Enigmatic Fossil Footprints from the Sundance Formation (Upper Jurassic) of Bighorn Canyon National Recreation Area,Wyoming

The Sundance Formation (Middle-Upper Jurassic) of Wyoming is well known for pterosaur footprints. Two new partial trackways from the upper Sundance Formation of the Bighorn Canyon National Recreation Area (BICA) of north-central Wyoming are enigmatic. The trackways are preserved in rippled, flaser bedded, glauconitic sand and mud. The deposits were laid down in tidal flats, behind barrier islands, along the mesotidal Sundance Sea.

The best-preserved print of the primary trackway possesses four impressions: three shorter digits with negative rotation and an elongate, caudally-oriented mark. The primary trackway has low pace angulation. The combination of morphology and pace angulation matches neither tracks nor body fossils of horseshoe crabs, theropod dinosaurs, pterosaurs, crocodylomorphs, “lacertoids,” or mammaliforms. The secondary trackway, possibly consisting of undertracks, similarly possesses elongate caudal impressions but differs from the former by possessing four subparallel, cranially-oriented digits. These prints also do not closely resemble any of the aforementioned taxa. While the secondary trackway does not lend itself to conclusion, the primary track maker could have been either an injured, pathologic pterosaur or a pterosaurian taxon otherwise unknown from the ichnological record.     

Pterosaur tracks and the terrestrial ability of pterosaurs

Unusual tracks of a quadrupedal animal with a three-digit (occasionally four-digit) manus print and four-digit pes print were first interpreted as those of pterosaurs in the 1950s. In the 1980s these tracks were reinterpreted as crocodilian, but new material shows that the original identification was correct. Two features: evidence for elongate penultimate phalanges in digits two to four of the pes, and manus trackways up to three times the width of pes trackways, can only be attributed to pterosaurs. Recent improvements in understanding of pterosaur anatomy and functional morphology explain remaining difficulties regarding the interpretation of ich-nites such as the orientation of the manus digits and the absence of some expected ichnological features. Pteraichnus and Pteraichnus-like tracks show that, when grounded, some, perhaps all, pterosaurs were plantigrade, quadrupedal, and had a semi-erect stance and gait. This is consistent with some functional interpretations of pterosaur anatomy and resolves a long-running debate regarding the terrestrial ability of this group.     

Locomotor kinetics on sloped arboreal and terrestrial substrates in a small quadrupedal mammal

Small animals must be capable of moving on a wide variety of surfaces; thus, examining the mechanics of locomotion on a wide variety of substrates is necessary to understand how the animal can utilize its habitat. Therefore, locomotor kinetics are examined on arboreal and terrestrial sloped substrates in the marsupial Monodelphis domestica (gray short-tailed opossum). Substrate reaction forces were measured as opossums moved across four trackways: 30 degrees upslope and 30 degrees downslope trackways, which were flat ("terrestrial") or cylindrical ("arboreal"). Regardless of substrate slope, medial limb forces were measured on arboreal trackways and usually lateral limb forces on terrestrial trackways. Otherwise the general patterns of vertical and craniocaudal forces and impulses were similar between same-sloped terrestrial and arboreal trackways. Some significant modifications to these gross patterns occurred: on the arboreal upslope trackway, hindlimbs supported more body weight than on the terrestrial uphill, possibly because hindlimbs were more stably positioned on the upslope arboreal trackway than forelimbs. Furthermore, the difference between fore- and hindlimbs with respect to craniocaudal impulses was less on the arboreal sloped trackways. In conclusion, kinetic patterns can usually be explained by body weight support roles and by the placement of the limbs on the arboreal trackway.     

The phylogenetic position of the Pterosauria within the Archosauromorpha

In recent years the hypothesis that pterosaurs were the major sister-group of dinosaurs and a closely-linked hypothesis that pterosaurs evolved flight from the ground up have gained general acceptance. A cladistic analysis of the Archosauromorpha using characters presented by previous workers results in a single most parsimonious tree with the Pterosauria as the major sister-group of the Dinosauria. However, that sister-group relationship is supported only by a suite of hindlimb characters that are correlated with bipedal digitigrade locomotion in dinosaurs. In pterosaurs the characters have been interpreted as correlates of bipedal cursorial locomotion, arboreal leaping, or involvement of the hindlimb in the wing. The homology of those characters in dinosaurs and pterosaurs cannot be supported. Reanalysis of the data after exclusion of those hindlimb characters results in most parsimonious trees with the Pterosauria as the sister-group of the Erythrosuchidae + Proterochampsidae + Euparkeria + Archosauria, in that order. This sister-group relationship is supported by a diverse assemblage of functionally independent skeletal characters from all regions of the skeleton. The results of the analysis cast doubt on the hypothesis that pterosaurs evolved flight from the ground up.     

A New Pterosaur Tracksite from the Jurassic Summerville Formation,Near Ferron,Utah

Pterosaur tracks (cf. Pteraichnus) from the Summerville Formation of the Ferron area of central Utah add to the growing record of Pteraichnus tracksites in the Late Jurassic Summerville Formation and time-equivalent, or near time-equivalent, deposits. The site is typical in revealing high pterosaur track densities, but low ichnodiversity suggesting congregations or “flocks” of many individuals. Footprint length varies from 2.0 to 7.0 cms. The ratio of well-preserved pes:manus tracks is about 1:3.4. This reflects a bias in favor of preservation of manus tracks due to the greater weight-bearing role of the front limbs, as noted in other pterosaur track assemblages. The sample also reveals a number of well-preserved trackways including one suggestive of pes-only progression that might be associated with take off or landing, and another that shows pronounced lengthening of stride indicating acceleration.

One well-preserved medium-sized theropod trackway (Therangospodus) and other larger theropod track casts (cf. Megalosauripus) are associated with what otherwise appears to be a nearly monospecific pterosaur track assemblage. However, traces of a fifth pes digit suggest some tracks are of rhamphorynchoid rather than pterodactyloid origin, as usually inferred for Pteraichnus. The tracks occur at several horizons in a thin stratigraphic interval of ripple marked sandstones and siltstones. Overall the assemblage is similar to others found in the same time interval in the Western Interior from central and eastern Utah through central and southern Wyoming, Colorado, northeastern Arizona, and western Oklahoma. This vast “Pteraichnus ichnofacies,” with associated saurischian tracks, remains the only ichnological evidence of pre-Cretaceous pterosaurs in North America and sheds important light on the vertebrate ecology of the Summerville Formation and contiguous deposits.     

First record of a pterosaur landing trackway

The terrestrial progression of pterosaurs, the flying reptiles of the Mesozoic Era, has been debated for over two centuries. The recent discovery of quadrupedal pterodactyloid pterosaur tracks from Late Jurassic sediments near Crayssac, France, shows that the hindlimbs moved parasagittally, as in mammals, birds and other dinosaurs, and the hypertrophied forelimbs could make tracks both close to the body wall and far outside it. Their manus tracks are unique in form, position and kinematics, which would be expected because the forelimbs were used for flight. Here, we report the first record of a pterosaur landing track, which differs substantially from typical walking trackways. The individual landed on both hind feet in parallel fashion, dragged its toes slightly as it left the track, landed again almost immediately and placed the hindfeet parallel again, then placed its forelimbs on the ground, took another short step with both hindlimbs and adjusted its forelimbs, and then began to walk off normally. The trackway shows that pterosaurs stalled to land, a reflection of their highly developed capacity for flight control and manoeuverability.     

Functional morphology of cercopithecoid primate metacarpals

The primate fossil record suggests that terrestriality was more common in the past than it is today, particularly among cercopithecoid primates. Whether or not a fossil primate habitually preferred terrestrial substrates has typically been inferred from its forelimb anatomy. Because extant large-bodied terrestrial cercopithecine monkeys utilize digitigrade hand postures during locomotion, being able to identify if a fossil primate habitually adopted digitigrade postures would be particularly revealing of terrestriality in this group. This paper examines the functional morphology of metacarpals in order to identify osteological correlates of digitigrade versus palmigrade hand postures. Linear measurements were obtained from 324 individuals belonging to digitigrade and palmigrade cercopithecoid species and comparisons were made between hand posture groups. Digitigrade taxa have shorter metacarpals, relative to both body mass and humerus length, than palmigrade taxa. Also, digitigrade taxa tend to have metacarpals with smaller dorsoventral diameters, relative to the product of body mass and metacarpal length, compared to palmigrade taxa. The size and shape of the metacarpal heads do not significantly differ between hand posture groups. Multivariate analyses suggest that metacarpal shape can only weakly discriminate between hand posture groups. In general, while there are some morphological differences in the metacarpals between hand posture groups, similarities also exist that are likely related to the fact that even digitigrade cercopithecoids can adopt palmigrade hand postures in different situations (e.g., terrestrial running, arboreal locomotion), and/or that the functional demands of different hand postures are not reflected in all aspects of metacarpal morphology. Therefore, the lack of identifiable adaptations for specific hand postures in extant cercopithecoids makes it difficult to determine a preference for specific habitats from fossil primate hand bones.     

A new non-pterodactyloid pterosaur from the Late Jurassic of southern Germany

Background

The ‘Solnhofen Limestone’ beds of the Southern Franconian Alb, Bavaria, southern Germany, have for centuries yielded important pterosaur specimens, most notably of the genera Pterodactylus and Rhamphorhynchus. Here we describe a new genus of non-pterodactyloid pterosaur based on an extremely well preserved fossil of a young juvenile: Bellubrunnus rothgaengeri (gen. et sp. nov.).

Methodology/Principal Findings

The specimen was examined firsthand by all authors. Additional investigation and photography under UV light to reveal details of the bones not easily seen under normal lighting regimes was completed.

Conclusions/Significance

This taxon heralds from a newly explored locality that is older than the classic Solnhofen beds. While similar to Rhamphorhynchus, the new taxon differs in the number of teeth, shape of the humerus and femur, and limb proportions. Unlike other derived non-pterodacytyloids, Bellubrunnus lacks elongate chevrons and zygapophyses in the tail, and unlike all other known pterosaurs, the wingtips are curved anteriorly, potentially giving it a unique flight profile.     

New hand bones of Hadropithecus stenognathus: implications for the paleobiology of the Archaeolemuridae

A partial, associated skeleton of Hadropithecus stenognathus (AHA-I) was discovered in 2003 at Andrahomana Cave in southeastern Madagascar. Among the postcranial elements found were the first hand bones (right scaphoid, right hamate, left first metacarpal, and right and left fifth metacarpals) attributed to this rare subfossil lemur. These hand bones were compared to those of extant strepsirrhines and catarrhines in order to infer the positional adaptations of Hadropithecus, and they were compared to those of Archaeolemur in order to assess variation in hand morphology among archaeolemurids. The scaphoid tubercle does not project palmarly as in suspensory and climbing taxa, and the hamate has no hook at all (just a small tubercle), which also points to a poorly developed carpal tunnel. There is a distinctive, radioulnarly directed "spiral" facet for articulation with the triquetrum that is most similar in orientation to that of more terrestrial primates (i.e., Lemur catta, Papio, and Gorilla). The first metacarpal is very reduced and represents only 48% of the length of metacarpal V, as in Archaeolemur, which suggests that pollical grasping of arboreal supports was not important. Compared to Archaeolemur, the shaft of metacarpal V is gracile, and the head has no dorsal ridge and lacks characteristics functionally associated with digitigrade, extended metacarpophalangeal joint postures. Proximally, the articular facet for the hamate is oriented more dorsally. Thus, the carpometacarpal joint V appears to have a distinctive hyperextended set, which has no analog among living or extinct primates. The carpals of Hadropithecus are diagnostic of a pronograde, arboreal and terrestrial (although not digitigrade) locomotor repertoire that typifies Lemur catta and some Old World monkeys. No clinging, suspensory, or climbing specializations that characterize indriids or lorises can be found in the hand of this subfossil lemur. The hand of Hadropithecus likely had similar ranges of movement at the radiocarpal and midcarpal joints as of those of pronograde primates, such as lemurids, for which the hand is held in a more extended, pronated, and neutral (i.e., showing less ulnar deviation) position during locomotion in comparison to that of vertical clingers or slow climbers. Although highly autapomorphic, the hand of Hadropithecus resembles that of its sister taxon, Archaeolemur, in having a very reduced pollex and an articular facet on the scaphoid for a sizeable prepollex. These unusual hand features reinforce the monophyly of the Archaeolemuridae.     

Climbing,brachiation, and terrestrial quadrupedalism: Historical precursors of hominid bipedalism

The vertical-climbing account of the evolution of locomotor behavior and morphology in hominid ancestry is reexamined in light of recent behavioral, anatomical, and paleontological findings and a more firmly established phylogeny for the living apes. The behavioral record shows that African apes, when arboreal, are good vertical climbers, and that locomotion during traveling best separates the living apes into brachiators (gibbons), scrambling/climbing/brachiators (orangutans), and terrestrial quadrupeds (gorillas and chimpanzees). The paleontological record documents frequent climbing as an ancestral catarrhine ability, while a reassessment of the morphology of the torso and forelimb in living apes and Atelini suggests that their shared unique morphological pattern is best explained by brachiation and forelimb suspensory positional behavior. Further, evidence from the hand and foot points to a terrestrial quadrupedal phase in hominoid evolution prior to the adoption of bipedalism. The evolution of positional behavior from early hominoids to hominids appears to have begun with an arboreal quadrupedal-climbing phase and proceeded though an orthograde, brachiating, forelimb-suspensory phase, which was in turn followed by arboreal and terrestrial quadrupedal phases prior to the advent of hominid bipedality. The thesis that protohominids climbed down from the trees to become terrestrial bipeds needs to be reexamined in light of a potentially long history of terrestriality in the ancestral protohominid. © 1996 Wiley-Liss, Inc.     

Assessing arboreal adaptations of bird antecedents: testing the ecological setting of the origin of the avian flight stroke

The origin of avian flight is a classic macroevolutionary transition with research spanning over a century. Two competing models explaining this locomotory transition have been discussed for decades: ground up versus trees down. Although it is impossible to directly test either of these theories, it is possible to test one of the requirements for the trees-down model, that of an arboreal paravian. We test for arboreality in non-avian theropods and early birds with comparisons to extant avian, mammalian, and reptilian scansors and climbers using a comprehensive set of morphological characters. Non-avian theropods, including the small, feathered deinonychosaurs, and Archaeopteryx, consistently and significantly cluster with fully terrestrial extant mammals and ground-based birds, such as ratites. Basal birds, more advanced than Archaeopteryx, cluster with extant perching ground-foraging birds. Evolutionary trends immediately prior to the origin of birds indicate skeletal adaptations opposite that expected for arboreal climbers. Results reject an arboreal capacity for the avian stem lineage, thus lending no support for the trees-down model. Support for a fully terrestrial ecology and origin of the avian flight stroke has broad implications for the origin of powered flight for this clade. A terrestrial origin for the avian flight stroke challenges the need for an intermediate gliding phase, presents the best resolved series of the evolution of vertebrate powered flight, and may differ fundamentally from the origin of bat and pterosaur flight, whose antecedents have been postulated to have been arboreal and gliding.     

Pterosaur Stance and Gait and the Interpretation of Trackways

The tracks ascribed to pterosaurs from the Late Jurassic limestones at Crayssac, France, must be pterosaurian because the manus prints are so far outside those of the pes, the pes print is four times longer than wide, and the manus prints appear to preserve distinct traces of a posteromedially directed wing-finger. These tracks are different in important ways from previously described Pteraichnus trackways, which have been variably considered pterosaurian, crocodilian, or indeterminate. No Pteraichnus (sensu stricto: those not from Crayssac) tracks have diagnostic features of pterosaurs and in none can a complete phalangeal or digital formula be reconstructed; however, all published Pteraichnidae tracks fulfill the criteria of poor preservation, and some have some diagnostic features of crocodile tracks. Reconstructions of pterosaurs walking in pteraichnid tracks do not fit those tracks well, but crocodiles do. In contrast, the Crayssac tracks demonstrate the erect stance and parasagittal gait previously reconstructed for pterosaurs. They also demonstrate that the footfall pattern was not as in typical reptiles (LH-RF-RH-LF), but that the manus must have been raised before the next forward step of the ipselateral foot (LH-LF-RH-RF), suggesting that the quadrupedal pattern was secondary. The metatarsus in pterosaurs was set low at the beginning of a stride, as it is in crocodilians and basal dinosaurs. The diagnosis of the Ichnofamily Pteraichnidae comprises features of possible crocodilian trackmakers, but not of possible pterosaurian trackmakers. Trackways considered for attribution to pterosaurs should show (1) manus prints up to three interpedal widths from midline of body, and always lateral to pes prints, (2) pes prints four times longer than wide at the metatarso-phalangeal joint, and (3) penultimate phalanges longest among those of the pes.     

The arboreal leaping theory of the origin of pterosaur flight

Three theories about the origin of flight in pterosaurs have been proposed: 1) the arboreal parachuting theory (passive falling from trees leading to gliding and eventually to powered flight); 2) the cursorial theory (bipedal running and leaping leading directly to powered flight); and 3) the arboreal leaping theory (active leaping between branches and trees leading to powered flight). The available evidence as to the functional morphology of pterosaurs, and in particular their hindlimb, is reviewed and used to test the three theories. Pterosaurs were well suited for arboreality and their hindlimb morphology argues against cursoriality, but supports an arboreal leaping lifestyle for early pterosaurs or their immediate ancestors.     

The functional anatomy of the forelimb of some African viverridae (Carnivora)

The functional morphology of the forelimbs of the following African Viverridae was studied, Atilax paludinosus, Bdeogale crassicauda, Civettictis civetta, Genetta genetta, G. tigrina, Helogale parvula, Herpestes ichneumon, H. sanguineus, Ichneumia albicauda, Mungos mungo, Nandinia binotata. Their locomotory behaviour has been previously studied and described and is related to morphological differences. The osteology of all the species and the myology of three species is described. The species have been assigned to primary locomotor categories on the basis of their locomotion. These are 1, climbing, arboreal walking; 2, arboreal and terrestrial walking and jumping; 3, general terrestrial walking and scrambling; and 4, trotting. In the climbing arboreal walking category the most distinctive morphological adaptations are powerful flexors and extensors as well as a flexible plantigrade manus with retractile claws. In the arboreal and terrestrial walking category the shoulder, elbow and carpal joints are flexible and the manus has retractile claws, though the flexor and extensor musculature is insufficiently developed for controlled climbing. The trotting category is characterised by a high humero-radial index and a rigid antibrachium. The foot is digitigrade with the claws short and stout. Species in the general walking and scrambling category show many differences in the morphology of their feet, even though the proximal parts of the forelimb appear similar. Due to the restricted nature of the adaptations, these species have been assigned to secondary locomotor categories. Morphological characters typical of the locomotor categories are summarized in the discussion.     

Devonian tetrapod trackways and trackmakers; a review of the fossils and footprints

The earliest tetrapods are known from the Upper Devonian. Their remains are becoming better known from increasing numbers of specimens, localities, environments and ichnofossils. Each of the eight (or possibly nine) genera now represented by skeletal fossils is reviewed in its sedimentological, faunal and stratigraphic context, with an assessment of what might be inferred about the habitus and locomotory capabilities of each. Fossil trackways and their interpretations are then re-examined in the context of the known body forms, and consideration given to the degree of fit between the skeletal fossils, the trackways and their interpretations. The currently known Devonian tetrapods are unlikely to have made any of the known tracks, unless they were produced under water. Neither the skeletal fossils nor the trackways show good evidence of terrestrial locomotion among Devonian tetrapods. When the fossil material and recent phylogenetic analyses are taken in combination, it appears that neither tetrapods nor limbs with digits are likely to have arisen before the Frasnian. This should be borne in mind in palaeoecological studies of these animals.     

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.     

Nonplantigrade Foot Posture: A Constraint on Dinosaur Body Size

Dinosaurs had functionally digitigrade or sub-unguligrade foot postures. With their immediate ancestors, dinosaurs were the only terrestrial nonplantigrades during the Mesozoic. Extant terrestrial mammals have different optimal body sizes according to their foot posture (plantigrade, digitigrade, and unguligrade), yet the relationship of nonplantigrade foot posture with dinosaur body size has never been investigated, even though the body size of dinosaurs has been studied intensively. According to a large dataset presented in this study, the body sizes of all nonplantigrades (including nonvolant dinosaurs, nonvolant terrestrial birds, extant mammals, and extinct Nearctic mammals) are above 500 g, except for macroscelid mammals (i.e., elephant shrew), a few alvarezsauroid dinosaurs, and nondinosaur ornithodirans (i.e., the immediate ancestors of dinosaurs). When nonplantigrade tetrapods evolved from plantigrade ancestors, lineages with nonplantigrade foot posture exhibited a steady increase in body size following Cope’s rule. In contrast, contemporaneous plantigrade lineages exhibited no trend in body size evolution and were largely constrained to small body sizes. This evolutionary pattern of body size specific to foot posture occurred repeatedly during both the Mesozoic and the Cenozoic eras. Although disturbed by the end-Cretaceous extinction, species of mid to large body size have predominantly been nonplantigrade animals from the Jurassic until the present; conversely, species with small body size have been exclusively composed of plantigrades in the nonvolant terrestrial tetrapod fauna.     

Gigantism among Late Jurassic limulids: New ichnological evidence from the Causses Basin (Lozère,France) and comments on body-size evolution among horseshoe crabs

An abundant ichnological material composed of xiphosuran trackways and isolated traces was discovered in Upper Jurassic limestones from the Causses Basin (Causse Méjean, Lozère, France). The morphology of the imprints supports their identification as Kouphichnium isp. In contrast to the most frequent case, the trackways are composed of omnipresent pusher imprints sometime associated with leg traces, but with no telson mark. We argue that this pattern reflects actual surface traces rather than an incomplete set of undertracks. The size distribution of the sampled ichnites is broadly bimodal. This is best explained by sexual dimorphism, a phenomenon frequently observed in modern xiphosurans. Analysis of the trace fossils further suggests that several growth stages are recorded and that the horseshoe crabs were walking in a protected and flat environment like a lagoon. This area, certainly close to a mating ground, was occasionally affected by a continental influence. The biometric study of the tracks suggests a gigantic size for the trackmakers whose body may have reached 84 cm in length. This discovery complements the few reports on other gigantic horseshoe crabs in the Jurassic of Western Europe, thus casting doubt on the postulated increase in body size from the Palaeozoic to the Recent. Furthermore, a literature review shows that there are still major gaps in the record of limulid body-fossils and tracks. Thus, neither of these archives can be taken at face value for quantifying the body-size evolution of horseshoe crabs.     

Cope's Rule in the Pterosauria, and differing perceptions of Cope's Rule at different taxonomic levels

The remarkable extinct flying reptiles, the pterosaurs, show increasing body size over 100 million years of the Late Jurassic and Cretaceous, and this seems to be a rare example of a driven trend to large size (Cope's Rule). The size increases continue throughout the long time span, and small forms disappear as larger pterosaurs evolve. Mean wingspan increases through time. Examining for Cope's Rule at a variety of taxonomic levels reveals varying trends within the Pterosauria as a whole, as pterodactyloid pterosaurs increase in size at all levels of examination, but rhamphorhynchoid pterosaurs show both size increase and size decrease in different analyses. These results suggest that analyses testing for Cope's Rule at a single taxonomic level may give misleading results.