Tracking the Pliensbachian-Toarcian Karoo firewalkers: Trackways of quadruped and biped dinosaurs and mammaliaforms.
ABSTRACT: The Karoo igneous rocks represent one of the largest continental flood basalt events (by volume) on Earth, and are not normally associated with fossils remains. However, these Pliensbachian-Toarcian lava flows contain sandstone interbeds that are particularly common in the lower part of the volcanic succession and are occasionally fossiliferous. On a sandstone interbed in the northern main Karoo Basin, we discovered twenty-five tridactyl and tetradactyl vertebrate tracks comprising five trackways. The tracks are preserved among desiccation cracks and low-amplitude, asymmetrical ripple marks, implying deposition in low energy, shallow, ephemeral water currents. Based on footprint lengths of 2-14 cm and trackway patterns, the trackmakers were both bipedal and quadrupedal animals of assorted sizes with walking and running gaits. We describe the larger tridactyl tracks as "grallatorid" and attribute them to bipedal theropod dinosaurs, like Coelophysis, a genus common in the Early Jurassic of southern Africa. The smallest tracks are tentatively interpreted as Brasilichnium-like tracks, which are linked to synapsid trackmakers, a common attribution of similar tracks from the Lower to Middle Jurassic record of southern and southwestern Gondwana. The trackway of an intermediate-sized quadruped reveals strong similarities in morphometric parameters to a post-Karoo Zimbabwean trackway from Chewore. These trackways are classified here as a new ichnogenus attributable to small ornithischian dinosaurs as yet without a body fossil record in southern Africa. These tracks not only suggest that dinosaurs and therapsids survived the onset of the Drakensberg volcanism, but also that theropods, ornithischians and synapsids were among the last vertebrates that inhabited the main Karoo Basin some 183 Ma ago. Although these vertebrates survived the first Karoo volcanic eruptions, their rapidly dwindling habitat was turned into a land of fire as it was covered by the outpouring lavas during one of the most dramatic geological episodes in southern Africa.
Project description:LIDAR-based analyses of the first theropod dinosaur trackways known from the state of Arkansas, USA are reported. The trackways were found on a limestone bedding plane in the Albian De Queen Formation in an active gypsum quarry. Because limited access precluded thorough field study, fieldwork focused on preserving the entire site digitally with ground-based LIDAR, and detailed measurements were later taken digitally from point cloud data. The site contains eight tridactyl trackways associated with sauropod trackways and numerous isolated tracks. Although there appear to be two different tridactyl morphotypes, we show that the tracks are all likely from a single species of trackmaker. We apply a simple method of estimating substrate consistency by comparing the differences between true track dimensions and apparent track dimensions. The tridactyl tracks at the southern end of the site are preserved with significantly greater differences in true vs. apparent dimensions and are shallower than the rest of the tridactyl tracks at the site, which we interpret as the result of outward expansion of the soft tissues of the foot upon contact with a firm substrate. We interpret the firm substrate as having high bulk density and high shear strength, which also explain associated manus-only sauropod tracks. We show that the tridactyl tracks are likely from theropod trackmakers and that footprint lengths, trackway paces, stride lengths, and pace angulations of the De Queen trackways are statistically indistinguishable from equivalent measurements of theropod trackways in the Glen Rose Formation. The Glen Rose tracks are attributed to the large-bodied theropod, Acrocanthosaurus and we likewise attribute the De Queen tracks to Acrocanthosaurus, which is known from skeletal remains in temporally equivalent units and from the mine itself.
Project description:A new ichnospecies of a large theropod dinosaur, Megalosauripus transjuranicus, is described from the Reuchenette Formation (Early-Late Kimmeridgian, Late Jurassic) of NW Switzerland. It is based on very well-preserved and morphologically-distinct tracks (impressions) and several trackways, including different preservational types from different tracksites and horizons. All trackways were excavated along federal Highway A16 near Courtedoux (Canton Jura) and systematically documented in the field including orthophotos and laserscans. The best-preserved tracks were recovered and additional tracks were casted. Megalosauripus transjuranicus is characterized by tridactyl tracks with clear claw and digital pad impressions, and notably an exceptionally large and round first phalangeal pad on the fourth digit (PIV1) that is connected to digit IV and forms the round heel area. Due to this combination of features, M. transjuranicus clearly is of theropod (and not ornithopod) origin. M. transjuranicus is compared to other Megalosauripus tracks and similar ichnotaxa and other unassigned tracks from the Early Jurassic to Early Cretaceous. It is clearly different from other ichnogenera assigned to large theropods such as Eubrontes-Grallator from the Late Triassic and Early Jurassic or Megalosauripus-Megalosauropus-Bueckeburgichnus and Therangospodus tracks from the Late Jurassic and Early Cretaceous. A second tridactyl morphotype (called Morphotype II) is different from Megalosauripus transjuranicus in being subsymmetric, longer than wide (sometimes almost as wide as long), with blunt toe impressions and no evidence for discrete phalangeal pad and claw marks. Some Morphotype II tracks are found in trackways that are assigned to M. transjuranicus, to M.? transjuranicus or M. cf. transjuranicus indicating that some Morphotype II tracks are intra-trackway preservational variants of a morphological continuum of Megalosauripus transjuranicus. On the other hand, several up to 40 steps long trackways very consistently present Morphotype II features (notably blunt digits) and do not exhibit any of the features that are typical for Megalosauripus (notably phalangeal pads). Therefore, it is not very likely that these tracks are preservational variants of Megalosauripus transjuranicus or Megalosauripus isp. These trackways are interpreted to have been left by an ornithopod dinosaur. The high frequency of large theropod tracks in tidal-flat deposits of the Jura carbonate platform, associated on single ichnoassemblages with minute to medium-sized tridactyl and tiny to large sauropod tracks has important implications for the dinosaur community and for paleoenvironmental and paleogeographical reconstructions. As with most other known occurrences of Megalosauripus tracks, M. transjuranicus is found in coastal settings, which may reflect the preference of their theropod trackmakers for expanded carbonate flats where food was abundant.
Project description:BACKGROUND: The Las Cerradicas site (Tithonian-Berriasian), Teruel, Spain, preserves at least seventeen dinosaur trackways, some of them formerly attributed to quadrupedal ornithopods, sauropods and theropods. The exposure of new track evidence allows a more detailed interpretation of the controversial tridactyl trackways as well as the modes of locomotion and taxonomic affinities of the trackmakers. METHODOLOGY/PRINCIPAL FINDINGS: Detailed stratigraphic analysis reveals four different levels where footprints have been preserved in different modes. Within the tridactyl trackways, manus tracks are mainly present in a specific horizon relative to surface tracks. The presence of manus tracks is interpreted as evidence of an ornithopod trackmaker. Cross-sections produced from photogrammetric digital models show different depths of the pes and manus, suggesting covariance in loading between the forelimbs and the hindlimbs. CONCLUSIONS/SIGNIFICANCE: Several features (digital pads, length/width ratio, claw marks) of some ornithopod pes tracks from Las Cerradicas are reminiscent of theropod pedal morphology. This morphological convergence, combined with the shallow nature of the manus tracks, which reduces preservation potential, opens a new window into the interpretation of these tridactyl tracks. Thus, trackmaker assignations during the Jurassic-Cretaceous interval of purported theropod trackways may potentially represent ornithopods. Moreover, the Las Cerradicas trackways are further evidence for quadrupedalism among some basal small- to medium-sized ornithopods from this time interval.
Project description:The historically-famous Lotus Fortress site, a deep 1.5-3.0-meter-high, 200-meter-long horizonal notch high up in near-vertical sandstone cliffs comprising the Cretaceous Jiaguan Formation, has been known since the 13th Century as an impregnable defensive position. The site is also extraordinary for having multiple tetrapod track-bearing levels, of which the lower two form the floor of part of the notch, and yield very well preserved asseamblages of ornithopod, bird (avian theropod) and pterosaur tracks. Trackway counts indicate that ornithopods dominate (69%) accounting for at least 165 trackmakers, followed by bird (18%), sauropod (10%), and pterosaur (3%). Previous studies designated Lotus Fortress as the type locality of Caririchnium lotus and Wupus agilis both of which are recognized here as valid ichnotaxa. On the basis of multiple parallel trackways both are interpreted as representing the trackways of gregarious species. C. lotus is redescribed here in detail and interpreted to indicate two age cohorts representing subadults that were sometimes bipedal and larger quadrupedal adults. Two other previously described dinosaurian ichnospecies, are here reinterpreted as underprints and considered nomina dubia. Like a growing number of significant tetrapod tracksites in China the Lotus Fortress site reveals new information about the composition of tetrapod faunas from formations in which the skeletal record is sparse. In particular, the site shows the relatively high abundance of Caririchium in a region where saurischian ichnofaunas are often dominant. It is also the only site known to have yielded Wupus agilis. In combination with information from other tracksites from the Jiaguan formation and other Cretaceous formations in the region, the track record is proving increasingly impotant as a major source of information on the vertebrate faunas of the region. The Lotus Fortress site has been developed as a spectacular, geologically-, paleontologically- and a culturally-significant destination within Qijiang National Geological Park.
Project description:Large well-preserved crocodylomorph tracks from the Lower Cretaceous (? Aptian) Jinju Formation of South Korea, represent the well-known crocodylomorph ichnogenus Batrachopus. The Korean sample includes multiple, narrow-gauge, pes-only trackways with footprint lengths (FL) 18-24?cm, indicating trackmaker body lengths up to ~3.0?m. Surprisingly, the consistent absence of manus tracks in trackways, with well-preserved digital pad and skin traces, argues for bipedal trackmakers, here assigned to Batrachopus grandis ichnosp. nov. No definitive evidence, either from pes-on-manus overprinting or poor track preservation, suggests the trackways where made by quadrupeds that only appear bipedal. This interpretation helps solve previous confusion over interpretation of enigmatic tracks of bipeds from younger (? Albian) Haman Formation sites by showing they are not pterosaurian as previously inferred. Rather, they support the strong consensus that pterosaurs were obligate quadrupeds, not bipeds. Lower Jurassic Batrachopus with foot lengths (FL) in the 2-8?cm range, and Cretaceous Crocodylopodus (FL up to ~9.0?cm) known only from Korea and Spain registered narrow gauge trackways indicating semi-terrestrial/terrestrial quadrupedal gaits. Both ichnogenera, from ichnofamily Batrachopodidae, have been attributed to Protosuchus-like semi-terrestrial crocodylomorphs. The occurrence of bipedal B. grandis ichnosp. nov. is evidence of such adaptations in the Korean Cretaceous.
Project description:An ichnological and sedimentological study of the El Frontal dinosaur tracksite (Early Cretaceous, Cameros basin, Soria, Spain) highlights the pronounced intra-trackway variation found in track morphologies of four theropod trackways. Photogrammetric 3D digital models revealed various and distinct intra-trackway morphotypes, which reflect changes in footprint parameters such as the pace length, the track length, depth, and height of displacement rims. Sedimentological analyses suggest that the original substrate was non-homogenous due to lateral changes in adjoining microfacies. Multidata analyses indicate that morphological differences in these deep and shallow tracks represent a part of a continuum of track morphologies and geometries produced by a gradient of substrate consistencies across the site. This implies that the large range of track morphologies at this site resulted from similar trackmakers crossing variable facies. The trackways at the El Frontal site present an exemplary case of how track morphology, and consequently potential ichnotaxa, can vary, even when produced by a single trackmaker.
Project description:Ichniotherium tracks with a relatively short pedal digit V (digit length ratio V/IV < 0.6) form the majority of yet described Late Carboniferous to Early Permian diadectomorph tracks and can be related to a certain diadectid clade with corresponding phalangeal reduction that includes Diadectes and its close relatives. Here we document the variation of digit proportions and trackway parameters in 25 trackways (69 step cycles) from nine localities and seven further specimens with incomplete step cycles from the type locality of Ichniotherium cottae (Gottlob quarry) in order to find out whether this type of Ichniotherium tracks represents a homogeneous group or an assemblage of distinct morphotypes and includes variability indicative for evolutionary change in trackmaker locomotion. According to our results, the largest sample of tracks from three Lower Permian sites of the Thuringian Forest, commonly referred to I. cottae, is not homogeneous but shows a clear distinction in pace length, pace angulation, apparent trunk length and toe proportions between tracks from Bromacker quarry and those from the stratigraphically older sites Birkheide and Gottlob quarry. Three Late Carboniferous trackways of Ichniotherium with relatively short pedal digit V from Haine's Farm, Ohio, and Alveley near Birmingham, United Kingdom, that have been referred to the ichnotaxa "Baropus hainesi," "Megabaropus hainesi" and "Ichniotherium willsi," respectively, share a marked outward rotation of foot imprints with respect to walking direction. Apart from this feature they are in many aspects similar to the Birkheide and Gottlob records of I. cottae. With the possible exception of the Maroon Formation (Early Permian, Colorado) sample, Early Permian Ichniotherium trackways with a relatively short pedal digit V fall into the morphological spectrum of the three well defined "Hainesi-Willsi," "Birkheide-Gottlob" and "Bromacker" morphotypes. With their more obtuse pace angulations and higher body-size-normalized pace and stride lengths the Bromacker type tracks imply higher walking speeds of their trackmakers compared to all other Ichniotherium tracks. More generally, a trend towards higher locomotion capability from the last common ancestor of all Ichniotherium producers to the last common ancestor of all "Ichniotherium with relatively short pedal digit V" and from the latter to the trackmakers of the mid-Early Permian Bromacker type can be deduced-with the reservation that overall sample size is relatively small, making this scenario a preliminary assessment. Whether the presumed advancements represent a more general pattern within diadectomorphs remains open until the non-European Ichniotherium trackway record improves. Ichnotaxonomic implications are discussed.
Project description: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 degrees ), 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.
Project description:The new ichnospecies Paleohelcura araraquarensis isp. nov. is described from the Upper Jurassic-Lower Cretaceous Botucatu Formation of Brazil. This formation records a gigantic eolian sand sea (erg), formed under an arid climate in the south-central part of Gondwana. This trackway is composed of two track rows, whose internal width is less than one-quarter of the external width, with alternating to staggered series, consisting of three elliptical tracks that can vary from slightly elongated to tapered or circular. The trackways were found in yellowish/reddish sandstone in a quarry in the Araraquara municipality, São Paulo State. Comparisons with neoichnological studies and morphological inferences indicate that the producer of Paleohelcura araraquarensis isp. nov. was most likely a pterygote insect, and so could have fulfilled one of the ecological roles that different species of this group are capable of performing in dune deserts. The producer could have had a herbivorous or carnivorous diet or been part of the fauna of omnivores, being able to adopt herbivorous, carnivorous, and saprophagous diets when necessary. In modern dune deserts, some species of pterygote insects are detritivores (like Tenebrionidae), relying on organic matter that accumulated among the sand grains of the dunes during dry periods with no plant growth. The presence of additional burrows suggests that the Botucatu paleodesert would have had a detritivorous fauna like this. Based on the interpretation of the ichnofossil producers, it was possible to reconstruct the food web of this paleodesert. All the omnivorous and herbivorous invertebrates and the herbivorous ornithopod dinosaurs made up the primary consumers. These animals were, in turn, the food source for bigger carnivorous or omnivorous animals unable to feed on detritus, like arachnids, possible predatory insects, mammaliaforms, and theropod dinosaurs. The highest trophic level was occupied by larger theropod dinosaurs and mammaliaforms, which, because of their size, could prey upon a wide range of animals. The producer of Paleohelcura araraquarensis isp. nov. could have been a primary consumer if it were an omnivorous detritivore or a herbivore, or a secondary consumer if it were produced by a predatory insect or an omnivore relying on animal biomass. The description of this new trackway expands the knowledge on the faunal composition of the Botucatu paleodesert and provides insights into the ecological relationships in ancient deserts. The presence of these arthropod trackways in Mesozoic eolian deposits helps to trace a continuity between Paleozoic and post-Paleozoic desert ichnofaunas, further reinforcing a single Octopodichnus-Entradichnus Ichnofacies for eolian deposits.
Project description:A new dinosaur tracksite in the Vale de Meios quarry (Serra de Aire Formation, Bathonian, Portugal)preserves more than 700 theropod tracks. They are organized in at least 80 unidirectional trackways arranged in a bimodal orientation pattern (W/NW and E/SE). Quantitative and qualitative comparisons reveal that the large tridactyl, elongated and asymmetric tracks resemble the typical Late Jurassic-Early Cretaceous Megalosauripus ichnogenus in all morphometric parameters. Few of the numerous tracks are preserved as elite tracks while the rest are preserved as different gradients of modified true tracks according to water content, erosive factors, radial fractures and internal overtrack formations. Taphonomical determinations are consistent with paleoenvironmental observations that indicate an inter-tidal flat located at the margin of a coastal barrier. The Megalosauripus tracks represent the oldest occurrence of this ichnotaxon and are attributed to large megalosaurid dinosaurs. Their occurrence in Vale de Meios tidal flat represents the unique paleoethological evidence of megalosaurids moving towards the lagoon, most likley during the low tide periods with feeding purposes.