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Patrones De Diversidad, Distribución Y Dominancia De Plantas Leñosas En Los Bosques Montanos Del Parque Nacional Madidi, Bolivia

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Patrones De Diversidad, Distribución Y Dominancia De Plantas Leñosas En Los Bosques Montanos Del Parque Nacional Madidi, Bolivia Patrones de diversidad, distribución y dominancia de plantas leñosas en los bosques montanos del Parque Nacional Madidi, Bolivia Gabriel Arellano Tesis doctoral Madrid, 2013 Universidad Autónoma de Madrid Facultad de Ciencias, Departamento de Biología Patrones de diversidad, distribución y dominancia de plantas leñosas en los bosques montanos del Parque Nacional Madidi, Bolivia Tesis doctoral Gabriel Arellano Torres Director: Dr. Manuel J. Macía Facultad de Ciencias, Departamento de Biología Madrid, 2013 Agradecimientos A Pablo Solíz, con quien trabajé codo con codo durante toda mi estancia en Bolivia, tanto en La Paz como en el campo, y sin cuya ayuda impagable y generosa no se podría haber realizado este trabajo. A mi director de tesis, Manuel J. Macía, por su paciencia, colaboración y apoyo constante en todas las fases de este trabajo. Al resto del Proyecto Madidi, muy especialmente a Peter M. Jørgensen y al personal en La Paz: Alfredo F. Fuentes, Leslie E. Cayola, Maritza Cornejo y M. Isabel Loza. La presente tesis es fruto de su colaboración e implicación directa en algunas de las fases más críticas del trabajo. Al personal del Herbario Nacional de Bolivia, especialmente a Stephan G. Beck, Mónica Moraes, Rosa I. Meneses, Narel Y. Paniagua y Freddy S. Zenteno. La multitud de detalles en que me ayudaron durante mis dos años en Bolivia sería demasiado larga de enumerar aquí. A Victoria Cala, del Departamento de Geoquímica y Química Agrícola de la Universidad Autónoma de Madrid, por su buena disposición a enfrentarse con un volumen de trabajo casi el triple que lo acordado. A Melecio Sullca, D. Raúl Huasurco “Chaolín”, D. Justo Salas “Zorro”, D. Honorato Pinto “Don Honorio”, D. Santiago Mamani, Aníbal Mamani, Beto Apaza- Mariaca, D. Juan Mamani, D. Severino Coda “Cipriano”, Martín Chacón, D. Rogelio Mamani, Vladimir Chura “la Princesa” y Juan Carlos Mamani “el Mariposa”. Todos ellos fuertes en campo; levantaron puentes, abrieron caminos y encendieron fuego bajo la lluvia. Sin su habilidad, inteligencia y compromiso, esta investigación no hubiera sido posible. A Óscar Quiroga "el Pulguita". Él no era fuerte ni habilidoso, pero sí muy divertido. A Daniel Rubín de Celis, por múltiples hazañas. De entre todas destaca su inesperada aparición con cien panes y mantequilla mientras mi equipo de trabajo, Pablo y yo huíamos de Culi tras cuarenta días en campo plagados de penurias, tábanos y errores. A Alain Messili, que aportó nombres, teléfonos, mapas reales e imaginarios, y algunas otras claves. 3 A Ramón Morales, por su supervisión en el Real Jardín Botánico (Consejo Superior de Investigaciones Científicas). También agradezco la paciencia y profesionalidad del personal gerente y administrativo del Real Jardín Botánico, la Dirección General de Universidades e Investigación (Comunidad de Madrid), los Departamentos de Ecología y Biología (Universidad Autónoma de Madrid), el Herbario Nacional de Bolivia (Universidad Mayor de San Andrés) y el Missouri Botanical Garden. A Pedro Armesilla, Amaya Herrero, Ignacio Palomo y Rut Caparrós, por ayudarme con trámites burocráticos en España mientras estaba en Bolivia y ofrecerme alojamiento durante mis estancias en Madrid durante aquel período. Esta tesis ha sido realizada durante un contrato de cuatro años como técnico de apoyo a la investigación en el Real Jardín Botánico, con fondos aportados por la Comunidad de Madrid. El trabajo en Bolivia durante 2009 y 2010 empleó fondos del proyecto Inventario florístico de los bosques montanos del Parque Nacional Madidi (Bolivia), lo mismo que el análisis de las muestras de suelo, financiado también por el proyecto Florística y papel de los factores edáficos en la composición de los bosques: extensión metodológica de los bosques tropicales a los bosques templados. Ambos proyectos los financió el Centro de Estudios de América Latina, una fundación conjunta entre el Banco Santander y la Universidad Autónoma de Madrid. Para la realización de esta tesis también se han empleado datos de otros investigadores, que forman o formaron parte del Proyecto Madidi, con financiación de la Comunidad de Madrid, el programa ENRECA (Agencia Danesa de Desarrollo Internacional), National Science Foundation (USA), National Geographic Society (USA), Andrew W. Mellon Foundation (USA), Christopher y Sharon Davidson, y la familia Taylor. 4 Índice Página Capítulo 1: Introducción general 9 Marco teórico: diversidad, distribución y dominancia de plantas leñosas 12 Dominancia e hipótesis de la oligarquía 13 Recambio de especies: procesos determinísticos vs. procesos neutros 14 Zona de estudio: región Madidi 15 Datos empleados, colaboración con el Proyecto Madidi 15 Estructura y objetivos de la tesis 18 Referencias 20 Capítulo 2: A standard protocol for woody plant inventories and soil characterization using 0.1-ha plots in tropical forests 25 Abstract 27 Introduction 29 Plot-level protocol 34 Plot shape and distance measurements 34 Plot description data 35 Measurements of dbh 35 Multiple stems 36 Stem height 36 Soil sampling and laboratory analysis 36 Plot metadata and data storage 38 Site-level protocol 39 Acknowledgments 40 References 40 Capítulo 3: Local and regional dominance of woody plants along an elevational gradient in a tropical montane forest of northwestern Bolivia 45 Abstract 47 Introduction 49 Methods 51 Sampling design and study area 51 Data collection 51 Data analysis 52 Results 53 Floristic composition 53 Overlap of important taxa vs. non-important taxa 55 Correlations for the importance of taxa between sites 55 5 Página Discussion 56 Floristic patterns 56 Dominance across elevations 57 Dominance within elevational ranges 58 Acknowledgments 58 References 59 Capítulo 4: Commonness patterns and the size of the species pool along a tropical elevational gradient: insights using a new quantitative tool 63 Abstract 65 Introduction 67 Methods 69 Study region and floristic data 69 Characterization of species commonness 70 Mean commonness at the local scale: measurement and determinants 70 Delimitation of the set of common species 71 Patterns of common species at the local scale 72 Comparison between forest types and characterization of the region 73 Comparison between hh criterion and Pitman et al. (2001, 2013) perspective on commonness 73 Results 73 Commonness at the local scale 73 Commonness at larger scales 74 Delimitation of the set of common species below 500 m 76 Discussion 76 Acknowledgments 79 References 79 Appendix A: R code for hp and hh calculation 83 Appendix B: Common species below 500 m 84 Capítulo 5: Niche breadth of oligarchic species in Amazonian and Andean rainforests 87 Abstract 89 Introduction 91 Methods 93 Study area and sampling design 93 Floristic data collection 94 Environmental data collection 94 Data analysis 97 Results 99 Floristic data 99 Oligarchy and niche breadth 99 Amazonian vs. Andean oligarchies 102 6 Página Discussion 106 The NBH considering the total niche size 106 The NBH considering particular environmental variables 107 Amazonian vs. Andean rainforests 107 Acknowledgments 108 References 109 Capítulo 6: Potential determinants of rarity and commonness of woody plants in different types of tropical forests 113 Abstract 115 Introduction 117 Methods 118 Study region and sampling design 118 Floristic data 118 Plant traits 119 Rarity and commonness measures 120 Statistical analysis 120 Results 121 General results 121 Rarity classification and relationship between local commonness and landscape commonness 121 Relationship between local commonness, landscape commonness and traits 121 Relationship between the four rarity classes and traits 123 Discussion 125 Community-level patterns 125 Relationship between plant traits and local and landscape-scale commonness 127 Species with different habits are apportioned very differently into rarity classes 128 Conclusions 129 Acknowledgments 129 References 130 Capítulo 7: Disentangling species turnover of woody plants in a tropical forest: from local to regional scales 133 Abstract 135 Introduction 137 Methods 138 Study region and floristic data 138 Environmental data 138 Definition of sub-regions at different spatial scales 139 Response and explanatory matrices 139 Hierarchical variation partitioning 140 7 Página Relationship between explained variation and spatial scale 141 Results 143 Discussion 147 Total explained variation decreases with increasing spatial scale 147 Processes along spatially structured environmental gradients are scale-dependent, whereas processes along un-structured environmental gradients are not 149 Effects of environment are more important than those of space across scales 150 Climate and soils both contribute to the environmental effect structuring floristic assemblages 151 Conclusions 151 Acknowledgments 152 References 152 Capítulo 8: Conclusiones generales 157 Dominancia 159 Recambio de especies 160 Referencias 161 Apéndices 163 Apéndice 1: Lista de especies inventariadas 165 Apéndice 2: Características generales de las parcelas 205 Apéndice 3: Características edáficas de las parcelas 207 8 CAPÍTULO 1 Introducción general 9 10 Los bosques tropicales de los Andes son ecosistemas de excepcional interés porque presentan (a) una alta riqueza de especies, (b) un alto nivel de endemismo, y (c) un alto número de especies amenazadas. Estos tres criterios son clave para optimizar la conservación de la biodiversidad a escala global y solo se simultanean en menos del 0,2% de la superficie terrestre (Orme et al. 2005). Distintos procesos explican por qué se encuentra aquí una gran
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