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Terrestrial and Freshwater Turtles of Early Cretaceous Australia

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Terrestrial and Freshwater Turtles of Early Cretaceous Australia TERRESTRIAL AND FRESHWATER TURTLES OF EARLY CRETACEOUS AUSTRALIA ELIZABETH T. SMITH Research Associate The Australian Museum, Sydney A thesis submitted in fulfillment of the requirements for the degree of Doctor of Philosophy February 2009 Vertebrate Palaeontology Laboratory School of Biological, Earth and Environmental Sciences University of New South Wales, Sydney ABSTRACT An unusual turtle fauna from Lightning Ridge, New South Wales (Albian) reveals that Australian turtles had a more extensive Mesozoic history than previously indicated. Reevaluation of several primitive groups provides novel information on turtle evolution in the southern hemisphere. Seven turtle taxa are identified at Lightning Ridge. Two are Testudines indet. and two indeterminate chelid groups are evinced by isolated elements. Three new taxa are assigned to the new family Spoochelyidae in the superfamily Meiolanoidea. Spoochelys ormondea n. fam., gen. et sp., Sunflashemys bartondracketti n. gen. et sp. and Opalania baagiwayamba n. gen. et sp. are predominantly land-living turtles with high-domed shells and short manus and pes. The sister-group relationship with the Meiolaniidae, supported by a suite of cranial and postcranial synapomorphies, increases the stratigraphic range of the horned turtles by around ~ 50 my. Primitive structures in Spoochelys (postparietal, supratemporal and interpterygoid vacuity), occur with derived features that are variably developed across Triassic and Jurassic turtles. Phylogenetic analysis precariously resolves the meiolanoids as sister group to a clade containing Palaeochersis and Proterochersis. Limited pleurodiran attributes suggest that meiolanoids may be pleurodiromorphs, closer to primitive pleurodires than to cryptodires. As basal ‘side-necked’ turtles, the Lightning Ridge meiolanoids permit first insights into cranial and postcranial progressions in pleurodiran stem taxa. Evidence of diverse meiolanoids in Early Cretaceous Australia and ancient radiations of meiolanoid-like turtles in southern Pangea, suggest that the horned turtles are a Triassic group and that the dichotomy between Pleurodira and Cryptodira occurred well before the Late Triassic. Early Cretaceous chelids at Lightning Ridge occur at higher palaeolatitude than in South America. The temporal range of Australian chelids is extended by ~ 50 my, demonstrating that chelids had a Jurassic history in Australia, with broad diversifications across the polar supercontinent. The palaeoecological setting of Lightning Ridge is comprehensively described for the first time. Diverse invertebrates and vertebrates include terrestrial, freshwater aquatic and rare marine forms that are anomalous at this near-polar palaeolatitude (~65-70oS). The anachronistic occurrence in Early Cretaceous Australia of distinctive radiations of ‘Triassic-type’ turtles, and other relic groups, implies prolonged intervals of biogeographic isolation in the eastern provinces of Pangea. Keywords: Early Cretaceous Australia; Lightning Ridge; Meiolanoidea; Chelidae. ACKNOWLEDGEMENTS My sincere appreciation and thanks to my supervisors Dr Michael Archer and Dr Suzanne Hand, University of New South Wales, for expertise, wisdom and encouragement when it was most needed; Henk Godthelp for his enthusiasm for search and research at Lightning Ridge and for the braincase of cf. Warkalania from Riversleigh; Robin Beck for perseverance and utmost patience (PAUP); Karen Black and Anna Sainsbury for molding and casting the Spook skull and braincase; Dr Thomas Rich, National Museum of Victoria; Lesley and Gerry Kool and Dr Patricia Vickers-Rich, Monash Science Centre, Melbourne; Dr Alex Ritchie and Robert Jones of the Australian Museum, Sydney; Dr Ralph Molnar and Joanne Wilkinson of the Queensland Museum, Brisbane; Dr Ben Kear; Dr Susan Turner; Dr Jeffrey Stilwell, James Bresnahan, Rodney Berrell - all helped in various capacities, and deserve my gratitude. Dr Gene Gaffney offered advice, goodwill, reprints and instruction, sent photographs of the Niolamia type skull and provided access to type material of Chubutemys. Dr P M Datta, Subhash Sen and Asish Kumar Ray at the Geological Survey of India, Kolkata, permitted me to examine, sketch and photograph the type material of Indochelys spatulata at short notice, and graciously. Thanks are due to Walgett Shire Council for supporting the Australian Opal Centre project; and to the Lightning Ridge Bowling Club for helping with fossil acquisitions. I am indebted – the science of palaeontology is indebted - to the opal miners of Lightning Ridge. Palaeontology at Lightning Ridge is a community enterprise, relying heavily on the energetic and generous involvement of the opal miners. Over the past twenty five years, hundreds of opalised fossils have been made available to the Australian Museum, Sydney, and the Australian Opal Centre, Lightning Ridge. Thankyou to Peter and Brett Barton who donated the Sunflash Turtle to the Australian Museum under the Federal Government’s Cultural Gifts Program. Peter and Brett met the Sunflash Turtle underground on the Coocoran and allowed me to collect further material. Dave Roussel, Rod Abels and my daughter Clytie helped to rescue the turtle from the Tyrone’s site. Warmest thanks to many other CGP donors - Bob Sutherland, Neroli and Stephen Bevan, Imo and Louise Stein, Rob and Debbie Brogan, Ken and Marie Lindquist, Dave Roussel and Lalja Petersson, John Cucuk, Stephen Turner, George and Bill Molder, Graeme and Christine Thomson, Clytie Smith, Peter and Vicki Drackett, Joe Walker, Paul Burza, Brian Senior, Dave Sanders, Stewart-Tranter Brown, David and Greg Lane, Colin and Marie Fletcher, Marcel and Sam Miltenburg, Adrian Boot, Dave Barclay, Matthew Goodwin, Mick and Doris Cooke, Jack Fahey and Emilia Katajisto. Praise to the valiant - Ormie Molyneux who loaned the Spook skull and braincase for study over many years, with commendable fortitude and good humour; Ed Long and Henny Kunzelman, Nils and Bea Tape, Bob Cropp, Dave and Alan Galman, Chris Underwood, Ken Hudson, Les Price, Fred Mallouk, Ian Cops, Peter Hall, Dusan, Brownie, Andrew Lindsay, Jo Babic, Virgil, Peter and Lisa Carroll, Andrew Cody, Barefoot Mick Dundee, New Blue from Kellies Two, Doug and Loppy, Martin, Crain Johannson, Slim, Michelle and Gussy Knee, Eric the Viking, Greg Bateup, Hydro Tony, Richard Wagner, James Haverhoek, Peter McKenzie, Geoff Peady, Stan the Man, Greg Johnston, Arthur and Joe Molyneux, Larry White, Richard Slip, Max Caslick, Orange Joe, Silly Pinkies, Smith Boys, Chris Gawthorpe, Graeme Lester, Brian Casement, Fraser Island George, Rod McCracken, Joe Belicka, Laurie Kree, John McCabe, Ken Stephenson, Darren Mack, Bill Kotru, Bob Barrett, Drago Panich, Franz Roehleder, Lennie Cram, Butch McFadden, The Warlords, Darryl Ferguson, Norm, Stardust Clive, Alf White, Col Duff, Vic Morgan, Toby and Matthew Egan, Sali Money, Alan Summers, Mick and Donna Lund, Michael and Rebel Matson, Marilyn Milas, Anne-Marie Deane, Kerry Reid, Murray Gatt, Lizard, June Barker and Rose Fernando. Thankyou one and all. Margie and Warwick Schofield, Tony and Karola, Shovel, Graeme Anderson, Barbara Moritz, Bev Ogle, Roland Beckett, Gwen Jenkins, and the little Scotties and cohorts from next door, contributed in many special ways. Jenni Brammall helped with powerpoints, photographs, layout of figures for chapters four, five and seven, general support and commitment. Thanks also to Tiggen, my Mum and Dad, Rohan, Brinny, David and Penny, for keeping this slow old turtle on track. My husband Robert photographed, photoshopped and held the fort. Gurukangaroo doesn’t believe in turtles but he sacrificed at least two trips to India for this. My eternal gratitude. TABLE OF CONTENTS Page ABSTRACT Acknowledgements Table of contents Abbreviations CHAPTER ONE 1 INTRODUCTION CHAPTER TWO 13 Spoochelys ormondea n. fam., gen. et sp. (Meiolanoidea: Spoochelyidae), an archaic meiolaniid-like land turtle from Lightning Ridge, New South Wales, Australia CHAPTER THREE 55 Sunflashemys bartondrackettii n. gen. et sp. (Meiolanoidea: Spoochelyidae), a primitive swamp turtle from the opal fields of Lightning Ridge, New South Wales, Australia CHAPTER FOUR 90 Opalania baagiiwayamba n. gen. et sp. (Meiolanoidea: Spoochelyidae), a near-polar land turtle from the Early Cretaceous of Lightning Ridge, New South Wales, Australia CHAPTER FIVE 106 New cranial material of a meiolaniid cf. Warkalania from the Miocene of Riversleigh, North Queensland CHAPTER SIX 116 Redescription and reinterpretation of the Liassic turtle Indochelys spatulata Datta et al. 2000 from Maharashtra, India CHAPTER SEVEN 129 Redescription of the Casterton steinkern, Chelycarapookus arcuatus Warren 1969, from the Albian of western Victoria CHAPTER EIGHT 141 Australia’s oldest chelid pleurodires from Lightning Ridge, New South Wales - first evidence of chelids from Mesozoic near-polar Australia CHAPTER NINE 151 Phylogenetic analysis - non-marine turtles of Early Cretaceous Australia, and relationships of the superfamily Meiolanoidea CHAPTER TEN 216 The Lightning Ridge fossil flora and fauna – a diverse high- latitude, warm climate biota from Early Cretaceous Australia CHAPTER ELEVEN 294 Palaeoecology of terrestrial and freshwater turtles of Lightning Ridge, New South Wales CHAPTER TWELVE 301 Biogeography of non-marine turtles of Early Cretaceous Australia CHAPTER THIRTEEN 308 Possible Triassic origin for the horned turtles (Testudines: Meiolanoidea) and implications for turtle relationships CHAPTER FOURTEEN 312 CONCLUSIONS REFERENCES
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