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研究生: 戴芮琪
Rikke Beckmann Dahl
論文名稱: 鯊魚皮齒的演化與功能形態
Evolution and Functional Morphology of Shark Dermal Denticles
指導教授: 沈聖峰
Shen, Sheng-Feng
陳柏宇
Chen, Po-Yu
口試委員: Peter Rask Moller
Peter Rask Moller
黃仁磐
Huang, Jen-Pan
紀凱容
Chi, Kai-Jung
沈聖峰
Shen, Sheng-Feng
陳柏宇
Chen, Po-Yu
口試日期: 2022/11/29
學位類別: 博士
Doctor
系所名稱: 生命科學系
Department of Life Science
論文出版年: 2023
畢業學年度: 111
語文別: 英文
論文頁數: 92
英文關鍵詞: dermal denticles, shark, evolution, functional morphology
研究方法: 實驗設計法
DOI URL: http://doi.org/10.6345/NTNU202300371
論文種類: 學術論文
相關次數: 點閱:74下載:1
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  • In this Ph.D. study, I examine the morphology and function of dermal denticles, both as single entities and as an interacting armour across sharks. Further, I explore the tempo and mode of denticle disparity on a macroevolutionary scale.
    In chapter 2, I use deep learning to objectively quantify denticle disparity and establish ten distinct morphogroups on which all my experiments are based. I further devised a protocol for studying the functional properties of the morphogroups, based on nanoindentation, electron probe micro-analysis, and computational fluid dynamics. The experiments yielded functional profiles relating mechanical properties, elemental composition, and swimming performance. I found seven of the ten morphogroups to include ridged denticles, reflecting the evolutionary success of this particular design. Further, I find that ridged denticles may have been key in a radiation event within the ground sharks (Carcharhiniformes) 166 MYA.
    In chapter 3, I redirect the focus back to the living, interacting sharks. Here, I expand the number of morphogroups from ten to 17 in order to fully appreciate denticle disparity across individual sharks. I apply our understanding of denticle functionality from Chapter 2, and find that each body region is equipped with a set of denticle morphogroups designed to mitigate the physical challenges related to its particular lifestyle. I conclude that denticles of the snout, and fin edge regions exhibit low disparity and relate a “general” function, similar across species. Whereas, denticles from the special, trunk, and fin surface regions display high disparity and express a “special” function linked to lifestyle.
    Future endeavours should focus on optimizing sampling strategies to include non-lethal methods, and compile all available data on an online database to facilitate and mitigate conservation management of unknown species involved in the illegal shark finning trade.

    TABLE OF CONTENTS ACKNOWLEDGEMENTS…………………………………………………………………...ii ABSTRACT…………………………………………………………………………………...v TABLE OF CONTENTS……………………………………………………………………..vi LIST OF TABLES……………………………………………………………………..….…..x LIST OF FIGURES…………………………………………………………………………xiii LIST OF MOVIES………………………………………………………………………..…xix Chapter 1 – General Introduction 1.1 What are dermal denticles? ……………………………………………………….………1 1.2 A history of dermal denticle research……………………………………………….…….1 1.3 Why study dermal denticles? …………………………………………………….……….2 1.4 Objectives…………………………………………………….……………………………3 1.5 Thesis outline…………………………………………………….………………………..4 Figures…………………………………………………….…………………………………...4 Chapter 2 – Sharks with Ridged Dermal Denticles Have a Higher Diversity 2.1 Abstract…………………………………………………….………………………..…….5 2.2 Introduction……………………………………………………...…….…………………..5 2.3 Materials and Methods……………………………………………..…………..….………7 2.3.1 Sample collection………………………………………….…………….………7 2.3.2 Scanning electron microscopy (SEM) ………………………………..…………8 2.3.3 Morpho-space construction with deep learning model…………....……….……8 2.3.4 Nanoindentation (NI) and electron probe microanalysis (EPMA)……………..10 2.3.5 Micro-CT…………………………………………………….…………...…….13 2.3.6 3D-models…………………………………………………….………….…….13 2.3.7 Computational fluid dynamics (CFD) …………………………………………14 2.3.8 Ancestral state reconstruction (ASR) ……………………….…………………15 2.3.9 Diversification rate analysis……………………………………………………15 2.3.10 Functional trait analysis ………………………………………………………17 2.4 Results…………………………………………………….…………….………………..18 2.4.1 Deep learning constructs the morpho-space of dermal denticles for delineating morphogroups …………..……………………………………………………………18 2.4.2 Ancestral state reconstruction and diversification rate of morphogroups…..….19 2.4.3 Form and function of dermal denticles…………………………………………21 2.4.3.1 Morphogroup 1, 3…………………………………………………………….22 2.4.3.2 Morphogroup 4, 5, 7………………………………………………………….22 2.4.3.3 Morphogroup 6, 8, 9, 10…...…………………………………………………23 2.5 Discussion…………………………………………………….…………………….……23 2.5.1 Evolutionary history of dermal denticles………………………………………23 2.5.2 The active swimmers (grps 1, 3-8)……………………………………………..25 2.5.3 The coral reef (grps 4, 5, 7)………………………………………………….…26 2.5.4 The deep sea (grps 6, 8, 9, 10)………………………………………………….27 Acknowledgements…………………………………………………….…………………….28 Figures…………………………………………………….………………………………….29 Supplementary Materials……………………………………………….…………….………34 Notes……………………………………………………………..…..……….………34 Figures…………………………………………………………….………….………45 Tables……………………………………………………………..………….………62 Chapter 3 - Dermal Denticles Provide a Spectrum of Functions Across the Shark 3.1 Abstract…………………………………………………….……………………….……64 3.2 Introduction……………………………………………………………………....………64 3.3 Materials and Methods…………………………………………………………...………66 3.3.1 Sample collection…………………………………………………...….………66 3.3.2 Scanning electron microscopy (SEM) …………………………………………66 3.3.3 Morphogroup formation…………………………………………….………….67 3.3.4 Micro-CT and 3D-models………………………………………...……………67 3.3.5 Principal component analysis (PCA) ………………………………….………68 3.4 Results…………………………………………………….……………..…………….…68 3.4.1 Denticles vary across the individual……………………………………………68 3.4.2 Denticles reflect lifestyle………………………………………….……………69 3.5 Discussion………………………………………………………………………………..70 3.5.1 Low disparity regions……………………………………………………..……70 3.5.1.1 The snout and fin edge regions…………….…………………………70 3.5.2 High disparity regions………………………………………………………….71 3.5.2.1 The special region (eyes, mouth, gills, claspers)……..………………71 3.5.2.2 The trunk and fin surface regions……………….……………………72 3.5.3 Denticles reflect lifestyle……………………………………………………….72 3.5.3.1 The active swimmer……………………………….…………………72 3.5.3.2 The deep sea……………………………………………………….…73 3.5.3.3 The benthic dweller……………………………………..……………74 3.5.3.4 Concluding remarks…………………………………….……………74 Acknowledgements……………………………………………………………….….………75 Figures…………………………………………………….…………………………….……76 Tables………………………………………………………………………………..……….79 Chapter 4 – General Discussion 4.1 Summary of findings………………………………………………….………….………80 4.2 Future directions………………………………………………………………….………80 References…………………………………………………………………...……….………82

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