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Jack Oyston Thesis Final Bind University of Bath PHD Comparative Insights Into Convergent Evolution Oyston, Jack Award date: 2018 Awarding institution: University of Bath Link to publication Alternative formats If you require this document in an alternative format, please contact: [email protected] General rights Copyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. • Users may download and print one copy of any publication from the public portal for the purpose of private study or research. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying the publication in the public portal ? Take down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Download date: 03. Oct. 2021 University of Bath PHD Comparative Insights Into Convergent Evolution Oyston, Jack Award date: 2018 Awarding institution: University of Bath Link to publication General rights Copyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. • Users may download and print one copy of any publication from the public portal for the purpose of private study or research. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying the publication in the public portal ? Take down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Download date: 15. May. 2019 Comparative Insights Into Convergent Evolution Jack William Oyston A thesis submitted for the degree of Doctor of Philosophy University of Bath Department of Biology and Biochemistry February 2018 COPYRIGHT Attention is drawn to the fact that copyright of this thesis rests with the author. A copy of this thesis has been supplied on condition that anyone who consults it is understood to recognise that its copyright rests with the author and that they must not copy it or use material from it except as permitted by law or with the consent of the author This thesis may be made available for consultation within the University Library and may be photocopied or lent to other libraries for the purposes of consultation with effect from......................... Signed on behalf of the Faculty of Science………………………… 1 Summary Evolution has traditionally been seen as an open-ended, adirectional process. However, the ubiquity of convergent evolution instead suggests that a hierarchy of physical and biological constraints shape evolution. This thesis examines the empirical evidence for convergent evolution’s impact on macroevolutionary patterns. One corollary of convergent evolution is the tendency for animal groups to reach maximum morphological disparity early in their evolutionary histories. An analysis of plants confirms that bottom heavy disparity profiles (Centre of Gravity < 0.5) are not unique to animals. This pattern is most easily explained by character exhaustion, with repetition of character states becoming increasingly probable as evolutionary time increases. However, in a sample of 93 extinct clades, no correlation between character exhaustion and disparity profile shape was found. Instead, ecological or genetic constraints likely limit organism form. Convergent evolution can introduce noise to morphological phylogenies. Anecodotal evidence in mammals shows molecular phylogenies are more congruent with biogeography than their morphological forbears, suggesting ecological constraint could be driving morphological convergence which confounds phylogeny. Similar patterns are common in other clades. In a systematic study of 48 plant and animal clades, the significant majority (70%) of molecular trees were more consilient with biogeographical distributions than their morphological counterparts. Genetic constraint might also limit evolutionary possibility and drive convergent evolution. If so, genome duplications, by introducing genetic redundancy, are expected to be associated with evolutionary novelty and diversification. An analysis of 356 sister clade pairs shows polyploids contain significantly more species than their non-polyploid counterparts. Whilst a direct link between morphological and genetic constraints has yet to be identified genetic constraints are likely to play an important role in the diversification of species. Finally, the importance of these findings for the study of convergent evolution and evolutionary processes in general is discussed, along with future lines of research. 2 Acknowledgements There are many people I’m very much indebted to for being able to make it this far. Firstly, I’d like to thank my primary supervisor, Prof. Matthew Wills, for his unwavering support during this project. During difficult times he readily gave both his time and considerable expertise, helping to make the past four years enjoyable and memorable. I would also like to thank my secondary supervisor Dr. Mark Wilkinson for his unique and valuable insights into evolutionary biology and many engaging conversations. I’d like to thank the John Templeton Foundation for their generous funding allowing me to pursue this research and reach this point. I would also like to thank all the members, past and present, of the Biodiversity Lab at the University of Bath. Drs. Tim Astrop, Katie Davies, Kate Ashbrook and James Sciberras all played an instrumental role in welcoming me to the department and helping me settle into my new academic life, as well as being good friends to this day. I would like to thank Polly Russel and Catherine Klein especially for their friendship and for making the dull days of data-crunching more bearable. I’d also like to thank the academic staff of the Department of Biology and Biochemistry. Special thanks go to Dr. Nick Longrich, Dr. Daniel Field, Prof. Tamas Szekely and Dr. Alex Jeffries for providing particularly useful discussions and ideas. My collaborators at other institutions, Dr. Pete Wagner, Dr. Martin Hughes and Dr. Sylvain Gerber all contributed to my publications and offered their support and advice during my PhD. Lastly, I would like to thank my friends and family who have been so helpful and understanding during these hectic past years. I would not have been able to make it as far as I have without their support and love. 3 Contents 1 Introduction ............................................................................................................ 11 1.1 Understanding Evolutionary Pattern & Process ........................................... 11 1.1.1 Similarities, Differences & Darwinism ............................................................. 11 1.1.2 The Neo-Darwinian Framework ......................................................................... 12 1.1.3 Evolution & Genetics ........................................................................................... 13 1.1.4 Macroevolutionary Patterns, Predictability & Gould’s ‘Tape of Life’ ...... 15 1.1.5 Summary ................................................................................................................. 17 1.2. Convergent Evolution .................................................................................... 17 1.2.1 Defining & Characterising Convergent Evolution ........................................ 17 1.2.1.1 General Agreement ....................................................................................... 18 1.2.1.2 Pattern & Process Based Definitions ....................................................... 18 1.2.1.3 Parallelism & Convergence ........................................................................ 19 1.2.2 Types Of Convergence ........................................................................................ 19 1.2.2.1 Functional Convergence ............................................................................. 20 1.2.2.2 Structural Convergence ............................................................................... 21 1.2.2.3 Mechanistic Convergence ........................................................................... 22 1.2.3 Convergent Adaptive Radiations...................................................................... 23 1.2.3.1 Island Scale ..................................................................................................... 24 1.2.3.2 Continent Scale .............................................................................................. 24 1.2.4 Convergence At Different Organisational Levels ........................................ 25 1.2.5 Summary ................................................................................................................. 26 1.3. Quantifying Convergence .............................................................................. 26 1.3.1 Quantifying Degree Of Similarity & Difference ............................................. 26 1.3.2 Frequency Based Measures .............................................................................
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