Ecology of Macroalgal Wrack on Exposed Sandy Beaches
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ECOLOGY OF MACROALGAL WRACK ON EXPOSED SANDY BEACHES ECOLOGIA DE LOS ARRIBAZONES DE MACROALGAS EN PLAYAS EXPUESTAS TESIS DOCTORAL FRANCISCO BARREIRO ROMANO VIGO 2013 Departamento de Ecología y Biología Animal Tesis Doctoral Ecology of macroalgal wrack on exposed sandy beaches Ecología de los varamientos de macroalgas en playas arenosas expuestas Francisco Barreiro Romano Directores Mariano Lastra Valdor Jesús López Pérez Vigo 2013 Departamento de Ecoloxía e Bioloxía Animal Os doutores Mariano Lastra Valdor e Jesús López Pérez, profesores titulares do Departamento de Ecoloxía e Bioloxía Animal da Universidade de Vigo CERTIFICAN: Que a presente memoria de doutoramento titulada “Ecology of macroalgal wrack on exposed sandy beaches”, presentada por Francisco Barreiro Romano para optar o grado de Doutor pola Universidade de Vigo, realizouse baixo a nosa dirección no Departamento de Ecoloxía e Bioloxía Animal da Universidade de Vigo, e considerando que cumpre os requisitos necesarios para ser considerada como un traballo de Tese Doutoral, autorizamos a súa presentación ante o Consello do Departamento e a Comisión de Posgrado. E para que así conste, emitimos e asinamos a presente certificación, en Vigo a 09 de Maio do 2013. Fdo. Mariano Lastra Valdor Fdo. Jesús López Pérez Edificio de Ciencias Experimentales Campus As Lagoas Marcosende 36310 VIGO Ecology of macroalgal wrack on exposed sandy beaches Dissertation presented in fulfillment of the requirements for the degree of Doctor by the University of Vigo PhD thesis Francisco Barreiro Romano Vigo 2013 TABLE OF CONTENTS 1. CHAPTER I. GENERAL INTRODUCTION 6 1.1. SANDY BEACHES .................................................................................................. 888 1.2. PHYSICAL CHARACTERIZATION ............................................................................... 9 1.2.1. Structural beach components ................................................................................ 9 1.2.2. Main influential factors in a beach system ............................................................ 12 1.2.3. Beach morphodynamics ...................................................................................... 13 1.3. ECOLOGY OF SANDY BEACHES .............................................................................. 18 1.3.1. Influence of the physical environment in the beach macrofauna ............................ 20 1.3.2. Zonation patterns on sandy beaches .................................................................... 22 1.3.3. Beach connectivity .............................................................................................. 24 1.4. Wrack ..................................................................................................................... 25 1.5. The socio-economic value of sandy beaches ........................................................ 27 1.6. Background of this project .................................................................................... 28 1.7. Approach taken in this thesis ................................................................................ 30 1.8. Aims of this thesis.................................................................................................. 31 1.9. Literature cited ...................................................................................................... 33 2.CHAPTER II. ANNUAL CYCLE OF WRACK SUPPLY TO SANDY BEACHES: EFFECT OF THE PHYSICAL ENVIRONMENT 40 1. ABSTRACT ................................................................................................................. 42 2.2. INTRODUCTION ..................................................................................................... 43 2.3. MATERIALS AND METHODS .................................................................................. 45 2.3.1. Field sampling design .......................................................................................... 45 2.3.2. Statistical analysis ............................................................................................... 47 2.4. RESULTS ................................................................................................................. 49 2.4.1. Beach characteristics and wrack deposition .......................................................... 49 2.4.2. Time-scale variability in wrack composition and biomass ....................................... 50 2.5. DISCUSSION ........................................................................................................... 57 2.6. ACKNOWLEDGEMENTS ......................................................................................... 60 2.7. LITERATURE CITED ................................................................................................. 61 3. CHAPTER III. COUPLING BETWEEN MACROALGAL INPUTS AND NUTRIENTS OUTCROP IN EXPOSED BEACHES 64 3.1. ABSTRACT .............................................................................................................. 66 3.2. INTRODUCTION ..................................................................................................... 67 3.3. MATERIALS AND METHODS .................................................................................. 69 3.3.1 Field sampling design ........................................................................................... 69 3.3.2. Nutrient analysis ................................................................................................. 71 3.3.3. Statistical analysis ............................................................................................... 72 3.4. RESULTS ................................................................................................................. 73 3.4.1. Beach characteristics ........................................................................................... 73 3.4.2. Dissolved nutrients and wrack biomass ................................................................ 74 3.4.3. Intertidal pore water (IPW) .................................................................................. 75 3.4.4. Surf zone water (SZW) ......................................................................................... 77 3.5. DISCUSSION ........................................................................................................... 79 3.6. ACKNOWLEDGMENTS ............................................................................................ 84 3.7. LITERATURE CITED ................................................................................................. 84 4. CHAPTER IV. THE INFLUENCE OF ALLOCHTHONOUS INPUTS ON THE MACROFAUNAL ASSEMBLAGES IN OCEAN-EXPOSED SANDY BEACHES 90 4.1. Abstract .................................................................................................................. 92 4.2. Introduction ........................................................................................................... 93 4.3. Material and methods ........................................................................................... 95 4.3.1. Field sampling design .......................................................................................... 95 4.3.2. Statistical analysis ............................................................................................... 97 4.4. Results .................................................................................................................... 98 4.4.1. Species richness and total abundance ................................................................... 98 4.4.2. Community composition ................................................................................... 101 5. Discussion ............................................................................................................... 104 4.6. ACKNOWLEDGMENTS ......................................................................................... 109 3.7. LITERATURE CITED ............................................................................................... 110 5. CHAPTER V. EFFECT OF MECHANICAL GROOMING ON THE MACROFAUNAL COMMUNITY OF EXPOSED SANDY BEACHES 116 5.1. Abstract ................................................................................................................ 118 5.2. Introduction ......................................................................................................... 119 5.3. Material and methods ......................................................................................... 121 5.3.1. Field sampling design ........................................................................................ 121 5.3.2. Statistical analysis ............................................................................................. 123 5.4. Results .................................................................................................................. 124 5.4.1. Number of species and density .......................................................................... 124 5.4.2. Community composition ................................................................................... 126 5.4.3. Specie-site association ...................................................................................... 128 5.5. Discussion ............................................................................................................ 130 5.6. Acknowledgements ............................................................................................. 134 5.7. Literature cited