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Pedro Sánchez Palomino VARIACIÓN DEMOGRAFICA Y MORFOLÓGICA DE ESPECIES DE ROEDORES CRICETIDOS EN ECOSISTEMAS ALTOANDINOS DE LA CORDILLERA ORIENTAL COLOMBIANA Pedro Sánchez Palomino Universidad Nacional de Colombia Facultad de Ciencias, Departamento de Biología Bogotá, Colombia 2017 VARIACIÓN DEMOGRAFICA Y MORFOLÓGICA DE ESPECIES DE ROEDORES CRICETIDOS EN ECOSISTEMAS ALTOANDINOS DE LA CORDILLERA ORIENTAL COLOMBIANA Pedro Sánchez Palomino Tesis presentada como requisito parcial para optar al título de: Doctor en Ciencias-Biología Directora Ph.D. Olga L. Montenegro Instituto de Ciencias Naturales Programa de Doctorado en Ciencias-Biologia Línea Ecología Grupo en Conservación y Manejo de Vida Silvestre Universidad Nacional de Colombia Facultad De Ciencias, Departamento de Biologia Bogotá, Colombia 2017 Dedico este trabajo a la memoria de mi madre Fermina Palomino q.e.p.d (1933-2017) Dedico este trabajo a mi familia; en especial y con mucho amor a mi esposa Elizabeth Mesa y a mis hijos Nicolás y Sebastían, quienes fueron mi motivo y mi inspiración. Agradecimientos Agradezco profundamente a las siguientes instituciones y personas quienes de diversas formas y de manera muy importante contribuyeron a la concepción y desarrollo de la presente tesis, así como a varios estudios asociados a esta. El Programa de Doctorado en Biología del Departamento de Biología, Facultad de Ciencias de la Universidad Nacional de Colombia me brindó la licencia de estudio, el apoyo y la formación académica, así como el espacio y el tiempo para desarrollar y debatir ideas con mis compañeros, con mis colegas y con mis profesores Marta L. Bueno, Luis Fernando García, Diego Giraldo, Lauren Raz, Olga L. Montenegro, Joao Muñoz, Camilo García, Argenis Bonilla y Gabriel Guillot. El Jardín Botánico de Bogotá José Celestino Mutis mediante el Programa de Estímulos a la Investigación Thomas van der Hammen financió parte de algunas salidas de campo. Parques Nacionales Naturales de Colombia y el Parque Nacional Natural Chingaza me dieron el permiso de investigación y me dieron valioso apoyo administrativo y logístico para desarrollar la fase de campo de este proyecto. El Grupo en Conservación y Manejo de Vida Silvestre del Instituto de Ciencias Naturales, al cual pertenezco, me brindó soporte logístico, materiales, equipo, bibliografía e interesantes y motivantes discusiones académicas y prácticas. Los Funcionarios del Parque Nacional Natural Chingaza, la Doctora Paulina Castro Lalinde en calidad de Directora del PNN Chingaza, así como los Biólogos Miguel Suárez (q.e.p.d) y Diana Díaz Lezama Coordinadores de Investigaciones, y el Biólogo Luis Guillermo Agradecimientos V Linares coordinador de fauna, me brindaron su amistad y apoyo incondicional en aspectos logísticos para el trabajo en terreno en el campo, así como el uso de las oficinas y el laboratorio en las instalaciones del PNN Chingaza. Las y los asistentes de campo pertenecientes a las comunidades de la zona, aledañas al Parque en el Municipio de Choachí -Mauren Andrea Hortua Raigozo, Nancy Yaneth Hortúa Pulido, Beatríz Andrea Raigoso Hortua, Mauricio Raigos, Oscar Raigoso- fueron de gran ayuda y compañía para obtener la información de base de una manera técnica y confiable; en todo momento conté con su compañía y ayuda. Los estudiantes Lina García, Diana Marcela González, Catalina Cárdenas, María Camila Calderón, Luisa Fernanda Lievano, Darwin Morales y Felipe Suárez hoy ya graduados como Biólogos, graduados o a punto de lograrlo en la Maestría en Biología de la Universidad Nacional de Colombia; o algunos desarrollando estudios de Maestría o Doctorado en varios países, me acompañaron y me ayudaron como asistentes en el campo, laboratorio u en trabajo de oficina. Reconozco con gratitud la ayuda y enseñanzas técnicas de los Biólogos Jesualdo A. Fuentes y Lilian Alba en morfometría geométrica; de Eduardo Rudas y Jorge Contreras en aspectos de fotografía; y la ayuda desinteresada y profesional de los Biólogos Carlos Vargas, Jorge Contreras y María Mónica Henao en el trabajo botánico y el estudio de la vegetación. Agradezco a los Botánicos especialistas del Instituto de Ciencias Naturales de la Universidad Nacional de Colombia, quienes me ayudaron a identificar material botánico, profesores Diego Giraldo-Cañas (Poaceae), Edgar Linares (plantas no vasculares) y Julio Betancur (Bromeliaceae). Las sesiones de trabajo con mi Directora de Tesis, Profesora Olga Lucia Montenegro Ph.D., fueron escenario para la importante discusión y ajuste de ideas, comentarios y correcciones de documentos que facilitaron varias fases de la investigación desde la concepción del proyecto, el desarrollo de análisis y la concreción de las ideas centradas en los temas principales de estudio. VI Variación demografica y morfológica de especies de roedores cricetidos en ecosistemas altoandinos de la Cordillera Oriental colombiana Mi colega y amigo profesor Biólogo Hugo Fernando López me brindó su apoyo incondicional durante todo el trabajo desde la concepción del proyecto, la fase de campo, los análisis de laboratorio y la fase de escritura del documento final; fueron muy importantes su impulso afectivo, su compañía moral y su ayuda en varios aspectos técnicos y académicos. Con profunda gratitud, admiración, amor y respeto agradezco a mi colega, amiga, compañera y esposa Bióloga M.Sc. Elizabeth Mesa González por su incondicional y gran apoyo humano, por sus consejos y su ayuda en la búsqueda y manejo de información técnica y bibliográfica, revisión de textos, análisis de datos y un sinfín de largas tareas en las que de manera juiciosa y detallada me contribuyó de forma muy profesional. Finalmente agradezco a mis hijos Nicolás y Sebastián Sánchez Mesa, por entender mis estados de ánimo, por aguantar y respetar tantas horas de ausencia, así como tantos juegos y diálogos aplazados; fueron y son mi gran motivo, en mi intención de mostrarles un camino. Resumen y Abstract VII Resumen Variación Demográfica y Morfológica de Especies de Roedores Cricétidos en Ecosistemas Altoandinos de la Cordillera Oriental Colombiana. Uno de los grandes interrogantes actuales de la Biología es la forma y los mecanismos mediante los cuales se originan las especies. En este sentido, algunas propuestas apuntan hacia mecanismos ecológicos como posibles factores que promueven o producen de forma importante la especiación. Tales mecanismos se engloban en la hipótesis ecológica de la especiación, la cual podría ocurrir en alopatría o en simpatría. Se propone entonces que mediante tales mecanismos, las presiones selectivas debidas a factores bióticos o abióticos o ambos, originarían separación en uno o varios ejes del nicho ecológico entre variedades, razas o morfos las cuales causarían variación morfológica, demográfica, fisiológica o comportamental. Estas presiones y la adaptación a diferentes ambientes o diferentes ejes del nicho ecológico eventualmente originarían nuevas especies por selección disruptiva. Los roedores sigmodontinos son un excelente modelo para examinar la hipótesis ecológica de la especiación debido a sus características biológicas, ecológicas e historia evolutiva. Este grupo tiene una reciente radiación adaptativa al parecer relacionada con presiones de selección originadas por la historia geológica y orogénica de la Cordillera de los Andes. Estos procesos generaron nuevos ambientes los cuales jugaron un papel muy relevante en la biogeográfia de los roedores sigmodontinos en Suramérica. Si la especiación ecológica es un mecanismo que opera en este grupo, se esperaría que aparezcan diferencias morfológicas intraespecíficas entre poblaciones viviendo en ambientes diferentes. Igualmente, como una forma de adaptación, es posible que también existan diferencias demográficas intraespecificas en poblaciones viviendo en ambientes contrastantes. Por lo tanto, en el presente estudio se analizan las variaciones inter e intraespecíficas en las características demográficas y de conformación morfológica craneal de dos especies VIII Variación demografica y morfológica de especies de roedores cricetidos en ecosistemas altoandinos de la Cordillera Oriental colombiana filogenéticamente hermanas, los roedores altoandinos Thomasomys laniger y Thomasomys niveipes (Cricetidae, Sigmodontinae). Como modelo de estudio, poblaciones de estas dos especies brindan oportunidades de evaluar la consistencia de la hipótesis de la especiación ecológica en condiciones de simpatría y alopatría. En este contexto, se estudiaron las variaciones demográficas y de conformación morfológica craneal de dos poblaciones de cada una de las dos especies de Thomasomys en el Parque Nacional Natural Chingaza. Este parque alberga ambientes cerrados de bosque altoandino y abiertos de páramo, así como áreas mixtas en donde confluyen estos dos tipos de ambiente (mosaico). Este tipo de ambientes contrastantes permitió el estudio de poblaciones simpátricas en el mosaico y alopátricas en el bosque y el páramo. Para los estudios demográficos se usaron técnicas de captura-recaptura, análisis de matrices de transición y análisis de elasticidades de las tasas vitales y de los grupos de edad o tamaño. Se encontraron variaciones en varios parámetros demográficos entre poblaciones simpátricas y alopátricas de las dos especies. Se discuten posibles efectos de presiones selectivas originadas en el hecho de vivir en el ambiente abierto del páramo o en el ambiente cerrado del bosque. Para estudiar las variaciones morfológicas en la conformación craneal se usaron técnicas de morfometría geométrica de hitos discretos en dos dimensiones. Los
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