Developmental, ecological and biological drivers of macroevolutionary trajectories of postcranial morphological diversification in bats Camilo López-Aguirre A thesis in fulfillment of the requirements for the degree of Doctor of Philosophy School of Biological, Earth and Environmental Sciences Faculty of Science August 2020 3 4 5 6 7 Table of Contents List of abbreviations............................................................................................... 12 Acknowledgments ................................................................................................. 13 List of figures ......................................................................................................... 16 List of tables .......................................................................................................... 20 Chapter 1: General introduction……………………………………………………………………………. 21 References .................................................................................................................... 24 Chapter 2: Postcranial heterochrony, modularity, integration and disparity in the prenatal ossification in bats (Chiroptera)………………………………………………………………..44 Abstract .............................................................................................................................................. 44 Introduction ....................................................................................................................................... 45 Materials and methods ................................................................................................. 52 Data collection ................................................................................................................................... 52 Sequence heterochrony ..................................................................................................................... 55 Metric growth .................................................................................................................................... 56 Developmental modularity ................................................................................................................ 57 Developmental disparity and integration .......................................................................................... 58 Results .......................................................................................................................... 60 Sequence heterochrony ..................................................................................................................... 60 Developmental growth ...................................................................................................................... 62 Developmental modularity ................................................................................................................ 63 Disparity and integration ................................................................................................................... 64 Discussion ..................................................................................................................... 67 Sequence heterochrony and developmental growth ........................................................................ 67 Developmental modularity ................................................................................................................ 72 Disparity and integration ................................................................................................................... 74 Conclusions ................................................................................................................... 77 References .................................................................................................................... 78 Chapter 3: Prenatal allometric trajectories and the developmental basis of postcranial phenotypic diversity in bats (Chiroptera) .............................................. 89 Abstract ........................................................................................................................ 89 Introduction .................................................................................................................. 90 Materials and Methods ................................................................................................. 96 Data collection ................................................................................................................................... 96 Ontogenetic allometric trajectories ................................................................................................... 97 Evolution of ontogenetic allometric trajectories ............................................................................... 99 Results ........................................................................................................................ 100 Ontogenetic allometry ..................................................................................................................... 100 Allometric space analysis............................................................................................. 105 Evolution of ontogenetic allometric trajectories ............................................................................. 106 Discussion ................................................................................................................... 109 Conclusions ................................................................................................................. 115 8 References .................................................................................................................. 116 Chapter 4: Prenatal developmental trajectories of fluctuating asymmetry in bat humeri ............................................................................................................................. 132 Abstract ...................................................................................................................... 133 Introduction ................................................................................................................ 134 Methods ..................................................................................................................... 140 Sampling ........................................................................................................................................... 140 Data collection ................................................................................................................................. 141 Estimation of asymmetry ................................................................................................................. 142 Data analysis .................................................................................................................................... 144 Results ........................................................................................................................ 145 Symmetric and asymmetric shape variation ................................................................................... 145 Longitudinal and cross-sectional asymmetry .................................................................................. 146 Discussion ................................................................................................................... 149 Conclusions ................................................................................................................. 153 References .................................................................................................................. 154 Chapter 5: Variation in cross-sectional shape and biomechanical properties of the bat humerus under Wolff’s law ............................................................................................................................. 160 Abstract ...................................................................................................................... 160 Introduction ................................................................................................................ 161 Materials and methods ............................................................................................... 165 Sample description and digitisation................................................................................................. 165 Extraction of landmark and biomechanical data ............................................................................. 167 Ecological and phylogenetic characterisation of species ................................................................ 169 Statistical analyses ........................................................................................................................... 169 Results ........................................................................................................................ 171 Scaling of biomechanical and shape data ........................................................................................ 171 Ecological differences in morpho- and mechanospaces ................................................................. 172 Discussion ................................................................................................................... 176 Scaling of biomechanical and shape data ........................................................................................ 177 Ecological differences in shape and