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Redalyc.Filogenia De Hongos Roya (Uredinales) En La Zona Andina Revista de Biología Tropical ISSN: 0034-7744 [email protected] Universidad de Costa Rica Costa Rica Zuluaga, Catalina; Buriticá, Pablo; Marín, Mauricio Filogenia de hongos roya (Uredinales) en la zona andina colombiana mediante el uso de secuencias del ADN ribosomal 28S Revista de Biología Tropical, vol. 59, núm. 2, junio, 2011, pp. 517-540 Universidad de Costa Rica San Pedro de Montes de Oca, Costa Rica Disponible en: http://www.redalyc.org/articulo.oa?id=44922367001 Cómo citar el artículo Número completo Sistema de Información Científica Más información del artículo Red de Revistas Científicas de América Latina, el Caribe, España y Portugal Página de la revista en redalyc.org Proyecto académico sin fines de lucro, desarrollado bajo la iniciativa de acceso abierto Filogenia de hongos roya (Uredinales) en la zona andina colombiana mediante el uso de secuencias del ADN ribosomal 28S Catalina Zuluaga1, Pablo Buriticá2 & Mauricio Marín1 1. Universidad Nacional de Colombia sede Medellín, Laboratorio de Biología Celular y Molecular, Cra 64 x Calle 65, Autopista Norte, Medellín, Colombia; [email protected], [email protected] 2. Universidad Nacional de Colombia sede Medellín, Departamento de Ciencias Agronómicas, Cra 64 x Calle 65, Autopista Norte, Medellín, Colombia; [email protected] Recibido 16-IV-2010. Corregido 10-IX-2010. Aceptado 26-X-2010. Abstract: Phylogenetic analysis of rust fungi (Uredinales) from the Colombian Andean region using 28S ribosomal DNA sequences. Rust fungi (Uredinales, Basidiomycetes) are one of the most diverse and economi- cally important plant-obligated parasites. Taxonomy of this group has been under revision during the last years using molecular techniques to define phylogenetic relationships. In this study we evaluated the phylogenetic affinities of a group of 40 rust fungi obtained from different plants in the Colombian Andean region using sequence analysis of the 28S ribosomal DNA, specifically D1/D2 domains. Comparisons were undertaken with sequences of rust fungi from around the world deposited in the GenBank database. An alignment of sequences was used to build a phylogenetic tree through Maximum parsimony analysis. Our results support the taxo- nomical validity of families Pucciniaceae, Phakopsoraceae, Phragmidiaceae, Pileolariaceae, Mikronegeriaceae, Coleosporiaceae and Cronartiaceae, while Pucciniosiraceae represents redundant taxa with Pucciniaceae. The analyses indicated that Uropyxidaceae, Raveneliaceae, Chaconiaceae and Pucciniastraceae correspond to poly- phyletic families. Melampsoraceae appear to be a basal taxon to the Uredinales. Information obtained in this study will be useful to incorporate a higher number of sequences from tropical rust fungi within global efforts to redefine the taxonomy of order Uredinales. Additionally, we propose to give priority to future phylogenetic studies of taxa: Gerwasia, Hemileia, Phragmidium, Prospodium, Puccinia and Uromyces, genera that include a high number of rust fungi from the tropics. Rev. Biol. Trop. 59 (2): 517-540. Epub 2011 June 01. Key words: maximum parsimony, rDNA, rust fungi, sequence analyses. Los hongos roya (Urediniomycetes, Uredi- sus hospedantes; sin embargo, también pueden nales) constituyen uno de los grupos de hongos causar hipertrofias e hiperplasias, escobas de más numeroso, diverso y de amplia distribución bruja y formación de pseudoflores (Cummins mundial (Buriticá 2003a); son parásitos obliga- & Hiratsuka 2003). dos (holobiótrofos) de un amplio rango de El orden Uredinales está conformado por plantas incluidas helechos, coníferas y angios- 13 familias, 163 géneros y unas 7 000 espe- permas, con las cuales han coevolucionado, que cies (Hawksworth et al. 2001); aunque se adaptan su ciclo de vida a las condiciones eco- estima que el grupo puede contener de 20 000 lógicas del hábitat de sus hospedantes (Buriticá a 24 000 especies (Buriticá 2003b). Reciente- 2001, Buriticá 2003b, Cummins & Hiratsuka mente, Bauer et al. (2006) en su estudio de los 2003, Zuluaga et al. 2009). Los Uredinales Basidiomycetes con septos simples, sugirieron son conocidos como royas por la inducción de renombrar las categorías suprafamiliares en pústulas que contienen esporas con apariencia que se ubican los hongos roya como orden de un polvillo herrumbroso sobre los tejidos de Pucciniales de la clase Pucciniomycetes. Los Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 59 (2): 517-540, June 2011 517 hongos roya pueden producir hasta cinco esta- ADNr en hongos roya recolectados principal- dos espóricos que se designan como: esper- mente en Europa y que pusieron de manifiesto mogonio (espermacios e hifas receptivas) (O), que los géneros Puccinia, Pucciniastrum, The- aeciospora (I), uredospora (II), teliospora (III) kopsora y Uromyces son polifiléticos, mientras y basidiospora (IV) (Hahn 2000). Cada estado que Chrysomyxa, Coleosporium, Cronartium, espórico es morfológica y funcionalmente dife- Gymnosporangium, Melampsora, Phrag- rente dentro del ciclo de vida de las especies de midium y Tranzschelia, representan taxones royas (Hawksworth et al. 2001, Cummins & monofiléticos. La condición polifilética de los Hiratsuka 2003). géneros Puccinia y Uromyces fue confirma- La clasificación de los Uredinales a nivel da posteriormente por Van der Merwe et al. genérico y supragenérico ha sido basada casi (2007) quienes usaron análisis de secuencias exclusivamente en la morfología de los telio- de los genes del factor de elongación 1α y soros y teliosporas. Inicialmente dos fami- β-tubulina 1. Un estudio similar fue realizado lias fueron definidas por las características por Wingfield et al. (2004), pero mediante aná- morfológicas de sus teliosporas: Melampsora- lisis de secuencias de la subunidad pequeña del ceae con teliosporas no pediceladas (teliosoros ADNr en 64 especies de 12 familias de royas; subepidermales, cubiertos) y Pucciniaceae con en este caso se encontró que los géneros que teliosporas pediceladas (teliosoros erumpen- presentan estados aeciales sobre gimnospermas tes) (Dietel 1928). Posteriormente, Gaumann están filogenéticamente distantes de los hongos (1949) adicionó las familias Pucciniastraceae roya que desarrollan dicho estado sobre plan- (teliosoros inmersos), Cronartiaceae (telioso- tas angiospermas; además se determinó que ros columnares), Chrysomyxaceae (teliosoros la condición de autoicismo/heteroicismo no cupulares) y Coleosporiaceae (basidio interno). representa un carácter taxonómico válido y que Cummins & Hiratsuka (1983) propusieron la familias como Pucciniaceae y Pucciniastraceae definición de 14 familias para el orden Uredi- representan taxones polifiléticos. nales, las cuales luego redujeron a 13, una vez Aime (2006) utiliza la estrategia de com- unidas las familias Raveneliaceae y Sphaero- binar datos de ambas subunidades ribosomales, phragmiaceae (Cummins & Hiratsuka 2003). al realizar un estudio tendiente a evaluar el La taxonomía de hongos se ha fundamen- soporte filogenético de las familias del orden tado tradicionalmente en el empleo de carac- Uredinales propuestas por Cummins & Hirat- teres morfológicos. En muchos casos estas suka (2003) con base en caracteres morfoló- características son variables, que producen gicos. De las trece familias propuestas, ocho información insuficiente o poco precisa que (Coleosporiaceae, Melampsoraceae, Mikrone- conduce a la generación de clasificaciones no geriaceae, Phakopsoraceae, Phragmidiaceaeae, naturales. En las últimas décadas, la secuencia- Pileolariaceae, Pucciniaceae y Raveneliaceae) ción de regiones ribosomales y funcionales ha están bien soportadas por el análisis basado en jugado un papel fundamental en la inferencia secuencias, mientras que tres son redundantes de las relaciones evolutivas entre individuos y (Cronartiaceae, Pucciniastraceae y Pucciniosi- grupos de individuos (Sugiyama 1998, Blac- raceae), y en dos de ellas, su condición no pudo kwell et al. 2006). Sin embargo, el número de ser definida (Chaconiaceae y Uropyxidaceae). estudios que emplean estas herramientas mole- A pesar del alto número de especies de culares para la realización de estudios taxo- royas presentes en los trópicos, el nivel de nómicos en hongos roya es muy reducido, en conocimiento que se tiene de estos hongos en comparación con aquellos realizados en otros esta región es incipiente, tanto desde el punto grupos de hongos (Zuluaga et al. 2009), siendo de vista biológico como en lo relacionado a son los más destacados aquellos realizados por su clasificación taxonómica. Esta situación Maier et al. (2003, 2007), basados en la secuen- contrasta con la gran importancia económi- ciación de una porción de la región 28S del ca que presentan algunas royas en diversos 518 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 59 (2): 517-540, June 2011 agroecosistemas tropicales y con el potencial bioprospectivo de especies que podrían ser utilizadas como biorreguladores de plantas arvenses. Una de las regiones neotropicales de las que se posee menor conocimiento sobre este grupo de organismos, es la zona andina que comprende una franja altitudinal entre 2 000 y 4 600m.s.n.m, nicho ecológico que ofrece con- diciones físicas y biológicas muy particulares, y específicamente para el caso de hongos roya incluye fundamentalmente especies con ciclos de vida reducido, poco abordadas en los estu- Fig. 1. Departamentos andinos de Colombia en los cuales se dios micológicos mundiales que utilizan herra- obtuvieron las muestras de royas analizadas en este estudio. mientas moleculares
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