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1.
《Journal of anatomy》2017,231(4):515-531
Forelimb reduction occurred independently in multiple lineages of theropod dinosaurs. Although tyrannosaurs are renowned for their tiny, two‐fingered forelimbs, the degree of their reduction in length is surpassed by abelisaurids, which possess an unusual morphology distinct from that of other theropods. The forelimbs of abelisaurids are short but robust and exhibit numerous crests, tubercles, and scars that allow for inferences of muscle attachment sites. Phylogenetically based reconstructions of the musculature were used in combination with close examination of the osteology in the Malagasy abelisaurid Majungasaurus to create detailed muscle maps of the forelimbs, and patterns of the muscular and bony morphology were compared with those of extant tetrapods with reduced or vestigial limbs. The lever arms of muscles crossing the glenohumeral joint are shortened relative to the basal condition, reducing the torque of these muscles but increasing the excursion of the humerus. Fusion of the antebrachial muscles into a set of flexors and extensors is common in other tetrapods and occurred to some extent in Majungasaurus. However, the presence of tubercles on the antebrachial and manual elements of abelisaurids indicates that many of the individual distal muscles acting on the wrist and digits were retained. Majungasaurus shows some signs of the advanced stages of forelimb reduction preceding limb loss, while also exhibiting features suggesting that the forelimb was not completely functionless. The conformation of abelisaurid forelimb musculature was unique among theropods and further emphasizes the unusual morphology of the forelimbs in this clade.  相似文献   

2.
Ornithischia (the 'bird-hipped' dinosaurs) encompasses bipedal, facultative quadrupedal and quadrupedal taxa. Primitive ornithischians were small bipeds, but large body size and obligate quadrupedality evolved independently in all major ornithischian lineages. Numerous pelvic and hind limb features distinguish ornithischians from the majority of other non-avian dinosaurs. However, some of these features, notably a retroverted pubis and elongate iliac preacetabular process, appeared convergently in maniraptoran theropods, and were inherited by their avian descendants. During maniraptoran/avian evolution these pelvic modifications led to significant changes in the functions of associated muscles, involving alterations to the moment arms and the activation patterns of pelvic musculature. However, the functions of these features in ornithischians and their influence on locomotion have not been tested and remain poorly understood. Here, we provide quantitative tests of bipedal ornithischian muscle function using computational modelling to estimate 3D hind limb moment arms for the most complete basal ornithischian, Lesothosaurus diagnosticus. This approach enables sensitivity analyses to be carried out to explore the effects of uncertainties in muscle reconstructions of extinct taxa, and allows direct comparisons to be made with similarly constructed models of other bipedal dinosaurs. This analysis supports some previously proposed qualitative inferences of muscle function in basal ornithischians. However, more importantly, this work highlights ambiguities in the roles of certain muscles, notably those inserting close to the hip joint. Comparative analysis reveals that moment arm polarities and magnitudes in Lesothosaurus, basal tetanuran theropods and the extant ostrich are generally similar. However, several key differences are identified, most significantly in comparisons between the moment arms of muscles associated with convergent osteological features in ornithischians and birds. Craniad migration of the iliofemoralis group muscles in birds correlates with increased leverage and use of medial femoral rotation to counter stance phase adduction moments at the hip. In Lesothosaurus the iliofemoralis group maintains significantly higher moment arms for abduction, consistent with the hip abduction mode of lateral limb support hypothesized for basal dinosaurs. Sensitivity analysis highlights ambiguity in the role of musculature associated with the retroverted pubis (puboischiofemoralis externus group) in ornithischians. However, it seems likely that this musculature may have predominantly functioned similarly to homologous muscles in extant birds, activating during the swing phase to adduct the lower limb through lateral rotation of the femur. Overall the results suggest that locomotor muscle leverage in Lesothosaurus (and by inference basal ornithischians in general) was more similar to that of other non-avian dinosaurs than the ostrich, representing what was probably the basal dinosaur condition. This work thereby contradicts previous hypotheses of ornithischian-bird functional convergence.  相似文献   

3.
Bipedalism evolved multiple times within archosaurs, and relatively shorter forelimbs characterize both crocodyliforms and nonavian dinosaurs. Analysis of a comprehensive ontogenetic sequence of specimens (embryo to adult) of the sauropodomorph Massospondylus has shown that bipedal limb proportions result from negative forelimb allometry. We ask, is negative forelimb allometry a pattern basal to archosaurs, amplified in certain taxa to produce bipedalism? Given the phylogenetic position of extant crocodylians and their relatively shorter forelimb, we tested the hypothesis that prevalent negative forelimb allometry is present in Alligator mississippiensis from a sample of wild specimens from embryonic to adult sizes. Long bone lengths (humerus, radius, ulna, femur, tibia, fibula, third metapodials) were measured with their epiphyseal cartilage intact at all sizes. Our results show an overall isometric pattern for most elements regressed on femur length, humerus length, or total limb length. However, negative allometry was prevalent for the ulna, and the third metapodials scale with positive allometry embryonically. These data suggest that the general forelimb proportions in relation to the hindlimb do not change significantly with increasing size in A. mississippiensis. The negative allometry of the ulna and embryonicaly positive allometry of the third metapodials appears to be related to maintaining the functional integrity of the limbs. We show that this pattern is different from that of the sauropodomorph Massospondylus, and we suggest that if bipedalism in archosaurs is tied, in part, to negative forearm allometry, it was either secondarily lost through isometric scaling, or never developed in the ancestor of A. mississippiensis. Anat Rec, 292:787–797, 2009. © 2009 Wiley‐Liss, Inc.  相似文献   

4.
Mass extinctions change global ecosystems, and the end-Triassic mass extinction was hypothesized to have precipitated the rise of dinosaur dominance, with dinosaurs filling resource zones of eliminated large-bodied reptilian lineages. This replacement has been explicitly hypothesized for theropod dinosaurs, and the eastern North American theropod footprint record suggests an increase in maximum body size across the Triassic-Jurassic boundary. Without taking ontogenetic stage in account, the maximum size of the rare large Triassic theropods worldwide supports this hypothesis, with the size of the largest individuals corresponding to the largest Triassic theropod tracks. However, both morphological data and histological examination suggest that known large-bodied Triassic theropods are represented by immature individuals still growing rapidly at the time of death, indicating that the maximum body size of Triassic theropods was much larger than that a strict reading of the body fossil record would suggest. The size increase recorded in the sediments of eastern North America is not part of a global trend. Instead of a simple ecological replacement of non-dinosaurian archosaurs by dinosaurs, the rise in theropod dinosaurian ecological dominance was an extended process across the end of the Late Triassic into the Jurassic. Anat Rec, 303:1158–1169, 2020. © 2019 Wiley Periodicals, Inc.  相似文献   

5.
The crocodile‐line basal suchian Poposaurus gracilis had body proportions suggesting that it was an erect, bipedal form like many dinosaurs, prompting questions of whether its pedal proportions, and the shape of its footprint, would likewise “mimic” those of bipedal dinosaurs. We addressed these questions through a comparison of phalangeal, digital, and metatarsal proportions of Poposaurus with those of extinct and extant crocodile‐line archosaurs, obligate or facultatively bipedal non‐avian dinosaurs, and ground birds of several clades, as well as a comparison of the footprint reconstructed from the foot skeleton of Poposaurus with known early Mesozoic archosaurian ichnotaxa. Bivariate and multivariate analyses of phalangeal and digital dimensions showed numerous instances of convergence in pedal morphology among disparate archosaurian clades. Overall, the foot of Poposaurus is indeed more like that of bipedal dinosaurs than other archosaur groups, but is not exactly like the foot of any particular bipedal dinosaur clade. Poposaurus likely had a digitigrade stance, and its footprint shape could have resembled grallatorid ichnotaxa, unless digit I of the foot of Poposaurus commonly left an impression. Anat Rec, 297:1022–1046, 2014. © 2014 Wiley Periodicals, Inc.  相似文献   

6.
Neuroanatomical reconstructions of extinct animals have long been recognized as powerful proxies for palaeoecology, yet our understanding of the endocranial anatomy of dromaeosaur theropod dinosaurs is still incomplete. Here, we used X-ray computed microtomography (µCT) to reconstruct and describe the endocranial anatomy, including the endosseous labyrinth of the inner ear, of the small-bodied dromaeosaur, Velociraptor mongoliensis. The anatomy of the cranial endocast and ear were compared with non-avian theropods, modern birds, and other extant archosaurs to establish trends in agility, balance, and hearing thresholds in order to reconstruct the trophic ecology of the taxon. Our results indicate that V. mongoliensis could detect a wide and high range of sound frequencies (2,368–3,965 Hz), was agile, and could likely track prey items with ease. When viewed in conjunction with fossils that suggest scavenging-like behaviours in V. mongoliensis, a complex trophic ecology that mirrors modern predators becomes apparent. These data suggest that V. mongoliensis was an active predator that would likely scavenge depending on the age and health of the individual or during prolonged climatic events such as droughts.  相似文献   

7.
Poposaurus gracilis is a bipedal pseudosuchian archosaur that has been poorly understood since the discovery of the holotype fragmentary partial postcranial skeleton in 1915. Poposaurus. gracilis is a member of Poposauroidea, an unusually morphologically divergent clade of pseudosuchians containing taxa that are bipedal, quadrupedal, toothed, edentulous, and some individuals with elongated thoracic neural spines (i.e., sails). In 2003, a well preserved, fully articulated, and nearly complete postcranial skeleton of P. gracilis was discovered with some fragmentary cranial elements from the Upper Triassic Chinle Formation of Grand Staircase-Escalante National Monument of southern Utah, USA. The aim of this work is to describe the osteology of this specimen in detail and compare P. gracilis to other closely related pseudosuchian archosaurs. The open neurocentral sutures throughout the majority of the vertebral column, the small size of this individual, and the presence of seven evenly spaced cyclic growth marks in the histologically sectioned femur indicate that this specimen was a skeletally immature juvenile, or subadult when it died. The pes of P. gracilis contains multiple skeletal adaptations and osteological correlates for soft tissue structures that support a hypothesis of digitigrady for this taxon. When coupled with the numerous postcranial characters associated with cursoriality, and the many anatomical traits convergent with theropod dinosaurs, this animal likely occupied a similar ecological niche with contemporaneous theropods during the Late Triassic Period. Anat Rec, 303:874–917, 2020. © 2019 American Association for Anatomy  相似文献   

8.
Therizinosaurs are unusual theropods from the Upper Cretaceous of Asia and North America. North American representatives include Falcarius utahensis from central Utah, Nothronychus mckinleyi from west central New Mexico, and N. graffami from southern Utah. Nothronychus was quite large, with well-developed forelimbs and pectoral girdle. In many respects, however, these structures were typical for conventional carnivorous theropods, although therizinosaurs have been hypothesized to be herbivorous using anatomical and functional inferences. There is no indication of increased range of motion within the forelimbs, as might be predicted for derived non-avian theropods. The muscular anatomy of the pectoral girdle and forelimb of Nothronychus is reconstructed using visible muscle scars, data from extant birds and crocodilians, and models for other theropods. The osteology and inferred musculature is a mosaic of primitive and derived characters for theropods. A fossa pneumotricipitales may have been present in the proximal humerus. There was a well-developed fossa brachialis in the distal humerus. The epicleidium of the furcula is deflected, reflecting either taphonomic deformation or possibly accommodation of M. supracoracoideus in a triosseal canal, but such a development has yet to be described in any non-avian theropod. In many respects, the other muscular results were quite similar to those inferred for dromaeosaurs.  相似文献   

9.
For many vertebrates the lung is the largest and lightest organ in the body cavity and for these reasons can greatly affect an organism's shape, density, and its distribution of mass; characters that are important to locomotion. In this paper non-respiratory functions of the lung are considered along with data on the respiratory capacities and gas exchange abilities of birds and crocodilians to infer the evolutionary history of the respiratory systems of dinosaurs, including birds. From a quadrupedal ancestry theropod dinosaurs evolved a bipedal posture. Bipedalism is an impressive balancing act, especially for tall animals with massive heads. During this transition selection for good balance and agility may have helped shape pulmonary morphology. Respiratory adaptations arising for bipedalism are suggested to include a reduction in costal ventilation and the use of cuirassal ventilation with a caudad expansion of the lung into the dorsal abdominal cavity. The evolution of volant animals from bipeds required yet again a major reorganization in body form. With this transition avian air sacs may have been favored because they enhanced balance and agility in flight. Finally, I propose that these hypotheses can be tested by examining the importance of the air sacs to balance and agility in extant animals and that these data will enhance our understanding of the evolution of the respiratory system in archosaurs.  相似文献   

10.
Birds have lost and modified the musculature joining the pectoral girdle to the skull and hyoid, called the pectoral extrinsic appendicular and infrahyoid musculature. These muscles include the levator scapulae, sternomandibularis, sternohyoideus, episternocleidomastoideus, trapezius, and omohyoideus. As non-avian theropod dinosaurs are the closest relatives to birds, it is worth investigating what conditions they may have exhibited to learn when and how these muscles were lost or modified. Using extant phylogenetic bracketing, osteological correlates and non-osteological influences of these muscles are identified and discussed. Compsognathids and basal Maniraptoriformes were found to have been the likeliest transition points of a derived avian condition of losing or modifying these muscles. Increasing needs to control the feather tracts of the neck and shoulder, for insulation, display, or tightening/readjustment of the skin after dynamic neck movements may have been the selective force that drove some of these muscles to be modified into dermo-osseous muscles. The loss and modification of shoulder protractors created a more immobile girdle that would later be advantageous for flight in birds. The loss of the infrahyoid muscles freed the hyolarynx, trachea, and esophagus which may have aided in vocal tract filtering.  相似文献   

11.
Early Miocene sloths are represented by a diversity of forms ranging from 38 to 95 kg, being registered mainly from Santacrucian Age deposits in southern‐most shores of Patagonia, Argentina. Their postcranial skeleton differs markedly in shape from those of their closest living relatives (arboreal forms of less than 10 kg), Bradypus and Choloepus. In order to gain insight on functional properties of the Santacrucian sloths forelimb, musculature was reconstructed and a comparative, qualitative morphofunctional analysis was performed, allowing proposing hypotheses about biological role of the limb in substrate preferences, and locomotor strategies. The anatomy of the forelimb of Santacrucian sloths resembles more closely extant anteaters such as Tamandua and Myrmecophaga, due to the robustness of the elements, development of features related to attachment of ligaments and muscles, and conservative, pentadactylous, and strong‐clawed manus. The reconstructed forelimb musculature was very well developed and resembles that of extant Pilosa (especially anteaters), although retaining the basic muscular configuration of generalized mammals. This musculature allowed application of powerful forces, especially in adduction of the forelimb, flexion and extension of the antebrachium, and manual prehension. These functional properties are congruent with both climbing and digging activities, and provide support for proposed Santacrucian sloths as good climbing mammals, possibly arboreal or semiarboreal, being also capable diggers. Their climbing strategies were limited, thus these forms relied mainly on great muscular strength and curved claws of the manus to move cautiously on branches. Anat Rec, 2013. © 2012 Wiley Periodicals, Inc.  相似文献   

12.
The origin of the avian hand, with its reduced and fused carpals and digits, from the five‐fingered hands and complex wrists of early dinosaurs represents one of the major transformations of manus morphology among tetrapods. Much attention has been directed to the later part of this transition, from four‐ to three‐fingered taxa. However, earlier anatomical changes may have influenced these later modifications, possibly paving the way for a later frameshift in digit identities. We investigate the five‐ to four‐fingered transition among early dinosaurs, along with changes in carpus morphology. New three‐dimensional reconstructions from computed tomography data of the manus of the Triassic and Early Jurassic theropod dinosaurs Coelophysis bauri and Megapnosaurus rhodesiensis are described and compared intra‐ and interspecifically. Several novel findings emerge from these reconstructions and comparisons, including the first evidence of an ossified centrale and a free intermedium in some C. bauri specimens, as well as confirmation of the presence of a vestigial fifth metacarpal in this taxon. Additionally, a specimen of C. bauri and an unnamed coelophysoid from the Upper Triassic Hayden Quarry, New Mexico, are to our knowledge the only theropods (other than alvarezsaurs and birds) in which all of the distal carpals are completely fused together into a single unit. Several differences between the manus of C. bauri and M. rhodesiensis are also identified. We review the evolution of the archosauromorph manus more broadly in light of these new data, and caution against incorporating carpal characters in phylogenetic analyses of fine‐scale relationships of Archosauromorpha, in light of the high degree of observed polymorphism in taxa for which large sample sizes are available, such as the theropod Coelophysis and the sauropodomorph Plateosaurus. We also find that the reduction of the carpus and ultimate loss of the fourth and fifth digits among early dinosaurs did not proceed in a neat, stepwise fashion, but was characterized by multiple losses and possible gains of carpals, metacarpals and phalanges. Taken together, the high degree of intra‐ and interspecific variability in the number and identities of carpals, and the state of reduction of the fourth and fifth digits suggest the presence of a ‘zone of developmental variability’ in early dinosaur manus evolution, from which novel avian‐like morphologies eventually emerged and became channelized among later theropod clades.  相似文献   

13.
14.
Therizinosaurs are highly modified, probably herbivorous, theropods from the Upper Cretaceous of Asia and North America. They are characterized by an extensively pneumatized axial skeleton, and in the derived forms, an incipiently opisthopubic pelvis. The evolution of such a pelvis is expected to be associated with extensive modification of the muscular system. The muscular system is reconstructed using observed muscle scars, reconstructions of the theropods Staurikosaurus and Tyrannosaurus, the ornithischian Maiasaura, and extant crocodilians and birds. The results indicate convergence with birds and ornithischian dinosaurs, such that the retroverted pubis in some maniraptorans can be regarded as analogous with the postacetabular bar in ornithischians. Functional implications also make derived therizinosaurs, such as Nothronychus, in some respects convergent with birds as the pubis is retroverted, becoming fused with the ischium, a laterally flaring synsacrum, and an avian-like pes.  相似文献   

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17.
Herbivorous dinosaurs exhibited diverse cranial feeding mechanisms. Although osteological, microwear, and biomechanical research has revealed some of this diversity, the evolutionary reorientation of cranial musculature throughout nonavian herbivorous Dinosauria and its influence on feeding mechanisms requires more study. Here, cranial muscle reconstructions in herbivorous dinosaurs are reviewed and informative anatomical characters are compared across 142 dinosaur genera (84 ornithischians, 36 sauropodomorphs, and 22 herbivorous nonavian theropods), both through examination of specimens and literature. Traits include those relating to the temporal region, adductor chamber, palate, and mandibular attachments, such as the coronoid elevation and retroarticular process. Findings reveal many combinations of anatomical traits influencing a diversity of feeding mechanisms. Some primarily more orthal feeders, including herbivorous theropods, nonsauropod sauropodomorphs, basal ornithischians, and derived stegosaurs (which also show varying degrees of coinciding slight palinal motion and long-axis hemimandibular rotation), possess traits indicative of more prominent temporal musculature and moderately sized palatal musculature. However, orthal feeding sauropods and pachycephalosaurs possess traits indicative of greatly reduced, low-angled temporal musculature, and enhanced palatal musculature producing a primarily vertical, orthal feeding vector. Among ankylosaurs, hadrosaurids, and neoceratopsians, a rostrolabial temporal muscle expansion is present (with a tall coronoid elevation in hadrosaurids and ceratopsids) for greater temporal muscle support and mechanical advantage for complex palinal feeding motions. This also aids in long-axis hemimandibular rotation against the predentary in hadrosaurs and ankylosaurs. This diversity in cranial muscle architecture provides an informative spectrum of numerous adaptations acquired given the evolution of various anatomical constraints in the skull. Anat Rec, 303:1104–1145, 2020. © 2019 American Association for Anatomy  相似文献   

18.
Piatnitzkysauridae were Jurassic theropods that represented the earliest diverging branch of Megalosauroidea, being one of the earliest lineages to have evolved moderate body size. This clade's typical body size and some unusual anatomical features raise questions about locomotor function and specializations to aid in body support; and other palaeobiological issues. Biomechanical models and simulations can illuminate how extinct animals may have moved, but require anatomical data as inputs. With a phylogenetic context, osteological evidence, and neontological data on anatomy, it is possible to infer the musculature of extinct taxa. Here, we reconstructed the hindlimb musculature of Piatnitzkysauridae (Condorraptor, Marshosaurus, and Piatnitzkysaurus). We chose this clade for future usage in biomechanics, for comparisons with myological reconstructions of other theropods, and for the resulting evolutionary implications of our reconstructions; differential preservation affects these inferences, so we discuss these issues as well. We considered 32 muscles in total: for Piatnitzkysaurus, the attachments of 29 muscles could be inferred based on the osteological correlates; meanwhile, in Condorraptor and Marshosaurus, we respectively inferred 21 and 12 muscles. We found great anatomical similarity within Piatnitzkysauridae, but differences such as the origin of M. ambiens and size of M. caudofemoralis brevis are present. Similarities were evident with Aves, such as the division of the M. iliofemoralis externus and M. iliotrochantericus caudalis and a broad depression for the M. gastrocnemius pars medialis origin on the cnemial crest. Nevertheless, we infer plesiomorphic features such as the origins of M. puboischiofemoralis internus 1 around the “cuppedicus” fossa and M. ischiotrochantericus medially on the ischium. As the first attempt to reconstruct muscles in early tetanurans, our study allows a more complete understanding of myological evolution in theropod pelvic appendages.  相似文献   

19.
Theropod dinosaurs were relatively scarce in the Late Cretaceous ecosystems of southeast Brazil. Instead, hypercarnivorous crocodyliforms known as baurusuchids were abundant and probably occupied the ecological role of apex predators. Baurusuchids exhibited a series of morphological adaptations hypothesized to be associated with this ecological role, but quantitative biomechanical analyses of their morphology have so far been lacking. Here, we employ a biomechanical modelling approach, applying finite element analysis (FEA) to models of the skull and mandibles of a baurusuchid specimen. This allows us to characterize the craniomandibular apparatus of baurusuchids, as well as to compare the functional morphology of the group with that of other archosaurian carnivores, such as theropods and crocodylians. Our results support the ecological role of baurusuchids as specialized apex predators in the continental Late Cretaceous ecosystems of South America. With a relatively weak bite force (~600 N), the predation strategies of baurusuchids likely relied on other morphological specializations, such as ziphodont dentition and strong cervical musculature. Comparative assessments of the stress distribution and magnitude of scaled models of other predators (the theropod Allosaurus fragilis and the living crocodylian Alligator mississippiensis) consistently show different responses to loadings under the same functional scenarios, suggesting distinct predatory behaviors for these animals. The unique selective pressures in the arid to semi-arid Late Cretaceous ecosystems of southeast Brazil, which were dominated by crocodyliforms, possibly drove the emergence and evolution of the biomechanical features seen in baurusuchids, which are distinct from those previously reported for other predatory taxa.  相似文献   

20.
Background: Avian forelimb (wing) contains only three digits, and the three‐digit formation in the bird forelimb is one of the avian‐specific limb characteristics that have been evolutionarily inherited from the common ancestral form in dinosaurs. Despite many studies on digit formation in the chick limb bud, the developmental mechanisms giving rise to the three‐digit forelimb in birds have not been completely clarified. Results: To identify which cell populations of the early limb bud contribute to digit formation in the late limb bud, fate maps of the early fore‐ and hindlimb buds were prepared. Based on these fate maps, we found that the digit‐forming region in the forelimb bud is narrower than that in the hindlimb bud, suggesting that some developmental mechanisms on the anterior‐most region lead to a reduced number of digits in the forelimb. We also found temporal differences in the onset of appearance of the ANZ (anterior necrotic zone) as well as differences in the position of the anterior edge of the AER. Conclusions: Forelimb‐specific events in the anterior limb bud are possible developmental mechanisms that might generate the different cell fates in the fore‐ and hindlimb buds, regulating the number of digits in birds. Developmental Dynamics 243:741–752, 2014. © 2014 Wiley Periodicals, Inc.  相似文献   

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