An articulated titanosaur from Patagonia (Argentina): New evidence of neosauropod pedal evolution

Most titanosaur dinosaurs are represented by incomplete skeletal elements lacking articulated pes. An exceptionally preserved specimen from the Late Campanian–Early Maastrichtian strata of Patagonia (Argentina) provides new data on pedal morphology and the evolutionary trends of these huge dinosaurs. This finding is one of the few articulated titanosaur pes known in the world, and shows a phalangeal formula of 2-2-2-2-0. The first three digits possess sickle-shaped claws and the articular facets of ungual phalanges, suggesting mobility in horizontal and vertical planes. A comparative analysis of available record suggests that titanosaurs had a progressive reduction of size and number of pedal phalanges in digits III and IV during the Late Cretaceous.     

First report of dinosaurian claws from the Late Triassic of India

The Late Triassic Tiki Formation has yielded five isolated nearly complete claws or ungual phalanges from a fossil locality, which are described in detail and compared with other Late Triassic tetrapods. Of these, four ungual phalanges are slender, asymmetric, ventrally recurved, transversely compressed, and contain deep collateral grooves on either side, a low median keel on the proximal articular surface and a prominent proximoventral flexor tubercle showing their high similarity to the theropod dinosaurs. The remaining claw is unlike that of any theropods in terms of high robusticity and near symmetry. However, as in dinosaurs it is ventrally recurved and contains deep lateral grooves, a small flexor tubercle, lateromedially extended proximal articular surface with a distinct median keel and is considered as belonging to an indeterminate dinosaur. Although it is not possible to ascertain whether the unguals belong to a single taxon or multiple taxa, this new find points towards the presence of small dinosaurs in the Late Triassic Tiki fauna.     

An Early Pleistocene human pedal phalanx from Swartkrans,SKX 16699, and the antiquity of the human lateral forefoot

In order to assess the antiquity of derived human lateral (lesser) toe morphology, the SKX 16699 Early Pleistocene pedal proximal phalanx from Swartkrans (South Africa) was compared to samples of pedal phalanges attributed to Pliocene/Pleistocene australopithecines, Homo naledi and Late Pleistocene Homo. In contrast to australopith lateral phalanges, the SKX 16699 phalanx exhibits an absolutely (and probably relatively) short length, limited plantar diaphyseal curvature, proximal-to-midshaft and mid-dorsoplantar flexor sheath insertions, and a marked proximodorsal orientation of the metatarsal facet. SKX 16699 is intermediate between the australopith phalanges and later Homo ones in its modest dorsal diaphyseal curvature and mid-dorsoplantar metatarsophalangeal collateral ligament insertion areas. Its diaphyseal robustness is similar to that of Homo phalanges, but overlaps the range of later australopith ones. This combination of features and the close morphological affinities of SKX 16699 to later Homo proximal pedal phalanges suggest the emergence of a distinctly human lateral forefoot by the initial Early Pleistocene.     

辽宁早白垩世义县组一原始鸟脚类恐龙

初步记述了采自辽西地区早白垩世义县组新的原始鸟脚类恐龙化石材料,并依此建立一新属新种上园热河龙（Jeholosaurus shangyuanensis gen. et sp. nov.）。化石产于北票市上园镇陆家屯义县组下部第一段灰白色凝灰质砂岩中,同一层位产出过大量鹦鹉嘴龙化石。上园热河龙的主要鉴定特征包括6个前上颌齿,鼻骨背面发育小孔,前齿骨约为前上颌骨主体长度的1.5倍、未发育股骨前髁间沟、骨不在一平面上,第三趾趾节中第四节最长。上园热河龙具有一些真鸟脚类恐龙的近裔性状,比如眶前孔小,方骨孔大,位于方颧骨侧面,外下颌孔缺失。另外,上园热河龙的股骨近端形态非常接近进步的鸟脚类恐龙。但是上园热河龙发育有6个前上颌齿,上下颌关节处与齿列位于同一水平线,前上颌齿列与上颌齿列位于同一水平线,这些原始特征未见于已知鸟脚类恐龙。上园热河龙确切系统分类位置需要进一步确定。 上园热河龙是义县组中发现的第二种鸟臀类恐龙,增加了这类恐龙在热河生物群中的分异度。     

A quantitative method for inferring locomotory shifts in amniotes during ontogeny,its application to dinosaurs and its bearing on the evolution of posture

Evolutionary transitions between quadrupedal and bipedal postures are pivotal to the diversification of amniotes on land, including in our own lineage (Hominini). Heterochrony is suggested as a macroevolutionary mechanism for postural transitions but understanding postural evolution in deep time is hindered by a lack of methods for inferring posture in extinct species. Dinosaurs are an excellent natural laboratory for understanding postural transitions because they demonstrate at least four instances of quadrupedality evolving from bipedality, and heterochronic processes have been put forward as an explanatory model for these transitions. We extend a quantitative method for reliably inferring posture in tetrapods to the study of ontogenetic postural transitions using measurements of proportional limb robusticity. We apply this to ontogenetic series of living and extinct amniotes, focusing on dinosaurs. Our method correctly predicts the general pattern of ontogenetic conservation of quadrupedal and bipedal postures in many living amniote species and infers the same pattern in some dinosaurs. Furthermore, it correctly predicts the ontogenetic postural shift from quadrupedal crawling to bipedal walking in humans. We also infer a transition from early ontogenetic quadrupedality to late-ontogenetic bipedality in the transitional sauropodomorph dinosaur Mussaurus patagonicus and possibly in the early branching ceratopsian Psittacosaurus lujiatunensis but not in the sauropodomorph Massospondylus carinatus. The phylogenetic positions of these ontogenetic shifts suggest that heterochrony may play a role in the macroevolution of posture, at least in dinosaurs. Our method has substantial potential for testing evolutionary transitions between locomotor modes, especially in elucidating the role of evolutionary mechanisms like heterochrony.     

Middle phalanx skeletal morphology in the hand: Can it predict flexor tendon size and attachments?

Specific sites on the palmar diaphysis of the manual middle phalanges provide attachment for the flexor digitorum superficialis (FDS) tendon. It has been assumed in the literature that lateral palmar fossae on these bones reflect locations for these attachments and offer evidence for relative size of the flexor tendon. This assumption has led to predictions about relative FDS muscle force potential from sizes of fossae on fossil hominin middle phalanges. Inferences about locomotor capabilities of fossil hominins in turn have been drawn from the predicted force potential of the flexor muscle. The study reported here provides a critical first step in evaluating hypotheses about behavioral implications of middle phalangeal morphology in fossil hominins, by testing the hypothesis that the lateral fossae reflect the size of the FDS tendon and the location of the terminal FDS tendon attachments on the middle phalanx. The middle phalangeal region was dissected in 43 individuals from 16 primate genera, including humans. Qualitative observations were made of tendon attachment locations relative to the lateral fossae. Length measurements of the fossae were tested as predictors of FDS tendon cross-sectional area and of FDS attachment tendon lengths. Our results lead to the conclusion that the hypothesis must be rejected, and that future attention should focus on functional implications of the palmar median bar associated with the lateral fossae.     

First dinosaur tracks from the Arabian Peninsula

Background

The evolutionary history of Mesozoic terrestrial vertebrates from the Arabian Peninsula is virtually unknown. Despite vast exposures of rocky outcrops, only a handful of fossils have yet been described from the region. Here we report a multi-taxon dinosaur track assemblage near Madar village, 47 km north of Sana''a, Republic of Yemen. This represents the first dinosaur tracksite from the Arabian Peninsula, and the only multi-taxon dinosaur ichnosite in the Middle East.

Methodology/Findings

Measurements were taken directly from trackway impressions, following standard ichnological conventions. The presence of bipedal trackmakers is evidenced by a long series of pes imprints preserving smoothly rounded posterior margins, no evidence of a hallux, bluntly rounded digit tips and digital divarication angles characteristic of ornithopod dinosaurs. Nearby, eleven parallel quadrupedal trackways document a sauropod herd that included large and small individuals traveling together. Based on the morphology of manus impressions along with a narrow-gauged stance, the quadrupedal trackways were made by non-titanosauriform neosauropods. Additional isolated tracks and trackways of sauropod and ornithopod dinosaurs are preserved nearby.

Conclusions/Significance

Taken together, these discoveries present the most evocative window to date into the evolutionary history of dinosaurs of the Arabian Peninsula. Given the limited Mesozoic terrestrial record from the region, this discovery is of both temporal and geographic significance, and massive exposures of similarly-aged outcrops nearby offer great promise for future discoveries.     

Rate heterogeneity, ancestral character state reconstruction, and the evolution of limb morphology in Lerista (Scincidae, Squamata)

Rates of phenotypic evolution derive from numerous interrelated processes acting at varying spatial and temporal scales and frequently differ substantially among lineages. Although current models employed in reconstructing ancestral character states permit independent rates for distinct types of transition (forward and reverse transitions and transitions between different states), these rates are typically assumed to be identical for all branches in a phylogeny. In this paper, I present a general model of character evolution enabling rate heterogeneity among branches. This model is employed in assessing the extent to which the assumption of uniform transition rates affects reconstructions of ancestral limb morphology in the scincid lizard clade Lerista and, accordingly, the potential for rate variability to mislead inferences of evolutionary patterns. Permitting rate variation among branches significantly improves model fit for both the manus and the pes. A constrained model in which the rate of digit acquisition is assumed to be effectively zero is strongly supported in each case; when compared with a model assuming unconstrained transition rates, this model provides a substantially better fit for the manus and a nearly identical fit for the pes. Ancestral states reconstructed assuming the constrained model imply patterns of limb evolution differing significantly from those implied by reconstructions for uniform-rate models, particularly for the pes; whereas ancestral states for the uniform-rate models consistently entail the reacquisition of pedal digits, those for the model incorporating among-lineage rate heterogeneity imply repeated, unreversed digit loss. These results indicate that the assumption of identical transition rates for all branches in a phylogeny may be inappropriate in modeling the evolution of phenotypic traits and emphasize the need for careful evaluation of phylogenetic tests of Dollo's law.     

A NEW, LARGE ORNITHOMIMID FROM THE CRETACEOUS DINOSAUR PARK FORMATION OF ALBERTA, CANADA: IMPLICATIONS FOR THE STUDY OF DISSOCIATED DINOSAUR REMAINS

Abstract:  Only two ornithomimid genera, Ornithomimus and Struthiomimus , are currently known from the Upper Cretaceous of North America. However, a number of ornithomimid elements from Alberta's Dinosaur Park Formation (Upper Campanian), cannot be assigned to either Ornithomimus or Struthiomimus . These bones, including a frontal, caudal vertebrae, and unguals of the manus and the pes, come from animals significantly larger than any previously known Judithian ornithomimid. The frontal exhibits several unusual features, including transverse expansion over the prefrontals, and extreme reduction of the supratemporal fossae. Caudal vertebrae are characterized by neural arches that are posteriorly shifted and transversely expanded. Manual unguals possess a highly concave articular surface, a flexor tubercle divided by a sulcus, and a broad claw. Pedal unguals display highly concave articular surfaces, and a ridge-like flexor tubercle dividing a deep ventral fossa. Although it is difficult to know whether these elements represent a single taxon, this is currently the most parsimonious hypothesis. This study demonstrates how isolated dinosaur bones can extend our knowledge of dinosaur faunas.     

Ontogenetic allometry of the digital rays of the leopard gecko (Gekkota: Eublepharidae; Eublepharis macularius)

The role of allometry in producing the variation in autopodial morphology observed among the lizards is not well understood. Allometry of metapodial and digit lengths in the manus and pes of the primitively padless gekkotan (Eublepharis macularius) is explored using maximum‐likelihood repeated‐measures ANCOVAs with body length as the covariate. Estimated variance–covariance matrices differed significantly within and between autopodia, and integration was stronger among the metapodials than the digits. The first metapodial and the first digit of each autopodium exhibit the strongest covariances with each of the remaining components in each variance–covariance matrix, suggesting that the lengths of the first rays are important for allometric integration of both manus and pes. Metapodials scale isometrically and digits negatively allometrically; both display allometric heterogeneity among themselves in both autopodia. Both autopodia exhibit changes in proportion over the ontogenetic size range, attributable to variation in scaling among the components of the rays. Allometric coefficients do not vary among pedal digits, despite differences in phalanx number, although phalanx number is associated with differences in slope in the manual digits. This is suggestive of heterogeneity in allometry among the manual phalanges, which thus may be associated with variation in phalanx length within gekkotan digits.     

Dynamic Locomotor Capabilities Revealed by Early Dinosaur Trackmakers from Southern Africa

Background

A new investigation of the sedimentology and ichnology of the Early Jurassic Moyeni tracksite in Lesotho, southern Africa has yielded new insights into the behavior and locomotor dynamics of early dinosaurs.

Methodology/Principal Findings

The tracksite is an ancient point bar preserving a heterogeneous substrate of varied consistency and inclination that includes a ripple-marked riverbed, a bar slope, and a stable algal-matted bar top surface. Several basal ornithischian dinosaurs and a single theropod dinosaur crossed its surface within days or perhaps weeks of one another, but responded to substrate heterogeneity differently. Whereas the theropod trackmaker accommodated sloping and slippery surfaces by gripping the substrate with its pedal claws, the basal ornithischian trackmakers adjusted to the terrain by changing between quadrupedal and bipedal stance, wide and narrow gauge limb support (abduction range = 31°), and plantigrade and digitigrade foot posture.

Conclusions/Significance

The locomotor adjustments coincide with changes in substrate consistency along the trackway and appear to reflect ‘real time’ responses to a complex terrain. It is proposed that these responses foreshadow important locomotor transformations characterizing the later evolution of the two main dinosaur lineages. Ornithischians, which shifted from bipedal to quadrupedal posture at least three times in their evolutionary history, are shown to have been capable of adopting both postures early in their evolutionary history. The substrate-gripping behavior demonstrated by the early theropod, in turn, is consistent with the hypothesized function of pedal claws in bird ancestors.     

Latest Cretaceous hadrosauroid (Dinosauria: Ornithopoda) remains from Bulgaria

Disarticulated dinosaur bones have been discovered in a fossiliferous lens in the Labirinta Cave, southwest of the town of Cherven Bryag, in NW Bulgaria. This cave is formed within marine limestones belonging to the Kajlâka Formation of Latest Cretaceous age. Associated fossils and Sr isotopy suggest that the fossiliferous sediments belong to the uppermost part of the Upper Maastrichtian. The dinosaur bones discovered in this lens include the distal portion of a left femur, a right tibia, the proximal part of a right fibula, a left metatarsal II, the second or third phalanx of a left pedal digit IV, the proximal end of a second metacarpal, and a caudal centrum. All the bones undoubtedly belong to ornithopod dinosaurs and more accurately to representatives of the hadrosauroid clade. All belong to small-sized individuals, although it cannot be assessed whether they belong to juveniles or small-sized adults, pending histological analyses. Hadrosauroid remains have already been discovered in Late Maastrichtian marine sediments from western, central and eastern Europe, reflecting the abundance of these dinosaurs in correlative continental deposits. Indeed, hadrosauroids were apparently the dominating herbivorous dinosaurs in Eurasia by Late Maastrichtian time.     

Dinosaur killer claws or climbing crampons?

Dromaeosaurid theropod dinosaurs possess a strongly recurved, hypertrophied and hyperextensible ungual claw on pedal digit II. This feature is usually suggested to have functioned as a device for disembowelling herbivorous dinosaurs during predation. However, modelling of dromaeosaurid hindlimb function using a robotic model and comparison of pedal ungual morphology with extant analogue taxa both indicate that this distinctive claw did not function as a slashing weapon, but may have acted as an aid to prey capture.     

Telling Apart Ornithopod and Theropod Trackways: A Closer Look at a Large,Late Jurassic Tridactyl Dinosaur Trackway at Serwah,Republic of Yemen

A large bipedal tridactyl dinosaur trackway from the Late Jurassic of Serwah, near Madar, Arhab district, Republic of Yemen, has been attributed to an ornithopod trackmaker. As the distinction between theropod and ornithopod dinosaurs can pose a challenge, we present additional data to support and reconfirm the previous attribution.     

A new specimen of the azhdarchoid pterosaur Tapejara wellnhoferi

A new specimen of the Early Cretaceous azhdarchoid Tapejara wellnhoferi is described from the Romualdo Member of the Santana Formation, NE Brazil, providing the first detailed account of the postcranial skeleton. Although limited in its preservation, the osteology is typical of other azhdarchoid pterosaurs from these deposits and represents a juvenile animal with a relatively small wing span of < 1.5 m. The ratios of the pedal elements are identical to those noted for larger, indeterminate azhdarchoids of the Nova Olinda Member of the Crato Formation, where the unguals of the pes are greatly enlarged relative to those of the ornithocheiroids that co-inhabited the Santana lagoon. The ratios of these elements suggest that, as part of a larger suite of characters, these animals were likely better adapted for life on the ground than their ornithocheiroid relatives.     

Description and interpretation of interphalangeal lines in tetrapods

A previously unnoticed geometric pattern is present in the extremities of all tetrapods. Sets of straight and typically uninterrupted hinge lines pass through neighboring interphalangeal joints and across ungual tips. Four sets of these lines appear in basal polydactyl tetrapods, two medial sets, a transverse set and a lateral set. The two medial sets merge in primitive pentadactyl tetrapods. The resulting three line sets persist in later taxa, even when digits shrink and disappear. Primitively and typically the lines in each set are more or less parallel, but lines may converge, merge and shift as phalanges disappear or phalangeal proportions change. Confirming this geometric pattern, complex interphalangeal joint surfaces typically align with hinge lines and pad divisions parallel them. In addition, unguals rarely cross extensions of hinge lines and longer unguals may divert medially or laterally rather than cross them. Exceptions occur most commonly on ungual II. Line sets may exist because phalanges appear to flex and extend most efficiently in unison. Hinge line patterns appear to identify clades so they may, to a limited extent, be used taxonomically. Hinge lines also have predictive value in that missing phalanges, including unguals, can be reconstructed with confidence using hinge lines as size guides. Correct digit spread and metapodial configuration can also be determined in extinct taxa by seeking the appearance of continuous interphalangeal hinge lines in tested reconstructions.     

Locomotion in ornithischian dinosaurs: an assessment using three‐dimensional computational modelling

Ornithischian dinosaurs were primitively bipedal with forelimbs modified for grasping, but quadrupedalism evolved in the clade on at least three occasions independently. Outside of Ornithischia, quadrupedality from bipedal ancestors has only evolved on two other occasions, making this one of the rarest locomotory transitions in tetrapod evolutionary history. The osteological and myological changes associated with these transitions have only recently been documented, and the biomechanical consequences of these changes remain to be examined. Here, we review previous approaches to understanding locomotion in extinct animals, which can be broadly split into form–function approaches using analogy based on extant animals, limb‐bone scaling, and computational approaches. We then carry out the first systematic attempt to quantify changes in locomotor muscle function in bipedal and quadrupedal ornithischian dinosaurs. Using three‐dimensional computational modelling of the major pelvic locomotor muscle moment arms, we examine similarities and differences among individual taxa, between quadrupedal and bipedal taxa, and among taxa representing the three major ornithischian lineages (Thyreophora, Ornithopoda, Marginocephalia). Our results suggest that the ceratopsid Chasmosaurus and the ornithopod Hypsilophodon have relatively low moment arms for most muscles and most functions, perhaps suggesting poor locomotor performance in these taxa. Quadrupeds have higher abductor moment arms than bipeds, which we suggest is due to the overall wider bodies of the quadrupeds modelled. A peak in extensor moment arms at more extended hip angles and lower medial rotator moment arms in quadrupeds than in bipeds may be due to a more columnar hindlimb and loss of medial rotation as a form of lateral limb support in quadrupeds. We are not able to identify trends in moment arm evolution across Ornithischia as a whole, suggesting that the bipedal ancestry of ornithischians did not constrain the development of quadrupedal locomotion via a limited number of functional pathways. Functional anatomy appears to have had a greater effect on moment arms than phylogeny, and the differences identified between individual taxa and individual clades may relate to differences in locomotor performance required for living in different environments or for clade‐specific behaviours.     

“Keep your feet on the ground”: Simulated range of motion and hind foot posture of the Middle Jurassic sauropod Rhoetosaurus brownei and its implications for sauropod biology

The biomechanics of the sauropod dinosaur pes is poorly understood, particularly among the earliest members of the group. To date, reasonably complete and articulated pedes in Early Middle Jurassic sauropods are rare, limited to a handful of taxa. Of these, Rhoetosaurus brownei, from eastern Australia, is currently the only one from the Gondwanan Middle Jurassic that preserves an articulated pes. Using Rhoetosaurus brownei as a case exemplar, we assessed its paleobiomechanical capabilities and pedal posture. Physical and virtual manipulations of the pedal elements were undertaken to evaluate the range of motion between the pedal joints, under both bone-to-bone and cartilaginous scenarios. Using the results as constraints, virtual reconstructions of all possible pedal postures were generated. We show that Rhoetosaurus brownei was capable of significant digital mobility at the osteological metatarsophalangeal and distal interphalangeal joints. We assume these movements would have been restricted by soft tissue in life but that their presence would have helped in the support of the animal. Further insights based on anatomy and theoretical mechanical constraints restricted the skeletal postures to a range encompassing digitigrade to subunguligrade stances. The approach was extended to additional sauropodomorph pedes, and some validation was provided via the bone data of an African elephant pes. Based on the resulting pedal configurations, the in-life plantar surface of the sauropod pes is inferred to extend caudally from the digits, with a soft tissue pad supporting the elevated metatarsus. The plantar pad is inferred to play a role in the reduction of biomechanical stresses, and to aid in support and locomotion. A pedal pad may have been a key biomechanical innovation in early sauropods, ultimately resulting in a functionally plantigrade pes, which may have arisen during the Early to Middle Jurassic. Further mechanical studies are ultimately required to permit validation of this long-standing hypothesis.     

From extant to extinct: locomotor ontogeny and the evolution of avian flight

Evolutionary transformations are recorded by fossils with transitional morphologies, and are key to understanding the history of life. Reconstructing these transformations requires interpreting functional attributes of extinct forms by exploring how similar features function in extant organisms. However, extinct-extant comparisons are often difficult, because extant adult forms frequently differ substantially from fossil material. Here, we illustrate how postnatal developmental transitions in extant birds can provide rich and novel insights into evolutionary transformations in theropod dinosaurs. Although juveniles have not been a focus of extinct-extant comparisons, developing juveniles in many groups transition through intermediate morphological, functional and behavioral stages that anatomically and conceptually parallel evolutionary transformations. Exploring developmental transitions may thus disclose observable, ecologically relevant answers to long puzzling evolutionary questions.     

Geometrical changes of knee ligaments and patellar tendon during passive flexion

Patterns of fibre elongation and orientation for the cruciate and collateral ligaments of the human knee joint and for the patellar tendon have not yet been established in three-dimensions. These patterns are essential for understanding thoroughly the contribution of these soft tissues to joint function and of value in surgical treatments for a more conscious assessment of the knee status. Measurements from 10 normal cadaver knees are here reported using an accurate surgical navigation system and consistent anatomical references, over a large flexion arc, and according to current recommended conventions. The contours of relevant sub-bundles were digitised over the corresponding origins and insertions on the bones. Representative fibres were calculated as the straight line segments joining the centroids of these attachment areas. The most isometric fibre was also taken as that whose attachment points were at the minimum change in length over the flexion arc. Changes in length and orientation of these fibres were reported versus the flexion angle. A good general repeatability of intra- and inter-specimens was found. Isometric fibres were found in the locations reported in the literature. During knee flexion, ligament sub-bundles slacken in the anterior cruciate ligament, and in the medial and lateral collateral ligaments, whereas they tighten in the posterior cruciate ligament. In each cruciate ligament the two compounding sub-bundles have different extents for the change in fibre length, and also bend differently from each other on both tibial planes. In the collateral ligaments and patellar tendon all fibres bend posteriorly. Patellar tendon underwent complex changes in length and orientation, on both the tibial sagittal and frontal planes. For the first time thorough and consistent patterns of geometrical changes are provided for the main knee ligaments and tendons after careful fibre mapping.