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Evolutionary Progression of the Iconic Australasian Kangaroos, Rat-Kangaroos, and Their Fossil Relatives (Marsupialia: Macropodiformes)

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Evolutionary Progression of the Iconic Australasian Kangaroos, Rat-Kangaroos, and Their Fossil Relatives (Marsupialia: Macropodiformes) Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 1624 Evolutionary Progression of the Iconic Australasian Kangaroos, Rat-Kangaroos, and their Fossil Relatives (Marsupialia: Macropodiformes) WENDY DEN BOER ACTA UNIVERSITATIS UPSALIENSIS ISSN 1651-6214 ISBN 978-91-513-0216-4 UPPSALA urn:nbn:se:uu:diva-339593 2018 Dissertation presented at Uppsala University to be publicly examined in Hamberg, Geocentrum, Department of Earth Sciences, Villavägen 16, Uppsala, Friday, 9 March 2018 at 10:00 for the degree of Doctor of Philosophy. The examination will be conducted in English. Faculty examiner: Dr. Christine Janis. Abstract den Boer, W. 2018. Evolutionary Progression of the Iconic Australasian Kangaroos, Rat-Kangaroos, and their Fossil Relatives (Marsupialia: Macropodiformes). Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 1624. 105 pp. Uppsala: Acta Universitatis Upsaliensis. ISBN 978-91-513-0216-4. The exceptionally diverse macropodiform’s (kangaroos, rat-kangaroos and their fossil allies) currently have a fossil record that spans from the late Oligocene to the Holocene with an Australasian widespread fossil occurence. The origins of the macropodiforms are believed to have been during the Eocene possibly having split from the Phalangeridae. This is largely based on molecular data as there is a complete lack of macropodiform fossil material prior to the late Oligocene leaving the origins of the macropodiforms to be largely speculative. Thus, late Oligocene rat-kangaroo dental fossil elements associated with Palaeopotorous priscus (which shares characteristics observed in both phalangerid and macropodiforms) were examined to get a better insight into the potential origins of the macropodiforms. The results obtained suggested that P. priscus is currently the most basal macropodiform known. Furthermore, due to the absence of adequate macropodiform post-cranial material, the functional eco-morphological interpretation of various macropodiform fossil taxa has been based largely on cranial and dental characteristics. Consequently, the examination of Miocene Balbaridae kangaroo’s (Balbaroo nalima and an untaxonomised balbarid) post-cranial elements was undertaken, suggesting an array of locomotion types (similar to living macropodiforms) and a likely persistent arboreal and quadrupedal lifestyle. In addition, the Pleistocene Protemnodon anak’s post-cranial material are examined, proposing a probable combination of quadrupedal bounding and a slow walking gait alternative to the eminent fast bipedal saltation seen in almost all extant macropodiforms. Keywords: macropodiforms, kangaroos, rat-kangaroos, fossil ancestors, post-cranial, eco- morphology, systematic, phylogeny Wendy den Boer, Department of Earth Sciences, Palaeobiology, Villav. 16, Uppsala University, SE-75236 Uppsala, Sweden. © Wendy den Boer 2018 ISSN 1651-6214 ISBN 978-91-513-0216-4 urn:nbn:se:uu:diva-339593 (http://urn.kb.se/resolve?urn=urn:nbn:se:uu:diva-339593) For Violetta - Lyza List of Papers This thesis is based on the following papers, which are referred to in the text by their Roman numerals. I Black, K.H., Travouillon, K.J., den Boer, W., Kear, B.P., Cooke, B.N., & Archer, M. (2014) A New Species of the Basal ‘‘Kangaroo’’ Balbaroo and a Re-Evaluation of Stem Macropo- diform Inter relationships. PLoS ONE, 9(11): 1 - 30. II den Boer, W., & Kear, B.P. (in press). Is the fossil rat- kangaroo Palaeopotorous priscus the most basally branching stem Macropodoid? Journal of Vertebrate Paleontology. III den Boer, W., Campione, N., & Kear, B.P. (submitted). Climb- ing adaptations, locomotory disparity and ecological conver- gence in ancient stem ‘kangaroos’. Proceedings of the Royal Society B. IV den Boer, W., & Kear, B.P. (manuscript still in draft format and was prepared for Journal of Vertebrate Paleontology). Comparative Postcranial Osteology of the Plio-Pleistocene ‘Gi- ant Wallaby’ Protemnodon anak, and its Implications for Lo- comotion. V den Boer, W., & Kear, B.P. (manuscript still in draft format and was prepared for Australian Journal of Zoology). The Sys- tematics of Fossil Kangaroos, Wallabies, Rat-Kangaroos, and their Related Stem Clades (Macropodiformes). Reprints were made with permission from the respective publishers. 5 Contents - Introduction – What are the Macropodiformes? ........................................... 9 - The Cenozoic Climate, Environmental and Geographical History of Australia ................................................................................................. 11 - Research Aims ............................................................................................ 16 PAPER I ....................................................................................................... 17 - Post-Cranial Fossil Remains of a Miocene Kangaroo Balbaroo nalima (Balbaridae, Balbaroo) Suggesting Quadrupedal Locomotion Strategies ................................................................................ 17 - Description and Functional Eco-Morphological Interpretation of Balbaroo nalima’s Post-Cranial Material ............................................... 20 - Conclusion .................................................................................................. 26 PAPER II ..................................................................................................... 28 - Phylogenetic and Morphometric Assessment of the Late Oligo- cene Rat-Kangaroo Palaeopotorous priscus ............................................... 28 - Material and Methods ................................................................................. 29 - Results and Conclusion .............................................................................. 30 PAPER III ................................................................................................... 34 - Arboreal Features Revealed in Miocene Balbarid Kangaroo Post-Cranial Fossil Elements ...................................................................... 34 - Description and Functional Interpretation of the Balbarid Fossil Elements ...................................................................................................... 36 - Material and Methods ................................................................................. 37 - Results ........................................................................................................ 40 - Conclusion .................................................................................................. 47 PAPER IV .................................................................................................... 50 - Description and Functional Interpretation of New Cranial and Post-Cranial Fossil Material of the ‘Giant Extinct Wallaby’ Protemnodon anak ...................................................................................... 50 - Material and Methods Linear Morphometric Analysis............................... 51 - Results and Conclusion Linear Morphometric Analysis ............................ 52 - Description of Protemnodon anak’s Cranial, Dentary and Dental Fossil Material ............................................................................................ 52 7 - Comparison of Protemnodon anak’s Cranial, Dentary and Dental Fossil Material ............................................................................................ 58 - Description and Functional Interpretations of Protemnodon anak’s Post-Cranial Fossil Material ....................................................................... 60 - Forelimb Elements ............................................................... 60 - Ribs, Pelves, and Vertebrae Elements.................................. 65 - Hind Limb Elements ............................................................ 69 Conclusion .................................................................................................... 80 - Cranial Material ................................................................... 80 - Post-Cranial Material ........................................................... 80 PAPER V ..................................................................................................... 83 - Systematic Revision of the Fossil Macropodiformes ................................. 83 - Future Objectives ........................................................................................ 88 - Svensk Sammanfattning ............................................................................. 89 - Acknowledgements .................................................................................... 92 - References .................................................................................................. 93 Introduction – What are the Macropodiformes? The Macropodiformes (kangaroos, rat-kangaroos and their fossil relatives) are the most diverse radiation of marsupials on Earth, adopting various different ecological roles throughout Austral- ia, Papua New Guinea, Indonesia, and on some surrounding islands (Burke et al. 1998). The current fossil record of the macropodiforms dates back to the Late Oligocene, as until now no macropodiform fos- sils prior to the Late Oligocene have been uncovered in Australasia (Rich 1996, Godthelp et al. 1999). This leaves the origins of the macropodiform super-family to remain relatively unrequited. Howev- er, based on molecular data, the origins of this mosaic super-family are believed to have occurred
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