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FILOGENIA MOLECULAR Y BIOGEOGRAFÍA HISTÓRICA DE Oreobolus (CYPERACEAE)

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FILOGENIA MOLECULAR Y BIOGEOGRAFÍA HISTÓRICA DE Oreobolus (CYPERACEAE) FILOGENIA MOLECULAR Y BIOGEOGRAFÍA HISTÓRICA DE Oreobolus (CYPERACEAE) JULIANA CHACON PINILLA UNIVERSIDAD DE LOS ANDES FACULTAD DE CIENCIAS DEPARTAMENTO DE CIENCIAS BIOLÓGICAS BOGOTA D. C. 2003 FILOGENIA MOLECULAR Y BIOGEOGRAFÍA HISTÓRICA DE Oreobolus (CYPERACEAE) JULIANA CHACON PINILLA Trabajo de Grado para optar al título de Biólogo Director Santiago Madriñán, Ph. D. Profesor Asistente del Departamento de Ciencias Biológicas Universidad de los Andes Codirector Favio González, Ph. D. Profesor del Departamento de Biología Universidad Nacional de Colombia ii AGRADECIMIENTOS Quisiera agradecer al profesor Santiago Madriñán, director de esta tesis, por hacer posible la realización de esta investigación y por su continuo interés y dedicación hacia la misma. Gracias por transmitirme el interés por el estudio de las plantas y la evolución, y por la energía, la paciencia y el apoyo incondicional que me brindó desde la difícil etapa de laboratorio hasta la ejecución del manuscrito final. Al profesor Favio González, codirector de esta tesis, por sus valiosos comentarios y sugerencias. También quisiera agradecer a Julio Betancur por las muestras de Oreobolus que amablemente cedió para ser utilizadas en este trabajo. A Mark W. Chase y a mis compañeros en Kew Gardens, por abrirme las puertas del laboratorio Jodrell y por su constante interés y sugerencias. A mis compañeros del Laboratorio de Botánica & Sistemática de la Universidad de los Andes; Rafa, Alfredo, Jorge y Adriana, por sus sonrisas y su amistad. A Andrés por estar siempre ahí y ser un apoyo incondicional en todos los momentos. Finalmente quisiera dar las gracias por todo a mis padres, mis hermanos y a mi grandiosa familia…a ellos les dedico este trabajo. iii TABLA DE CONTENIDO RESUMEN...........................................................................................................................v INTRODUCCION.............................................................................................................. 1 Distribución geográfica de Oreobolus ............................................................................ 1 Taxonomía del género Oreobolus................................................................................... 3 Estudios sistemáticos de Oreobolus realizados por Mora-Osejo (1987) y Seberg (1988) ......................................................................................................................................... 6 Comparación de la hipótesis de Mora-Osejo (1987) y la de Seberg (1988)................... 6 MATERIALES Y MÉTODOS......................................................................................... 10 Muestreo taxonómico.................................................................................................... 10 Secuenciación del ADN ................................................................................................ 10 Análisis filogenético...................................................................................................... 12 Calibración del reloj molecular..................................................................................... 14 RESULTADOS................................................................................................................. 16 DISCUSION ..................................................................................................................... 19 Relaciones filogenéticas del género Oreobolus............................................................ 19 Biogeografía histórica de Oreobolus............................................................................. 21 CONCLUSION................................................................................................................. 26 LITERATURA CITADA.................................................................................................. 28 FIGURAS Y TABLAS ..................................................................................................... 31 iv RESUMEN Oreobolus (Cyperaceae) es un género de plantas que crecen formando cojines cerca de pantanos de las montañas de Centro y Suramérica, Islas del Pacífico Sur, Australia y el archipiélago de Hawaii. Su hábito peculiar es el resultado de las ramificaciones acrotónicas, justo por debajo de la inflorescencia, carácter compartido con el género monotípico Schoenoides y con unas pocas especies del género Costularia. Schoenoides oligocephalus y Oreobolus comparten además la presencia de un pseudopeciolo en la lámina foliar. Previas hipótesis sobre las relaciones filogenéticas de las especies de Oreobolus han sido propuestas por Mora-Osejo en 1987 y Seberg en 1988, el primero limitándose a las especies americanas y el segundo abarcando la totalidad del género. Las incongruencias entre dichas filogenias han generado hipótesis biogeográficas diferentes que intentan explicar los patrones de distribución del género. Mediante la secuenciación del gen nuclear ITS y de los genes plastidiales trnL y rbcL, se propone una filogenia molecular de Oreobolus, la cual se compara con las hipótesis filogenéticas propuestas previamente. Los resultados del análisis molecular generaron el siguiente árbol: (C. laxa ((O. pectinatus, O. strictus), (S. oligocephalus (O. furcatus (O. oxycarpus, O. distichus)))), (O. pumilio (O. obtusangulus (O. ecuadorensis (O. cleefii (O. goeppingeri, O. venezuelensis)))))). Este resultado demuestra que S. oligocephalus no es el grupo hermano de Oreobolus, como propone Seberg, sino que se encuentra embebido en el clado de especies de este género provenientes del Pacífico. En las diferentes filogenias obtenidas se presentan dos clados; uno conformado por las especies suramericanas (clado suramericano) y otro conformado por la especies del Pacífico Sur, v Australia y Hawaii (clado Pacífico). Oreobolus distichus (especie australiana) hace parte del clado Pacífico, posición que conlleva a una hipótesis biogeográfica diferente a la de Seberg, según la cual esta especie forma parte del clado suramericano. La topología del clado suramericano y la posición de O. obtusangulus como grupo hermano de las demás especies suramericanas, sugieren una progresión geográfica de algunas especies hacia el norte de Suramérica, apoyando la hipótesis biogeográfica de Mora-Osejo. La evidencia molecular indica que O. obtusangulus ssp. unispicus no es una subespecie de O. obtusangulus, validando el estatus de especie de O. cleefii sensu Mora-Osejo. La presencia de O. furcatus en Hawaii permitió la calibración de un reloj molecular basada en la edad mínima de surgimiento de éste archipiélago hace mínimo 5,1 millones de años. La distribución de esta especie en Hawaii es el resultado de eventos de dispersión a larga distancia a partir de Australia u otras islas del Pacífico Sur. vi INTRODUCCION El género Oreobolus R. Br. (Familia Cyperaceae) está conformado por un conjunto de plantas que crecen formando cojines debido a la innovación basal de los brotes justo por debajo de la inflorescencia (carácter c' en Seberg 1988, figura 1). Esta característica es compartida con otras especies como Schoenoides oligocephalus (W. M. Curtis) O. Seberg, y con las especies del género Costularia subgénero Chamaedendron de Kükenthal, más C. brevicaulis (Seberg 1988). Schoenoides y Oreobolus forman un grupo monofilético ya que, además del tipo de innovación mencionada, comparten otras características como la filotaxis espiro- ortodística, la presencia de pseudopecíolo (carácter e' en Seberg 1988), que puede ser una sinapomorfia para el grupo Schoenoides+Oreobolus, y la presencia de cuerpos de sílica piriformes, cónicos o en forma de domo. Además poseen un tipo de tépalos escamosos ubicados en dos verticilos alternos, los cuales están presentes en todas las especies de Schoenoides y Oreobolus, excepto en O. ambiguus donde dichos tépalos tienen forma de cerda (Seberg 1988). Oreobolus aparentemente es un género monofilético que se separa de Schoenoides debido a que presenta espiguillas unifloras ligeramente comprimidas (carácter f' en Seberg 1988), compuestas de tres o cuatro glumas según la especie. Distribución geográfica de Oreobolus Oreobolus presenta un patrón de distribución complejo, que no ha podido explicarse con certeza. La distribución de este género se conoce como disyunta ya que se encuentra en dos regiones ampliamente separadas del hemisferio sur (Brown & Lomolino 1998). La primera región comprende los páramos, punas y jalcas de la cordillera de los Andes en 2 Suramérica (Colombia, Venezuela, Ecuador, Perú, Chile y Argentina), con algunas poblaciones aisladas en Brasil (Sierra de la neblina), y parte de Centroamérica (Panamá y Costa Rica). La segunda comprende las praderas alpinas de algunas islas del Pacífico Sur (Nueva Zelandia, Tasmania, Nueva Guinea, Borneo, Sumatra, Filipinas), Australia y Hawaii (Seberg 1989; 1991). Oreobolus presenta una interesante historia filogenética y biogeográfica similar a la de otras plantas como Nothofagus (familia Fagaceae) y algunos miembros de las familias Proteaceae, Winteraceae y Gesneriaceae (Seberg 1988). Varios grupos de animales también presentan este tipo de distribución, como por ejemplo ciertos dípteros de las familias Chironomidae y Sciadoceridae, algunos invertebrados de las familias Lepidoptera y Hemiptera (Seberg 1988), y algunos vertebrados como las ranas Pipinae y Xenopinae, los peces Galaxioidea y varios grupos de marsupiales (Brown & Lomolino 1998). La coincidencia en cuanto a la distribución de dichos grupos ha inspirado a varios investigadores para explicar
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