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TESIS Salgado-Negret I EFECTOS DE LA FRAGMENTACIÓN SOBRE LA DISTRIBUCIÓN DE ESPECIES ARBÓREAS EN EL PARQUE NACIONAL FRAY JORGE: IMPORTANCIA DE LOS ATRIBUTOS ECOFISIOLÓGICOS II PONTIFICIA UNIVERSIDAD CATÓLICA DE CHILE FACULTAD DE CIENCIAS BIOLÓGICAS PROGRAMA DOCTORADO EN CIENCIAS BIOLÓGICAS MENCIÓN ECOLOGÍA EFECTOS DE LA FRAGMENTACIÓN SOBRE LA DISTRIBUCIÓN DE ESPECIES ARBÓREAS EN EL PARQUE NACIONAL FRAY JORGE: IMPORTANCIA DE LOS ATRIBUTOS ECOFISIOLÓGICOS Por BEATRIZ EUGENIA SALGADO NEGRET Tesis presentada a la Facultad de Ciencias Biológicas de la Pontificia Universidad Católica de Chile para optar al grado académico de Doctor en Ciencias Biológicas mención Ecología Dirigida por: Dr. Juan José Armesto Dra. Fernanda Pérez Noviembre, 2013 Santiago, Chile III Agradecimientos Quiero comenzar agradeciendo a la Comisión Nacional de Investigación Científica y Tecnológica (CONICYT, Chile) y al Instituto de Ecología y Biodiversidad (IEB) por el apoyo financiero para realizar este doctorado y el trabajo de investigación. Quiero agradecer a mi tutor principal Juan Armesto, por apostar a ciegas y permitirme ser parte de su equipo, por sus invaluables enseñanzas y apoyo incondicional. A todos los miembros de su Laboratorio por todas las tertulias e increíbles discusiones. A Fernanda Pérez por ser una excelente guía, por las eternas discusiones teóricas, por todas las salidas de campo, pero sobretodo por convertirse en una gran amiga y confidente. Fefita, eres de los grandes regalos que me llevo de Chile…gracias por todo! A Fernando Valladares y su equipo por adoptarme por meses en su laboratorio y acogerme como un miembro más del equipo. Gracias por todos los análisis y discusiones que mejoraron este documento. A Pablo Marquet, Javier Figueroa y Martín Carmona por sus aportes y comentarios que mejoraron esta propuesta desde sus inicios. A Mylthon Jimenez-Castillo y su equipo por enseñarme el mundo de la hidráulica. Gracias especiales a Paulina Lobos. A Juan Monardez, por su fiel compañía, por las deliciosas cenas y discusiones en compañía del mejor vino. Juan mil gracias por presentarme un ecosistema maravilloso. A Aurora Gaxiola y Daniel Stanton por las múltiples charlas planteando hipótesis y discutiendo resultados…. sus comentarios mejoraron enormemente los manuscritos y su compañía fue un gran apoyo. A Felipe Albornoz, Rafaella Canessa, Carmen Ossa, Daniel Salinas, Patricio Valenzuela e Isabel Mujica por su invaluable apoyo en campo y laboratorio y por hacer de las salidas de campo paseos repletos de risas y complicidad. A Mariela Aguilera y Ximena Alvarez por todas las reuniones, discusiones y tertulias alrededor de la fisiología de las plantas…hubo momentos brillantes…gracias queridas! A mis compañeras de batalla y familia en Chile: Lidia Mansur, Sabrina Clavijo, Daniela Rivera y Carmencha Ossa…. no habría sido lo mismo sin ustedes. A Leo por todo su apoyo durante tantos años. IV A Carolina Alcázar, Olga Caro y Carolina Useche por ser mis terapeutas en la distancia…gracias por todo el apoyo. A mi familia en Colombia….por darme la libertad de soñar y por estar al pie del cañón….mil gracias por estar siempre tan cerca a pesar de la distancia. A todas aquellas personas que no he nombrado pero que hicieron parte de este logro. Muchas gracias. V Tabla de contenidos Lista de abreviaturas ............................................................ VIII Resumen .................................................................................IX Introducción General ……………….………………………..………1 Estructura de la tesis………………………… .................................. 6 Área de estudio ............................................................................ 7 Visión general….. ........................................................................ 9 Referencias .................................................................................. 11 Capítulo I Estrategias divergentes de tolerancia a la sequía explican la distribución de especies arbóreas a través de un gradiente de humedad dependiente de neblina en un bosque lluvioso templado Abstract ......................................................................................... 17 Introduction .................................................................................. 18 Materials and Methods ................................................................. 20 Results .......................................................................................... 25 Discussion.................................................................................... 27 Acknowledgements ………………………………………………31 References .................................................................................... 32 Tables........................................................................................... 38 Figures ......................................................................................... 41 VI Capítulo II Variación en rasgos funcionales explica la distribución de Aextoxicon punctatum a través de un fuerte gradiente de humedad en un bosque fragmentado dependiente de neblina Abstract ........................................................................................ 47 Introduction .................................................................................. 48 Materials and Methods ................................................................. 51 Results ........................................................................................... 54 Discussion ..................................................................................... 56 Acknowledgements…………………………………………………….. 59 References ..................................................................................... 60 Tables............................................................................................ 67 Figures .......................................................................................... 70 Online supplemental materials ……………………………………74 Conclusiones Generales Conclusiones ................................................................................ 78 Anexo I Simetría de los parches de bosque depende de la dirección de los recursos limitantes Abstract ........................................................................................ 84 Introduction .................................................................................. 85 Materials and Methods ................................................................. 88 Results ........................................................................................... 90 Discussion ..................................................................................... 91 Acknowledgements……………………..………………………………. 95 VII References ..................................................................................... 96 Tables ........................................................................................100 Figures .................................................................................... 102 VIII Lista de abreviaturas AMAX = Photosynthetic rate; Tasa fotosíntesis gs = Stomatal conductance; Conductancia estomática Hv = Huber value; Valor Huber Ks = Sapwood-specific hydraulic conductivity LA = Leaf area; Área foliar LMA = Leaf mass area; Relación masa: área de la hoja PLC = Percentage of loss conductivity; Porcentaje de pérdida de conductividad RWC tlp = Relative water content at turgor loss point; Contenido relativo de agua al punto de pérdida de turgor SD = Stomatal density; Densidad estomática TD = Trichome density; Densidad de tricomas VD = Vessel density; Densidad de vasos VDi = Vessel diameter, Diámetro de vasos π0 = Solute potential at full turgor; Potencial de solutes a full turgor πtlp = Water potential at turgor loss; Potencial hídrico al punto de pérdida de turgor ɛ = Bulk modulus of elasticity; Modulo de elasticidad ψPD = Leaf water potentials predawn; Potencial hídrico al amanecer ψMD = Leaf water potentials at midday; Potencial hídrico al medio día IX Resumen El estudio de los rasgos funcionales y mecanismos fisiológicos que determinan la tolerancia de las especies a la sequía y su habilidad para competir por agua es fundamental para entender su distribución a través de gradientes de humedad y predecir su respuesta al cambio global, donde la fragmentación del hábitat y el cambio de uso del suelo son los principales motores de cambio. En este sentido, los bosques dependientes de neblina en las regiones semiáridas del mundo son un buen modelo de estudio para entender las respuestas de las especies al incremento en la aridez y la fragmentación del hábitat. En esta tesis se estudiaron los mecanismos fisiológicos que explican los patrones contrastantes de distribución observados a través de gradientes de humedad generados por la neblina costera en las tres principales especies arbóreas Aextoxicon punctatum , Drimys winteri y Myrceugenia correifolia que coexisten en los fragmentos de bosque del Parque Nacional Fray Jorge, en la región semiárida en Chile. Se identificó un continuo de estrategias en el uso de agua explicando la distribución de las especies a través del gradiente de humedad a pequeña escala. Drimys winteri , una especie restringida al núcleo húmedo, mostró rasgos que permiten un eficiente transporte de agua y ganancia de carbono; en contraste, Myrceugenia correifolia , especie que domina los bordes secos de sotavento, presentó rasgos que promueven la conservación del agua y menores tasas de intercambio de gases, así como menor potencial hídrico al punto de pérdida de turgor. La especie con amplia distribución
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