The Dietary and Thermoregulatory Role of Blubber As Revealed by Fatty Acids ______Abstract: Blubber Was a Crucial Adaptation for Mammals Living in Water
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The dietary and thermoregulatory role of blubber as revealed by fatty acids Alicia I. Guerrero Vega Evolution and Ecology Research Centre School of Biological, Earth and Environmental Sciences University of New South Wales A thesis in fulfilment of the requirements for the degree of Doctor of Philosophy July 2017 ______________________________________________________________________________________________ THE UNIVERSITY OF NEW SOUTH WALES Thesis/Dissertation Sheet Surname or Family name: Guerrero Vega First name: Alicia Other name/s: Isabel Abbreviation for degree as given in the University calendar: PhD School: School of Biological, Earth and Environmental Sciences Faculty: Faculty of Sciences Title: The dietary and thermoregulatory role of blubber as revealed by fatty acids ______________________________________________________________________________________________ ______________________________________________________________________________________________ Abstract: Blubber was a crucial adaptation for mammals living in water. Blubber serves as an energy reservoir, where surplus energy is deposited in the form of fatty acids (FAs). Most FAs are obtained from an animal’s diet (dietary FAs), thus a predator’s FA signature has the potential to provide dietary information. However, some FAs in the predator may be synthesised or modified intrinsically to fulfil physiological demands (non-dietary FAs), which complicates dietary studies. The aim of this thesis was to understand how FAs signatures relate to the dietary and thermoregulatory roles of blubber. I analysed FAs of leopard and crabeater seals to determine how FAs vary across the blubber depth and how this variability influenced dietary interpretations. I found that blubber is not uniform; where some FAs are abundant in the outer (superficial) layer others are dominant in the inner (deepest) layer. This suggests that the inner layer has a dietary role whereas the outer layer has a more structural role. The FA signatures from the predator’s inner layer resembled more closely those of their prey than the FAs in the outer layer. Trophic predictions were clearer when using only the dietary FAs rather than all FAs; this indicates that there are other factors influencing the metabolism of the non-dietary FAs. To examine if a mammal’s thermoregulatory requirements impact the shifts in FAs, I conducted a meta-analysis including 48 mammals from terrestrial, semi-aquatic, and fully-aquatic environments. I found that the FAs of aquatic mammals are more highly desaturated than those of terrestrial mammals. Higher desaturation helps reduce heat loss and ensures that blubber remains flexible in cold environments. FA desaturation is correlated with latitude and fur density in semi-aquatic mammals. Thus, they increase FA desaturation when living in colder habitats and when they have sparser fur. I compared the FAs of three sympatric Antarctic seals and found that FA desaturation changes as seals grow, which suggests that the thermal efficiency of blubber develops with age. In order to obtain better results in dietary studies, the effect of thermal habitat, fur density and age, on the metabolism of FAs must be considered. ______________________________________________________________________________________________ ______________________________________________________________________________________________ Declaration relating to disposition of project thesis/dissertation I hereby grant to University of New South Wales or its agents the right to archive and to make available my thesis or dissertation in whole or in 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 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 dissertation. I also authorise University Microfilms to use the 350 word abstract of my thesis in Dissertation Abstracts International. ………………………………………….. ………………………………………….. ………………………………………….. Signature Witness Date ______________________________________________________________________________________________ ______________________________________________________________________________________________ FOR OFFICE USE ONLY Date of completion of requirements for Award: ______________________________________________________________________________________________ ii 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 …………………………………………….............. Date : …………………………………………….............. iii 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 ……………………………………………........................... Date ……………………………………………........................... 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 ……………………………………………........................... Date ……………………………………………........................... A mi madre, Teresa, por su amor incondicional y por creer en mí desde el día uno iv Acknowledgements Firstly, I have to thank Tracey for her excellent work as a supervisor. I have only good words to say; thank you for being so supportive during my whole PhD and for believing in me and helping me even before we met. You gave me the opportunity to work in Antarctica, which was a dream come true for me. I will always appreciate it. I would like to thank all members of The Mammal Lab at UNSW, for their help and support, their always constructive feedback on the early versions of my manuscripts, and those occasional chats over lunch. Thank you Kobe, Marlee, Naysa, Gary, Ricardo and Cat. A big thank you to the enthusiastic Team ‘Mamíferos’ from Base Primavera: Sebastián Poljak, Magalí Bobinac, Pedro Carlini, Juan Galliari and Gastón Lo Coco. Thanks for sharing the Antarctic adventure with me, for your generosity and for everything you taught me. A special ‘gracias’ to my dear Chilean gang, especially Romi, Dani and Kimie. The whole process would have not been the same without our coffee get-togethers. Being able to complain about the PhD life with you guys has been a therapy and kept me going during the tough periods. A huge thank you to my big family in Chile. Aunties, uncles, cousins; they are so many that I could not name them here. We are physically separated, but I have felt their love every single day since day one in Australia. Especial thanks to my dear Sol, thanks for your support and for being always there to cheer me up. v Lastly, thanks mom, dad and brother. I could never thank you enough for your love and support. I know it was hard to let me go, especially for you mom, but as always you put me first and supported me no matter what. Thanks for believing in me since day one and for being always there for me. vi Preface Some of the chapters of my thesis have been published, or are in press, elsewhere: Chapter 2 has been previously published as: Guerrero, A.I., Negrete, J., Márquez, M.E., Mennucci, J., Zaman, K., Rogers, T.L., 2016. Vertical fatty acid composition in the blubber of leopard seals and the implications for dietary analysis. J. Exp. Mar. Biol. Ecol. 478, 54-61. (Appendix 1). AIG and TLR conceived the study. JN, MEM and JM collected the samples. KZ conducted the laboratory analyses. AIG and TLR wrote the paper. Chapter 3 has been previously published as: Guerrero, A. I., Rogers, T.L., 2017. Blubber fatty acid composition and stratification in the crabeater seal, Lobodon carcinophaga. J. Exp. Mar. Biol. Ecol. 491: 51-57 (Appendix 2) AIG and TLR conceived the study. AIG collected the samples and conducted laboratory analyses. AIG and TLR wrote the paper. vii General abstract Blubber was a crucial adaptation for mammals living in water. Blubber serves as an energy reservoir,