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Life History Changes with the Colonisation of Land by Fish

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Life history changes with the colonisation of land by fish Edward Richard Murray Platt Supervised by: Terry Ord THESIS SUBMITTED FOR THE DEGREE OF MASTER OF PHILOSOPHY Evolution and Ecology Research Centre School of Biological, Earth and Environmental Sciences Faculty of Science University of New South Wales March 2014 THE UNIVERSITY OF NEW SOUTH WALES Thesis/Dissertation Sheet Surname or Family name: Platt First name: Edward Other name/s: Richard Murray Abbreviation for degree as grven an the University calendar: MPhil School: Biological, Earth and Environmental Sciences F acuity: Science Title:Mr My thesis addressed two questions: whether survival was inferred to have improved for fish that moved onto land, and what the relative role of predation and density were for detenmining life history variation among populations within one of these land species. For the first question I used life history theory to examine whether survival was inferred to have improved in two fish families which have independently made the transition onto land: Gobiidae and Blenniidae. I examined growth and various aspects of reproductive investment among terrestrial and aquatic species, finding that differences varied according to the level of independence from water. This was consistent with improved survival for certain age classes on land. Nevertheless, the details of life history change differed in each family, with the greatest increases in survival implied for early age classes in Blenniidae, but older age classes in Gobiidae. This suggests fundamental differences in the way the colonization of land occurred in each family. For the second question I investigated the consequences of predataon and density on life hastory variation among frve populations of the Pacific leaping blenny A/ficus amoldorum. This frsh lives out of the water on rocks in the splash zone of Guam, where it is vulnerable to predation and likely sensitive to changes in population density which impact resource availability. I found the extent to which populations invested in reproductaon was inversely related to predation rate, while growth rate appeared to vary largely in response to population density. The differences in fife history among populations were also likely to be plastic an the Pacific leaping blenny. My results revealed the interaction between predation and density can lead to complex outcomes in life history, and inferred that plastic life history traits could allow populations to persist in new environments. For example, plasticity in life history may have been an important facilitator for these fish to make the transition onto land in the first place. As a whole, my thesis demonstrates how life history theory, by identifying differences in survival between species and populations respectively, provides a window into the ecological conditions impacting an organism. Declaration relating to disposition of project thesis/dissertation 1 hereby grant to the University of New South Wales or its agents the right to archive and to make available my thesis or dissertation in whole or m part in the University libraries in all fonms of media, now or here after known, subject to the provisions of the Copyright Act 1968. I retain all property rights, such as patent rights. I also retain the right to use in future works (such as articles or books) all or part of this thesis or dissertataon. 1 also authorise University Microfilms to use the 350 word abstract of my thesis in Dissertation Abstracts International (this is applicable to doctoral theses only). 3??/;."f/~ .? ..1.?. ........ Date The University recognises that there may be exceptional circumstances requiring restrictions on copying or conditions on use. Requests for restriction for a period of up to 2 years must be made in writing. Requests for a longer period of restriction may be considered in exceptaonal cirrumstances and reauire the approval of the Dean of Graduate Research. FOR OFFICE USE ONLY Date of completion of requirements for Award: iii ORIGINALITY STATEMENT 'I hereby declare that this submission is my own work and to the best of my knowledge it contains no materials previously published or written by another person, or substantial proportions of material which have been accepted for the award of any other degree or diploma at UNSW or any other educational institution, except where due acknowledgement is made in the thesis. Any contribution made to the research by others, with whom I have worked at UNSW or elsewhere, is explicitly acknowledged in the thesis. I also declare that the intellectual content of this thesis is the product of my own work, except to the extent that assistance from others in the project's design and conception or in style, presentation and linguistic expression is acknowledged.' Signed . h:.". 'fl . ,. ~ ... ................ Date ....3~;ftolS ...... .... ..................... i COPYRIGHT STATEMENT 'I hereby grant the University of New South Wales or its agents the right to archive and to make available my thesis or dissertation in whole or part in the University libraries in all forms of media, now or here after known, subject to the provisions of the Copyright Act 1968. I retain all proprietary rights, such as patent rights. I also retain the right to use in future works (such as articles or books) all or part of this thesis or dissertation. I also authorise University Microfilms to use the 350 word abstract of my thesis in Dissertation Abstract International (this is applicable to doctoral theses only). I have either used no substantial portions of copyright material in my thesis or I have obtained permission to use copyright material; where permission has not been granted I have applied/will apply for a partial restriction of the digital copy of my thesis or dissertation.' signed ..... i'~ .. j?J!? .. ?.. .. .......................... Date ... ... 3;1~<?15 ....................................... AUTHENTICITY STATEMENT 'I certify that the Library deposit digital copy is a direct equivalent of the final officially approved version of my thesis. No emendation of content has occurred and if there are any minor variations in formatting, they are the result of the conversion to digital format.' Signed .. d.:,zt; ..~ .......................... 01 Date ... 3flj?:: 5 ........................................ ii Abstract My thesis addressed two questions: whether survival was inferred to have improved for fish that moved onto land, and what the relative role of predation and density were for determining life history variation among populations within one of these land species. For the first question I used life history theory to examine whether survival was inferred to have improved in two fish families which have independently made the transition onto land: Gobiidae and Blenniidae. I examined growth and various aspects of reproductive investment among terrestrial and aquatic species, finding that differences varied according to the level of independence from water. This was consistent with improved survival for certain age classes on land. Nevertheless, the details of life history change differed in each family, with the greatest increases in survival implied for early age classes in Blenniidae, but older age classes in Gobiidae. This suggests fundamental differences in the way the colonization of land occurred in each family. For the second question I investigated the consequences of predation and density on life history variation among five populations of the Pacific leaping blenny Alticus arnoldorum. This fish lives out of the water on rocks in the splash zone of Guam, where it is vulnerable to predation and likely sensitive to changes in population density which impact resource availability. I found the extent to which populations invested in reproduction was inversely related to predation rate, while growth rate appeared to vary largely in response to population density. The differences in life history among populations were also likely to be plastic in the Pacific leaping blenny. My results revealed the interaction between predation and density can lead to complex outcomes in life history, and inferred that plastic life history traits could allow populations to persist in new environments. For example, plasticity in life history may have been an important facilitator for these fish to make the transition onto land in the first place. As a whole, my thesis demonstrates how life history theory, by identifying iv differences in survival between species and populations respectively, provides a window into the ecological conditions impacting an organism. v Table of Contents Page CHAPTER 1: Introduction 1 CHAPTER 2: Predictable life history changes reveal enhanced 19 survival for marine fish living on land ABSTRACT 20 INTRODUCTION 21 MATERIALS AND METHODS 29 RESULTS 39 DISCUSSION 53 LITERATURE CITED 59 CHAPTER 3: Population variation in the life history of a land fish 68 reveals the interacting impact of predation and conspecific density ABSTRACT 69 INTRODUCTION 70 MATERIALS AND METHODS 78 RESULTS 85 DISCUSSION 94 vi LITERATURE CITED 97 CHAPTER 4: Conclusion 105 APPENDIX 110 CHAPTER 2 110 CHAPTER 3 129 ACKNOWLEDGEMENTS 131 vii LIST OF TABLES MAIN TEXT Page CHAPTER 2 Table 1: Life history predictions for different changes in survival. 25 Table 2: Parameter values of various goby life history traits in 43 response to moving onto land. Table 3: Parameter values of various blenny life history traits in 48 response to moving onto land. CHAPTER 3 Table 1: AIC table of predictors of population
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