Ichnology Research Papers - Academia.edu (original) (raw)

Dinosaur tracks are biogenic, sedimentary structures and not body fossils or biological objects in the common sense. They result from the complex interaction of the kinematics of the trackmaker, its foot anatomy, and the substrate properties, and from taphonomic processes acting prior to the incorporation of the tracks into the sedimentary record. The objective of this work is an interdisciplinary study of a large sample of dinosaur tracks and trackways linking sedimentology with vertebrate ichnology, palaeontology, and palaeoecology.

Excellent conditions are provided by the Late Jurassic (Kimmeridgian) Chevenez—Combe Ronde tracksite, which is one of several tracksites located on the future course of the Transjurane highway near Porrentruy (Canton Jura, NW Switzerland). Here, eight superimposed dinosaur track-bearing surfaces were systematically excavated level-by-level within a 0,65 m thick laminite interval, unearthing almost 1400 dinosaur tracks. The main track level, located at the base of the interval, is the most diverse ichnoassemblage composed of 14 trackways of tiny (Pes Length < 25 cm) and small (25 cm < PL < 50 cm) sauropods and 43 trackways of minute (PL < 10 cm), small (10 cm < PL < 20 cm), and medium-sized
(20 cm < PL < 30 cm) bipedal, tridactyl dinosaurs.

The main issues are: (1) identification of true tracks, undertracks, and overtracks, and their relationships with substrate properties, their link with the exposure index, and their utility in the reconstruction of the palaeoenvironment; (2) implications of the main track level ichnoassemblage for dinosaur behaviour, the terrestrial palaeoecosystem, and vertebrate ichnofacies; (3) relationships between variability in trackway patterns and configurations with locomotion speed, behaviour, and substrate properties as well as implications for locomotion capabilities; (4) Quantification and relevance of sauropod trackway gauge; and (5) interpretation of manus-dominated and pes-only sauropod trackways.

The approach is first actualistic by studying human footprints and processes acting during their formation and preservation on modern tidal-flats. In these highly structured environments, microbial mats are ubiquitous, strongly facies-specific, and occupy a key position during and after footprint formation. Undertracks readily form in biolaminated sediment, whilst underprints and deep tracks are common in unlaminated, water-saturated sediment. Most consolidated vertebrate tracks are affected by taphonomic processes, including renewed and/or repeated growth of microbial mats leading to the formation of modified true tracks, internal overtracks (track fills), and overtracks.

The sauropod tracks and the encasing laminite interval of the Combe Ronde site are then subject of detailed sedimentological and taphonomical analyses. This discloses the sediment properties at the time of track formation and reveals the processes modifying the tracks during subaerial exposure and integrating them into the sedimentary record. Track morphology, associated track features, and sedimentary features can be linked with the exposure index, identifying the palaeoenvironment as a supratidal flat not located in close proximity to a coastline. These flats were susceptible for track recording only during short periods after wetting due to a rainy period or due to occasional storms. Longer periods of subaerial exposure prior to burial are indicated by the presence of internal overtracks and/or overtracks, and rapid covering up is indicated by the lack of overtracks on top of tracks with large displacement rims. Cross-sections of sauropod tracks provide insight into the consolidation history of the substrate prior to track formation and into the walking dynamics of dinosaurs, confirming that sauropods put their hindfeet in a pronounced plantigrade way on the ground.

The level-by-level superimposition of the studied surfaces enables to identify true tracks, undertracks, and overtracks. The best-defined true tracks (anatomical morphotypes) of the main track level are then used for ichnotaxonomy and trackmaker identification, and the detailed analyses of trackway parameters, including trackway gauge, provide insight into the locomotion capabilities of dinosaurs.

The best-defined minute and small tridactyl tracks can be assigned to the ichnogenus Carmelopodus, extending it from the Middle Jurassic into the Late Jurassic. These tracks were likely left by a small theropod dinosaur similar in size to Compsognathus or Juravenator. The medium-sized tridactyl tracks of morphotype II exhibit some of the typical features of the ichnogenus Therangospodus (attributed to large and robust theropods) but also some of ornithopod ichnotaxa.

The sauropod trackways show a wide range of patterns and configurations but are all medium- to wide-gauge. Therefore, they are assigned tentatively to the ichnogenus Brontopodus attributed to derived “brachiosaurid” or “titanosaurid” dinosaurs. The variability of the trackways reflects the general locomotion capabilities of the trackmakers and is an expression of individual walking style and behaviour, which may be related to substrate properties. Trackway patterns (the degree of manus overprinting by the pes) and different trackway configurations including trackway gauge are not only related to locomotion speed, and they provide no evidence of a relationship with ontogeny.

The gauge of sauropod trackways can be quantified with the pes trackway ratio and the here defined [WAP/PL]-ratio (Width of the pes Angulation Pattern / Pes Length). Gauge is possibly related to the substrate and the behaviour of the trackmaker adapting to it, but this does not change the overall medium-gauge to wide-gauge appearance of the trackways. The manus-dominated and pes-only sauropod trackways of the Combe Ronde site are explained by trackmakers exerting more pressure on the manus than the pes, and by overprinting of the manus by the pes, respectively.

The alignment of trackways on the main track level shows no evidence of a nearby shoreline and of interactions between the different groups of dinosaurs. It indicates gregarious behaviour amongst tiny and small sauropods, and suggests that minute and small bipedal dinosaurs were frequent visitors on the supratidal flats.

The ichnoassemblage of the main track level is the first one found in the Jura Mountains displaying abundant minute and small tridactyl tracks. This is also typical for the other Ajoie ichnoassemblages, which further exhibit tracks of tiny to large (up to 1,1 m PL) sauropods, and tracks of medium-sized to large (up to 0,8 m PL) bipedal dinosaurs. Sauropod trackways include narrow-gauge and wide-gauge trackways indicating the presence of “basal” and derived sauropods. This suggests that dwarfed insular animals can be excluded as trackmakers of the tiny and small sauropod trackways of the Ajoie ichnoassemblages and the Combe Ronde tracksite and that the Jura carbonate platform was connected with the landmasses of the London-Brabant Massif and the Massif Central during periods of emersion. Dinosaurs used the Jura carbonate platform for the establishment of in situ, predominantly saurischian dinosaur populations, but also as a migration corridor between the massifs.

Because the Ajoie ichnoassemblages are dominated by small tridactyl tracks, they differ from other Jurassic tetrapod ichnofacies in carbonate settings, notably from the Brontopodus ichnofacies. In the case of those ichnoassemblages commonly attributed to the Brontopodus ichnofacies, the lack or rareness of small tridactyl tracks may indicate the absence of small trackmakers in those palaeoenvironments or unsuitable conditions for the formation and preservation of small tracks.

This study highlights the benefits of systematic and interdisciplinary analyses of dinosaur tracks, which disclose variations related to behaviour and to differences in substrate. This allows recognizing anatomical morphotypes and trackway configurations representative of typical trackmaker behaviour. The latter can then also be used in ichnotaxonomical classification. Similar approaches should be in the focus of future work and performed on the other tracksites and ichnoassemblages of the Ajoie. Together with the evidence from other tracksites of the Jura Mountains, this will contribute towards a better understanding of the terrestrial palaeoenvironments and palaeogeography, and of dinosaur palaeoecology and palaeobiogeography on the Jura carbonate platform.