The complexity of tracking stegosaurs and their gregarious behavior.

Stegosaur tracks were unknown until the identification of Deltapodus more than 20 years ago. Currently, the Iberian Peninsula, especially Teruel Province, is one of the areas globally with the most occurrences of these tracks. However, their identification, based on the global record, is problematic due to their similarities with sauropod tracks. A review of the largest number of analyzed Deltapodus tracks globally, including the holotype of D. ibericus and a description of new occurrences, has been carried out. Our research shows substantial morphological variations, but all the studied tracks can be considered D. ibericus based on the manus morphology and the morphometric data. These variations are related to substrate differences and/or different dynamic foot postures (possibly ontogenetically related) during locomotion, as evidenced by changes within the same trackway. We provide detailed comparisons via 3D modeling with sauropod tracks, and our data show that they generally have proportionally longer manus and wider pes because of the differences in the metapodial bones. The scarcity of stegosaur trackways in the fossil record has prevented the identification of gregarious behavior in this group of herbivorous dinosaurs. Two of the studied tracksites show evidence of this behavior, being the only examples among stegosaurs described thus far in the fossil record.

The largest European theropod dinosaurs: remains of a gigantic megalosaurid and giant theropod tracks from the Kimmeridgian of Asturias, Spain.

The Kimmeridgian Vega, Tereñes and Lastres formations of Asturias have yielded a rich vertebrate fauna, represented by both abundant tracks and osteological remains. However, skeletal remains of theropod dinosaurs are rare, and the diversity of theropod tracks has only partially been documented in the literature. Here we describe the only non-dental osteological theropod remain recovered so far, an isolated anterior caudal vertebra, as well as the largest theropod tracks found. The caudal vertebra can be shown to represent a megalosaurine megalosaurid and represents the largest theropod skeletal remain described from Europe so far. The tracks are also amongst the largest theropod footprints reported from any setting and can be assigned to two different morphotypes, one being characterized by its robustness and a weak mesaxony, and the other characterized by a strong mesaxony, representing a more gracile trackmaker. We discuss the recently proposed distinction between robust and gracile large to giant theropod tracks and their possible trackmakers during the Late Jurassic-Berriasian. In the absence of complete pedal skeletons of most basal tetanurans, the identity of the maker of Jurassic giant theropod tracks is difficult to establish. However, the notable robustness of megalosaurine megalosaurids fits well with the described robust morphotypes, whereas more slender large theropod tracks might have been made by a variety of basal tetanurans, including allosaurids, metriocanthosaurids or afrovenatorine megalosaurids, or even exceptionally large ceratosaurs. Concerning osteological remains of large theropods from the Late Jurassic of Europe, megalosaurids seem to be more abundant than previously recognized and occur in basically all Jurassic deposits where theropod remains have been found, whereas allosauroids seem to be represented by allosaurids in Western Europe and metriacanthosaurids in more eastern areas. Short-term fluctuations in sea level might have allowed exchange of large theropods between the islands that constituted Europe during the Late Jurassic.

A walk in the maze: variation in Late Jurassic tridactyl dinosaur tracks from the Swiss Jura Mountains (NW Switzerland).

BACKGROUND: Minute to medium-sized (footprint length (FL) less than 30 cm) tridactyl dinosaur tracks are the most abundant in the Late Jurassic tracksites of Highway A16 (Reuchenette Formation, Kimmeridgian) in the Jura Mountains (NW Switzerland). During excavations, two morphotypes, one gracile and one robust, were identified in the field. Furthermore, two large-sized theropod ichnospecies (Megalosauripus transjuranicus and Jurabrontes curtedulensis) and an ornithopod-like morphotype (Morphotype II) have recently been described at these sites. METHODS: The quality of morphological preservation (preservation grade), the depth of the footprint, the shape variation, and the footprint proportions (FL/footprint width (FW) ratio and mesaxony) along the trackways have been analyzed using 3D models and false-color depth maps in order to determine the exact number of small to medium-sized morphotypes present in the tracksites. RESULTS: The study of footprints (n = 93) recovered during the excavations has made it possible to identify and characterize the two morphotypes distinguished in the field. The gracile morphotype is mainly characterized by a high FL/FW ratio, high mesaxony, low divarication angles and clear, sharp claw marks, and phalangeal pads (2-3-4). By contrast, the robust morphotype is characterized by a lower FL/FW ratio, weaker mesaxony, slightly higher divarication angles and clear, sharp claw marks (when preserved), whereas the phalangeal pads are not clearly preserved although they might be present. DISCUSSION: The analysis does not allow the two morphotypes to be associated within the same morphological continuum. Thus, they cannot be extramorphological variations of similar tracks produced by a single trackmaker. Comparison of the two morphotypes with the larger morphotypes described in the formation (M. transjuranicus, J. curtedulensis, and Morphotype II) and the spatio-temporal relationships of the trackways suggest that the smaller morphotypes cannot reliably be considered as small individuals of any of the larger morphotypes. The morphometric data of some specimens of the robust morphotype (even lower values for the length/width ratio and mesaxony) suggest that more than one ichnotaxon might be represented within the robust morphotype. The features of the gracile morphotype (cf. Kalohipus isp.) are typical of "grallatorid" ichnotaxa with low mesaxony whereas those of the robust morphotype (cf. Therangospodus isp. and Therangospodus? isp.) are reminiscent of Therangospodus pandemicus. This work sheds new light on combining an analysis of variations in footprint morphology through 3D models and false-color depth maps, with the study of possible ontogenetic variations and the identification of small-sized tridactyl ichnotaxa for the description of new dinosaur tracks.

Ichnological evidence of Megalosaurid Dinosaurs Crossing Middle Jurassic Tidal Flats.

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.

A reappraisal of the Middle Triassic chirotheriid Chirotherium ibericus Navás, 1906 (Iberian Range NE Spain), with comments on the Triassic tetrapod track biochronology of the Iberian Peninsula.

Triassic vertebrate tracks are known from the beginning of the 19th century and have a worldwide distribution. Several Triassic track ichnoassemblages and ichnotaxa have a restricted stratigraphic range and are useful in biochronology and biostratigraphy. The record of Triassic tracks in the Iberian Peninsula has gone almost unnoticed although more than 25 localities have been described since 1897. In one of these localities, the naturalist Longinos Navás described the ichnotaxon Chirotherium ibericus in 1906.The vertebrate tracks are in two sandy slabs from the Anisian (Middle Triassic) of the Moncayo massif (Zaragoza, Spain). In a recent revision, new, previously undescribed vertebrate tracks have been identified. The tracks considered to be C. ibericus as well as other tracks with the same morphology from both slabs have been classified as Chirotherium barthii. The rest of the tracks have been assigned to Chirotheriidae indet., Rhynchosauroides isp. and undetermined material. This new identification of C. barthii at the Navás site adds new data to the Iberian record of this ichnotaxon, which is characterized by the small size of the tracks when compared with the main occurrences of this ichnotaxon elsewhere. As at the Navás tracksite, the Anisian C. barthii-Rhynchosauroides ichnoassemblage has been found in other coeval localities in Iberia and worldwide. This ichnoassemblage belongs to the upper Olenekian-lower Anisian interval according to previous biochronological proposals. Analysis of the Triassic Iberian record of tetrapod tracks is uneven in terms of abundance over time. From the earliest Triassic to the latest Lower Triassic the record is very scarce, with Rhynchosauroides being the only known ichnotaxon. Rhynchosauroides covers a wide temporal range and gives poor information for biochronology. The record from the uppermost Lower Triassic to the Middle Triassic is abundant. The highest ichnodiversity has been reported for the Anisian with an assemblage composed of Dicynodontipus, Procolophonichnium, Rhynchosauroides, Rotodactylus, Chirotherium, Isochirotherium, Coelurosaurichnus and Paratrisauropus. The Iberian track record from the Anisian is coherent with the global biochronology proposed for Triassic tetrapod tracks. Nevertheless, the scarcity of track occurrences during the late Olenekian and Ladinian prevents analysis of the corresponding biochrons. Finally, although the Iberian record for the Upper Triassic is not abundant, the presence of Eubrontes, Anchisauripus and probably Brachychirotherium is coherent with the global track biochronology as well. Thus, the Triassic track record in the Iberian Peninsula matches the expected record for this age on the basis of a global biochronological approach, supporting the idea that vertebrate Triassic tracks are a useful tool in biochronology.

Intra-trackway morphological variations due to substrate consistency: the El Frontal dinosaur tracksite (Lower Cretaceous, Spain).

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.

Discriminating between medium-sized Tridactyl Trackmakers: tracking Ornithopod tracks in the base of the Cretaceous (Berriasian, Spain).

BACKGROUND: Recent work on the Jurassic-Cretaceous transition of the Iberian Range (Spain) has opened a new window onto the interpretation of the trackmakers of some medium-sized tridactyl tracks. The ichnotaxon Therangospodus oncalensis has been described in the Huérteles Formation (Berriasian) and is one of the classical tracks from the area assigned to medium-sized theropods. METHODOLOGY/PRINCIPAL FINDINGS: A review of the type locality of Therangospodus oncalensis (Fuentesalvo tracksite) and other tracksites from the Huérteles Formation (Berriasian) has yielded new information on the morphology, gait and trackmaker identity of the aforementioned ichnospecies. The new data suggest that the trackmaker is an ornithopod rather than a theropod on the basis of the length/width ratio, the anterior triangle length-width ratio, the short steps, the round to quadrangular heel pad impression and the probable manus impressions. CONCLUSIONS/SIGNIFICANCE: T. oncalensis shows similarities with various tracks from the Berriasian of Europe assigned to Iguanodontipus. The ichnotaxonomical status of this ichnospecies is here considered as Iguanodontipus? oncalensis due to the current state of knowledge of the ichnotaxonomy of medium-sized ornithopod tracks. This reassessment of I? oncalensis also has two significant implications for the palaeoecology of the faunas during the deposition of the Huérteles Formation: 1- the high number and percentage of theropod tracks would be lower than previous papers have suggested. 2- the gregarious behaviour described in the type locality (Fuentesalvo) would be among ornithopods instead of theropods.

Manus track preservation bias as a key factor for assessing trackmaker identity and quadrupedalism in basal ornithopods.

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.