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Functional Morphology of the Avian Cervical Column

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Functional Morphology of the Avian Cervical Column The Surrogate Arm: Functional Morphology of the Avian Cervical Column Thesis submitted in accordance with the requirements of the University of Liverpool for the degree of Doctor in Philosophy by Ryan Derek Marek September 2019 1 Table of Contents Abstract ................................................................................................................................... 7 Acknowledgements .............................................................................................................. 8 Chapter 1: Introduction ...................................................................................................... 11 References ............................................................................................................................. 20 Chapter 2: Background ...................................................................................................... 33 The vertebrate axial column and regionalisation ...................................................... 33 The avian cervical column: a model for regionalisation.......................................... 40 Geometric morphometrics: a quantitative proxy for regionalisation .................... 47 Allometric scaling in the avian cervical column ........................................................ 56 Kinematics of the Avian Craniocervical System ......................................................... 60 Introduction to musculoskeletal kinematics ............................................................. 60 Kinematics of the avian cervical column ................................................................... 62 Anatomy of the avian cervical column ........................................................................ 70 Introduction to animal bone and avian cervical vertebrae osteology .................. 70 Myology ........................................................................................................................ 75 Summary ......................................................................................................................... 107 References ...................................................................................................................... 109 Chapter 3: Patterns of avian cervical regionalisation ................................................. 135 Introduction ................................................................................................................... 135 Methods .......................................................................................................................... 141 Specimen information and 3D digitisation ............................................................. 141 Morphological analysis of 3D landmarks ................................................................ 143 Analysing morphological regionalisation in the avian cervical column.............. 145 Results ............................................................................................................................. 148 Q1: Is the number of cervical regions across extant Aves constant? .................. 148 2 Q2: Are cervical regions defined by specific morphological traits? Do these traits vary across Aves in accordance with external factors? ......................................... 151 ..................................................................................................................................... 153 Q3: Do these traits vary across Aves in accordance with external factors? ....... 154 ......................................................................................................................................... 160 Q4: Do variations in cervical region size across Aves correlate with external factors? ........................................................................................................................ 161 Discussion ...................................................................................................................... 163 Cervical regions are defined by specific morphologies which don’t respond to external factors: a ‘general pattern’ ......................................................................... 164 Shape change across the cervical column deviates from the ‘general pattern’ for extreme ecologies ..................................................................................................... 168 Number of vertebrae per cervical region does not correlate with external factors ..................................................................................................................................... 173 Evolution of regionalisation in archosaurs ............................................................. 176 Conclusions .................................................................................................................... 178 References ...................................................................................................................... 179 Chapter 4: Neck length and head mass variation ....................................................... 188 Introduction ................................................................................................................... 188 Methods ......................................................................................................................... 191 Defining gross morphological parameters: head mass, body mass and neck length .......................................................................................................................... 191 ..................................................................................................................................... 193 Hypothesis testing ..................................................................................................... 194 Results ............................................................................................................................. 196 Q1. How does neck length scale with body mass across extant Aves? .............. 196 Q2. How does head mass scale with body mass across extant Aves? ................ 198 Q3. Do ecological factors correlate with neck length and head mass allometry? ..................................................................................................................................... 198 Q4. How does neck length vary with head mass across extant Aves? ............... 199 3 Discussion ....................................................................................................................... 201 Perspectives on the study of the vertebrate cervical column .............................. 201 How does neck length and head size scale with body mass across extant Aves? Do ecological factors correlate with neck length and head size? ....................... 206 How does neck length vary with head mass across extant Aves? Do ecological factors correlate with this relationship? .................................................................. 212 Conclusions and future work ...................................................................................... 214 References ...................................................................................................................... 215 Chapter 5: Variation in regional vertebral morphology and region length ........... 228 Introduction ................................................................................................................... 228 Methods .......................................................................................................................... 231 Defining functional factors: head mass, body mass and neck length ................. 231 Geometric morphometrics ........................................................................................ 234 Hypothesis testing ..................................................................................................... 235 Results ............................................................................................................................. 237 Q1: Does vertebral morphology correlate with functional factors? .................... 237 Q2: Do ecological factors have any correlations with morphology? .................. 240 Q3: How do functional factors and ecological factors correlate with region length? ......................................................................................................................... 242 Discussion ....................................................................................................................... 245 Does vertebral morphology correlate with functional factors? Do ecological factors have any correlations with morphology? .................................................. 245 How do functional factors and ecological factors correlate with region length? ...................................................................................................................................... 251 Conclusions .................................................................................................................... 255 References ...................................................................................................................... 256 Chapter 6: Variation in cervical muscle architecture and muscle mass................... 268 Introduction ................................................................................................................... 268 4 Material and methods .................................................................................................
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