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Morphological Variation and Ichnotaxonomy of Dinosaur Tracks: Linking Footprint Shapes to Substrate And

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Morphological Variation and Ichnotaxonomy of Dinosaur Tracks: Linking Footprint Shapes to Substrate And ADVERTIMENT. Lʼaccés als continguts dʼaquesta tesi queda condicionat a lʼacceptació de les condicions dʼús establertes per la següent llicència Creative Commons: http://cat.creativecommons.org/?page_id=184 ADVERTENCIA. El acceso a los contenidos de esta tesis queda condicionado a la aceptación de las condiciones de uso establecidas por la siguiente licencia Creative Commons: http://es.creativecommons.org/blog/licencias/ WARNING. The access to the contents of this doctoral thesis it is limited to the acceptance of the use conditions set by the following Creative Commons license: https://creativecommons.org/licenses/?lang=en Morphological variation and ichnotaxonomy of dinosaur tracks: linking footprint shapes to substrate and trackmaker's anatomy and locomotion Novella Razzolini PhD Thesis Universitat Autònoma de Barcelona Facultat de Ciències Departament de Geologia 2016 Front cover illustration by Dr. Oriol Oms: reconstruction of Megalosaurids crossing tidal flats Morphological variation and ichnotaxonomy of dinosaur tracks: linking footprint shapes to substrate and trackmaker's anatomy and locomotion Memoir presented by Novella Razzolini in order to apply for the degree of Doctor in Geology by the Universitat Autònoma de Barcelona, Departament de Geologia, complemented under the tutelage of Dr. Oriol Oms and the supervision of: - Dr. Àngel Galobart Lorente, Institut Català de Paleontologia Miquel Crusafont - Dr. Bernat Vila i Ginestí, Institut Català de Paleontologia Miquel Crusafont Dr. Àngel Galobart Lorente Dr. Bernat Vila i Ginestí Novella Razzolini WORK FUNDED BY: - FPI grant BES-2012-051847 associated to the project CGL2011-30069-C02-01 (Ministerio de Ciencia e Innovación, Spain) - Ministerio de Economia y Competitividad (Spain) who funded abroad interniships at: - EEBB-I-14-09084: Royal Veterinary College of London (UK) - EEBB-I-15-09494: Museu Nacional de História Natural e da Ciência (Portugal) - EEBB-I-16-11441: Naturhistorisches Museum (Basel, Switzerland) - Research developed in CERCA program. Generalitat de Catalunya - Institut Català de Paleontologia “Miquel Crusafont” Actually, everything that can be known has a Number; for it is impossible to grasp anything with the mind or to recognize it without this Philolaus, greek pythagorean and presocratic philosopher CONTENTS ABSTRACT/Resum 1 PART 1. INTRODUCTION Chapter 1. Structure of the dissertation and goals 9 Chapter 2. Account of work undertaken 2.1 Motivation and background 15 2.2 Context of localities 17 2.3 Material and Methods 23 PART 2. TRACK MORPHOLOGICAL VARIATION AND INTERPRETATION Chapter 3. Ichnological context 3.1. Historical background 37 3.2. Documenting, describing and quantifying tracks 42 Chapter 4. Difficulties in interpreting tracks 4.1 Track morphological variation and trackmaker’s anatomy 53 4.2 Track morphological variation and trackmaker’s locomotion 57 4.3 Track morphological variation and substrate 63 PART 3. TRACK MORPHOLOGICAL VARIATION AND TRACKMAKER’S ANATOMY Chapter 5. Pes shape variation in an ornithopod dinosaur trackway (Lower Cretaceous, NW Spain): New evidence of an antalgic gait in the fossil track record 5.1 Author contribution 75 77 5.2 Article Chapter 6. Megalosauripus transjurani ichnosp. nov. A new Late Jurassic theropod ichnotaxon from NW Switzerland 6.1 Author contribution 89 6.2 Article 91 PART 4. TRACK MORPHOLOGICAL VARIATION AND TRACKMAKER’S LOCOMOTION Chapter 7. Ichnological evidence of Megalosaurid dinosaurs crossing Middle Jurassic tidal flats 7.1 Author contribution 193 7.2 Article 195 Chapter 8. Crossing slopes: unusual trackways of recent birds and implications for tetrapod footprint preservation 8.1 Author contribution 219 8.2 Article 221 PART 5. TRACK MORPHOLOGICAL VARIATION AND SUBSTRATE Chapter 9. Intra-trackway morphological variations due to substrate consistency: the El Frontal dinosaur tracksite (Lower Cretaceous, Spain). 9.1 Author contribution 241 9.2 Article 243 Chapter 10. First attempts of laboratory-controlled simulations of vulture footprints in heterogeneous substrates 261 GENERAL DISCUSSION 277 GENERAL CONCLUSION 287 Summary statements 288 Future directions 289 REFERENCES 293 ACKNOWLEDGEMENTS 313 ANNEX FOR CHAPTER 6 319 ABSTRACT Abstract/Resum Abstract The study of tridactyl dinosaurs tracks from a morphological perspective is here presented in the form of a new look and approach to the classic ichnology. The mechanisms that mostly control and affect track morphology during its formation have been thoroughly analysed, so that the ultimate goal is to provide extensive quantitative data to discuss the main foundation of morphological variation registered in individual tracks, trackways and large sets of trackways. The compendium of this PhD collected six different studies from distinct geographical (Spain, Portugal, Morocco and Switzerland) and geological frames (from Middle Jurassic to present) and the proposal of a preliminary approach in a laboratory controlled-experiment. This experiment aims to extrapolate and determine all parameters (water content, substrate properties and composition, grain size, porosity) intervening in the track formation process on heterogeneous substrates during the mechanical indentation of a vulture foot cast. The Cameros basin (NW Spain) offered two Early Cretaceous different scenarios for this thesis, the re-interpretation of a long ornithopod trackway on homogeneous substrate at the Barranco de La Canal tracksite and the new study of four previously unpublished theropod trackways inter-crossing a heterogeneous substrate at the El Frontal tracksite. The locality visited within the Argana basin (Morocco) provided neoichnological observations that are consider important for the identification of misinterpretations of ichnopathologies and sloping surfaces in the fossil record. These localities showed two different intra- trackway morphological variation patterns defined as alternate, which depends on the limb dynamics and foot anatomy of the trackmaker and continuous, which depends on the substrate consistency change along the tracking surface. The Lusitanian basin (Central-West Portugal) presented the previously known but unpublished Middle Jurassic quarry of Vale de Meios which underscores the importance of analyzing all types of track preservations in order to recognize the average morphology and that there should not be an assumption that vertebrate ichnotaxa are confined to specific ages or geographic regions. The Jura Carbonate platform (NW Switzerland) encompassed six Late Jurassic tracksites, which together with 49 trackways and 397 tracks provided the new and unpublished material for the description of a new ichnospecies and the discussion of track morphological variations, introducing the possibility that classic ichnoassociations might be the result of preservational variants of the same trackmaker. These two studies showed that when comparing multiple trackways on the same site or ichnoassemblage, taxonomical diversity and behavioural changes have to be considered together with substrate conditions and limb dynamics. Three-dimensional technologies have been the support and tool for all the quantitative analysis undertaken. LiDAR scans have been always complemented with a close range photogrammetry in order to give the highest morphological details and to provide a precise and systematic quantification of the track morphological variations recorded. 1 Abstract/Resum Resum La present tesi presenta un estudi de les petjades tridàctiles de dinosaure a partir d’una perspectiva morfològica que representa una nova visió i aproximació a la icnologia clàssica. S’han analitzat en profunditat els mecanismes que principalment controlen i afecten la morfologia de les petjades durant la seva formació, amb l’objectiu final d’aportar dades àmplies i quantitatives per discutir el fonament principal de la variació morfològica registrada en petjades individuals, rastres i conjunts de rastres. El compendi d’aquesta tesi recull sis estudis diferents de diversos marcs geogràfics (España, Portugal, Marroc i Suïssa) i geològics (del Juràssic Mitjà a l’actualitat) i la proposta d’un enfocament preliminar en un experiment controlat de laboratori. Aquest experiment pretén extrapolar i determinar tots els paràmetres (contingut d’aigua, propietats i composició del substrat, mida de gra, porositat) que intervenen en el procés de formació d’una petjada en substrats heterogenis durant la penetració mecànica d’un motlle de peu de voltor. Per aquesta tesi la conca de Cameros (Nord-oest d’Espanya) ofereix dos escenaris diferents del Cretaci Inferior: la reinterpretació d’un rastre llarg d’ornitòpode sobre un substrat homogeni al jaciment amb petjades de Barranco de La Canal, i el nou estudi de quatre rastres inèdits de teròpodes creuant-se en un substrate heterogeni al jaciment de petjades d’El Frontal. La localitat de la conca de l’Argana (Marroc) ha proporcionat observacions neoicnològiques que es consideren importants a l’hora d’identificar interpretaciones errònies d’icnopatologies i superfícies en pendent en el registre fòssil. Aquestes localitats mostren diferents patrons de variació morfològica intrarastre definits com alternat, el qual depèn de la dinàmica de les extremitats i l’anatomia del peu de l’animal productor; i continu, el qual depèn de la consistència del substrat al llarg de la superfície on es produeixen les petjades. La conca lusitana (centre-oest de Portugal) conté la localitat ja coneguda però inèdita del Juràssic Mitjà de Vale de Meios, l’estudi de la qual
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