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.

BackgroundLarge ornithopod tracks are known from the Upper Jurassic to the uppermost Cretaceous rocks of all continents but Antarctica. They include the tracks historically called Iguanodon footprints, iguanodontid footprints, hadrosaur/hadrosaurid footprints, and other large ornithopod tracks that have been used to define ichnotaxa. More than 40 ichnospecies based on large ornithopod tracks have been defined, but the validity of many of them is questionable.Methodology/Principal Findings34 ichnogenera and 44 ichnospecies have been analysed in this work. Many of them are considered to be invalid because they have been defined on the basis of poorly preserved tracks without diagnostic features, have an inadequate diagnosis, or are based on temporal and/or geographical criteria. Only eight ichnospecies belonging to the ichnogenera Caririchnium, Iguanodontipus and Hadrosauropodus are here regarded as valid.Conclusions/SignificanceThe monospecific ichnogenus Iguanodontipus (I. burreyi) is characterized by a small, rounded heel and elongate, narrow digit impressions. Its distribution is limited to the Berriasian-Valanginian of Europe. Caririchnium consists of four ichnospecies (C. magnificum [type ichnospecies], C. kortmeyeri, C. billsarjeanti and C. lotus) with a large, rounded heel and short, wide digit impressions. This ichnogenus ranges from the Berriasian-Hauterivian to the Aptian-Albian of South America, North America, Asia and Europe. Finally, Hadrosauropodus (three ichnospecies: H. langstoni [type ichnospecies], H. leonardii and H. kyoungsookimi) shows a large, bilobed heel and short, wide digit impressions. It is known from the Aptian-Albian to the Maastrichtian of North America, Asia and Europe. The ichnofamily Iguanodontipodidae includes large iguanodontian tracks characterized mainly by mesaxonic, tridactyl and subsymmetrical pes tracks that are as wide as (or wider than) long and have one pad impression in each digit and one in the heel. Its distribution is confidently limited to the Cretaceous of Europe, Asia, North America and South America.

Disarticulated and incomplete remains from a new diminutive ornithopod are described. They come from the Cameros Basin in the north of Spain and were collected from the red clays of the Castrillo de la Reina Formation, ranging from Upper Barremian to Lower Aptian. The new ornithopod described here is slender and one of the smallest ever reported. An up-to-date phylogenetic analysis recovers this taxon as a basal iguanodontian. Its unique combination of characters makes it more derived than slender ornithopods like Hyphilophodon and Gasparinisaura, and bring very interesting insights into the basal iguanodontian phylogeny. Though possessing a minimum of three premaxillary teeth, this taxon also bears an extensor ilio-tibialis groove on the distal part of its femur. Moreover, its dentary and maxillary teeth are unique, remarkably similar to those regarded as having a “rhabdomorphan” affinity. This unknown taxon is suggested to be a stem taxon within Rhabdodontidae, a successful clade of basal iguanodonts from the Late Cretaceous of Europe. The Gondwanan ornithopods share the strongest affinities with this family, and we confirm Muttaburrasaurus as a sister taxon of the Rhabdodontidae within a newly defined clade, the Rhabdodontomorpha.

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Fossil associations from the middle and upper Eocene (Bartonian and Priabonian) sedimentary succession of the Pamplona Basin are described. This succession was accumulated in the western part of the South Pyrenean peripheral foreland basin and extends from deep-marine turbiditic (Ezkaba Sandstone Formation) to deltaic (Pamplona Marl, Ardanatz Sandstone and Ilundain Marl formations) and marginal marine deposits (Gendulain Formation). The micropalaeontological content is high. It is dominated by foraminifera, and common ostracods and other microfossils are also present. The fossil ichnoasssemblages include at least 23 ichnogenera and 28 ichnospecies indicative of Nereites, Cruziana, and ?Scoyenia-Mermia ichnofacies. Body macrofossils of about 80 taxa corresponding to macroforaminifera, sponges, corals, bryozoans, brachiopods, annelids, molluscs, arthropods, echinoderms and vertebrates have been identified. Both the number of ichnotaxa and of species (e. g. bryozoans, molluscs and condrichthyans) may be considerably higher. Body fossil assemblages are comparable to those from the Eocene of the Nord Pyrenean area (Basque Coast), and also to those from the Eocene of the west-central and eastern part of South Pyrenean area (Aragon and Catalonia). At the European scale, the molluscs assemblages seem endemic from the Pyrenean area, although several Tethyan (Italy and Alps) and Northern elements (Paris Basin and Normandy) have been recorded. Palaeontological data of studied sedimentary units fit well with the shallowing process that throughout the middle and late Eocene occurs in the area, according to the sedimentological and stratigraphical data.Se describen las asociaciones fósiles del Eoceno medio y superior (Bartoniense y Priaboniense) de la sucesión sedimentaria de la Cuenca de Pamplona. Esta sucesión se acumuló en la parte oeste de la Cuenca de antepaís periférica surpirenaica y comprende desde turbiditas marinas profundas (Formación Areniscas de Ezkaba) hasta depósitos deltaicos (Formaciones Margas de Pamplona, Areniscas de Ardanatz y Margas de Ilundain) y marinos marginales (Formación Gendulain). El contenido micropaleontológico es alto y está dominado por los foraminíferos. Los fósiles de ostrácodos y otros organismos son también abundantes. Las asociaciones de icnofósiles incluyen al menos 23 icnogéneros y 28 ichnoespecies, indicativas de las icnofacies de Nereites, Cruziana, y ?Scoyenia-Mermia. Se han identificado fósiles corporales de unos 80 taxones, correspondientes a macroforaminíferos, esponjas, corales, briozoos, braquiópodos, anélidos, moluscos, artrópodos, equinodermos y vertebrados. El número de icnotaxones y de especies (p. e. briozoos, moluscos gasterópodos, bivalvos y peces condríctios) puede ser considerablemente mayor. Las asociaciones de fósiles corporales son comparables a las del Eoceno del Área norpirenaica (Costa vasca) y de las partes central y este del Área surpirenaica (Aragón y Cataluña). A escala europea, la asociación de moluscos parece endémica del Área pirenaica,...

Here, we report the recent discovery of 87 Neandertal footprints on the Southwest of the Iberian Peninsula (Doñana shoreline, Spain) located on an upper Pleistocene aeolian littoral setting (about 106 ± 19 kyr). Morphometric comparisons, high resolution digital photogrammetric 3D models and detailed sedimentary analysis have been provided to characterized the footprints and the palaeoenvironment. The footprints were impressed in the shoreline of a hypersaline swamped area related to benthic microbial mats, close to the coastline. They have a rounded heel, a longitudinal arch, relatively short toes, and adducted hallux, and represent the oldest upper Pleistocene record of Neandertal footprints in the world. Among these 87 footprints, 31 are longitudinally complete and measure from 14 to 29 cm. The calculated statures range from 104 to 188 cm, with half of the data between 130 and 150 cm. The wide range of sizes of the footprints suggests the existence of a social group integrated by individuals of different age classes but dominated, however, by non-adult individuals. The footprints, which are outside the flooded area are oriented perpendicular to the shoreline. These 87 footprints reinforce the ecological scenario of Neandertal groups established in coastal areas.