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Linea En Sistematica Efrain FACULTAD DE CIENCIAS PROGRAMA BIOLOGIA DOCTORADO EN BIOLOGIA- LINEA EN SISTEMATICA EFRAIN REINEL HENAO BAÑOL ANÁLISIS FILOGENÉTICO DEL GRUPO DALLA CYPSELUS EVANS (1955) (HESPERIIDAE: HETEROPTERINAE) CON BASE EN CARACTERES MORFOLÓGICOS Y MOLECULARES BOGOTA 25 DE NOVIEMBRE DE 2016 EFRAIN REINEL HENAO BAÑOL ANÁLISIS FILOGENÉTICO DEL GRUPO DALLA CYPSELUS EVANS (1955) (HESPERIIDAE: HETEROPTERINAE) CON BASE EN CARACTERES MORFOLÓGICOS Y MOLECULARES Tesis presentada al programa de Biología, Doctorado en Biología línea sistemática de la Universidad Nacional de Colombia como requisito parcial para la obtención del título de Doctor en Ciencias Biológicas Director: Dr. JOHN DOUGLAS LYNCH Bogotá-D.C 2016 EFRAIN REINEL HENAO BAÑOL ANÁLISIS FILOGENÉTICO DEL GRUPO DALLA CYPSELUS EVANS (1955) (HESPERIIDAE: HETEROPTERINAE) CON BASE EN CARACTERES MORFOLÓGICOS Y MOLECULARES Profesor. Dr. JONH DOUGLAS LYNCH ________________________________________________ Tesis aprobada como requisito parcial para obtar al grado de Doctor en Ciencias Biologicas- Sistemática. En el programa de Biologia de la Universidad Nacional de Colombia. Bogotá, 23 de noviembre de 2016 A mis hijos…Carlos Alejandro, Emmanuel y Samara. “Dado que en principio, una matriz de datos que contiene caracteres para diferentes minerales se puede analizar con un programa, para obtener un dendrograma, la aplicación de técnicas cladísticas por sí sola, no crea un análisis filogenético”. Wägele, 2004 “Dios en su infinita misericordia nos permite tomar desiciones, pero esas desiciones pueden ser adecuadas o no según los criterios humanos, sin embargo cada decisión tiene sus efectos, está en nuestras manos tomar las mejores desiciones para que las consecuencias sean en nuestro propio beneficio, caso contrario pagaremos por nuestros errores” A Dios gracias por permitirnos tomar nuestras propias desiciones aunque a veces no sean las mejores. AGRADECIMIENTOS En primer lugar, me gustaría agradecer a mi profesor-director Dr. John Douglas Lynch por sus grandes enseñanzas y su tiempo dedicado que permitieron y apoyaron mi formación personal e intelectual además de sus críticas constructivas sobre sistemática filogenética. A el Profesor Gonzalo Andrade-C quien confió en mí y asesoro en algunas actividades al inicio del doctorado. Al Dr. Andrew Warren “Andy” por su tiempo, dedicación, compañía y apoyo en las actividades de mi pasantía en el MGCL, además de sus buenas conversaciones, buen humor y buenas recoletas en los alrededores de Gainesville- Florida (USA), a mis profesores Olaf H. H. Mielke, Mirna Casagrande y Eduardo Carneiro por su confianza, compañía y asesoría durante mi pasantía en la Universidad Federal do Parana (Brasil) su motivación constante permitió continuar y sacar adelante este doctorado. Al Instituto de Ciencias Naturales de la Universidad Nacional de Colombia en especial al Dr. Carlos E. Sarmiento y Gary Stiles, quienes inicialmente me facilitaron el espacio de trabajo y los equipos durante la permanencia de mis estudios. A partir de allí fueron los doctores Fernando Fernández y Eduardo Flórez quienes permitieron mi instancia dentro de este espacio...mil y mil gracias. A mi profe “Gary” Stiles por toda su colaboración, comentarios, buenas charlas y excelentes recolectas de ejemplares del género Dalla en su reserva. Al profesor German Amat (directorde ICN), que siempre tuvo palabras de aliento en los momentos de desánimo. A COLCIENCIAS-COLFUTURO por la financiación de mis estudios doctorales dentro de la convocatoria doctorados nacionales-567. A Conservación Internacional por hacerme parte de su grupo de trabajo en el proyecto “Diversidad de la cuenca alta del Rio Bogota” especialmente a Carlos Andres Paez y José Vicente Rodriguez, quienes amablemente depositaron los ejemplares de Lepidoptera en el ICN-MHN-L para poder ser estudiados en este trabajo. Muchas personas contribuyeron con información para el desarrollo de esta tesis (espero no omitir a nadie) a quienes deseo agradecer inmensamente: Dra. Blanca Huertas (BMHN), Ernst Brockmann y Jean-Claude Petit, Gabriel Rodríguez, Juan Guillermo Jaramillo y Kim Garwood quienes ofrecieron imágenes que sirvieron y orientaron sobre la identificación de algunas especies. Salidas de campo tuvieron apoyo en su debido momento de Cristóbal Ríos-Málaver, Fabián G. Gaviria, Aymer Andrés Vásquez y Carlos Prieto. Finalmente a Julian A. Salazar, José Ignacio Vargas, Jean F. Le Crom, contribuyeron con el préstamo de ejemplares, Katherine Domingue y la Dra Sandra I. Uribe ayudaron en la secuenciación de las especies utilizadas en la parte molecular. A los auxiliares del laboratorio del ICN, Carlos Chaparro y Claudia Peña, su ayuda desinteresada fue su mejor regalo, A Luz Angela Carrillo y Olga Lucia Arenas las secretarias del posgrado y del ICN, quienes con su trabajo diligente siempre nos brindaron su apoyo y colaboración. Al Dr. Dimitri Forero (Museo Javeriano de Historia Natural “Lorenzo Uribe”, Pontificia Universidad Javeriana, Bogotá y Claudia Medina (Instituto Alexander von Humboldt- Villa de Leyva- Boyaca), por permitir el ingreso y el uso de la información de los ejemplares a su cargo. A mis compañeros y amigos del Laboratorio de Estudios de Lepidoptera Neotropical: Fábio dos Santos, Fernando Dias, Gabriela Leviski, Lucy Salik, Luziany Santos, Ricardo Siewert, Thamara Zacca y esposo, su agradable convivencia, apoyo, discusiones, recoletas, etc., dejaron huelle en mi vida. Agradecimiento especial a Diego Dolibaina por sus discusiones y enseñanzas que contribuyeron grandemente en mi compresión sobre la familia Hesperiidae. A mis compañeros y profesores de los seminarios de investigación por sus sugerencias y correcciones: Dra. Dolors Armenteras, René Lopez y Rafael Moreno. A Paola Triviño, Carlos Gantiva que contribuyeron con material utilizado en este trabajo y Michel bejarano quien estuvo presente en la mayor parte de esta tesis y coloboro en muchas de sus actividades... A Marvin Anganoy, Raul Pedroza, Juan Pablo López por sus comentarios, discusiones y entretenidas charlas. A todos mis familiares, especialmente mis padres Efraín y Disleny, mis hermanos Tata, Alejo y Carlos por su amor y apoyo incondicional. Por ultimo inmensa gratitud a mi esposa Luz Nancy por su amor, cariño y comprensión. No puedo dejar pasar la oportunidad de agradecer a la “Santisima trinidad” por todos los favores recibidos, mi vida y mis estudios. RESUMEN El género Dalla fue establecido por Mabille en 1904, con 47 especies, de las cuales Lyndsy, 1921 seleccionó a Cyclopides eryonas Hewitson, 1877 como especie tipo. Antes, Felder & Felder, (1867) establecieron el género Eumesia con sólo una especie, E. semiargentea como tipo. Sin embargo, Evans (1955) notó que los dos nombres genéricos eran sinónimos y rechazo a Eumesia debido a que el holotipo de E. semiargentea es un espécimen de un hespérido al que se le adjunto la cabeza de un satírino (Steinhauser, 1991). Este género se distribuye en el neotrópico con mayor concentración de especies en las regiones andinas. Actualmente cuenta con 96 especies descritas y ha sido subdivido en 7 grupos de especies con base en caracteres del ala (Evans, 1955). En esta investigación se desarrolló un análisis filogenético para el género Dalla (Hesperiidae: Heteropterinae), con base principalmente en caracteres morfológicos y se evalua la monofilia del grupo. Adicionalmente se hace enfasis en el grupo Dalla cypselus Evans (1955), donde se estudian las relaciones y se pone a prueba su monofilia, debido a ser el grupo de especies mas grande propuesto por Evans (1955), presentar amplia distribución en el neotropico y algunas especies son diciles de identificar e incluso varias especies están aun sin nombrar. Por ultimo se realiza una aproximación filogenética para 10 especies del género con datos morfológicos y moleculares. Los cladogramas resultantes muestran algunas agrupaciones poco soportadas por sinapomorfias. La monofilia del grupo Dalla cypselus Evans (1955), es soportada por caracteres de los órganos genitales y son la base para continuar futuros estudios en el grupo. Con base en el estudio morfológico de las estructuras de los órganos genitales se propone un nuevo reagrupamiento de las especies para el grupo denominado “cypselus” y se exploran las relaciones, mediante un análisis filogenético. Se describe una nueva especie y se propone cambiar el estatus de dos subespecie a especie con base en los resultados de la revisión taxonómica, datos de las colecciones biológicas y los resultados de la filogenia. Palabras clave: Filogenia, Género Dalla, Hesperiidae, Heteropterinae. ABSTRACT The Dalla genus was established by Mabille in 1904, with 47 species, of which Lyndsy, 1921 selected Cyclopides eryonas Hewitson, 1877 as species type. Before, Felder and Felder, 1867 established Eumesia with genus with only a species, E. semiargentea as species type. Nevertheless, Evans (1955) noticed that both generic names were synonymous and I reject Eumesia due to the fact that the holotipo of E. semiargentea plated is a specimen of a skypper to which I attach the head of a satyrini (Steinhauser, 1991). This genus is distributed in the neotrópico by major concentration of species in the Andean regions. It possesses 96 described species and has been I subdivide in 7 groups of species with base in characters of the wing (Evans, 1955). In this investigation an phylogenetic analysis was developed for the Dalla genus (Hesperiidae: Heteropterinae), with base principally on morphologic characters and the monophyly of the group is evaluated. In addition,
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