CRYPTIC DIVERSITY OF A GLOSSOPTERIS FOREST: THE PERMIAN PRINCE CHARLES MOUNTAINS FLORAS, ANTARCTICA by Ben James Slater A thesis submitted to the University of Birmingham for the degree of DOCTOR OF PHILOSOPHY School of Geography, Earth and Environmental Sciences College of Life and Environmental Sciences University of Birmingham September 2013 University of Birmingham Research Archive e-theses repository This unpublished thesis/dissertation is copyright of the author and/or third parties. The intellectual property rights of the author or third parties in respect of this work are as defined by The Copyright Designs and Patents Act 1988 or as modified by any successor legislation. Any use made of information contained in this thesis/dissertation must be in accordance with that legislation and must be properly acknowledged. Further distribution or reproduction in any format is prohibited without the permission of the copyright holder. ABSTRACT The Toploje Member chert is a Roadian to Wordian autochthonous– parautochthonous silicified peat preserved within the Lambert Graben, East Antarctica. It preserves a remarkable sample of terrestrial life from high-latitude central Gondwana prior to the Capitanian mass extinction event from both mega- and microfossil evidence that includes cryptic components rarely seen in other fossil assemblages. The peat layer is dominated by glossopterid and cordaitalean gymnosperms and contains sparse herbaceous lycophytes, together with a broad array of dispersed organs of ferns and other gymnosperms. The peat also hosts a wide range of fungal morphotypes, Peronosporomycetes, rare arthropod remains and a diverse coprolite assemblage. The fungal and invertebrate-plant interactions associated with various organs of the Glossopteris plant reveal the cryptic presence of a ‘component community’ of invertebrate herbivores and fungal saprotrophs centred around the Glossopteris organism, and demonstrate that a multitude of ecological interactions were well developed by the Middle Permian in high-latitude forest mires. Comparisons of coal maceral data from co-occurring coal seams with quantitative analyses of the silicified peat constituents reveals that while silicified peats provide an unparalleled sample of the organisms forming these coal deposits, they do not necessarily reflect the constituents that ultimately dominate the coal maceral volume. ACKNOWLEDGEMENTS I am deeply grateful to: Jason Hilton, my PhD supervisor for his years of support, encouragement, scientific guidance and the many invaluable discussions about the project in Staff House. Stephen McLoughlin, my PhD supervisor, for the years of guidance throughout every turn of the project, intellectual input, advice, great hospitality on visits to Stockholm and for the opportunity to work on such interesting material. All the staff, post-grads and students in Earth Sciences over the years who have made the whole post graduate experience so enjoyable. Special mention to the residents of the various post-grad offices over the last few years for all the welcome distractions, and to the former office 116 inhabitants whose many exploits have made the whole experience so memorable. Guy Harrington, Stephen Jones and Warren Eastwood for their constructive comments and ideas during panel meetings. All the staff in the Palaeobotany department at the Swedish Museum of Natural History who were a great help on visits to the museum. I am eternally grateful to my family for all their support and encouragement at every step of my studies. Thanks to Kristina for all her support throughout my final PhD year. Also thanks to Stella the dog for her relentless enthusiasm to play with a tennis-ball every waking hour which was a great distraction from thesis writing. On the funding and logistical side of things my thanks go to the School of Geography, Earth and Environmental Sciences and to the Natural Environment Research Council (NE/H5250381/1) for funding the project. Thanks also to the Synthesys programme of the EU for providing funding for a research visit to the NRM in Stockholm (SE-TAF-4827). Thanks also to Keiraville Konsultants Pty Ltd., Wollongong, Australia, who provided data on coal maceral proportions. Many thanks to Pollyanna von Knorring for the brilliant landscape illustration in Figure 6.9. Thanks to Else Marie Friis and Anna Lindström (NRM), Marco Stampanoni and Frederica Marone (Paul Sherrer Institute) for aid in the synchrotron tomography work (supported by the European Union FP6, project number 20100167, to P.C.J. Donoghue, S. Bengston and E.M. Friis and the European Community – Research Infrastructure Action under the FP7 ‘Capacities’ Specific Programme). The Australian Antarctic Division provided financial and logistical support for collection of the specimens via Antarctic Science Advisory Council Project 509. Thanks to Stephen McLoughlin, David Cantrill and Andrew Drinnan for making this research possible by collecting the material on two Antarctic expeditions in the early 1990s. CONTENTS Chapter 1: Introduction ............................................................................... Page 1 1.1 Terrestrial ecosystems of Permian Gondwana ................................................... 1 1.1.1 Gondwana and the Glossopteris flora ................................................. 1 1.1.2 Preservation of Gondwanan Permian terrestrial ecosystems .............. 2 1.1.3 Contents of the peats ........................................................................... 7 1.2 Aims of thesis .................................................................................................. 10 1.3 Structure of thesis ............................................................................................ 10 Chapter 2: Geological setting .............................................................................. 13 2.1 Palaeogeography .............................................................................................. 13 2.2 Lithostratigraphy and the Toploje Member chert ............................................ 16 2.3 Preservation ...................................................................................................... 18 Chapter 3: Guadalupian (Middle Permian) megaspores and sporangia from the Toploje Member chert, Bainmedart Coal Measures, Prince Charles Mountains, Antarctica .............. 20 3.1 Background ...................................................................................................... 20 3.2 Materials and methods ..................................................................................... 22 3.3 Systematic palaeobotany .................................................................................. 23 3.3.1 Duosporites lambertensis .................................................................. 23 3.3.2 Banksisporites antarcticus ................................................................ 27 3.3.3 Singhisporites hystrix ........................................................................ 31 3.3.4 Singhisporites mahanadiensis ........................................................... 46 3.3.5 Isolated fern sporangia ...................................................................... 46 3.4 Discussion ........................................................................................................ 53 3.4.1 Reproductive biology ........................................................................ 54 3.4.2 Fern sporangia ................................................................................... 55 3.4.3 Taphonomy ....................................................................................... 59 3.4.4 Palaeobiogeography .......................................................................... 62 3.5 Summary of findings ........................................................................................ 62 Chapter 4: Animal-plant interactions in a Middle Permian permineralised peat of the Bainmedart Coal Measures, Prince Charles Mountains, Antarctica .............................................................. 64 4.1 Background ...................................................................................................... 64 4.2 Materials and methods ..................................................................................... 68 4.3 Results .............................................................................................................. 69 4.3.1 Coprolites in Vertebraria and Australoxylon .................................... 70 4.3.2 Coprolites in leaves ........................................................................... 72 4.3.3 Coprolite within sporangium ............................................................ 73 4.3.4 Isolated large coprolites .................................................................... 74 4.3.5 Isolated small coprolites ................................................................... 75 4.3.6 Fungi-rich coprolites ......................................................................... 76 4.3.7 Coprolites containing pollen ............................................................. 77 4.3.8 Coprolites with coarse constituents .................................................. 78 4.3.9 Isolated spiral-ornamented coprolite ................................................. 79 4.3.10 Other evidence of animals .............................................................. 79 4.4 Discussion .......................................................................................................