Three-dimensional study of the Iberian red deer antler (Cervus elaphus hispanicus): application of geometric morphometrics techniques and other methodologies Débora Martínez Salmerón ADVERTIMENT. La consulta d’aquesta tesi queda condicionada a l’acceptació de les següents condicions d'ús: La difusió d’aquesta tesi per mitjà del servei TDX (www.tdx.cat) i a través del Dipòsit Digital de la UB (diposit.ub.edu) ha estat autoritzada pels titulars dels drets de propietat intel·lectual únicament per a usos privats emmarcats en activitats d’investigació i docència. No s’autoritza la seva reproducció amb finalitats de lucre ni la seva difusió i posada a disposició des d’un lloc aliè al servei TDX ni al Dipòsit Digital de la UB. No s’autoritza la presentació del seu contingut en una finestra o marc aliè a TDX o al Dipòsit Digital de la UB (framing). Aquesta reserva de drets afecta tant al resum de presentació de la tesi com als seus continguts. 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Three-dimensional study of the Iberian red deer antler (Cervus elaphus hispanicus): application of geometric morphometrics techniques and other methodologies Débora Martínez Salmerón PhD Thesis Barcelona, 2014 Facultad de Biología Departamento de Biología Animal Programa de Doctorado de Biodiversidad Three-dimensional study of the Iberian red deer antler (Cervus elaphus hispanicus): application of geometric morphometrics techniques and other methodologies Memoria presentada por Débora Martínez Salmerón para optar al título de Doctora por la Universidad de Barcelona Barcelona, 2014 VºBº de los directores: Director/Tutor Directora Adrià Casinos Pardos Concepción Azorit Casas Depto. Biología Animal Depto. Biología Animal, Universidad de Barcelona Biología Vegetal y Ecología Universidad de Jaén Contents Abstract/Resumen ................................................................................ 7 1. Introduction .................................................................................... 15 1.1 Geometric morphometrics brief history ............................... 15 1.1.1 Traditional morphometrics vs. Geometric morphometrics . 16 1.1.2 Shape space and dimensionality ....................................... 19 1.1.3 Shape components ........................................................... 20 1.2 Photogrammetric Interactive Measure Method (PhIMM) .. 21 1.3 Antlered deer ancestors ......................................................... 22 1.4 Cervus elaphus hispanicus: systematic aspects and distribution .................................................................................. 26 1.5 Economic interest of the Iberian red deer in Sierra Morena ........................................................................................ 27 1.6 Antler traits and structure .................................................... 29 1.7 Antlers vs. Horns ................................................................... 32 1.8 Objectives and justification .................................................. 33 2. Material and methods ..................................................................... 39 2.1 Study area and material collection........................................ 22 2.2 Age determination ................................................................. 22 2.3 Data obtaining and treatment ............................................... 22 2.3.1 Landmarks and semilandmarks ........................................ 42 2.3.2 Antler landmark definition ............................................... 43 2.3.3 Centroid size and allometry .............................................. 44 2.3.4 Procrustes Superimposition (GPA) or Procrustes fit .......... 46 2.3.5 Multivariate statistics ....................................................... 47 2.3.6 Symmetry and asymmetry ................................................ 48 2.3.7 Integration and modularity ............................................... 51 2.3.8 Digitalization methods ..................................................... 53 1 Contents 2.4 Tools for the application of the PhIMM ............................... 54 3. Geometric morphometric analysis of antler development in Iberian red deer (Cervus elaphus hispanicus)................................................. 59 3.1 Abstract ................................................................................. 59 3.2 Introduction and objectives ................................................... 60 3.3 Material and methods ........................................................... 64 3.4 Results .................................................................................... 69 3.5 Discussion and conclusions .................................................... 78 3.5.1 How do the studied factors affect antler morphology? ...... 78 3.5.2 Thoughts on antler quality optimization............................ 82 3.5.3 Modularity in antler development? ................................... 85 3.5.4 Interactions between factors ............................................. 86 4. Asymmetry study of Iberian red deer antler (Cervus elaphus hispanicus) by using geometric morphometrics ................................. 93 4.1 Abstract ................................................................................. 93 4.2 Introduction and objectives ................................................... 94 4.3 Material and methods ........................................................... 98 4.4 Results .................................................................................. 104 4.4.1 Measurement error and FA-DA significance................... 104 4.4.2 Differences between the right and left antler sides .......... 105 4.4.3 Allometric test ............................................................... 107 4.4.4 PCA and MANOVA ...................................................... 107 4.4.5 Antler asymmetries on age class and capture year .......... 110 4.5 Discussion and conclusions .................................................. 113 4.5.1 Presence of DA and FA .................................................. 113 4.5.2 Directional Asymmetry .................................................. 114 4.5.3 Hunter beliefs ................................................................ 116 2 Contents 4.5.4 Influence of age and capture year over antler asymmetries ........................................................................... 118 5. Exploring integration and modularity in the antler of Iberian red deer (Cervus elaphus hispanicus) ..................................................... 125 5.1 Abstract ............................................................................... 125 5.2 Introduction and objectives ................................................. 126 5.2.1 Antler regeneration process ............................................ 128 5.2.2 Genetic basis of antler growth ........................................ 130 5.3 Material and methods ......................................................... 131 5.4 Results .................................................................................. 142 5.4.1 Allometric effect ............................................................ 142 5.4.2 H1: Complete Dataset - Right antler side vs. Left antler side .................................................................. 142 5.4.3 H2: Half antler dataset - Antler crown vs. Antler rest ...... 142 5.4.4 H3: Half antler dataset - Antler beam vs. Total tines ...... 143 5.4.5 H4: Half antler dataset - Eye tine vs. Antler rest ............. 144 5.4.6 H5: Half antler dataset - Trez tine vs. Antler rest ............ 144 5.5 Discussion and conclusions .................................................. 145 5.5.1 Complete antler dataset .................................................