A Study on Tenuibranchiurus and the Evolution of the Burrowing Clade of Australian Freshwater Crayfish
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A Study on Tenuibranchiurus and the Evolution of the Burrowing Clade of Australian Freshwater Crayfish Author Dawkins, Kathryn L Published 2015 Thesis Type Thesis (PhD Doctorate) School Griffith School of Environment DOI https://doi.org/10.25904/1912/1932 Copyright Statement The author owns the copyright in this thesis, unless stated otherwise. Downloaded from http://hdl.handle.net/10072/367987 Griffith Research Online https://research-repository.griffith.edu.au A study on Tenuibranchiurus and the evolution of the burrowing clade of Australian freshwater crayfish PhD Thesis Submitted in fulfilment of the requirements of the degree of Doctor of Philosophy February 2015 Kathryn L. Dawkins B.Sc. (Hons) School of Environment Griffith University Supervisors Professor Jane Hughes (GU) Professor Clyde Wild (GU) Dr. James Furse (GU) Abstract ABSTRACT The overall aim of this thesis was to investigate the morphological and molecular diversity within Tenuibranchiurus, and to utilise these data to further the understanding of the evolution of this freshwater crayfish and the other genera of the Australian burrowing clade from both a phylogenetic and biogeographic perspective. The genus Tenuibranchiurus occurs within coastal eastern Australia and currently represents the largest gap in knowledge within this clade of crayfish. It is also the only monotypic parastacid genus, containing the single species T. glypticus. Additionally, it has morphological, phylogenetic, and geographical attributes that are not exhibited by other members of the burrowing clade, or by other freshwater fauna found throughout its distributional range. Examination and clarification of these unique features has the potential to elucidate evolutionary processes determining the distribution and genetic structure of the genus, the burrowing clade, and the freshwater fauna of coastal eastern Australia. Examination and analysis of the morphological and molecular characteristics of Tenuibranchiurus identified the presence of two distinct genera, with the populations from Queensland representing Tenuibranchiurus and those from New South Wales representing a proposed new genus, Gen. nov.. The analyses also supported the recognition of six species within Tenuibranchiurus (including the previously recognised T. glypticus), and two within Gen. nov.. All species within both genera exhibited highly restricted and/or highly disjunct distributions. The molecular data suggest that species within both genera are highly structured spatially; this is likely the result of cyclical population retractions and expansions, and shifting distributions in response to changes in sea level and increased aridity altering available and accessible habitat. The present day distribution of Tenuibranchiurus and Gen. nov. species has been strongly influenced by historical climate events. Dispersal most likely occurred during periods of lowered sea level which would have exposed swampy coastal plains, creating a dispersal pathway for populations to use as they followed a moisture gradient as the sea receded. Genetic divergence, and subsequent speciation, likely occurred through vicariance as a result of population isolation, habitat heterogeneity, and/or subsequent sea level oscillations. These processes were estimated to have occurred over ~50 million years, commencing with the i Abstract splitting of an ancestral genus into Tenuibranchiurus and Gen. nov., and subsequent divergence resulting in the eight species present today. With the molecular diversity of Tenuibranchiurus and Gen. nov. clarified, the phylogenetic relationships between all of the burrowing clade genera were re-examined. However, the relationship between the three oldest genera (Engaewa, Engaeus sensu stricto, and E. lyelli) could not be resolved, probably as a result of almost simultaneous divergence. These genera were estimated to have diverged during the Cretaceous, followed by Gramastacus (also during the Cretaceous), Geocharax (Palaeocene), and Tenuibranchiurus and Gen. nov. (Eocene). Investigation of the biogeographic history of the burrowing crayfish suggests that the genera (and species within them) display highly concordant distributional patterns, appearing to be influenced largely by climate change and fluctuating sea level rather than by drainage architecture, which is the most commonly accepted freshwater paradigm. Therefore, the burrowing clade crayfish, and potentially other parastacid genera, can be viewed as ecologically removed from other freshwater taxa, requiring detailed historical climate and geological data in combination with dispersal models in order to reconstruct their biogeographic histories. ii Statement of Originality STATEMENT OF ORIGINALITY This work has not previously been submitted for a degree or diploma in any university. 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 in the thesis itself. (Signed) _______________________ Kathryn Dawkins iii Acknowledgements ACKNOWLEDGEMENTS To my principle supervisor Professor Jane Hughes, thank you for undertaking this project with me and giving me the freedom to mould and re-mould it how I chose over the many years. Thank you also for your support and advice. To my associate supervisors Professor Clyde Wild and Dr. James Furse, thank you for your proof-reading and helpful suggestions. Special thanks are given to Andrew Bentley and Quinton Burnham for providing additional sample material for my genetic analyses, and to Stephen Keable for helping me gain access to the Australian Museum collections. This research was funded by the Griffith School of Environment, and made possible with an Australian Postgraduate Award. Additional funding was supplied by the Australian Rivers Institute and Griffith Graduate Research School for conference attendance to present my work. All specimens from this study were collected under permits WITK08599510, WISP08599610, and TWB/01/2011 issued by the Department of Environment and Resource Management. I would also like to thank the plethora of people who have given me assistance whenever I ran into one of the many pitfalls that seem to surround molecular analyses on crayfish. Specifically, to Dan Schmidt, Bob (aka Andrew Bentley), Jesse Breinholt, Alicia Toon, Kathryn Real, and Jemma Somerville for answering countless emails on genetic techniques and lab questions, and allowing me to hassle you any time for explanations. A huge thank you also is warranted for Michael Arthur who never complained when I would turn up unannounced at his office door with a stack of statistical analyses to discuss. This project would never have been possible without my field lackeys; you were both my helpers and my sanity-keepers, so thank you to James Furse, Seanan Wild, Amanda Dawson, Diana Virkki, and Shane Howard for the many hours (and prescription sunglasses, Seanan) you sacrificed for me. I am eternally grateful for the support I received from my friends during this journey. To DD and Bones, you were my coffee support team and my lunch buddies, providing invaluable meaningless chit-chat to break up the hard slog that is a PhD. You were also (and still are) close friends and helped me through a lot. Manana and Seaman, you followed in our footsteps and decided to join a team of crazy people doing their PhD’s – for reasons I still do iv Acknowledgements not understand. You were also there as close friends and staunch supporters, and I cannot thank you enough for being there through this entire journey. You will always be my Joben and favourite woo-girl. I am also grateful to Jason Coughran for being my office buddy and friend when I moved to Perth for the last year and a half of my PhD. I always enjoyed talking about how time might not even be real, to then look up and realise we had just wasted an hour of it. Last, but not least, are my family. Mum and Dad, you supported me throughout my degree; from undergraduate, to Honours, through to a PhD that seemed like it would never end. Although the times I had to camp out at your house were because I had a lot of hard lab work ahead of me, the little things like packed lunches and treaties made it so much easier for me. Thank you also to Anne and Rohan who did their best to understand what I was doing and why I was doing it – I know that you will encourage Kate and Sarah to follow the path of science, no matter what you may say now. To my other set of parents, Kay and Geoff, thank you for putting up with both Quinton and I as we both struggled through the last steps of our PhD journey. It could not have been easy dealing with the daily mood-swings that are part and parcel of that, so I am grateful to you for sticking with us. Three of the most important supporters in this journey were in fact my puppies; Molly, Nikki, and Rowdy. You provided sympathetic ears when I needed a chat, and endless hours of fun and laughter – something not to be dismissed lightly during such a long journey. Finally, and my biggest thank you yet, is for my Q. We came into each other’s lives at perhaps the most difficult points – I don’t know any other couple that were crazy enough to try and finish two PhD’s at the same time, and I suspect there is a reason for that. You have been my biggest supporter, taking on the tasks of keeping me happy, sane, fed, and relaxed as if it were easy. You survived it through the many emotional ups and downs and the tears that came with both of these. No one will ever truly understand