Evolutionary Adaptations to Climate Change in Australian Flora
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Faculty of Science and Engineering Department of Environment and Agriculture Evolutionary Adaptations to Climate Change in Australian Flora Haylee Marie D’Agui This thesis is presented for the degree of Doctor of Philosophy of Curtin University March 2017 Declaration To the best of my knowledge and belief this thesis contains no material previously published by any other person except where due acknowledgement has been made. This thesis contains no material that has been accepted for the award of any other degree or diploma in any university. Signature………………………….. Date……………21/03/17……... ii Abstract Climate changes have been impacting ecosystems worldwide, causing shifts in the habitats suitable for plant species and threatening their continuing persistence. Mediterranean-type Ecosystems contain extraordinarily high levels of plant diversity and endemism, yet are vulnerable to climate and environmental changes. Southwest Western Australia is recognised as a global biodiversity hotspot, though has been experiencing decreased rainfall and increased drought over the past 40 years, causing great concerns for the long-term survival of its extraordinary plant diversity. Plants may respond to changes in climate by migrating to new habitats, persisting in their current habitat, or by adapting to changes through evolution. In this thesis, I investigated the potential of Southwest Western Australian plants to rapidly evolve to tolerate changes in climate, particularly decreased rainfall and increased temperature. I first assessed the floristic composition of a nature reserve in speciose kwongan heathland following four decades of decreased rainfall and increased temperature through comparison of a present day vegetation survey to historic surveys. In Chapter 3, I further investigated the potential of seed banks to mitigate the effects of decreased rainfall through comparison of the growth of seedlings descended from parents established in years of either average or below-average rainfall when subjected to drought and control treatments. In Chapter 4, I explored the potential of Banksia hookeriana (Proteaceae) to accumulate adaptive genetic variation in order to resist the effects of a drying climate through the assessment of the fitness of seedlings established from seed produced pre- and post- drought to determine whether post-drought seed displayed increased drought tolerance. Finally, in Chapter 5, I investigated whether adaptive genetic diversity in B. attenuata was affected by rainfall and temperature gradients, or altered fire regime through genomic scanning and environmental association analysis on plants distributed across its range to determine candidate genes associated with these environmental variables. Overall, results from my study suggest that the flora of Southwest Western Australia has been able to tolerate the changes in climate that have occurred so far, and that there may be potential for tolerance of further changes in climate. However changes in fire regime in the region may have a detrimental effect on the ability of some species to adapt and survive. iii Acknowledgements The completion of this thesis would not have been possible without the valuable contributions of the following people, to whom I wish to express my sincere thanks. I would first like to express my gratitude to my supervisors Dr Tianhua He, Professor Neal Enright, and Dr Richard Harris, for their ongoing support, advice, and patience throughout the past four years of my PhD, and without whose expertise, encouragement, and assistance this work would not have been possible. Thanks also to Dr Sim Lin Lim for his valuable advice and assistance with Banksia DNA sequencing and analyses. Thanks to the staff of the Department of Environment and Agriculture at Curtin University, for their assistance and provision of facilities. Special thanks to John Jackson, Lydia Kupsky, Liz Crompton, William Parkinson and Peta Tilbrook for their invaluable technical support. I also wish to thank the Australian Research Council for support for this project in the form of funding awarded to Tianhua He and Neal Enright (ARC DP13013029). Thanks to the current and past members of the Plant Ecology and Evolution Group at Curtin University; Dr Tianhua He, Dr Sim Lin Lim, William Fowler, Dr Sh-Hoob Mohamed El-Ahmir, Tran Duc Thien, Emmaline Yearsley, Casey Causley and Ha Huy Hoang for their friendship, support and assistance. Thanks to William Fowler, Liam Mallon and Alexa Tunmer for assistance with field and laboratory work. Thanks to my fellow PhD students, for their support, encouragement and friendship over the past four years. Special thanks to Dilanka Mihindukulasooriya, Ashleigh Wolfe and Araz Abdullah, for being there for me throughout all the ups and downs of the past year. Finally, I would like to express my sincere thanks to my family and friends, for their patience, support, and belief in me. Special thanks to my dear friend Jessica Eglinton, for her constant encouragement and support throughout this PhD and in life in general. Last, but not least, I would like to express my profound appreciation to my husband Liam Mallon, for his unwavering support and encouragement not only throughout this PhD, but in everything I do in life. iv Table of Contents Declaration ....................................................................................................................................................... ii Abstract ............................................................................................................................................................ iii Acknowledgements ....................................................................................................................................iv Table of Contents .......................................................................................................................................... v List of Figures .................................................................................................................................................. x List of Tables .................................................................................................................................................xv List of Abbreviations .............................................................................................................................xviii Chapter 1: An introduction to the kwongan vegetation of Southwest Australia and potential threats from climate change. ............................................................................................ 1 1.1 Climate change and plant response ............................................................................................. 1 1.1.1 Climate change over the past half century ......................................................................... 1 1.1.2 Plant responses to climate change ........................................................................................ 1 1.1.3 Evolutionary adaptations as a result of climate change ............................................... 2 1.2 Vegetation, ecology and adaptations in the Southwest Australian Floristic Region, a global biodiversity hotspot ................................................................................................................. 4 1.2.1 The Southwest Australian Floristic Region ........................................................................ 4 1.2.2 Mediterranean Type Ecosystems and climate change .................................................... 4 1.2.3 Southwest Australia and climate change ............................................................................ 5 1.2.4 Future species distributions – current projections and caveats ................................. 6 1.2.5 Fire and climate as drivers of ecological adaptation in SWA ...................................... 7 1.3 Eneabba Sandplain: Kwongan vegetation and current climate change .................... 10 1.3.1 Kwongan vegetation of the Eneabba sandplain ............................................................ 10 1.3.2 Eneabba study site and climate ........................................................................................... 12 1.4 Objectives and structure of this thesis .................................................................................... 14 1.5 References ........................................................................................................................................... 17 Chapter 2: Effect of decreasing rainfall on floristic composition of South Eneabba Nature Reserve, Eneabba, Western Australia. ........................................................................... 24 v 2.1 Abstract ................................................................................................................................................ 24 2.2 Introduction ....................................................................................................................................... 25 2.2.1 Climate change and the flora of Southwest Australia ................................................. 26 2.2.2 Value of historic data in climate change studies ........................................................... 27 2.3 Methods ................................................................................................................................................ 28 2.3.1 Study system...............................................................................................................................