from Europe, South America and China. In particular, China has yielded a number of potential basal tetanurans, but these have received little detailed treatment in the literature. Chief among these is Monolophosaurus jiangi, known from a single skeleton that includes a nearly complete and well‐preserved skull characterized by a bizarre cranial crest. Here, we redescribe the skull of Monolophosaurus, which is one of the most complete basal tetanuran skulls known and the only quality source of cranial data for Middle Jurassic Chinese theropods. The cranial crest is atomized into a number of autapomorphic features and several characters confirm the tetanuran affinities of Monolophosaurus. However, several features suggest a basal position within Tetanurae, which contrasts with most published cladistic analyses, which place Monolophosaurus within the more derived Allosauroidea. Cranial characters previously used to diagnose Allosauroidea are reviewed and most are found to have a much wider distribution among Theropoda, eroding an allosauroid position for Monolophosaurus and questioning allosauroid monophyly. The use of phylogenetic characters relating to theropod cranial crests is discussed and a protocol for future use is given. The systematic position of Guanlong wucaii is reviewed, and a basal tyrannosauroid affinity is upheld contrary to one suggestion of a close relationship between this taxon and Monolophosaurus. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 573–607.     

A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Bayan Mandahu,Inner Mongolia

Abstract: A new oviraptorid is described on the basis of a partial forelimb collected from the Upper Cretaceous redbeds of Bayan Mandahu, Inner Mongolia. Machairasaurus leptonychus, gen. et sp. nov. is diagnosed by slender, weakly curved manual unguals, reduced flexor tubercles, penultimate phalanges that are subequal in length to the preceding phalanges, and short, robust manual digits. Machairasaurus is found to be a member of the Ingeniinae, along with Ingenia yanshini, Heyuannia huangi, Conchoraptor gracilis, and Nemegtomaia barsboldi. Machairasaurus exhibits unusual proportions of the manus, suggesting that the manus was not primarily used to grasp prey. Instead, Machairasaurus and other oviraptorids are likely to have fed largely on plant material. The recognition of a previously unknown oviraptorid at Bayan Mandahu provides further evidence that the Bayan Mandahu dinosaur assemblage is distinct from that found at the Djadokhta Formation exposures at Bayn Dzak, Tugriken Shireh, and Ukhaa Tolgod. Given that these localities are separated by just a few hundred kilometres and represent similar palaeoenvironments, marked differences in the fauna suggest that the Bayan Mandahu Formation of Inner Mongolia is not coeval with the known Djadokhta localities in Mongolia.     

The evolution of cranial form and function in theropod dinosaurs: insights from geometric morphometrics

Theropod dinosaurs, an iconic clade of fossil species including Tyrannosaurus and Velociraptor, developed a great diversity of body size, skull form and feeding habits over their 160+ million year evolutionary history. Here, we utilize geometric morphometrics to study broad patterns in theropod skull shape variation and compare the distribution of taxa in cranial morphospace (form) to both phylogeny and quantitative metrics of biting behaviour (function). We find that theropod skulls primarily differ in relative anteroposterior length and snout depth and to a lesser extent in orbit size and depth of the cheek region, and oviraptorosaurs deviate most strongly from the "typical" and ancestral theropod morphologies. Noncarnivorous taxa generally fall out in distinct regions of morphospace and exhibit greater overall disparity than carnivorous taxa, whereas large-bodied carnivores independently converge on the same region of morphospace. The distribution of taxa in morphospace is strongly correlated with phylogeny but only weakly correlated with functional biting behaviour. These results imply that phylogeny, not biting function, was the major determinant of theropod skull shape.     

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.     

Dientes de Theropoda (Dinosauria: Saurischia) de la Formación Bajo Barreal, Cretácico Superior, Provincia del Chubut, Argentina

Theropod teeth from the Bajo Barreal Formation (Upper Cretaceous: Cenomanian-Turonian) at the “estancia Ocho Hermanos”, Chubut Province are described. Most of them show features of Abelisauroidea, a clade already represented in the Bajo Barreal Formation by Xenotarsosaurus bonapartei and other isolated remains including a left maxilla. Dromaeosauridae and Carcharodontosauridae represent the first record of these taxa for the Bajo Barreal Formation. Besides this, at least three theropod clades (Abelisauroidea, Carcharodontosauridae, and Dromaeosauridae) coexisted in central Patagonia during the Cenomanian-Turonian along with other theropods as Aniksosaurus darwini and Megaraptor sp.     

Theropod forelimb design and evolution

We examined the relationship between forelimb design and function across the 230-million-year history of theropod evolution. Forelimb disparity was assessed by plotting the relative contributions of the three main limb elements on a ternary diagram. Theropods were divided into five functional groups: predatory, reduced, flying, wing-propelled diving, and flighdess. Forelimbs which maintained their primitive function, predation, are similarly proportioned, but non-avian theropods with highly reduced forelimbs have relatively longer humeri. Despite the dramatically different forces imparted by the evolution of flight, forelimb proportions of basal birds are only slighdy different from those of their non-avian relatives. An increase in disparity accompanied the subsequent radiation of birds. Each transition to flightlessness has been accompanied by an increase in relative humeral length, which results from relatively short distal limb elements. We introduce theoretical predictions based on five biomechanical and developmental factors that may have influenced the evolution of theropod limb proportions.     

Genetic diversity and structure of the cycad Zamia loddigesii Miq. (Zamiaceae): implications for evolution and conservation

The genetic diversity and structure of four populations of the cycad Zamia loddigesii were studied throughout its range in Mexico. Allozyme electrophoresis of 15 loci was conducted. The mean number of alleles per locus was 1.80 ± 0.09, the percentage of polymorphic loci was 66.6 ± 5.4, and the expected heterozygosity was 0.266 ± 0.02. The results indicated that the genetic diversity was relatively higher, with respect to tropical tree species and other cycads. The genetic variation explained by differences among populations was 18%. On average, gene flow between paired populations was similar ( Nm  = 1.6) to other tropical forest trees and cycad species. Our results indicated that the geographical isolation among populations of Z. loddigesii generated allele loss, as well as a clinal variation in the frequencies of two loci ( MDH and MNR2 ), in relation to the latitudinal distribution of populations. The populations have become fragmented due to increasingly higher pressure of habitat conversion and disturbance. The importance of the establishment of sanctuaries and protected areas and a reduction in deforestation is highlighted in this research as a way of preserving the high genetic diversity of this and other endemic species.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 533–544.     

Phylogenetic eigenvectors and nonstationarity in the evolution of theropod dinosaur skulls

Despite the long‐standing interest in nonstationarity of both phenotypic evolution and diversification rates, only recently have methods been developed to study this property. Here, we propose a methodological expansion of the phylogenetic signal‐representation (PSR) curve based on phylogenetic eigenvectors to test for nonstationarity. The PSR curve is built by plotting the coefficients of determination R² from phylogenetic eigenvector regression (PVR) models increasing the number of phylogenetic eigenvectors against the accumulated eigenvalues. The PSR curve is linear under a stationary model of trait evolution (i.e. the Brownian motion model). Here we describe the distribution of shifts in the models R² and used a randomization procedure to compare observed and simulated shifts along the PSR curve, which allowed detecting nonstationarity in trait evolution. As an applied example, we show that the main evolutionary pattern of variation in the theropod dinosaur skull was nonstationary, with a significant shift in evolutionary rates in derived oviraptorosaurs, an aberrant group of mostly toothless, crested, birdlike theropods. This result is also supported by a recently proposed Bayesian‐based method (AUTEUR). A significant deviation between Ceratosaurus and Limusaurus terminal branches was also detected. We purport that our new approach is a valuable tool for evolutionary biologists, owing to its simplicity, flexibility and comprehensiveness.