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Filogenia De Markea Rich. (Solanaceae: Juanulloeae) Con Base En Datos De Secuencias De ADN Nuclear Y Del Cloroplasto

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Filogenia De Markea Rich. (Solanaceae: Juanulloeae) Con Base En Datos De Secuencias De ADN Nuclear Y Del Cloroplasto Filogenia de Markea Rich. (Solanaceae: Juanulloeae) con base en datos de secuencias de ADN nuclear y del cloroplasto Andrés Orejuela Ramírez Universidad Nacional de Colombia Facultad de Ciencias, Departamento de Biología Bogotá D.C., Colombia Marzo 2016 Filogenia de Markea Rich. (Solanaceae: Juanulloeae) con base en datos de secuencias de ADN nuclear y del cloroplasto Andrés Orejuela Ramírez Tesis presentada como requisito parcial para optar al título de: Magister en Ciencias-Biología Directora: Dra. Clara Inés Orozco Línea de Investigación: Sistemática Universidad Nacional de Colombia Facultad de Ciencias, Departamento de Biología Bogotá D.C., Colombia Marzo 2016 A la memoria de mi padre. A mi madre y mi hijo. Agradecimientos Agradezco muy especialmente a la profesora Clara Inés Orozco por sus enseñanzas, apoyo y constante guía durante todos mis años como estudiante, su paciencia y confianza a pesar de los altibajos me han permitido mejorar cada vez y aprender de errores y problemas. Quiero agradecer a la Universidad Nacional de Colombia y especialmente al Instituto de Ciencias Naturales y a todos sus profesores por el apoyo brindado. A la dirección de investigación, sede Bogotá de la Universidad Nacional (DIB), proyecto 13574, al programa Jóvenes investigadores e innovadores de COLCIENCIAS y al programa de estímulos a la investigación Thomas van der Hammen del Jardín Botánico de Bogotá por la financiación. A los curadores de los herbarios visitados, a las personas que me acompañaron en campo y a quienes facilitaron sus valiosas muestras para los estudios moleculares y fotografías en vivo de las especies que acompañan este documento. Agradezco especialmente a la Dra. Lynn Bohs y al Dr. Gregory Wahlert durante mi estadía en Utah, lugar en el que fue desarrollado la totalidad del trabajo de laboratorio, a Greg un gran agradecimiento por las enseñanzas y la ayuda con la secuenciación del gen GBSSI y de algunas otras secuencias de difícil amplificación. A la Dra. Gloria Barboza por apoyar este proyecto y por su hospitalidad durante mi visita a Córdoba, Argentina y a la Dra. Carmen Benítez de Rojas por sus sabias palabras y por ayudarme gentilmente con algunas muestras de Venezuela. Quiero agradecer también a quienes fueron mis compañeros del grupo de Solanáceas de Colombia durante varios años, Duban Canal, Jorge Mario Vélez, Angélica Alba y Gabriel Beltrán, gracias por su compañerismo durante numerosos congresos, cursos y salidas de campo. Agradezco de manera especial a mi padre, quien siempre me inculcó el valor del estudio, su reciente muerte ha sido un duro golpe para mí, a mi madre que ha sido un apoyo constante en todos los aspectos, a Rocio y a mi hijo Esteban, él me dio un norte y una razón para luchar. Igualmente, quiero agradecer a Jhoana Castillo, quien me ha brindado su amor y apoyo constante e incondicional, sin ella no habría sido posible la finalización de este trabajo. A mis compañeros del Jardín Botánico de Bogotá, Marcela Celis, Carlos Vargas, Janice Valencia, Diego Moreno, Angela Rodriguez y José Muñoz. Finalmente, quiero agradecer a todos mis amigos y a aquellas personas que de una u otra forma han sido participes de mi vida, a todas muchas gracias. Resumen 7 Resumen Markea (Solanaceae, Juanulloeae) es un género neotropical de lianas o arbustos epífitos, con aproximadamente 20 especies. La circunscripción actual del género es controversial y se basa en taxonomía tradicional y en estudios filogenéticos morfológicos. Para resolver las dudas sobre la monofilia de Markea se realizó un análisis filogenético bajo máxima parsimonia (MP) e inferencia bayesiana (IB) con información de dos marcadores nucleares (ITS y GBSSI) y dos del cloroplasto, (trnT–trnF y rps16–trnK). Se incluyeron en los análisis 18 especies de Markea y representantes de todos los géneros de la tribu Juanulloeae y de otros géneros de Solanaceae como Atropa, Lycium y Nicandra. Los resultados indicaron que Markea como ha sido circunscrito en el pasado no es monofilético, solo las especies M. coccinea, M. costanensis, M. formicarum y M. sessiliflora pertenecerían a Markea. Estas especies se distribuyen en el Escudo Guayanés, la Amazonía y la cordillera de la Costa de Venezuela. Las especies restantes, la mayoría de ellas distribuidas en los Andes, se agruparon en otros géneros de la tribu Juanulloeae. Se discuten los resultados del presente trabajo y sus implicaciones en la circunscripción de Markea, de los demás géneros de Juanulloeae y de la tribu en general. Por último, se caracterizan morfológicamente los clados obtenidos y se presentan algunas novedades taxonómicas. Palabras clave: filogenia, Hawkesiophyton, Juanulloeae, Markea, Solanaceae. Abstract 8 Abstract Markea (Juanulloeae, Solanaceae) is a Neotropical genus of epiphytic shrubs or vines, with ca. 20 species. Current circumscription of the genus is controversial and is based on traditional taxonomy or morphological phylogenetic studies. To resolve the doubtful monophyly of Markea, one phylogenetic analysis was performed using two nuclear markers (ITS and GBSSI) and two chloroplast regions (trnT–trnF and rps16–trnK) and analyzing the sequences obtained under Maximum parsimony and Bayesian inference. 18 species of Markea and species of all genera of the tribe Juanulloeae and other genera of Solanaceae as Atropa, Lycium and Nicandra were included in this study. It was found that genus Markea as has been previously defined is not monophyletic and only M. coccinea, M. costanensis, M. formicarum and M. sessiliflora must be kept in Markea, all species distributed in the Guiana Shield, the Amazonian basin or the Venezuelan Coastal Ranges System. The remaining species, most of them Andeans in distribution were grouped into different clades along the phylogeny of the tribe. The results and their implications for the circumscription of Markea, the remaining genera of Juanulloeae and the all tribe are discussed. Additionally, the clades obtained are morphologically characterized and some taxonomic novelties are presented. Keywords: Hawkesiophyton, Juanulloeae, Markea, molecular phylogeny, Solanaceae. Contenido 9 CONTENIDO Pág. INTRODUCCIÓN ........................................................................................................... 18 Reconocimiento morfológico de Markea......................................................................... 20 Problema de investigación ............................................................................................. 20 Objetivos ........................................................................................................................ 22 Objetivos específicos...................................................................................................... 22 1. MATERIALES Y MÉTODOS................................................................................... 23 Estudio de las especies: trabajo de herbario y campo ......................................... 23 Análisis filogenético ............................................................................................. 24 1.2.1 Representación de taxones.............................................................................. 24 1.2.2 Elección de marcadores moleculares............................................................... 27 1.2.3 Extracción, amplificación y secuenciación de ADN........................................... 30 1.2.4 Construcción de la matriz de secuencias, alineamiento y combinación de las fuentes de información................................................................................................ 33 1.2.5 Análisis de datos .............................................................................................. 36 2. RESULTADOS ....................................................................................................... 38 2.1 Máxima parsimonia (MP) ..................................................................................... 38 2.1.1 Análisis de MP para el marcador ITS ............................................................... 40 2.1.2 Análisis de MP para el marcador GBSSI .......................................................... 41 2.1.3 Análisis de MP para el marcador trnT–trnF ...................................................... 43 2.1.4 Análisis de MP para el marcador rps16–trnK.................................................... 44 2.1.5 Análisis de MP para la combinación de los marcadores nucleares................... 46 2.1.6 Análisis de MP para la combinación de los marcadores de cloroplasto............ 47 2.1.7 Análisis de MP para la combinación de los marcadores nucleares y del cloroplasto……………………………………………………………………………………...49 2.2 Inferencia bayesiana (IB) ..................................................................................... 51 2.2.1 Análisis de IB para el marcador ITS ................................................................. 51 2.2.2 Análisis de IB para el marcador GBSSI............................................................ 52 2.2.3 Análisis de IB para el marcador trnT–trnF ........................................................ 54 2.2.4 Análisis de IB para el marcador rps16–trnK ..................................................... 55 2.2.5 Análisis de IB para la combinación de los marcadores nucleares..................... 57 2.2.6 Análisis de IB para la combinación de los marcadores del cloroplasto ............. 58 Contenido 10 2.2.7 Análisis de IB para la combinación de los marcadores nucleares y del cloroplasto……………………………………………………………………………………...60
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