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Tesis Angel Benitez2 2016 – Tesis Doctoral Tesis - Chavez Universidad Rey Juan Carlos Rey Universidad Benitez Angel EFECTOS DE LA ALTERACIÓN ANTRÓPICA EN BOSQUES TROPICALES SOBRE LA DIVERSIDAD DE ORGANISMOS EPÍFITOS (LÍQUENES Y BRIÓFITOS) Angel Raimundo Benitez Chavez: Tesis Doctoral – 2016 Universidad Rey Juan Carlos Directores: María Prieto Álvaro y Gregorio Aragón Rubio Departamento de Biología y Geología, Física y Química Inorgánica EPÍFITOS (LÍQUENES Y BRIÓFITOS) Y (LÍQUENES EPÍFITOS TROPICALES SOBRE LA DIVERSIDAD DE ORGANISMOS ORGANISMOS DE DIVERSIDAD LA SOBRE TROPICALES EFECTOS DE LA ALTERACIÓN ANTRÓPICA EN BOSQUES ANTRÓPICA ALTERACIÓN LA DE EFECTOS TESIS DOCTORAL EFECTOS DE LA ALTERACIÓN ANTRÓPICA EN BOSQUES TROPICALES SOBRE LA DIVERSIDAD DE ORGANISMOS EPÍFITOS (LÍQUENES Y BRIÓFITOS) EFFECTS OF TROPICAL FORESTS DISTURBANCE ON EPIPHYTE DIVERSITY (LICHENS AND BRYOPHYTES) ANGEL RAIMUNDO BENITEZ CHAVEZ Directores: María Prieto Álvaro Gregorio Aragón Rubio Departamento de Biología y Geología, Física y Química Inorgánica UNIVERSIDAD REY JUAN CARLOS Junio de 2016 i ii Dra. María Prieto Álvaro profesora visitante y Dr. Gregorio Aragón Rubio profesor titular del Departamento de Biología y Geología, Física y Química Inorgánica de la Universidad Rey Juan Carlos, Certifican: Que los trabajos de investigación desarrollado en la memoria de tesis doctoral “Efectos de la alteración antrópica en bosques tropicales sobre la diversidad de organismos epífitos (Líquenes y Briófitos)” se han realizado bajo su supervisión y son aptos para ser presentados por Angel Raimundo Benitez Chavez ante el tribunal que en su día se consigne, para aspirar al Grado de Doctor por la Universidad Rey Juan Carlos. V°B° Directora de Tesis V°B° Director de Tesis Dra. María Prieto Álvaro Dr. Gregorio Aragón Rubio iii iv A mis padres Gregorio y Esperanza; y la mí amada Anita v vi ÍNDICE RESUMEN Agradecimientos 1 Resumen 3 Antecedentes 6 Objetivos y estructura de la tesis 15 Metodología general 17 Discusión general 22 Conclusiones 31 Lista de manuscritos 33 Afiliación de coautores 34 Referencias 35 CAPÍTULOS/CHAPTERS Capítulo I / Chapter I 51 Effects of tropical montane forest disturbance on epiphytic macrolichens Capítulo II / Chapter II 71 Large trees and dense canopies: key factors for maintaining high epiphytic diversity on trunk bases (bryophytes and lichens) in tropical montane forests Capítulo III / Chapter III 101 Functional traits of epiphytic lichens as indicators of forest disturbance and predictors of total richness in a tropical montane rainforest of Ecuador Capítulo IV / Chapter IV 143 Lichen diversity in tropical dry forest is influenced by host preference more that disturbance Capítulo V / Chapter V 171 Additions to the bryophyte flora of Ecuador 2 Capítulo VI / Chapter VI 189 More than one hundred new records of lichens from Ecuador vii viii AGRADECIMIENTOS Agradezco a la Universidad Técnica Particular de Loja (UTPL), Secretaria Nacional de Educación Superior, Ciencia, Tecnología e Innovación (SENESCYT) y la Universidad Rey Juan Carlos por brindar las herramientas y los recursos necesarios para el desarrollo de este trabajo de investigación. A todos los integrantes del departamento de Ciencias Naturales de la UTPL de la ciudad de Loja, de manera especial a Ana Arévalo, Yadira González, Gabriela Cevallos, Geovanny Cango, Fernando Gaona y Elizabeth Guzmán por su colaboración en el trabajo de campo. Mil gracias a la desaparecida sección de sistemática de la UTPL conformada por Carlos Naranjo, Fani Tinitana, Omar Cabrera, Nixon Cumbicus y Diego Marín por su apoyo y colaboración durante el desarrollo de este trabajo de investigación, es muy grato seguir contando con su amistad y apoyo, mil gracias amigos. Al Dr. Adrián Escudero, Dra. Isabel Martínez, Dr. José María Iriondo, Dr. Marcos Méndez y Dr. Luis Cayuela de la Universidad Rey Juan Carlos, por su disposición y colaboración desinteresada durante el desarrollo de este trabajo. Adicionalmente agradezco a Silvia Santamaría, Sonia Merinero, Jesús López y Samuel por su colaboración y apoyo durante las diferentes estancias doctorales. Bueno para mis directores de tesis, empezando con el Dr. Gregorio Aragón solo tengo tres palabras respeto, admiración y sobre todo agradecimiento ya que gracias a su calidad profesional y personal he podido empezar el duro pero gratificante camino de la investigación. Mil Gracias Goyo por el tiempo y dedicación que le pusiste a este trabajo y por darme la oportunidad de hacer investigación en tu campo de estudio. Sin sentimentalismos que yo no soy muy bueno para eso, pero la verdad tengo una gran admiración por el gran trabajo que realizas y la visión que tienes para hacer investigación. A la Dra. María Prieto Álvaro, por todo su apoyo incondicional durante el desarrollo de este trabajo, gracias por seguir confiando en mis capacidades a pesar que muchas de las veces ya sobrepasaba tu gran paciencia. Gracias por enseñarme a determinar los líquenes, sobre todo los costrosos, que ahora ya los llamo crustáceos. Mil gracias a los dos por confiar en mí y darme esta gran oportunidad de que hayan sido mi guía durante este largo camino. 1 Así mismo quiero agradecer infinitamente al Profesor Robbert Gradstein por su calidad personal y profesional en los diferentes momentos que compartimos durante diferentes salidas de campo e investigaciones que hemos desarrollado en el campo de la briología. Mil gracias profesor por darme la oportunidad y sobre todo por confiar en mis capacidades. Mil gracias a mi amada familia de España, la familia de Robert, Francisca y Gregorio. De manera especial a Fátima y Johnny por su gran hospitalidad y por hacerte sentir como en casa y sobre todo por apoyarme en los momentos más difíciles durante mis estancias en España. Especialmente quiero agradecer a mi querido y amado sobrino Ismael siempre serás como el hijo que sueño tener, gracias por esperarme siempre con la comida en la mesa y en las mañanas compartir los desayunos sin importante al hora que sea. Mil gracias Isma por todos los momentos compartidos que han sido geniales. A mis padres que se han convertido en los promotores y sobre todo los impulsores para poder llegar cumplir con esta meta. Además a toda mi familia y sobrinos que gracias a su alegría he podido seguir adelante. Finalmente a la persona que se ha convertido en la alegría y paz que encontraba en los momentos más difíciles de mi vida (Anita), ya que gracias a su apoyo incondicional en las etapas de campo, de laboratorio e investigación he podido cumplir con esta etapa de mi vida, mil gracias por estar siempre para mí y esperarme aunque no haya estado cuando tú me necesitabas debido a mis estancias en España. 2 RESUMEN La deforestación es una de las principales causas de pérdida de biodiversidad en los ecosistemas tropicales a nivel global. Los bosques montanos y estacionalmente secos están despareciendo a ritmos alarmantes por lo que han sido catalogados como los más amenazados y de prioridad para la conservación. Estos ecosistemas se caracterizan por sus altos niveles de diversidad y endemismo de flora y fauna, donde los epífitos son un grupo importante en términos de diversidad y funcionamiento. Dentro de los epífitos, una fracción significativa está representada por líquenes y briófitos que, por sus características fisiológicas relacionadas con la disponibilidad de agua y luz, se han convertido en organismos modelo para evaluar cambios en el ambiente. A pesar de ello, los estudios centrados en analizar los efectos de la disturbancia de estos ecosistemas sobre los epífitos no vasculares son muy limitados, de manera especial en los bosques estacionalmente secos. Por este motivo, el objetivo general de esta tesis doctoral se centra en analizar los efectos de la alteración de los bosques tropicales sobre la diversidad de los organismos epífitos (Iíquenes y briófitos), que permitan un mejor entendimiento de las respuestas de estos organismos a cambios en el ambiente. En estos dos ecosistemas (bosques montanos y estacionalmente secos), se identificaron manchas de bosques con diferentes tipos de manejo. Se definieron tres categorías de manejo para los bosques montanos, distribuidas en dos manchas de bosques primarios, dos de bosques secundarios y dos de bosques monoespecíficos de Alnus acuminata. Para los bosques estacionalmente secos se identificaron cuatro manchas de bosques correspondientes a dos categorías de manejo. En cada bosque se analizaron los cambios en la diversidad de epífitos no vasculares a nivel de árbol y parcela; y se identificaron los factores ambientales (altitud, pendiente y orientación) y de estructura forestal (cobertura del dosel arbóreo, diámetro de los árboles, especie del árbol y textura de la corteza de los árboles) que podrían limitar estas comunidades. Además, se analizaron los efectos de la alteración sobre la diversidad funcional de líquenes epífitos en los bosques montanos y se utilizaron las formas de crecimiento para predecir la riqueza total de líquenes epífitos. Los resultados muestran que la deforestación disminuye drásticamente la diversidad de epífitos no vasculares en bosques montanos y estacionalmente secos tropicales, donde los bosques primarios mantienen una mayor diversidad en comparación con los bosques secundarios. Los epífitos de sombra (ej. Leptogium, Lobaria, Sticta, Pannaria, 3 Plagiochila) adaptadas a condiciones de alta humedad fueron más afectadas por la pérdida de la cobertura arbolada, debido a cambios microclimáticos relacionados con la disponibilidad de humedad. Sin embargo los epífitos de sol
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