Andrés Meynard Pontificia Universidad Católica De
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1 PONTIFICIA UNIVERSIDAD CATÓLICA DE CHILE FACULTAD DE CIENCIAS BIOLÓGICAS PROGRAMA DE DOCTORADO EN CIENCIAS BIOLÓGICAS MENCIÓN ECOLOGÍA TESIS DOCTORAL GRADO DE ESPECIALIZACIÓN ECOLÓGICA Y FILOGEOGRAFÍA COMPARADA ENTRE EL HERBÍVORO ESPECIALISTA SCURRIA SCURRA Y SUS DOS ALGAS HOSPEDEROS EN LA COSTA CHILENA POR ANDRÉS MEYNARD Junio 2014 2 PONTIFICIA UNIVERSIDAD CATÓLICA DE CHILE FACULTAD DE CIENCIAS BIOLÓGICAS PROGRAMA DE DOCTORADO EN CIENCIAS BIOLÓGICAS MENCIÓN ECOLOGÍA GRADO DE ESPECIALIZACIÓN ECOLÓGICA Y FILOGEOGRAFÍA COMPARADA ENTRE EL HERBÍVORO ESPECIALISTA SCURRIA SCURRA Y SUS DOS ALGAS HOSPEDEROS EN LA COSTA CHILENA Tesis presentada a la Pontificia Universidad Católica de Chile como parte de los requisitos para optar al grado de Doctor en Ciencias Biológicas mención Ecología POR ANDRÉS MEYNARD Directores de Tesis: Sylvain Faugeron y Juan Correa M. Comisión de Tesis: Claudio Latorre David Véliz Evie Wieters Junio 2014 i Agradecimientos Quiero comenzar agradeciendo el financiamiento que hizo posible esta tesis de parte de la Comision Nacional de Investigación Científica y Tecnológica (CONICYT, Chile), y el Núcleo Milenio (Centro de Conservación Marina). A mi tutor Sylvain Faugeron por su transmisión de conocimiento y rigurosidad en el análisis científico de datos. Igualmente agradezco el conocimiento transmitido y los comentarios en el momento preciso por parte de mi co-tutor, Juan Correa Maldonado. A mis compañeros de laboratorio, Fernanda Araujo y Gabriel Montecinos por su ayuda y siempre transmitir buena onda.A Gioconda Peralta por su ayuda técnica en Biología Molecular y su buena disposición. Agradecido igualmente de todos aquellos que me ayudaron en el trabajo de laboratorio y sobre todo enel muestreo en terreno, sin lo cual no podría haber recolectado datos. Es decir, agradecido de mis amigos y colegas de laboratorio Juan Luis Allendes, Loretto Contreras- Porcia, Anibal Contreras-Porcia, Tatiana Manzur, Renato Borrás, Daniela Thomas, Paula Ayerdi, Jessica Beltrán, Verónica Flores, Daniella Mella, Santiago Andrade, Pedro Sepúlveda, y Florence Tellier. Agradecido igualmente por la ayuda teorica y consejosdel Dr. David Véliz de la U. de Chile, y de la Dra. Fernanda Pérez de la P. Universidad Católica de Chile. Gracias igualmente a mis padres y amigos por su apoyo. ii TABLE OF CONTENTS Agradecimientos...................................................................................................................... iii Table of contents......................................................................................................................iv Index of figures....................................................................................................... ...............vii Index of tables..........................................................................................................................x Resumen...................................................................................................................................xii Abstract...................................................................................................................................xiv CHAPTER I: GENERAL INTRODUCTION .................................................................... 1 1 Theoretical background on ecological specialization: Why are there so few specialist herbivores in marine environments? ................................................................................... 2 1.1 The specialist-generalist continuum in marine environments: ....................................... 2 1.2 Section I. Key concepts to define degrees of ecological specialization: ........................... 4 1.3 Section II. Main hypotheses on the origin of herbivore specialization in marine communities: ............................................................................................................................... 18 1.4 Biological model and main hypotheses of this thesis: .................................................... 24 2 Referencias:................................................................................................................. 29 CHAPTER II: ABUNDANCE AND DEGREE OF SELECTIVITY OF A SPECIALIST MARINE HERBIVORE CORRELATED TO AVAILABILITY OF MICROHABITATS OF SPECIFIC SHAPE AND INCREASING SIZE WITHIN ITS TWO HOST KELPS 38 1 Introduction: ............................................................................................................... 39 2 Material and Methods: ............................................................................................... 44 2.1 Study sites and data collection ....................................................................................... 44 2.1.1 Host selectivity: standardized degree of utilization according to host availability and structural complexity (Component 2 of ecological versatility) ................................................... 45 2.1.2 Relative performance in different hosts (Component 3 of ecological versatility) ........... 46 iii 3 Results: ........................................................................................................................ 47 3.1 Regional picture of component 2 ................................................................................... 47 3.1.1 Component 2: standardized degree of utilization according to host availability ............. 49 3.1.2 Component 2: standardized degree of utilization according to host structural complexity 51 3.1.3 Component 3: Relative performance in different hosts .................................................. 57 4 Discussion .................................................................................................................... 59 5 References: .................................................................................................................. 63 CHAPTER III:COMPARATIVE PHYLOGEOGRAPHY OF A SPECIFIC BUT RECENT ASSOCIATION BETWEEN A SPECIALIST MARINE HERBIVORE AND ITS HOST KELP IN THE SOUTHEAST PACIFIC COAST: CONCORDANT SPATIAL GENETIC STRUCTURES AND SIMILAR DEMOGRAPHIC TRAJECTORIES SINCE THE LAST GLACIAL MAXIMUM. .................................... 69 1 Introduction: ............................................................................................................... 70 2 MATERIAL AND METHODS .................................................................................. 77 2.1 Sampling for COI and AFLP genotyping ...................................................................... 77 2.2 DNA extraction and COI amplification ......................................................................... 78 2.3 AFLP genotyping ........................................................................................................... 78 2.4 Population genetic analyses: .......................................................................................... 80 2.5 Phylogeographic inferences: .......................................................................................... 82 2.6 Correlation of pairwise population genetic distances between S. scurra and L. spicata: 83 2.7 Searching for loci under selection in S. scurra: ............................................................. 83 3 RESULTS ................................................................................................................... 84 3.1 Identification of the true number of S. scurra populations and degree of similarity with Lessonia phylogeographic structure: ......................................................................................... 84 3.1.1 Genetic diversity of S. scurra revealed by the COI and AFLP’s: ................................... 84 3.1.2 Population genetic structure of S. scurra and its host L. spicata: ................................... 87 iv 3.2 Evaluation of contrasting demographic scenarios since the LGM in S. scurra populations .................................................................................................................................. 93 3.2.1 Reconstruction of the effective population sizes of S. scurra clusters using the COI: .... 93 3.2.2 Simulations of Biogeographic scenarios for S. scurra and L. spicata populations since the LGM: 95 3.3 Correlation of pairwise population genetic distances between S. scurra and L. spicata 101 3.4 Searching for loci under selection in S. scurra inhabiting L. spicata or D. antartica: . 103 4 DISCUSSION AND PERSPECTIVES: ....................................................................104 4.1 Common phylogeographic structure in S. scurra and its host kelp ............................ 104 4.2 Shared demographic history between S. scurra and its host kelp ............................... 108 4.3 Role of selection in the genetic structure of S. scurra .................................................. 110 4.4 Comparative phylogeography of S. scurra and L. spicata ........................................... 111 5 References: .................................................................................................................114 CHAPTER IV: FINAL DISCUSSION .............................................................................122 1 Some logistic issues and perspectives in the study of ubiquity and ecological versatility ...........................................................................................................................123 1.1 Concepts previously developed for terrestrial systems as criteria to evaluate the degree of specialization in the marine environment: ..........................................................................