This study concerns the formation, taphonomy, and preservation of human footprints in microbial mats of present-day tidal-flat environments. Due to differences in water content and nature of the microbial mats and the underlying sediment, a wide range of footprint morphologies was produced by the same trackmaker. Most true tracks are subjected to modification due to taphonomic processes, leading to modified true tracks. In addition to formation of biolaminites, microbial mats play a major role in the preservation of footprints on tidal flats. A footprint may be consolidated by desiccation or lithification of the mat, or by ongoing growth of the mat. The latter process may lead to the formation of overtracks. Among consolidated or (partially) lithified footprints found on present-day tidal flats, poorly defined true tracks, modified true tracks, and overtracks were most frequently encountered while unmodified and well-defined true tracks are rather rare. We suggest that modified true tracks and overtracks make up an important percentage of fossil footprints and that they may be as common as undertracks. However, making unambiguous distinctions between poorly defined true tracks, modified true tracks, undertracks, and overtracks in the fossil record will remain a difficult task, which necessitates systematic excavation of footprints combined with careful analysis of the encasing sediment.
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.
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.
Jurabrontes curtedulensis, a new ichnogenus and species of Late Jurassic giant theropod dinosaur track is described based on very well-preserved and morphologically-distinct tracks, all carefully excavated along federal highway A16 (Canton Jura, NW Switzerland). All trackways were systematically documented including parameter measurements, descriptions, outline drawings, orthophotos and laserscans. Jurabrontes is characterised by sub-equal track length and width, a small anterior triangle, weak mesaxony, three blunt digits (dII-III-IV) with pronounced (sub)triangular claw marks, a rounded heel, and clear phalangeal pad impressions. The combination of features of Jurabrontes is typical for a theropod (and not ornithopod) trackmaker. Jurabrontes is compared to other similar ichnotaxa and unnamed tracks of large theropods from the Early Jurassic to Late Cretaceous, from which it is clearly different. The sheer size of the largest tracks, that are amongst the largest worldwide and of similar size to Tyrannosauripus from the Late Cretaceous, suggests a 'megalosaurid' or large allosaurid theropod as a trackmaker. The presence of such large theropod tracks in tidal-flat deposits of the Jura carbonate platform and associated with small to large sauropod tracks has important palaeoecological implications for the dinosaur community and for palaeoenvironmental and palaeogeographical reconstructions.
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.
Force Base containing some of the largest tracts. Feral swine damage is one of the greatest threats to this wetland habitat. We conducted a multi-year study to evaluate the impacts of sport hunting and supplemental swine removal on damage to seepage slopes. Prior to initiation of removal in 2003, swine damage to seepage slopes in the portion of the base closed to hunting averaged 25.0%, over twice the 10.9% losses in the portion open to hunting. After less than one year of supplemental removal, damage in the closed-to-hunting area dropped to 7.2%. Although supplemental removal was not applied in the open hunting area, damage dropped significantly to 5.6%, statistically indistinguishable from the swine-controlled (closed) portion. After another year of removal, average damage in the closed hunting area dropped further to 5.6%, while the open hunting area dropped to 4.3%, again statistically indistinguishable. Even though removal was only applied to the area closed to hunting, it also produced damage reductions in the open hunting area, as swine were free to move among areas. Declines in damage following implementation of removal corresponded with large drops in swine population indices for the base. Economic valuations of seepage slope damage losses demonstrated substantial benefit-cost ratios for application of removal.
In 2002 a new dinosaur tracksite was discovered in calcareous laminites of early Late Kimmeridgian age along the future course of the "Transjurane" highway in Courtedoux, Canton Jura, Northern Switzerland. The site has an extraordinary scientific potential, as the laminites, which have been deposited in an intertidal to supratidal environment, contain at least 6 track-bearing levels in a total thickness of about 1 m. The laminites are being systematically excavated by the "Section de paléontologie" over an area of approximately 1500 m 2 . So far the main track level has been uncovered over an area of about 650 m 2 , which reveals 2 trackways of theropods and 17 trackways of sauropods. The sauropod tracks are the smallest known in the Kimmeridgian so far, and the trackways belong to the ichnogenus Parabrontopodus, which has been revealed for the first time in Switzerland. The tracksite belongs to the "Middle Kimmeridgian megatracksite" sensu Meyer (2000), and represents the most important dinosaur tracksite in Switzerland, perhaps with the potential for development into one of the world's largest sauropod tracksites. It will be protected in situ underneath an especially constructed highway-bridge, thus offering opportunities for future research and the development of an interpretative center for education and tourism.
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