Cursorial Ability Research Articles

The last ceratosaur of Asia: a new noasaurid from the Early Cretaceous Great Siberian Refugium.

The noasaurid ceratosaur Kiyacursor longipes gen. et sp. nov. is described based on a fragmentary skeleton including cervical vertebra, pectoral girdle, humerus and hind limbs from the Lower Cretaceous (Aptian) Ilek Formation at Shestakovo 1 locality in Western Siberia, Russia. This is the first ceratosaur from the Early Cretaceous of Asia, extending the stratigraphic range of Ceratosauria by 40 Myr on that continent. Kiyacursor shares unique hind limb proportions with Elaphrosaurus and Limusaurus, suggesting improved cursorial ability. These taxa show an ostrich-like specialization of the pes, with a large third metatarsal and greatly reduced second metatarsal. By contrast, all other fast running non-avian theropod dinosaurs have an arctometatarsalian pes, with the third metatarsal strongly reduced proximally. The new taxon lived in the Early Cretaceous ecosystem containing a number of other Jurassic relics, such as stem salamanders, protosuchian and shartegosuchid crocodyliforms, tritylodontid synapsids and docodontan mammaliaforms.

Anatomical Correlates of Cursoriality are Compromised by Body Size and Propensity to Burrow in a Group of Small Mammals (Lagomorpha)

Highly cursorial animals are specialised for fast, sustained running via specific morphological adaptations, notably including changes in limb segment length and mechanical advantage. Members of the order Lagomorpha (hares, rabbits and pikas) vary in cursorial ability; hares are generally highly cursorial, rabbits more frequently saltate, and pikas predominantly trot. Previous investigations of lagomorphs have identified anatomical trends correlated with this ‘cursoriality gradient’, however, the phylogenetic sampling of such investigations has been limited to three American species, namely the American pika (Ochotona princeps), brush rabbit (Sylvilagus bachmani), and black-tailed jackrabbit (Lepus californicus). Here, we expand the phylogenetic sample and body size range by including novel data from Australian samples of the European rabbit (Oryctolagus cuniculus) and European hare (L. europaeus), alongside unpublished data on the Eastern cottontail (S. floridanus). X-ray Computed Tomography and digital landmarking were used to capture proportions within the appendicular skeleton of ~ 40 specimens of each European species. In doubling the number of species studied, we find the previously-identified morphological gradients associated with cursorial behaviour are complicated when evaluated in the larger sample. The relative length and joint velocity of limbs was found to be lower than predicted in European rabbits and hares. Furthermore, we present a novel assessment of morphological integration in the lagomorph appendicular skeleton, finding between-limb covariation patterns that are generally similar to those of other mammals. Broadly, these results suggest cursoriality is only one of many selective forces driving lagomorph skeletal evolution, with variations in body size and fossoriality potentially having measurable impacts.

Osteomorphological features of the hind limb bones of Saiga antelope (Saiga tatarica).

The intralimb indices and calcaneal linear metrics are known as the reliable predictors of locomotor adaptation in artiodactyls. The osteological features of hindlimb in adult Saiga (Saiga tatarica) were described, and its correlation with cursoriality and habitat adaptation was discussed. Gross anatomy data showed Saiga owned the deep acetabulum as a broad lunate surface, the large acetabular anteversion, the well-developed ischiatic tuberosity and the prominent gluteal lines. It also presented the robust rough line and the strong gluteal tuberosity. A proximodistally elongated eminence located on the cranially distal tibia, which had not been found in goat. The tibial extensor groove was deep. The calcaneal tuberosity was robust. Digital anatomy data showed Saiga owned the higher metatarsal-femur ratio than forest musk deer and sheep. Comparing with wild bovids and sheep, Saiga presented a transitional variation in calcaneal form. The mean greatest length of the calcaneus (GLC) and the height of the sustentacular facet (HSF) in Saiga were shorter than that in sheep and longer than that in wild bovids respectively (F=587.492; F=10.264, p<.05). The wild bovids had longer cubonavicular facets than the other two groups (F=18.587, p<.05). The great metatarsal-femur ratio of Saiga implied a superior cursorial ability and high conservation confronting the different habitats. The calcaneal linear metrics might shed light on lifestyle-related functional adaptation over decades of short-term evolution in the semi-free range environment.

Shape variability in tridactyl dinosaur footprints: the significance of size and function

AbstractThe functional anatomy of the hindlimb of bipedal dinosaurs has been intensively studied. Yet, surprisingly little work has been done concerning functional adaptation of digits for terrestrial locomotion. While complete and articulated pes skeletons are scarce, pes shape is abundantly recorded by fossil footprints. We elucidate the significance of footprint shape and size for locomotion using a large sample (n = 303) of tridactyl dinosaur footprints from a broad range of geographical localities and time slots. Size and shape variation are characterized separately for theropods and ornithischians, the two principal trackmaker taxa. At smaller sizes, theropod footprints are best discriminated from ornithischian footprints by their smaller interdigital angle and larger projection of digit III; at larger sizes digital widths are effective discriminants. Ornithischian footprints increase in size from the Early Jurassic to the Late Cretaceous, a trend not observed in theropod footprints. Size and function are argued to be important determinants of footprint shape, and an attempt made to infer function from shape. Digit III projection and length‐to‐width ratio of the footprints are negatively correlated with size in both groups; digit impression width is positively correlated with size only in ornithischians. Digit III projection appears to be positively correlated with cursorial ability. Increased interdigital angles are associated with a decrease in digital width, possibly an adaptation for stability. Weak digit III projection and increased digital width are interpreted as adaptations for graviportality. Footprints yield great potential for the understanding of the functional morphology of dinosaur feet.

A mid-Cretaceous tyrannosauroid and the origin of North American end-Cretaceous dinosaur assemblages.

Late Cretaceous dinosaur assemblages of North America-characterized by gigantic tyrannosaurid predators, and large-bodied herbivorous ceratopsids and hadrosaurids-were highly successful from around 80 million years ago (Ma) until the end of the 'Age of Dinosaurs' 66 Ma. However, the origin of these iconic faunas remains poorly understood because of a large, global sampling gap in the mid-Cretaceous, associated with an extreme sea-level rise. We describe the most complete skeleton of a predatory dinosaur from this gap, which belongs to a new tyrannosauroid theropod from the Middle Turonian (~92 Ma) of southern Laramidia (western North America). This taxon, Suskityrannus hazelae gen. et sp. nov., is a small-bodied species phylogenetically intermediate between the oldest, smallest tyrannosauroids and the gigantic, last-surviving tyrannosaurids. The species already possesses many key features of the tyrannosaurid bauplan, including the phylogenetically earliest record of an arctometatarsalian foot in tyrannosauroids, indicating that the group developed enhanced cursorial abilities at a small body size. Suskityrannus is part of a transitional Moreno Hill (that is, Zuni) dinosaur assemblage that includes dinosaur groups that became rare or were completely absent in North America around the final 15 Myr of the North American Cretaceous before the end-Cretaceous mass extinction, as well as small-bodied forebears of the large-bodied clades that dominated at this time.

Oldest ctenodactyloid tarsals from the Eocene of China and evolution of locomotor adaptations in early rodents

BackgroundTamquammys has been considered one of the basal ctenodactyloid rodents, which has been documented in the earliest to middle Eocene (~ 56.0–48.5 Ma) in China. It was the most abundant and widespread rodent genus in the Erlian Basin (Nei Mongol, China) and dominated Arshantan small-mammal faunas of that region. Here for the first time we describe the morphology of the astragalocalcaneal complex in Tamquammys robustus (larger) and T. wilsoni, and interpret it against the background of locomotor adaptations of basal Euarchontoglires (rodents, lagomorphs, tree shrews, and primates).ResultsThe comparative morphology of the tarsal elements in Tamquammys robustus and T. wilsoni shows overall slenderness of the bones and their similarity to the tarsal elements of Rattus, a generalist species, and those of small rock squirrels (e.g. Sciurotamias). The two species differ slightly in their cursorial ability; smaller T. wilsoni shows some adaptations to climbing. The results of principal component analysis of the calcaneus and astragalus support this observation and place T. robustus in-between Rattus and ground/rock squirrel morphospace, and T. wilsoni closer to euarchontans, Tupaia and Purgatorius.ConclusionsThe morphology of the tarsal elements in Tamquammys indicates a generalist rodent morphotype with no particular adaptations to arboreality. We suggest that Tamquammys as a basal ctenodactyloid is closer to the ancestral astragalocalcaneal morphology of rodents than that of more derived North American paramyines of similar age. Overall similarity in Tamquammys tarsal elements structure to Purgatorius, a basal primate, may point to the antiquity of the tarsal structure in Tamquammys and a generally unspecialized foot structure in early Euarchontoglires.

Phylogenetic relationships of the Australian Oligo–Miocene ratite Emuarius gidju Casuariidae

In Australia, ratites (Aves: Palaeognathae) are represented in the extant fauna by the family Casuariidae with 1 species of emu Dromaius novaehollandiae and 1 cassowary Casuarius casuarius. The Australian fossil record reveals no other extinct ratite families but there are a number of other casuariid species. Most significant of these, due to its Oligo-Miocene age and because it is known from abundant material, is Emuarius gidju. Here, we describe additional material and confirm that the taxon had a temporal range of Late Oligocene to Middle Miocene (approximately 24-15 Ma). We reveal new morphological details, including notably that the species had relatively much smaller eyes than D. novaehollandiae, in addition to a less well-developed cursorial ability, as inferred from its pelvic limb. In these respects, Emuarius is similar to Casuarius and suggest that it was adapted to denser vegetation than the open woodlands and grasslands that characterise much of Australia today and to which D. novaehollandiae, with its large eyes and enhanced cursorial ability, is strongly adapted. Emuarius was compared to and found to be distinct from the poorly provenanced Australian fossil species C. lydekkeri. We conducted a phylogenetic analysis of morphological data that robustly shows that E. gidju is the sister taxon of Dromaius and together these taxa form a clade that is sister to Casuarius. This indicates that the evolution towards enhanced cursorality that characterises Dromaius took place after the divergence of the emu-cassowary lineages and was likely not the driving mechanism of this divergence. Comparisons between D. novaehollandiae and D. baudinianus revealed no qualitative skeletal differences and we suggest that the latter taxon is best considered to be an island dwarf that should be taxonomically recognized at a subspecific level only.

A new cursorial hyena from Tibet, and analysis of biostratigraphy, paleozoogeography, and dental morphology of Chasmaporthetes (Mammalia, Carnivora)

ABSTRACT Recent field work in the late Cenozoic Zanda Basin in southwestern Tibetan Plateau has provided new fossil evidence of vertebrate faunas spanning the late Miocene to Pleistocene, which represents new occurrences hitherto unknown in that region of Asia. In this paper we describe a new species of the cursorial hyaenid Chasmaporthetes, C. gangsriensis, sp. nov., from the Zanda Basin. Chasmaporthetes gangsriensis is smaller than other Plio-Pleistocene Eurasian records of the genus, and retains relatively wide premolars that are underdifferentiated in size. The m1 talonid has a trenchant hypoconid typical of Chasmaporthetes, but with a trigonid length-to-width ratio lower than all specimens referred to the genus. Metatarsal and phalangeal elements referred to C. gangsriensis are long and gracile, indicating cursorial abilities typical of Chasmaporthetes. With an age of early Pliocene (4.89–4.08 Ma), C. gangsriensis is morphologically the most basal Pliocene Chasmaporthetes in China, and is consistent with the ‘out of Tibet’ hypothesis for some Pleistocene megafauna. An analysis of nasal bone morphology revealed large intraspecific variation in extant spotted hyenas, showing that it is not a reliable criterion for species diagnosis in Chasmaporthetes. An evaluation of the biostratigraphic relationships among Asian and North American occurrences of Chasmaporthetes indicates that the genus first dispersed into the New World during the early Blancan North American land mammal age (NALMA) with likely ancestry close to the heterogeneous sample of C. lunensis in Eurasia. The possibility of a second dispersal is indicated by a mixed sample of specimens with significantly smaller p4/m1 length ratio than other Chasmaporthetes. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

The Late Triassic dinosauromorph Sacisaurus agudoensis (Caturrita Formation; Rio Grande do Sul, Brazil): anatomy and affinities

Abstract Silesauridae is an exclusively Triassic group of dinosauromorphs, knowledge on the diversity of which has increased dramatically in the last few years. Silesaurid relationships are still contentious, as a result in part of different homology statements, particularly regarding the typical edentulous mandible tip of these animals. One of the most complete silesaurids yet discovered is Sacisaurus agudoensis from the Caturrita Formation (Late Triassic: Norian) of Rio Grande do Sul, Brazil, represented by numerous isolated bones recovered from a single site. The anatomy of S. agudoensis is fully described for the first time here, and comparisons are provided to other basal dinosauromorphs. S. agudoensis is a small-bodied animal (less than 1 m in length) that possesses a dentition consisting of leaf-shaped crowns with large denticles in the carinae, a plesiomorphic propubic pelvis with an almost fully closed acetabulum, elongate distal hindlimbs suggesting well-developed cursorial ability, and a laterally projected outer malleolus in the tibia. All previous numerical phylogenies supported a non-dinosaur dinosauromorph affinity for Silesauridae, but the reanalysis of one of those studies suggests that a position within Dinosauria is not unlikely, with silesaurids forming the basal branch of the ornithischian lineage.

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.

The anatomy of the basal ornithischian dinosaur Eocursor parvus from the lower Elliot Formation (Late Triassic) of South Africa

Ornithischia is a morphologically and taxonomically diverse clade of dinosaurs that originated during the Late Triassic and were the dominant large-bodied herbivores in many Cretaceous ecosystems. The early evolution of ornithischian dinosaurs is poorly understood, as a result in part of a paucity of fossil specimens, particularly during the Triassic. The most complete Triassic ornithischian dinosaur yet discovered is Eocursor parvus from the lower Elliot Formation (Late Triassic: Norian–Rhaetian) of Free State, South Africa, represented by a partial skull and relatively complete postcranial skeleton. Here, the anatomy of Eocursor is described in detail for the first time, and detailed comparisons are provided to other basal ornithischian taxa. Eocursor is a small-bodied taxon (approximately 1 m in length) that possesses a plesiomorphic dentition consisting of unworn leaf-shaped crowns, a proportionally large manus with similarities to heterodontosaurids, a pelvis that contains an intriguing mix of plesiomorphic and derived character states, and elongate distal hindlimbs suggesting well-developed cursorial ability. The ontogenetic status of the holotype material is uncertain. Eocursor may represent the sister taxon to Genasauria, the clade that includes most of ornithischian diversity, although this phylogenetic position is partially dependent upon the uncertain phylogenetic position of the enigmatic and controversial clade Heterodontosauridae.

A new transitional sauropodomorph dinosaur from the Early Jurassic of South Africa and the evolution of sauropod feeding and quadrupedalism

Aardonyx celestae gen. et sp. nov. is described from the upper Elliot Formation (Early Jurassic) of South Africa. It can be diagnosed by autapomorphies of the skull, particularly the jaws, cervical column, forearm and pes. It is found to be the sister group of a clade of obligatory quadrupedal sauropodomorphs (Melanorosaurus + Sauropoda) and thus lies at the heart of the basal sauropodomorph-sauropod transition. The narrow jaws of A. celestae retain a pointed symphysis but appear to have lacked fleshy cheeks. Broad, U-shaped jaws were previously thought to have evolved prior to the loss of gape-restricting cheeks. However, the narrow jaws of A. celestae retain a pointed symphysis but appear to have lacked fleshy cheeks, demonstrating unappreciated homoplasy in the evolution of the sauropod bulk-browsing apparatus. The limbs of A. celestae indicate that it retained a habitual bipedal gait although incipient characters associated with the pronation of the manus and the adoption of a quadrupedal gait are evident through geometric morphometric analysis (using thin-plate splines) of the ulna and femur. Cursorial ability appears to have been reduced and the weight bearing axis of the pes shifted to a medial, entaxonic position, falsifying the hypothesis that entaxony evolved in sauropods only after an obligate quadrupedal gait had been adopted.

The intertarsal joint of the ostrich (Struthio camelus): Anatomical examination and function of passive structures in locomotion

The ostrich (Struthio camelus) is the largest extant biped. Being flightless, it exhibits advanced cursorial abilities primarily evident in its characteristic speed and endurance. In addition to the active musculoskeletal complex, its powerful pelvic limbs incorporate passive structures wherein ligaments interact with joint surfaces, cartilage and other connective tissue in their course of motion. This arrangement may enable energy conservation by providing joint stabilisation, optimised limb segment orientation and automated positioning of ground contact elements independently of direct muscle control. The intertarsal joint is of particular interest considering its position near the mid-point of the extended limb and its exposure to high load during stance with significant inertial forces during swing phase. Functional-anatomical analysis of the dissected isolated joint describes the interaction of ligaments with intertarsal joint contours through the full motion cycle. Manual manipulation identified a passive engage-disengage mechanism (EDM) that establishes joint extension, provides bi-directional resistance prior to a transition point located at 115 degrees and contributes to rapid intertarsal flexion at toe off and full extension prior to touch down. This effect was subsequently quantified by measurement of intertarsal joint moments in prepared anatomical specimens in a neutral horizontal position and axially-loaded vertical position. Correlation with kinematic analyses of walking and running ostriches confirms the contribution of the EDM in vivo. We hypothesise that the passive EDM operates in tandem with a stringently coupled multi-jointed muscle-tendon system to conserve the metabolic cost of locomotion in the ostrich, suggesting that a complete understanding of terrestrial locomotion across extinct and extant taxa must include functional consideration of the ligamentous system.

An almost complete skeleton of Metailurus parvulus (Carnivora, Felidae) from the late Miocene of Kerassia (Northern Euboea, Greece)

We describe a partial skeleton of Metailurus parvulus from the Turolian site of Kerassia 1 (Northern Euboea, Greece). The material, which consists of a mandible, the anterior and posterior limb-bone elements, some sternal bones and some vertebrae, is the most complete known of this species. The dental material is compared to specimens from Pikermi and Chomateri (Greece), and China. The limb-bones available offer us the possibility to discuss the status of some previously described specimens from Pikermi. The limb proportions indicate that M. parvulus had elongated posterior limbs relative to the anterior ones, which reflects developed jumping skills. M. parvulus had moderately developed cursorial abilities, intermediate between open and closed habitat felids, and probably frequented primarily relatively open woodlands.

A new emu (Dromaiinae) from the Late Oligocene Etadunna Formation

A new emu (Emuarius guljaruba, sp. nov.) is described from the Late Oligocene Etadunna Formation (Ngama Local Fauna), based on a complete tarsometatarsus. While exhibiting evidence of cursorial abilities advanced over those of cassowaries (Casuarius), this taxon was not as cursorially adapted as the living Emu (Dromaius novaehollandiae). This taxon is provisionally referred to the genus Emuarius, although a definite generic assignment cannot be made.

A crocodyliform from the Early Cretaceous Dinosaur Beds, northern Malawi

ABSTRACT In 1989 a crocodyliform having a mammal-like dentition was reported from the Early Cretaceous of Malawi, Africa. It shares with Notosuchus of South America the following derived features: jaw articulation that permitted longitudinal movement and the occipital condyle on the ventral surface of the braincase. The Malawi crocodyliform is distinct from Notosuchus in possessing multicusped posterior teeth, narrow and elongate prefrontals, no maxillo-palatal fenestrae, deeply overhung squamosal, and a longer, flat jaw articulation surface. The long, flat jaw articulation and tooth morphology suggest proal processing of food. The Malawi crocodyliform has cervical vertebrae with well developed hypapophyses and tall neural spines indicating strong neck muscles. The femur has a well developed medially directed head and posteriorly directed distal condyles. The crus projects from the tarsals vertically. These features suggest an erect posture and cursorial ability. The supracondylar ridges of the humerus are well developed for muscle attachment indicating strong rotation and retraction of the humerus. Articulated specimens suggest preservation in burrows.