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Filogenia Del Género Helvella (Ascomycota) Fidel Landeros Jaime

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Filogenia Del Género Helvella (Ascomycota) Fidel Landeros Jaime UNIVERSIDAD DE GUADAL.AJARA Centro Universitario de Ciencias· Biológicas y Agropecuarias Filogenia del género Helvella (Ascomycota) Tesis que para obtener el grado de Doctor en Ciencias en Biosist.emática, Ecología y Manejo de Recursos Naturales y Agrícolas Presenta Fidel Landeros Jaime DIRECTOR Dra. Laura Guzmán Dávalos Zapopan, Jalisco 8 de junio de 2012 UNIVERSIDAD DE GUADAL.AJARA Centro Universitario de Ciencias Biológicas y Agropecuarias Doctorado en Ciencias en Biosistemática, Ecología y Manejo de Recursos Naturales y Agrícolas Filogenia del género He/ve/la (Ascomycota) Por Fidel Landeros Jaime Tesis presentada como requisito parcial para obtener el grado de: Doctor en Ciencias en Biosistemática, Ecología y Manejo de Recursos Naturales y Agrícolas Aprobado por: Ha t ?J ; 6 / 2tJ1~ Fecha Director de Tesis e integrante del Jurado arón Rodríguez Contre a Asesor del Comité Particul Dra. María Teresa lturriaga de Capiello Fecha Asesora del Comité Partic lar e integrante del Jurado 4 ¿} J - ,, ,:UJ / ...( Dr. José Luis Nav rrete Heredia Fecha Integrante del Ju a o /{¿, -o ~-JO / ~ Dra. Marí Olivia Rodríguez Alcántar Fecha Integrante el Jurado Fecha Coordinadoraº'"· ~~ de la Orientación Tesis doctoral: Filogenia del género He/ve/la (Asco1nycota) Contenido CONTENIDO Resumen........................................................................................................... 2 Capítulo l. Introducción . .. ... 3 Capítulo II. Landeros F, Ittmiaga T, Guzmán-Dávalos L. 2012. Type studies in Helvella (Pezizales) l. Mycotaxon 119: 35-63 ........................................................................ 9 Capítulo III. Landeros F, Korf RP. 2012. Nomenclatura! notes 13. An incorrrect neotype designation and provision of a lectotype and an epitype for He/ve/la fi1sca. Mycotaxon 119: 431- 438 ·················································································································· 39 Capítulo IV. Landeros F, Iturriaga T, Rodríguez A, Vargas-Amado G, Guzmán-Dávalos L. 2012. He/ve/la (Pezizales: Ascomycota): phylogeny and taxonomy based on morphological data and LSU rDNA sequences. Fungal Diversity . ............ ......... .. 48 Capítulo V. Landeros F, Guzmán-Dávalos L. 2012. Revisión del género He/ve/la (Ascomycota: Fungí) en México. Revista Mexicana de Biodiversidad . ... .. .. 95 Capítulo VI. Discusión general y conclusiones . .. .. 120 Tesis doctoral: Filogenia del género He/vello (Ascon1ycota) Rcsutnen RESUMEN El género Helvel/a (Fnngi, Ascomycetes, Pezizales) se caracteriza por presentar apotecios en forma de copa o lobados. El estípite es liso, costillado o lagunoso, superficie glabra, subpubescente o pubescente. El excípulo está diferenciado en medular de textura intrincada y ectal de textura angular. Sus ascosporas son unigutulutadas (normalmente), lisas, ligeramente verrugosas o verrugosas. Existen tres propuestas de clasificación infragenérica basadas en características macromorfológicas y sólo en una de ellas utilizan además el tipo de asca. Para tener un mejor concepto y establecer los límites de las especies descritas de Helvella, se estudiaron 26 especímenes tipo. Como resultado, fue posible completar las descripciones originales. Se concluyó que H costifera es sinónimo de H. hyperborea. También fue posible reconocer a H. griseoa/ba de H. costifera, H. subglabra de H. atra, H. stevensii de H. latispora, y H. su/cata de H. lacunosa. Se designaron los lectotipos de H. a/bella, H. elastica, H. ephippi11111 y H. su/cata. Se corrigió la tipificación de H. jitsca. Por separado, la evaluación de las propuestas de clasificación que se han establecido para el género fue mediante un análisis filogenético, usando datos morfológicos y secuencias de ADN de la región 28S del DNA ribosomal. Se usaron 44 especímenes de 24 especies de He/ve/la y Wynel!a silvico/a como grupo externo para el análisis filogenético. La matríz combinando datos moleculares y mmfológicos generó árboles con mejores soportes, en comparación con la matríz sólo con datos moleculares. Sólo las secciones Elasticae, He/ve/la, Lacunosa y Leucome/aenae se recobraron de los análisis con soporte estadístico y/o probabilístico. Los resultados de este trabajo apoyan la división del género en He/vella subgénero He/ve/la y He/ve/la subgénero Leucomelaenae. Las características morfólogicas que lo apoyanb son el tipo de asca, pleurorrincosa para el primero y aporrincosa para el segundo. Por último, se hizo una revisión del género en México y se reconoce la presencia de 18 especies para el país. 2 Tesis doctoral: Filogenia del género !!e/vello (Asco1nycota) Capítulo l. Introducción Capítulo l. Introducción 3 Tesis doctoral: Filogenia del género He/ve/la (Asco1nycota) Capítulo l. Introducción INTRODUCCIÓN El presente trabajo es un estudio taxonómico y filogenético del género He/ve/la L. con base en datos morfológicos y secuencias de ADN. Se discuten las relaciones filogenéticas entre las especies y se comparan con las propuestas infragenéricas que han sido postuladas para el género. Se incluyen los resultados de los estudios macro y micromorfológicos de las especies estudiadas, siendo en la mayoría de los casos de ejemplares tipo. Además se aportan nuevas características microscópicas y reacciones microquímicas que deben de ser consideradas cuando se hacen estudios taxonómicos sobre Helvel/a. El trabajo consta de seis capítulos El primer capítulo es una introducción general al estudio de He/ve/la. Por su parte, el capíhilo II (Landeros et al. 2012) es la revisión de 16 especímenes tipo, y material adicional de 15 especies de Helvella. En él se argumenta que H. subglabra N.S. Weber es diferente de H. aira J. Konig, así como H. griseoalba N.S. Weber de H. costifera Nannf., mientras que H. hyperborea Harmaja debe de ser considerada un sinónimo de H. costifera. Se apoya que H. queletii Bres. es un sinónimo de H. solitaria P. Karst. de acuerdo con Hamaja (1977). Además, se hace una revisión de cie1tos conceptos (apotecio, costillado, hemiamiloide, lagunoso, píleo, pubescente, pruinoso, sulcado, velloso) que generan ambigüedad en su uso, y de los que debe haber una estandarización en su aplicación. Se incluye una clave sinóptica para la determinación de las especies estudiadas. El capítulo III fue publicado por Landeros y Korf (2012). Se argumenta que Abbott y Cunah (1997) propusieron incorrectamente un neotipo (el cual sólo puede ser seleccionado cuando no existe material original) para H. fi1sca Gillet. En la obra original donde se describió esta especie, Gillet ( 1879) publicó una figura, la cual automáticamente debe ser considerada el lectotipo. En el artículo se citan los artículos del Código de Nomenclatura Botánica de Viena que apoyan la designación del lectotipo para H. fitsca, y además se hace la designación de un epitipo (material que sirve como ayuda para interpretar cuando el material tipo designado para el taxon genera ambigüedad). El capítulo IV es un análisis filogenético y taxonómico de He/ve/la. En este capítulo se incluye la filogenia del género He/ve/la, con base en datos morfológicos y secuencias de ADN. También el estudio de ejemplares tipo que no se incluyeron en los capítulos II y III (Acetabula calyx Sacc., H. chinensis (Velen.) Nannf. & L. Holm, H. compressa (Snyder) N.S. Weber, H. confi1sa Hannaja, H. connivens Dissing & M. Lange, H. cupulijimnis Dissing & Nannf., H. melaleuca Rifai, H. pallidula N.S. Weber, Peziza debeauxii Roum. y P. Subclavipes W. Phillips & Ellis). Se analizan los caracteres macro y micromorfológicos que son impmtantes para el estudio taxonómico del género, se presenta una clave para 34 especies, se describen los taxones estudiados, y finalmente se hace la propuesta de dividir al género en He/ve/la subgénero He/ve/la y He/ve/la subgénero Leucomelaenae. A su vez, He/ve/la subg. Hevella puede dividirse en He/ve/la scción Helvella, He/ve/la sección Elasticae Dissing sensu Weber y He/ve/la sección Lacunosae Dissing. Mientras que He/vela sugb. Leucomelaenae sólo tiene una sección, He/ve/la sección Leucomelaenae Dissing sensu Weber. El capítulo V es una revisión del género He/ve/la (Ascomycota: Fungi) en México. Se hizo una selección de l 07 especímenes de los herbarios ENCB, FCME, IBUG, y XAL. Se hizo énfasis en los ejempalres referidos en las publicaciones. Después de estudiar el material, revisar los artículos originales, comparar con el material tipo, y consultar las descripciones originales, se reconoce la presencia de 18 especies de He/ve/la que en México: H. acetabulum (L.) Qué\., H. a/bella Qué\., H. aira, H. compressa, H. cost!fera, H. crispa (Scop.) Fr., H. cupuliformis, H. e/astica Bull., H. ephippium Lév., H. fibrosa (Wallr.) Korf, H. /actea Boud., H. lacunosa Afzel., H. leucomelaena (Pers.) Nannf., H. macropus (Pers.) P. Karst., H. pezizoides Afzel., H. solitaria, H. stevensii Peck y H. su/cata Afzel., de 4 Tesis doctoral: Filogenia del género Helvella (Asco1nycota) Capítulo I. Introducción las cuales Helvella compressa es un nuevo registro para el país. Además H. adhaerens Peck, H. corium (O. Weberb.) Massee, H. kichneri Manzi, H. latispora Boud., H. leucopus Pers., H. leucopus var. popu/ina I. Arroyo & Calonge, H. pityophila Boud., H. queletiana Sacc. & Traverso, H. sa/vatierrae Manzi, H. sogarii Manzi y H. subglabra no tienen el ejemplar de referencia o estaba incorrectamente dete1minado, en consecuencia, su presencia en México no puede ser confimrnda. Se incluyen comentarios taxonómicos de las especies que crecen en México y observaciones de las que no están presentes. Se presenta una clave
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