The morphology of fossil footprints is the basis of vertebrate footprint ichnology. However, the processes acting during and after trace fossil registration which are responsible for the final morphology have never been precisely defined, resulting in a dearth of nomenclature. Therefore, we discuss the concepts of ichnotaphonomy, ichnostratinomy, taphonomy, biostratinomy, registration and diagenesis and describe the processes acting on footprint morphology. In order to evaluate the morphological quality of tetrapod footprints, we introduce the concept of morphological preservation, which is related to the morphological quality of footprints (M-preservation, acronym MP), and distinguish it from physical preservation (P-preservation, acronym PP), which characterizes whether or not a track is eliminated by taphonomic and diagenetic processes. M-preservation includes all the morphological features produced during and after track registration prior to its study, and may be divided into substages (ichnostratinomic, registrational, taphonomic, stratinomic, diagenetic). Moreover, we propose an updated numerical preservation scale for M-preservation. It ranges from 0.0 (worst preservation) to 3.0 (best preservation); intermediate values may be used and specific features may be indicated by letters. In vertebrate footprint ichnotaxonomy, we regard the anatomy-consistent morphology and to a lesser extent the trackway pattern as the only acceptable ichnotaxobases. Only footprints showing a good morphological preservation (grade 2.0-3.0) are useful in ichnotaxonomy, whereas ichnotaxa based on poor morphological preservation (grade 0.0-1.5) are considered ichnotaphotaxa (nomina dubia) characterized by extramorphologies. We applied the preservation scale on examples from the Palaeozoic to the present time, including three ichnotaphotaxa and 18 anatomy-consistent ichnotaxa/morphotypes attributed to several vertebrate footprint producers. Results indicate the utility, feasibility and suitability of this method for the entire vertebrate footprint record in any lithofacies, strongly recommending its use in future ichnotaxonomic studies.

The collection and dissemination of vertebrate ichnological data is struggling to keep up with techniques that are becoming commonplace in the wider palaeontological field. A standard protocol is required to ensure that data is recorded, presented and archived in a manner that will be useful both to contemporary researchers, and to future generations. Primarily, our aim is to make the 3D capture of ichnological data standard practice, and to provide guidance on how such 3D data can be communicated effectively (both via the literature and other means) and archived openly and in perpetuity. We recommend capture of 3D data, and the presentation of said data in the form of photographs, false‐colour images, and interpretive drawings. Raw data (3D models of traces) should always be provided in a form usable by other researchers (i.e. in an open format). If adopted by the field as a whole, the result will be a more robust and uniform literature, supplemented by unparalleled availability of datasets for future workers.

BackgroundRecent 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 FindingsA 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.

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.

BackgroundThe 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 FindingsDetailed 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/SignificanceSeveral 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.

A new Iberian pleurosternid (Jurassic-Cretaceous transition, Spain) and first neuroanatomical study of this clade of stem turtlesA partial skeleton of a pleurosternid turtle (Paracryptodira), from the Jurassic-Cretaceous transition at the Spanish locality of Ágreda (Moncayo Region, Soria Province), is presented here. Its partial skull represents the third of this lineage to be recognized in the European record, with the previously known specimens corresponding to British species. The specimen of Pleurosternidae studied here is attributed to a new species, Pleurosternon moncayensis. This lineage of stem turtles is identified as the most abundant and diverse group of freshwater aquatic turtles in Europe for the stages adjacent to the Jurassic-Cretaceous transition. Its presence decreased radically at post-Berriasian levels, when freshwater lineages of Eucryptodira, of Asian origin, are identified as the dominant forms at these aquatic turtle faunas. The confirmation of Pleurosternidae as freshwater inhabitants is made here, through the first neuroanatomical study for this lineage. Thus, the neuroanatomical reconstruction of Pleurosternon moncayensis sp. nov. is the first to be carried out for a freshwater stem turtle, and it allows us to identify convergent adaptations with freshwater members of the crown Testudines.

Geometric morphometric methods applied to theropod tracks from the Huérteles Formation (Berriasian, Spain) are here shown to be invaluable for drawing comparisons between theropod tracks with different preservation modes (true tracks, shallow undertracks and natural casts) or differing in the preservation of anatomical features (e.g. digital pads). Principal components analysis and thin‐plate spline methods can quantitatively distinguish between the broad groups of tracks in a sample and establish the main differences between them. These methods offer a promising approach for estimating ichnodiversity, achieved by evaluating just the morphology of the tracks independent of other factors such as size. The theropod tracks of the Huérteles Formation can be classified into two broad groups: minute‐to‐medium‐sized gracile theropod tracks (Kalohipus bretunensis) and medium‐to‐large‐sized robust theropod tracks (Iberosauripus). The presence of a third group of more gracile medium‐to‐large‐sized theropod tracks (Megalosauripus) cannot be proven with certainty on the basis of the current data. These results indicate that the theropod ichnodiversity of the Huérteles Formation is probably lower than that estimated by means of conventional methods alone (e.g. qualitative description of the tracks) and that many of the described theropod morphotypes may represent extramorphological or ontogenetic variations of other morphotypes.

J., 2012: Ethological variations in gauge in sauropod trackways from the Berriasian of Spain. Lethaia, Vol. 45, Two sauropod trackways from the Early Cretaceous (Berriasian) of the Cameros Basin (Spain) show important variations in trackway gauge, along the same trackway. These variations seem to be associated with different behaviours; on the one hand with a significant variation in the direction of travel (turning phenomena) and on the other hand with a decrease in speed, probably associated with the properties of the substrate. These variations in sauropod trackways provide the trackway gauge debate with new data, supporting the hypothesis that the walking style and substrate properties may in some cases determine this sauropod character. The study of turning sauropod trackways improves our knowledge of sauropod locomotion.