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Pliocene Crocodilians of Chinchilla: Identification Using Dental Morphometrics

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Pliocene Crocodilians of Chinchilla: Identification Using Dental Morphometrics Christina Chiotakis Bachelor of Applied Science (Ecology) Supervised by Dr Matthew Phillips (QUT) and Dr Scott Hocknull (Queensland Museum) Submitted in fulfilment of the requirements for the degree of Master of Science (Research) Faculty of Science and Engineering Queensland University of Technology 2018 CHRISTINA CHIOTAKIS 08301166 Keywords: Chinchilla Local Fauna, Crocodilian, crocodile, Crocodylus, Pallimnarchus, Pliocene, Quinkana, teeth Abstract: The following thesis examines the Pliocene crocodilians of the Chinchilla Local Fauna. Crocodilians are an extremely diverse group of mostly amphibious, opportunistic hunters that include twenty-three extant species across the globe. A number of extinct species have also been described from Australia’s prehistory. Two species have been described from the Chinchilla Local Fauna, Pallimnarchus pollens and Quinkana sp. Teeth fossilise readily, and so make up the largest dataset of fossilised evidence of crocodilian species. For this study, the teeth were grouped visually, measured and scanned to created three-dimensional models for analysis. When graphed on a PCA, it is possible to distinguish two distinct groups of teeth. The two groups are differentiated based on overall shape of the tooth. One group contains conical teeth, the other laterally compressed teeth. Further analysis of the teeth was able to determine the approximate position of the teeth along the tooth row. Alveoli on available jaw specimens were also measured to determine if there was a way to match isolated teeth to them. However, when jaw specimens were measured the site shows potential for three different crocodilian taxa occurring within the Chinchilla Local Fauna. Further research will need to be conducted to determine the third potential species and correct positioning of isolated teeth along the tooth row. 1 CHRISTINA CHIOTAKIS 08301166 0.0 CONTENTS PAGE Section Page Number 0.0 Contents Page 2 0.1 List of Figures 3 0.2 List of Tables 6 0.3 List of Abbreviations 6 0.4 Glossary 6 Statement of Original Authorship 8 Acknowledgements 8 1.0 Introduction 9 1.1 Objectives 9 1.2 Significance 10 2.0 Literature Review 10 2.1 Crocodilians 10 2.2 Cenozoic Crocodilian Fossil Record 13 2.3 Plio-Pleistocene Extinction 18 2.4 Chinchilla Sands Local Fauna 20 2.5 Australian Crocodilians 25 2.5.1 Extant Australian Crocodilians 25 2.5.1.1 Crocodylus porosus 25 2.5.1.2 Crocodylus johnstoni 26 2.5.2 Extinct Australian Crocodilians 27 2.5.2.1 Pallimnarchus sp. 27 2.5.2.2 Quinkana spp. 28 2.6 Varanus priscus 29 2.7 Fossilised Teeth 30 2.8 Future Research into Extinct Crocodilians 32 2.9 Conclusion 33 3.0 Methodology 34 4.0 Ethics and Limitations 37 5.0 Results 37 5.1 Initial Dataset 37 5.1.1 Raw Data 38 2 CHRISTINA CHIOTAKIS 08301166 5.1.2 Principle Components Analysis 38 5.2 CT Scanned Dataset 41 5.2.1 Raw Data 41 5.2.2 Root Means Square Data Matrix 42 5.2.3 Principle Components Analysis 42 5.2.4 Non-Metric Multi-Dimensional Scaling 43 5.3 Caniniform and Molariform 45 5.3.1 Raw Data 46 5.3.2 Principle Components Analysis 46 5.3.3 Non-Metric Multi-Dimensional Scaling 48 5.4 Analysis Based Upon Ratio 52 5.4.1 Ratio of Mid-Height and Base Length versus 52 Width 5.4.2 Non-Metric Multi-Dimensional Scaling 55 5.5 Alveoli Measurements 58 5.5.1 Crocodylus sp. 58 5.5.2 Pallimnarchus sp. 59 5.5.3 Quinkana sp. 60 5.5.4 Unknown Species 61 6.0 Discussion 61 6.1 Future Research 69 7.0 Reference List 72 8.0 Appendices 87 9.0 Supplementary Figures 116 0.1 List of Figures Figure Number and Title Page Number 1 Ranges and Estimated Population Size of Extant 11 Crocodilian Species 2 Artists Impression of an Early Triassic Crocodilian 12 3 Possible Homo sapiens migration routes 19 4 Map of Main Queensland Fossils Sites 20 3 CHRISTINA CHIOTAKIS 08301166 5 Map of Crocodylus porosus Distribution across Australia 25 6 Map of Crocodylus johnstoni Distribution across 26 Australia 7 Location of Pliocene and Pleistocene Crocodilian bearing 28 Fossil Sites 8 Palaeogeography of the Giant Varanids 30 9 Examples of ziphodontid and conical Crocodilian Teeth 32 10 Diagram of Crocodilian Teeth 35 11 Principle Components Analysis with Crocodylus spp. 39 12 Principle Components Analysis without Crocodylus spp. 40 13 Principle Components Analysis with Crocodylus spp. 40 without Height 14 Principle Components Analysis without Crocodylus sp. 41 and Height 15 Principle Components Analysis of 3D Dataset 42 16 Principle Components Analysis of 3D Dataset without 43 Height 17 Non-Metric Multi-Dimensional Scaling of 3D Dataset 44 18 Non-Metric Multi-Dimensional Scaling with Teeth 45 19 Side and Base View of Teeth from Non-Metric Multi- 45 Dimensional Scaling Graph 20 Principle Components Analysis of Caniniform Teeth 46 21 Principle Components Analysis of Caniniform Teeth 47 without Height 22 Principle Components Analysis of Molariform Teeth 47 23 Principle Components Analysis of Molariform Teeth 48 without Height 24 Non-Metric Multi-Dimensional Scaling of Caniniform 49 Dataset 25 Non-Metric Multi-Dimensional Scaling with 50 Caniniform Teeth 26 Side and Base View of Caniniform Teeth from Non- 50 Metric Multi-Dimensional Scaling Graph 4 CHRISTINA CHIOTAKIS 08301166 27 Non-Metric Multi-Dimensional Scaling of Molariform 51 Dataset 28 Non-Metric Multi-Dimensional Scaling with 51 Molariform Teeth 29 Side and Base View of Molariform Teeth from Non- 52 Metric Multi-Dimensional Scaling Graph 30 Histogram Based Upon the Ratio of Mid-Height Length 53 vs Width 31 Histogram Based Upon the Ratio of Crown Base Length 53 vs Width 32 Scatterplot of Crown Base Length/Crown Base Width 54 vs Height 33 Scatterplot of Mid-Height Length/Width vs Height 54 34 Scatterplot of Mid-Height Length/Width vs Crown Base 55 Length/Width 35 Non-Metric Multi-Dimensional Scaling Coloured by 56 Ratios 36 Non-Metric Multi-Dimensional Scaling Coloured by 57 Ratios with Teeth 37 Side and Base View of Teeth from Non-Metric Multi- 57 Dimensional Scaling Graph 38 Alveoli Measurements of Crocodylus spp. 58 39 Alveoli Measurements of Pallimnarchus spp. 59 40 Alveoli Measurements of Quinkana spp. 60 41 Alveoli Measurements of Unknown species 61 42 Dentary Specimens of Modern Crocodylus porosus and 66 Crocodylus johnstoni 43 Fossilised Dentary Specimens 67 44 Quinkana Tooth from Mount Etna Deposits 69 45 Pallimnarchus sp. Dentary from the Darling Downs 71 46 Crocodilian Dentary from Winton 71 5 CHRISTINA CHIOTAKIS 08301166 0.2 List of Tables Table Number and Title Page Number 1 Cenozoic Crocodilian Species from Australia and 14 Oceania 2 Species belonging to the Chinchilla Local Fauna 21 0.3 List of Abbreviations PCA – Principle Components Analysis QM – Queensland Museum QMF – Queensland Museum Fossil CT (Scan) – Computed Tomography (Scan) MDS – Multi-Dimensional Scaling RMS – Root Means Square Mya – Million Years Ago 0.4 Glossary Labial – referring to cheek Lingual – referring to tongue Ziphodont – Laterally compressed, distally curved, serrated crowns (D’Amore and Bluenschine, 2009) Alveoli – Crypt in the jaw in which teeth are imbedded Osteoderm – Bony plate under the skin Maxilla – upper jaw Dentary – lower jaw Mekosuchine – Distinct group of Australia-western Pacific crocodilians belonging to the Eusuchia Eusuchian – Clade to which all extant crocodilians belong Pterygoid – Bone forming part of the palate in a skull Local Fauna – Species making up a spatially and 6 CHRISTINA CHIOTAKIS 08301166 temporally discrete fauna (e.g. Chinchilla Local Fauna 7 CHRISTINA CHIOTAKIS 08301166 Statement of Original Authorship: The work contained in this thesis has not been previously submitted to meet requirements for an award at this or any other higher education institution. To the best of my knowledge and belief, the thesis contains no material previously published or written by another person except where due reference is made. Signature: QUT Verified Signature Date: June 2018 Acknowledgements: I would like to thank my supervisors’ Dr Matthew Phillips and Dr Scott Hocknull for all their input and encouragement throughout the project. I extend this thank you to the Vertebrate Collection Managers Dr Andrew Amey and Dr Patrick Couper at the Queensland Museum for allowing me access to the modern crocodilian specimens. And thanks to my family and friends for their continued support throughout the completion of this research. I would also like to thank everyone at Geosciences, Queensland Museum, in particular, Kristen Spring for allowing me access to the collection, and Nikki Newman and Rochelle Lawrence for co-ordinating CT Scanning for the fossil teeth. A special thanks to Greenslopes Private Hospital and QX-ray for access to the CT Scanner for data collection, and the Phillips Lab Team, in particular, Carmelo Fruciano, for aiding with the statistical analysis. 8 CHRISTINA CHIOTAKIS 08301166 1.0 Introduction: This chapter outlines the objectives and significance of the research. This will be followed by a literature review of current and relevant scientific research to provide a background on the topic. The methodology will then be outlined followed by the results of the research. Finally, there will be a full analysis of all data collected, relating back to the relevant literature and potential future applications of the work. 1.1 Objectives The aim of this study is to find a way of identifying crocodilian species and species diversity for a single Local Fauna, the Chinchilla Local Fauna, by utilising the most abundant fossils available. Inspection of the Queensland Museum vertebrate fossil collection has identified teeth as one of the most common fossils found of crocodilians from the Chinchilla Local Fauna, due to their durability and constant shedding throughout the life of the animal.
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    https://doi.org/10.24199/j.mmv.1973.34.03 9 May 1973 FOSSIL VERTEBRATE FAUNAS FROM THE LAKE VICTORIA REGION, S.W. NEW SOUTH WALES, AUSTRALIA By Larry G. Marshall Zoology Department, Monash University, Victoria* Abstract Fossil vertebrate localities and faunas in the Lake Victoria region of S.W. New South Wales, Australia, are described. The oldest fossil bearing deposit, the late Pliocene or early Pleistocene Moorna Formation and associated Chowilla Sand have yielded specimens of Neoceratodus sp., Emydura macquarrii, several species of small dasyurid, specimens of Glaucodon cf. G. ballaratensis, a species of Protemnodon which compares closely with P. cf. P. otibandus from the late Pliocene or early Pleistocene Chinchilla Sand in SE. Queensland, specimens of Lagostrophus cf L. fasciatus, species of Petrogale, Macropus, Osphranter, Sthenurus, Bettongia, Diprotodon, Lasiorhinus, a peramelid, at least two species of pseudomyine rodents and a species of Rattus cf. R. lutreolus. It is shown that the holotype of Zygomaturus victoriae (Owen) 1872 may have been collected from these sediments. The late Pliocene or early Pleistocene Blanchetown Clay has yielded species of Neoceratodus, Thylacoleo, Phas- colonus, Bettongia, Sthenurus, a diprotodontid, and a rodent. The late Pleistocene Rufus Formation has yielded species of Dasycercus, Sarcophilus, Thylacinus, Phascolonus, Lasiorhinus, Bettongia, Procoptodon, Onychogalea, Macropus, and Leporillus. A large species of macropod and a species of Phascolonus were collected from the late Pleistocene Monoman Formation. The lunette on the E. side of Lake Victoria has yielded a large, diverse fauna of late Pleistocene —Holocene age, that includes such extinct species as Protemnodon anak, P. brehus, Procoptodon goliah, Sthenurus andersoni, S.